vvEPA
United States
Environmental Protection
Agency
Environmental Research
Laboratory
Duluth MN 55804
EPA 600 3-80 068
July 1980
Research and Development
A Guide to Freshwater
Mo Musks of the
Laurentian Great
Lakes with Special
Emphasis on the
Genus Pisidium
EP 600/3
80-068
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RESEARCH REPORTING SERIES
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EPA-600/3-80-068
July 1980
A GUIDE TO FRESHWATER MOLLUSKS
OF THE LAURENTIAN GREAT LAKES
WITH SPECIAL EMPHASIS ON THE GENUS PISIPIUM
by
Gerry L. Mackie
Department of Zoology
University of Guelph
Guelph, Ontario NlG 2W1
and
David S. White and Thomas W. Zdeba
Great Lakes Research Division
and School of Natural Resources
University of Michigan
Ann Arbor, Michigan 48109
Grant No. 805333
Project Officer
Nelson A. Thomas
Environmental Research Laboratory
Large Lakes Research Station
Grosse He, Michigan 48138
ENVIRONMENTAL RESEARCH LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U.S. ENVIRONMENTAL PROTECTION AGENCY
DULUTH, MINNESOTA 55804
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DISCLAIMER
This report has been reviewed by the Environmental Research Laboratory,
Office of Research and Development, Duluth, Minnesota, U.S. Environmental
Protection Agency, and approved for publication. Approval does not signify
that the contents necessarily reflect the views and policies of the U.S.
Environmental Protection Agency, nor does mention of trade names or commer-
cial products constitute endorsement or recommendation for use.
ii
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FOREWORD
Our nation's freshwaters are vital for all animals and plants, yet
our diverse uses of water—for recreation, food, energy, transportation,
and industry—physically and chemically alter lakes, rivers, and streams.
Such alterations threaten terrestrial organisms, as well as those living
in water. The Environmental Research Laboratory in Duluth, Minnesota
develops methods, conducts laboratory and field studies, and extrapolates
research findings
—to determine how physical and chemical pollution affects
aquatic life
—to assess the effects of ecosystem on pollutants
—to predict effects of pollutants on large lakes through
use of models
—to measure bioaccumulation of pollutants in aquatic
organisms that are consumed by other animals, including
man
This guide will aid in the identification of mollusks in the fresh-
water environment. To provide an accurate assessment as to the impact of
pollutants on the ecosystem, accurate identification of the organisms
inhabiting these zones is required. The taxonomy of mollusks will be
greatly enhanced through the descriptions provided in this guide.
Norbert Jaworski, Ph.D.
Director
Environmental Research Laboratory
Duluth, Minnesota
111
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ABSTRACT
Presented here are keys and notes on distribution and ecology for the
freshwater snails (Gastropoda), mussels (Unionidae), and fingernail clams
(Sphaeriidae) which have been collected from the Laurentian Great Lakes.
Including subspecies and forms, 121 taxa are discussed: 47 Gastropoda, 39
Unionidae, and 35 Sphaeriidae. Relations to substrate preferences and
pollution are summarized where known. Special emphasis is given to the
sphaeriid genus Pis-idiufn in discussions of morphological variability and
characters which will separate the species and forms both in the adult and
immature stages. Both drawings and scanning electron microscope
photomicrographs are presented for all sphaeriid taxa. A limited synonymy
has been compiled from publications and reports on the Laurentian Great Lakes
and the expanded reference section includes not only citations used in the
text but also publications and reports which may aid researchers in more
indepth studies of the fauna.
iv
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CONTENTS
Foreword ill
Abstract iv
Acknowledgments vi
I. Introduction 1
Sphaeriidae 3
Unionidae 4
Gastropoda 5
II. Checklist of Freshwater Mollusks in the Laurentian
Great Lakes 6
Gastropoda 6
Unionidae 6
Sphaeriidae 7
III. Key to Freshwater Mollusks of the Laurentian Great Lakes ... 8
Gastropoda 8
Unionidae 23
Sphaeriidae 36
IV. Ecology and Distribution of Gastropoda 97
V. Ecology and Distribution of Unionidae 101
VI. Ecology, Distribution, and Variation of Sphaeriidae 103
Synonymy 120
Glossary 124
References 130
Index to Species, Subspecies, and Forms 139
v
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ACKNOWLEDGMENTS
The authors are grateful to the National Museums of Canada and the
University of Michigan Museum of Zoology for providing lists of the Great
Lakes mollusks in their collections and for the loan of specimens-used in
the drawings and photographs. Leslie Newman has drawn most of the figures
of Gastropoda and Unionidae. Len Kalas read the original manuscript and
gave comments that contributed greatly toward the improvement of this draft.
We appreciate the help of George Te for the taxonomy of Great Lakes'
Physidae and Joel Lichty for technical assistance with identification and
data analysis of the Lake Michigan Pisidiwn.
The study was funded by grants from the Research Advisory Board,
University of Guelph, the National Research Council of Canada, Canada
Centre for Inland Waters, the Indiana and Michigan Power Company, and the
United States Environmental Protection Agency.
vi
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SECTION I
INTRODUCTION
In the Laurentian Great Lakes, considerable emphasis has been placed on
the use of benthic macroinvertebrates for the biological assessment of water
quality. The macroinvertebrate community, or parts of it, has been utilized
to determine areas of enrichment within the lakes (e.g., Cook and Powers,
1964; Carr and Hiltinen, 1965; Powers and Alley, 1967; Johnson and
Brinkhurst, 1971; Mozley and garcia, 1972; Kinney, 1972; Mozley and Alley,
1973; Mozley and Howmiller, 1977) and to relate species distributions to
physical-chemical parameters of the water or sediment (e.g., Henson and
Herrington, 1965; Johnson and Matheson, 1968; Adams and Kregar, 1969;
Hiltunen, 1969; Schneider et al., 1969). Others have investigated species
composition and productivity of the communities (e.g., Shelford and Boesel,
1942; Teter, 1960; Powers and Robertson, 1965, 1968; Thomas, 1966; Robertson
and Alley, 1966; Freitag et al., 1976).
Most of the studies cited above, to some extent, have attempted to
examine populations at the species level and most refer to the mollusk
community. The lack of up-to-date keys for Great Lakes' macroinvertebrates,
particularly the mollusks, has prevented more extensive analysis of community
dynamics. Many of the studies have not been consistant in the level of
identification, especially with the juvenile stages. By example, juvenile
Pisidiwrj collected from deeper portions of the lakes often are lumped under
Pisidium aonventus because it is the major profundal species. When this is
done, populations of Pisidiim lilljeborgi or Pisidiiffn easertanum, although
not common in very deep waters, may be overlooked. There are even greater
problems with immatures and variants from near-shore areas where mollusks
show the greatest densities and diversity.
This publication has been motivated by the need for a more comprehensive
key to the Sphaeriidae, notably the genus Pisidium, as they exist in the
Laurentian Great Lakes. Previous identification manuals (Herrington, 1962;
Burch, 1972, 1975a), are broad in scope and not designed to examine the
various life history stages nor the variability among more localized
populations. The above texts plus works by Kuiper (1956, 1962), Herrington
(1965), Heard (1966, 1969), and'Clarke (1973) should be consulted for greater
general treatments of distributions, ecology, and systematics. As a
supplement to the key, comparative notes are provided for adults and
immatures of the Great Lakes' Pisidiwn. It should be stressed that the
annotations are most applicable to specimens from the Great Lakes, and the
references listed above should be used for collections from other systems.
To complete the list of known freshwater mollusks from the Laurentian
Great Lakes, keys and short annotations are included for the Unionidae or
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freshwater mussels and the Gastropoda or freshwater snails. We hope that the
keys will allow workers to more readily identify specimens from the Great
Lakes as the often unwieldy number of species not found in these waters is
excluded. It is recognized that there is the distinct possibility of other
species either existing in, being introduced, or invading the Great Lakes
from tributaries. To aid in their identifications, the keys have been made
open-ended with couplets leading to more comprehensive references for
particular families and genera.
The final section of this manual is devoted to annotations of habitats,
distributions, and relative abundances. This information is highly
qualitative because there have been few good quantitative studies. Fig. 1
shows the areas of the Laurentian Great Lakes which have received the
majority of benthological attention. Clearly, even more qualitative surveys
are required before the basics of general distributions are fully known.
Little is known about large areas of Lakes Superior and Huron.
1
Figure 1. Outline map of the Laurentian Great Lakes showing areas (shaded)
which have had qualitative or quantitative surveys and for which
there are lists of freshwater mollusks at the species level.
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SPHAERIIDAE
The Sphaeriidae, or fingernail clams, are the numerically dominant
component of the open lake mollusk fauna, of the 37 distinct North American
species, 29 have been recorded from the Laurentian Great Lakes. Including
subspecies and forms, 35 taxa are present and included in the key and
checklist. Maximum densities and diversity occur in the 20-40 m depth
interval in silty-sand where there may be several hundred individuals per
square meter. Few species are found at depths greater than 50 m and only
P-isidium oonventus is known to inhabit the deepest parts of the lakes (Heard,
1963; Henson and Herrington, 1965; Robertson, 1967). Densities have been
shown to be enhanced by mild organic enrichment, but many species cannot
tolerate severer pollutants (Carr and Hiltunen, 1965); thus, there may have
been a general increase in the sphaeriid populations over the past few
decades.
Because of habitat, densities, and small sizes of most Sphaeriidae,
standard benthological techniques for collecting in open water have proven
adequate for clams when samples are washed through seives with mesh openings
of 0.400 mm or less (e.g., Petersen, Ekman, and Ponar grabs and various
coring devices; see Flannagan, 1970; Mozley and Howmiller, 1977; and Resh
1979 for review and criticisms of sampling methods). Several species of
SphaeTi-um and MuscuI'Lwn are associated with shallow, rocky areas and rooted
aquatic macrophytes, and special collection methods must be used (dip nets,
hand picking from vegetation, etc.). Sphaeriids are ovoviviparus, and some
adults may prematurely abort the young when distrubed or preserved. If
collections are to be used for life history studies, larger clams should be
preserved individually in vials.
For most general taxonomic work required in benthological studies, only
the shells need be examined; therefore, whole samples can be preserved in
70-80% ethyl alcohol or 4-10% formalin buffered by saturating with calcium
carbonate. Unbuffered formalin is slightly acidic and will dissolve shells
in a short period to time. Therefore, if formalin is the innitial
preservative (even if buffered), samples should be washed and transferred to
alcohol after a few days. Other preservatives which have proven successful
for long-term storage include 40% isopropyl alcohol and 1% propylene
phenoxetol. If soft anatomy is to be studied, specimens must be relaxed and
fixed before preservation (see Burch, 1972, 1975a, 1975b).
For the purposes of this key, the soft anatomy of sphaeriids may be
ignored because fairly reliable characters are found in differences between
hinge teeth and shell shape, even for immatures. A good stereomicroscope
with magnification to lOOx is needed to examine hinge teeth, and shells
should be examined dry.
Shells of sphaeriids tend to be yellowish or brownish when alive and
have little or no surface sculpture other than striae. The right valve
contains only one cardinal tooth and two pairs of lateral teeth. Conversely,
the left valve contains two cardinal teeth and only two lateral teeth (Figs.
117, 118). The three genera known from the Great Lakes can be distinguished
by the shells. In Sphaerium and Musculium the beaks are subcentral
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(Figs. 124, 132) or on the anterior side of center (Figs. 154, 162), while
in Pisidium the beaks are posterior of center (Figs. 117, 118). The shells
of Museulium generally differ from those of Sphaerium by having raised
umbonal caps and thinner shells (Figs. 132, 162).
UNIONIDAE
The Unionidae or freshwater mussels usually are confined to the
shallower or littoral portions of the Great Lakes where there is some
constant water movement (Walker, 1913; Goodrich and van der Schalie, 1932a,
1932b; Langlois, 1954; Stansbery, 1961; van der schalie, 1961; Wood, 1963;
Carr and Hiltunen, 1965; Wolfert and Hiltunen, 1968). While each of the
Laurentian Great Lakes has at least limited populations, the western basin
of Lake Erie has been noted for its extensive mussel beds. In the past few
decades, the populations in western Lake Erie have declined markedly and may
be limited in the open lake to small mumbers of Lampsilis radiata si.'i'icfuoi-dea
(Wood, pers. comm.). Reasons for declines in densities and diversity have
been attributed to several factors including physicochemical changes in the
open water and loss of host fishes for the parasitic glochidia or larval
stage of the mussels (Wood, 1963, per. comm.; Carr and Hiltunen, 1965;
Wolfert and Hiltunen, 1968). In the key we have included most of the recent
species; however, many of these may no longer be present in the Laurentian
Great Lakes but may be found in adjacent streams, rivers, and ponds.
When existing in a variety of habitats, unionids exhibit distinct forms
and morphological variations. This has been noted particularly in Lake Erie
where those species living in the wave zone are markedly different from
their counterparts from other bodies of water (Grier, 1918, 1920; Ball, 1922;
Brown et al., 1938; van der Schalie, 1941). The differences should be taken
into account when attempting to use the guide presented here, although we
have tried to illustrate typical Lake Erie forms and similar adjustments have
been made in the keys.
With their larger size and generally low densities, standard benthos
collection devices are of little help in surveys and quantitative studies.
Special dredges, crowfoot bars, SCUBA, and seines must be used to estimate
populations (see Starrett, 1971, for review of some commonly used methods).
Keys to the families (only Unionidae has been recorded from the
Laurentian Great Lakes), subfamilies, and tribes require the use of soft
anatomy. Most species, however, may be identified on the basis of hinge and
shell characters, many of which may be qualitative. Therefore, it has been
difficult historically to create workable keys to the unionid fauna. Even
with the fauna of the Great Lakes being somewhat limited, the key presented
here must make use of both shell characters and the soft anatomy of mature
specimens. Live mussels should be relaxed and fixed before preservation and
will be the easiest to identify. Empty shells in good condition may be
identified by going past the couplets where soft anatomy is used and then by
examining the figures. If difficulties remain, the guides of Clarke (1973)
and Burch (1973, 1975b) should be consulted.
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GASTROPODA
General distributions of most snails in the Laurentian Great Lakes are
not well known, and the systematics of many families and genera are in need
of much work. The list and keys presented here are as close to 'state-of-
the-art' as possible but in no way are definitive. As with unionids,
gastropods reach their greatest abundance and diversity in the near-shore,
rocky zones and areas of emergent vegetation, though a few species are
associated with the soft sediments of the open lake. The literal zones of
Lake Erie, Michigan, and the north shore of Ontario are composed of varying
combinations of mud, clay and sand which have a diverse mollusk fauna
including many species of snails. Much of the littoral zones of Lakes
Superior, Huron, and the southern shore of Ontario are quite rocky and the
fauna consequently is limited to gastropods.
Some aquatic pulmonates such as Physidae, Lymnaeidae, and the planorbid
Gyraulus cipeumstriatus estivate under logs, rocks, and leaves where they
may be collected by hand during dry periods. Several of the Physidae,
Lymnaeidae, Planorbidae, and Hydrobiidae are associated with submerged,
emergent, or floating gevetation and are collected by washing or shaking the
plants in a pail of warm water. Ancylidae must be gathered by hand since
they tend to cling tightly to smooth or "bottle like" surfaces of submerged
objects. Pleuroceridae and Viviparidae inhabit the rocky and softer
sediments of the open lake and standard benthological collecting techniques
may be adequate to estimate their populations. Very few quantitative studies
of snails are in the literature because of the variety of sampling methods
needed to examine the whole fauna.
With few exceptions, the snail fauna of the Laurentian Great Lakes may
be identified using shell characters alone. Preserved live specimens will
be easiest to identify particularly for the operculate species where the
operculum is a key character and it has remained attached. Although the
teeth of the radula are not diagnostic features of the keys, it may be
necessary to examine the radula for some species (e.g., the Lyrrmaea) thought
to be new to the Great Lakes (see Clarke, 1973 for methods).
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SECTION II
CHECKLIST OF FRESHWATER MOLLUSKS IN THE LAURENTIAN GREAT LAKES
The following list is based on museum records, benthological surveys,
and combines previously published and unpublished species lists.
GASTROPODA
Viviparidae
Campeloma deoisim (Say)
Campeloma rufum (Haldeman)
Viviparus geovgianus (Lea)
Valvatidae
Valvata bicarinata Lea
Valvata lewisi Currier
Valvata perdepressa Walker
Valvata pisoinalis (Miller)
Valvata sinoeva (Say)
Valvata tricarinata f. basalis
Vanatta
Valvata trioarinata f. pevconfusa
Walker
Valvata triearinata f. trioavinata
Say
Hydrobiidae
Amnicola limosa (Say)
Amni-oola walkeri Pilsbry
Bithynia tentaculata (Linnaeus)
Cineinnatia oinoinnatiensis
(Anthony)
Marstonia deoepta (Baker)
Probythinella lacustris (Baker)
Somatogyrus subglobosus (Say)
Pleuroceridae
Goniobasis livescens (Menke)
Pleurocera acuta (Rafinesque)
Lymnaeidae
Bul-imnea megasoma (Say)
Fossaria decampi (Streng)
Fossaria humilis Say
Fossaria obrussa Say
Lymnaea stagnalis appressa Say
Pseudosuocinea colwnella (Say)
Radix aupieulaT'La (Linnaeus)
Stagnioola oaperata (Say)
Stagnicola aatascopiwn f. oatasaop-ium
Say
Stagnicola aatasoopi-w f. nasoni
Baker
StagniGola emarginata (Say)
Stagnioola veflexa (Say)
Physidae
Fhysella gym-na sayi (Tappan)
Physella Integra (Haldeman)
Fhysella vinosa (Gould)
Pianorbidae
Gyraulus ciroumstriatus (Tryon)
Gyraulus deflectus (Say)
Gyraulus parvus (Say)
Helisoma anceps (Menke)
Helisoma oampanulatim (Say)
Helisoma oorpulentum (Say)
Helisoma trivolvis (Say)
Promenetus exaouous (Say)
Ancylidae
Ferrissia pavallela (Haldeman)
Ferrissia tarda (Say)
Laevapex fusous (Adams)
UNIONIDAE
(Lea)
Alasmidonta caloeolus
Amblema plioata Say
Anodonta oataraota Say
Anodonta grandis grandis Say
Anodonta grandis simpsoniana Lea
Anodonta irribecillis Say
Anodontoides fevussacianus (Lea)
CaTunculina parva (Barnes)
Cyolonaias tuberculata (Rafinesque)
Dysnomia sulcata Lea
Dysnomia triquetra (Rafinesque)
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Elliptic oomplanata (Lightfoot)
Elliptic dilatata (Rafinesque)
Fusoonaia flava (Rafinesque)
Lampsilis fasaiola Rafinesque
Lampsilis ovata (Say)
Lampsilis radiata radiata (Gmelin)
Lampsilis radiata siliquoidea
(Barnes)
Lasmigona aomplanata (Barnes)
Lasmigona oostata (Rafinesque)
Leptodea fragilis (Rafinesque)
Lignmia nasuta (Say)
Ligumia recta (Lamarck)
Obliquaria reflexa Rafinesque)
Obovaria olivaria (Rafinesque)
Obovaria subrotunda (Rafinesque)
Pleurobema aordatum f. ooccinewn
(Rafinesque)
Proptera alata (Say)
Proptera laevissima (Lea)
Ptyahobranchus fasciolare
(Rafinesque)
Quadrula nodulata Rafinesque
Quadmla pustulosa (Lea)
Quadrula quadrula (Rafinesque)
Simpsoniconcha ambigua (Say)
Strophitus undulatus (Say)
Truncilla donaeiformis (Lea)
Trunailla tvunoata Rafinesque
Villosa fabilis (Lea)
Villosa iris (Lea)
SPHAERIIDAE
Musaulium lacustre f. lacustre
(Miller)
Musaulium laoustre f. jayense Prime
Musaulium partumeiwn (Say)
Musaulium seouris (Prime)
Musaulium transversum (Say)
Pisidium adamsi Stimpson
Pisidium amnicum (Mliller)
Pisidium casertanum (Poli)
Pisidium compression Prime
Pisidium conventus Clessin
Pisidium dubium (Say)
Pisidium equilaterale Prime
Pisidium fallax Sterki
Pisidium ferrugineum Prime
Pisidium henslowanum (Sheppard)
Pisidium idahoense Roper
Pisidium lilljeborgi f. lilljeborgi
Esmark & Hoyer
Pisidium lilljeborgi f. cristatum
Sterki
Pisidium milium Held
Pisidium nitidum f. nitidum Jenyns
Pisidium nitidum f. pauperculatum
Sterki
Pisidium punotatum Sterki
Pisidium subtrunoatum Malm
Pisidium supinum Schmidt
Pisidium variabile Prime
Pisidium ventriaosum f. ventricosum
Prime
Pisidium -oentricosum f. rotundatum
Prime
Pisidium walkeri f. walkeri Sterki
Pisidium walkeri f. mainense Sterki
Sphaerium corneum (Linnaeus)
Sphaerium nitidum Westerlund
Sphaerium oooidentale (Prime)
Sphaerium rhomboideum (Say)
Sphaerium simile (Say)
Sphaerium striatinum (Lamarck)
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SECTION III
KEY TO THE FRESHWATER MOLLUSKS OF THE
LAURENTIAN GREAT LAKES
GASTROPODA
1 Shells whorled (Figs. 2, 3, 4) 2
Shells limpet-like (Fig. 5) ANCYLIDAE 9
«p«x —
protoconcK
spirt
tightly coiled
loosely coiltd
body whorl
maleitioni
plaited columolla
columtlla Umporloroto)
outtr lip
iptrturi
pirimmt
Figures 2-5. General shell shape and characters: 2- Lymnaeidae (Stagniaola
elodes); 3- Physidae (Physella sp.); 4- Planorbidae (Promenetus
exaeuous) ; 5- Ancylidae (Fewri-ssia pamllela) .
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2 (1)
3 (2)
Whorls in one plane, operculum absent (Fig. 4) .PLANORBIDAE.
Whorls in more than one plane, operculum present or absent
(Figs. 2, 5, 6)
Whorls sinistral (Fig. 3) PHYSIDAE.
Whorls dextral (Fig. 2)
12
3
21
4
ctnlliculiti iptrtun
piucitpiril opirculum
multitpiral •pirculvm
ci«cintric opucglum
carini
•mbilicil ilit
(pttttrilt)
nycispiril
•pifculim
Figures 6-9. General shell shape and characteristics: 6- Viviparidae
(Cconpelama deeisum); 7- Pleuroceridae (Goniobasts livescens);
8- Hydrobiidae (Arrmiaola limosa); 9- Valvatidae (Valvata trioarinata
f. triearinata).
4 (3) Without an operculum (Fig. 2); mantle cavity a lung ...
LYMNAEIDAE 24
Operculum present (Figs. 6, 7, 8, 9); with gills 5
5 (5) Spire only slightly elevated; operculum circular and
multispiral (Fig. 9) ..'. VALVATIDAE 37
Spire moderately to extremely elevated; operculum spiral
or concentric but not circular (Figs. 6, 7, 8) 6
6 (5) Operculum concentric (Fig. 6) 7
Operculum spiral (Figs. 7, 9) 8
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7 (6) Whorls flattened; umbilicus imperforate; adult shell
less than 13 mm high (Fig. 10): Bithynia tentaculata
Whorls inflated; umbilicus perforate; adult shell
greater than 15 mm high (Fig. 8) VIVIPARIDAE 45
Figure 10. Bithynia tentaculata (Hydrobiidae) showing concentric operculum.
8 (6)
9 (1)
Whorls flattened; umbilicus imperforate; adult shell
greater than 15 mm high (Fig. 7) PLEUROCERIDAE.
Whorls inflated; umbilicus perforate; adult shell less
than 12 mm high (Fig. 8) HYDROBIIDAE. . .
48
49
ANCYLIDAE. Apex smooth; shell with fine radial striae
on newer parts (Figs. 11, 12): Laevapex fusaus
Apex finely striate (may be eroded in older specimens)
shell with no striations on newer parts (Figs. 13-16) 10
Figures 11-16. Laevapex fuseus: 11- lateral aspect; 12- dorsal aspect,
Ferrissia parallela: 13- lateral aspect; 14- dorsal aspect.
Ferrissia tarda: 15- lateral aspect; 16- dorsal aspect.
10
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10 (9) Aperture with straight parallel sides; shell thin
(Figs. 5, 13, 14): Ferrissia parallela
Aperture ovate, sides curved; shell moderately thick;
apex may be pointed and recurved (Figs. 15, 16) 11
11 (10) Shell small, strongly elevated; apex dome-shaped;
peritreme ovoid (Figs. 15, 16): Fervissia tarda
Shell large, moderately elevated; apex acute; peritreme
broadly ovoid (see Basch, 1963; Hubendick, 1964;
Harman and Berg, 1971).
12 (2) PLANORBIDAE. Axial height 4 mm or more, diameter 7 mm or more
in post-juveniles; shell sinistral or dextral; aperture
length/shell diameter ration >0.51 or, if less, never
hirsute (Figs. 17-26) Helisoma
Axial height 1-4 mm, diameter less than 8 mm in post-
juveniles; shell dextral; aperture length/shell
diameter ratio <0.50 or, if greater, shell hirsute or
multicarinate (Figs. 27-33)
13
17
13 (12) Shell dextral; whorls flattened forming smooth-sided
funnel-shaped depressions on both sides of shell
(Figs. 17-19):
Shell sinistral; whorls rounded or flattened on one
side of shell (Figs. 21, 23, 26)
Helisoma anoeps
14
19
Figures 17-19. Helisoma anoeps: 17- dorsal aspect; 18- aperture aspect;
19- cross-section view.
14 (13) Aperture bell-shaped; whorls compressed with rounded
surfaces dorsally and ventrally (Figs. 20, 21):
Hel-isoma oampanulatim
Aperture flaring; whorls rounded on one side, flattened
on the other (Figs. 23, 24, 26) 15
11
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21
Figures 20-21.
view.
Helisoma eampanulatwn: 20- dorsal aspect; 21- cross-section
15 (14) Carinae present on body whorl (Fig. 25) 16
Carinae absent; whorls rounded ventrally and flattened
dorsally forming broad, flat surfaces, enclosed
by body whorl to create a flat-bottomed depression
(Figs. 22-24): Eelisoma trivolvis
Figures 22-24. Helisoma trivolvis: 22- dorsal aspect; 23- aperture aspect;
24- cross-section view.
16 (15) Carinae present at outer edge of upper and lower surfaces
of body whorl (Fig. 25); body whorl abaxially flattened,
either partly or completely (Fig. 26) (see Clarke, 1963
for subspecies): Eelisoma eoppulentum
Carinae present at least on early whorls and near centers
of both upper and lower surfaces; body whorl abaxially
rounded (see Clarke, 1973).
12
-------
26
Figures 25-26. Heli-soma oorpulentum: 25- dorsal aspect; 26- cross-section
view.
17 (12) Shell ornamented with denticles on inside with raised
growth lines; H:W ratio 1:2 or less or adult shell
less than 2 mm diameter (see Clarke, 1973; Harman and
Berg, 1971).
Shell smooth or with periostracal hairs or carinate or
H:W ratio greater than 1:2; adult diameter greater
than 2 mm diameter
18
18 (17) Shell extremely flattened, body whorl with a distinct
carina on outer circumference (Fig. 27): Promenetus exaauous
Shell not flattened (Figs. 29, 31, 33) 19
19 (18) Shell with hirsute periostracum and/or carina (Figs. 28,
29): Gyraulus deflectus
Shell without periostracal hairs and carina (Figs. 30-33) 20
29
Figures 27-29. Promenetus exacuous: 27-lateral aspect. Gyraulus
defleatus: 28- dorsal aspect; 29- lateral aspect.
13
-------
20 (19) Shell whitish or yellowish, semi-transparent; nearly
or entirely planospiral (Figs. 30, 31): Gyraulus c-ircumstriatus
Shell brownish, translucent but not transparent;
less planospiral, with dorsal and ventral aspects
distinctly different (Figs. 32, 33): Gyraulus parvus
31
33
Figures 30-33. Gyraulus ciraumstriatus: 30-dorsal aspect; 31- lateral
aspect. Gyraulus pawns: 32- dorsal aspect; 33- lateral aspect.
21 (3) PHYSIDAE. Shell elongate with very tall spire (Aplexa,
see Te, 1978).
Shell only moderately elevated above body whorl ....Physella...
22
22 (21) (This portion of key extracted from the recent revision by
Te, 1978, and includes only species he records from the
Great Lakes. Note that the key differs substantially from
Harman and Berg, 1971, and Te, 1975)
Shell symmetrical; whorls with only slight suture
indentations (Fig. 34): Physella gyvina sayi
Shell asymmetrical; whorls with distinct suture
indentations (Figs. 35, 36) 23
Figures 34-36. ^-Physella gyrina sayi; 35- Physella vinosa;
36- Physella integra.
14
-------
23 (22) Whorls strongly indented and rounded (Fig. 35);
shell reddish:
Whorls sharply indented and box-shaped (Fig. 36);
shell horn-colored:
Physella vinosa
Phusella integra
24 (4) LYMNAEIDAE. Shell narrow and elongate (W/L less
than 0.40) (Fig. 37) 25
Shell wider (W/L greater than 0.40) (Figs. 38-40) 26
25 (24) Shell 3 to 4 times higher than wide; columella
plaited (Fig. 37): Stagnisola reflexa
Shell more than 5 times higher than wide; columella
almost striaght (see Harman and Berg, 1971).
26 (24) Aperture more than 5 times higher than spire; shell
almost as wide as high (W/L greater than 0.90)
(Fig. 38): Radix auriaularia
Aperture less than 3 times higher than spire; shell
much higher than wide (W/L less than 0.70) (Figs. 39-41) 27
27 (26) Aperture more than 1.5 times higher than spire
(Figs. 41, 42) 28
Aperture less than 1.5 times higher than spire; shell
usually thin (Figs. 43-48) 29
28 (27) Shell solid and inflated; columella plait (Fig. 2)
moderately developed, poorly gyrate; adult shell less
than 0.50 mm high (Fig. 40): Stagniaola aatasaopium f. nasoni
Shell thin, only slightly inflated; columella plait,
clearly visible and gyrate but not well developed; adult
shell up to 0.75 cm high (Fig. 41): Pseudosuaainea columella
Figures 37-40. 37- Stagniaola reflexa; 38- Radix auriaularia:
39- Stagniaola oatasoopiwn f. aatasaopiim; 40- Stagniaola
oatasoopiwn f. nasoni.
15
-------
29 (27) Whorls loosely coiled; sutures deeply impressed 30
Whorls tightly coiled;, sutures shallow 35
30 (29) Body whorl near outer lip flattened laterally (Figs. 42, 43) 31
Body whorl near outer lip rounded (Figs. 44, 46) 32
31 (30) Whorls stongly shouldered; aperture narrowed
adapically; adult shell less than 12 mm (Fig. 42):
Fossari-a deeampi
Whorls weakly shouldered; aperture wider adapically;
adult shell less than 20 mm (Fig. 43): Fosscofla obrussa
32 (30) Spiral striae indistinct or absent (Fig. 44):
Fossaria hionilis
42
41
Figures 41-44. 41- Pseudosuecinea oolumel'La; 42- Fossaria deoampi;
43- Fosscocia obrussa; 44- Fossaria
33 (32) Body whorl with spiral, microscopic, blade-like
periostracal ridges; adult shell less than 15 mm
high (Fig. 45):
Body whorl with spiral striae but not blade like;
adult shell more than 15 mm high (Figs. 46, 47)
Stagn-loola caper>ata
34
34 (33) Inner lip of aperture strongly plaited with a small
umbilical chink; aperture not brownish or
purplish within (Fig. 46): Stagnioola emarginata
Inner lip of aperture gyrate plait and umbilical chink
absent; aperture brownish or purplish within
(Fig. 47): Bulirmea megasoma
35 (29) Spire elevated, whorls flattened; adults higher
than 35 mm (Fig. 48) : Lyrrmaea stagnalis appressa
Spire short, whorls more rounded; adults shorter
than 35 mm (Fig. 39) 36
16
-------
36 (35) Shell broad (W/L greater than 0.55); whorls not
malleated (Fig. 39): Stagnicola oatasooplum f. aatascopiwn
Shell narrower (W/L less than 0.55); whorls often
malleated (see Barman and Berg, 1971; Clarke 1973).
Figures 45-48. 45- Stagnicola caperata; 46- Stagnicola emarginata;
47- Bulirmea megasoma; 48- Lyrmaea stagnalis appressa.
37 (5) VALVATIDAE. Whorls angulate or carinate (Figs. 9, 49) 38
Whorls rounded and inflated (Figs. 51-54) 42
38 (37) Body whorl with 3 carinae (Fig. 49):
Valvata tficarinata f. tricarinata
Body whorl with 2 or fewer carinae (Fig. 50) 39
39 (38) Body whorl with 2 carinae (Fig. 50)
Body whorl with 1 carina (see Harman and Berg, 1971).
40
40 (39) Shell planospiral with nuclear whorl at same plane
as or sunken below following whorl (Fig. 50): Valvata bicarinata
Shell with a short spire (see Fig. 49) 41
41 (40) Upper and lower carinae present (see Fig. 49):
Valvata tricarinata f. peroonfusa
Upper and middle carinae present (see Fig. 49):
Valvata tricaipinata f. basalis
42 (37) Whorls with coarse growth lines (Fig. 51): Valvata lewisi
Whorls with fine growth lines (Figs. 52-54) 43
17
-------
Figures 49-51. ^-Valvata triaarinata f. trioarinata; 50- Valvata
biaarinata; 50- Valvata lewisi.
43 (42) Shell planospiral with nuclear whorl at same plane
as or sunken below following whorl (Fig. 52): Valvata perdepressa
Shell with a short spire (Figs. 53, 54) 44
44 (43) Spire conic (Fig. 53):
Spire attenuate (Fig. 54):
Valvata sincere
Valvata pisainalis
Figures 52-54. 52-Valvata perdepressa; 53- Valvata sinaera;
54- Vlavata piscinalis.
45 (7) VIVIPARIDAE. H:W ratio less than 1.35:1; whorls
inflated, outer lip usually straight (Figs. 55, 56) .-. .. 46
H:W ratio greater than 1.5:1 47
46 (45) Yellow-green with 4 brown bands on body whorl (Fig. 55):
Viviparus georg-ianus
Without brown bands; surface malleated; whorls
flattened on spire; large up to 70 mm high (Fig. 56):
Viviparus gaponiaus
47 (45) Apex of shell and interior aperture with a reddish color;
whorls of spire slightly impressed (Fig. 57): Campeloma rufiffn
Apex of shell and interior aperture without reddish color;
whorls of spire deeply impressed (Fig. 58): Campeloma decision
18
-------
Figures 55-58. 55- Viviparus georgianus; 56- Viviparus japonieus;
57- Campeloma rufim; 58- Campeloma deoiswn.
48 (8) PLEUROCERIDAE. Whorls very flat sided, often
shouldered, sutures shallow, aperture strongly
canaliculate (Fig. 59):
Whorls moderately flat sided, never shouldered,
sutures impressed, weakly canaliculate (Fig. 60):
Gonidbasis livesoens
Pleupooera aeuta
59
Figures 59-60. 59- Pleuvoaera aouta; 60- Goniobasis livesoens,
19
-------
49 (8) HYDROBIIDAE. Nuclear whorl elevated above second
whorl (Figs. 61-64) 50
Nuclear whorl level with second whorl or depressed
below it (Figs. 65, 66) 54
50 (49) Shell attenuate (Fig. 61) 51
Shell conic (Fig. 62) 52
51 (50) 4 or 5 whorls, umbilicus narrow or absent, columella
not reflected; 3 to 5 mm high (Fig. 61): Marstonia deoepta
6 or 7 whorls, umbilicus wide, columella reflected;
4 to 6 mm high (see Berry, 1943; Harman and Berg, 1971).
52 (50) Body whorl very large; adult shell higher than 9 mm
(Fig. 62): Somatogyrus subglobosus
Whorls increasing gradually in size; adult shells
less than 8 mm high (Figs. 63-66) 53
Figures 61-62. 61- Marstonia decepta; 62- Somatogyrus subglobosus.
53 (52) Adult shell 3 to 5 mm high; aperture broadly lacrimate
(Fig. 63): Cincinnatia Gincinnatiensis
Adult shell 1.5 to 2.5 mm high; aperture subcircular
(Fig. 64): Amnioola walkeri.
54 (49) Nuclear whorl at same plane as second whorl
(Fig. 65):
Nuclear whorl sunken below second whorl
(Fig. 66):
Amnioola limosa
Probythinella lacustris
20
-------
64
65
Figures 63-66. 63- Cineinnat-ia cinainnatiensis; 64- Amnieola walkeri;
65- Amnisola limosa; 66- Probythinella laaustris,
21
-------
ADDENDUM TO GASTROPOD KEY
At about the same time or possibly before the publication of the
Gastropoda keys used here, a guide the the Gastropoda of North America will
have been completed by Dr. John B. Burch of the University of Michigan
Museum and Department of Zoology. Burch?s work should be consulted for any
changes in systematics and synonymy of those species occurring in the
Laurentian Great Lakes. Both this study and that by Burch are being
published by the United States Environmental Protection Agency.
22
-------
All 4 dimibranchs serve as marsupia (i.e., appear
swollen in gravid females or water tubes numerous
with extra septa as seen in cross-section of the
females (Figs. 83, 84, 88-90) subfamily Arableminae 2
Only outer 2 demibranchs serve as marsupia
(Figs. 88-90) subfamily Unioninae 9
Beak or limbo
Ligament
Posterior slope —
Posterior ridge
Ventral margin
Umbo
Lateral
teeth
68
Interdentum
Pseudocardinal teeth
Figures 67-68. General features of a Unionidae shell: 67- exterior of right
valve; 68- interior of left valve.
23
-------
2 (1) Shell surface with distinct corrugations on posterior
slope (Fig. 69) 3
Shell surface without distinct corrugation on posterior
slope (Fig. 73) 5
3 (2) Shell surface with distinct pustules (see Burch, 1975b).
Shell surface without distinct pustules (Fig. 73) 4
4 (3) Adult shell small (less than 6 cm long); shell
corrugations fine (see Burch, 1975b).
Adult shell large (often up to 13 cm and sometimes
to 18 cm long); corrugations heavy (Fig. 69): Amblema plioata
5 (2) Shell surface pustulose (Figs. 70-72) 6
Shell surface smooth (Fig. 73) 8
6 (5) Shell lacking median sulcus on disc and umbonal
region (Figs. 71, 72) 7
Shell with median sulcus on surface which extends
from umbo to ventral margin; shell usually moderately
to heavily pustulose but occasionally smooth
(Fig. 70): Quadrula quadrula
Figures 69-70. 69- Amblema p1icata\ 70- Quadrula quadrula.
7 (6) Pustules on disc arranged in 2 divergent rows; shell
without green rays on umbonal region (Fig. 71): Quadmla nodulata
Pustules on disc evenly scattered over shell surface;
green rays present on umbonal region (Fig. 72);
some Lake Erie shells may be smooth: Quadrula pustulosa
8 (5) Shell triangular and high, inflated and heavy, with
very prominent and swollen beaks; posterior ridge
well developed (see Burch, 1975b)
Shell triangularly-rhomboidal or if triangular then
not as high as above; posterior ridge concave or
straight (Fig. 73): Fusconaia flava
24
-------
Figures 71-73. 71- Quadrula nodulata; 72- Quadmla pustulosa;
73- Fusconaia flava.
9 (1) Articulating hinge teeth absent or vestigial with only
pseudocardinal teeth present 10
Articulating hinge teeth present (Fig. 67) 17
10 (9) Pseudocardinal teeth absent with, at most, depressions
present 11
Pseudocardinal teeth present but vestigial (Fig. 67) 16
11 (10) Beak sculpture double-looped (Fig. 74) with lower apices
of loops elevated and nodulus and forming two short
radial rows on each valve (Figs 74, 75): Anodonta grand-is grandis
Beak sculpture single or double looped (Fig. 78) but
not nodulus - 12
12 (11) Beak raised, forming a smooth line along the
dorsal margin (Fig. 76): Anodonta irribeoillis
Beak distinctly raised from side view (Fig. 79) 13
Figures 74-76. Anodonta gvandis grandis: 74- dorsal view showing beak
sculpture; 75- lateral aspect. Anodonta imbesillis: 76- lateral aspect,
25
-------
13 (12) Beak sculpture single looped (Fig. 78), or faintly
double-looped 14
Beak sculpture distinctly double-looped (Fig. 74);
nacre white or iridescent (if pink see Burch, 1975b)
(Fig. 77): Anodonta aataraeta
14 (13) Major ridges of beak sculpture relatively coarse
(Figs. 78, 79); depression for pseudocardinal
teeth present but tooth blades or tubercles
absent: Strophitus undulatus
Major ridges of beak sculpture relatively fine 15
Figures 77-79. 77- Anodonta cataracta. Strophitus undulatus: 78- dorsal
view showing beak sculpture; 79- lateral aspect.
15 (14) Ridges of beak sculpture not parallel to concentric growth
lines of beak, but cross them obliquely (Fig. 80):
Anodontoides ferussacianus
Ridges of beak sculpture parallel to concentric growth
lines of beak (Fig. 81): Anodonta grandis simpsoniana
Figures 80-81. 80-Anodontoides fevussacianus; 81- Anodonta grandis
simpsoniana.
26
-------
16 (10) Pseudocardinal teeth thin and blade-like or posterior
slope with corrugated sculpture (see Burch, 1975b).
Pseudocardinal teeth tubercular and posterior
slope without corrugated sculpture (Fig. 82):
Simpsoniooncha ambigua
Figure 82. S-impsoni-ohocha amb-tgua.
17 (9) Gills of gravid females with secondary septa
dividing each water tube into 3 tubes (Fig. 83)
Septa and water tubes of gills undivided (Fig. 84)
18
21
—\f
83
84
Figures 83-84. Female gills: 83- division of water tubes by secondary
septa into three tubes; 84- unmodified gill^ glochidia shown in
one main water tube.
18 (17) Beak sculpture concentric; posterior end flatly truncate
(Fig. 85) (if rounded see Burch, 1975b): Alasmicbnta aalaeolus
Beak sculpture double-looped (Fig. 74) 19
19 (18) Posterior ridge with undulations; hinge teeth heavy
and rough 20
Posterior ridge without undulations; hinge teeth
smooth and more delicate (see Burch, 1975b).
20 (19) Shell with prominent wing; posterior slope with gentle
undulations (Fig. 86): Lasmigona aomplanata
Shell without wing; posterior slope strongly
corrugated (Fig. 87): Lasmigona oostata
27
-------
85
Figure 85. Alasmidonta calceolus.
Figures 86-87. 86- Lasmigona eomplanata; 87- Lasmigona eostata.
21 (17) Marsupium filling entire outer pair of gills (Fig. 88);
shells not sexually dimorphic
Marsupium confined to restricted regions of outer
demibranchs (Figs. 89, 90); shells commonly
sexually dimorphic (Figs 101, 102)
22
25
Figures 88-90. Marsupial gills: 88- outer section serving entirely as a
marsupium; 89 and 90- only restricted regions of outer gill serving
as a marsupium.
28
-------
22 (21) Shell surface sculpture with pustules (Fig. 91);
nacre purple: Cyolonaias tuberaulata
Shell surface without pustules; nacre with, pink,
or purple 23
23 (22) Shell triangular; beaks high and arched forward;
nacre white (Fig. 92): Pleuvobema aordatwn f. ooeeinewn
Shell elongate, rhomboidal (Figs. 93, 94); beaks
low, not arched; nacre purple, pink, or iridescent 24
Figures 91-92. 91- Cyclonaias tubereulata; 92 Pleurdbema oordatum
f. Gooaineim.
24 (23) Posterior ridge relatively close to dorsal margin
and bowed upward (Fig. 93):
Posterior ridge more median in position and nearly
straight (Fig. 94):
Elliptic) dilatata
Elliptic) oomplanata
Figures 93-94. 93- Elliptic dilatata', 94- Elliptic oomplanata.
29
-------
25 (21) Shell with single median row of large tubercles
(Fig. 95); tubercles alternate on each valve:
Shell without tubercles ,
ObliquaTia veflexa
26
26 (25) Shell elongate, L/H ratio greater than 2.0
(Figs. 96-98) 27
Shell higher, L/H ratio less than 2.0 (Figs. 99-106) 29
27 (26) Females with caruncle on inner edge of each side of
mantle in fromt of branchial opening; beak sculpture
of strong concentric ridges; adult shell less than
4 cm long (Fig. 96):
Females lack caruncles; beak structure of 1 or 2
subconcentric bars followed by 3 to 5 double-
looped bars; adult shell more than 5 cm long
(Figs. 97, 98)
Carunculina parva
28
Figures 95-96. 95- Obliquaria reflexa; 96- Carunaulina parva.
28 (27) Shell medium in size (adult less than 10 cm long),
usually compressed; posterior ridge extends to
near umbo; posterior slope usually concave; posterior
margin meets dorsal margin at angle forming a low wing;
periostracum olive green to olive brown (Fig. 97): Ligumia nasuta
Shell larger (adults greater than 10 cm long), usually
inflated; posterior ridge indistinct near posterior
margin of shell and is broadly rounded near umbo;
posterior slope usually not concave; without
dorso-posterior wing; periostracum dark green to
black (Fig. 98): Ligwnia recta
Figures 97-98. 97- Ligimia nasuta; 98- Ligumia reota.
30
-------
29 (26) Shell high-oval; teeth heavy; nacre white (Figs. 99, 100) 30
Shell elongate, subelliptical, subrhomboidal,
triangular, or oval; teeth heavy or thin
(Figs. 101-110) 31
30 (29) Beaks of shell high and arched strongly anteriorly
(Fig. 99): Obovaria olivaria
Beaks of shell lower, central in positon and not
strongly arched (Fig. 100): Obovaria subrotunda
100
Figures 99-100. 99- Obovaria olivcccia; 100- Obovaria subrotunda.
31 (29) Shell small (adults less than 6 cm long), solid,
strongly sexually dimorphic (females with a greatly
expanded projection where shell covers marsupial
area of gills); posterior end with radiating raised
ridges (Figs. 101-102) 32
Shell without above combination of characters
(Figs. 104-110) 33
32 (31) Rays discontinuous, especially on posterior ridge,
giving chevroned appearance (Figs. 101-102): Dysnomia triquetva
Rays continuous, shell high, inflated (Fig. 103): Dysnomia suloata
102
Figures 101-103. Dysnomia tTi,quet-r>a\ 101- female; 102- male.
Dysnomi,a suloata: 103- male.
31
-------
33 (31) Posterior ridge angular (Fig. 67) 34
Posterior ridge rounded or absent 36
34 (33) Shell compressed; beak cavities shallow; color rays
on shell with or without U-shaped markings
(Figs. 105, 106)
Shell inflated in upper half; beak cavities deep;
color rays present (without U-shaped markings)
(Fig. 104):
35
Lampsilis ovata
104
Figure 104. Lampsilis ovata.
35 (34) Posterior ridge sharp, distinct down to ventral
margin of shell; posterior slope very short and
very steep (Fig. 105): Trunoilla truncata
Posterior ridge angular but becoming round and
fading out near ventral margin of shell (Fig. 106):
Trunoilla donaciformis
105
106
Figures 105-106. 105- Trunoilla truncata; 106- Truncilla donaeiformis.
32
-------
36 (33) Pseudocardinal teeth well developed; dorso-posterior
wing present or absent (Figs. 108-112) 37
Pseudocardinal teeth (Fig. 68) poorly developed;
dorso-posterior wing conspicuous (Fig. 107):
Proptera laevissima and Leptodea fragilis*
*It is difficult to distinguish between these species
in old or worn shells. In live specimens, L. fragilis
has a yellow to light green shell with white to pink
nacre and may be abundant along the shores of
western Lake Erie. P. laevissima is rare in the
lakes, the shell is purple to brownish-purple, and
the nacre is very dark pink to dark purple.
37 (36) Well-developed wing present (Fig. 108):
Wing usually lacking, but if present very low
and poorly developed (Figs. 109, 110, 116)
Proptera alata
38
107
108
Figures 107-108. 107-Leptodea fragilis; 108- Proptera alata.
38 (37) Posterior mantle margin in front of branchial
opening undifferentiated; disc with broad, green
rays (Fig. 109): Ptychobranohus fasaiolare
Posterior mantle margins differentiated into
papillate or flap-like structures (Figs. Ill, 114);
disc with or without rays 39
109
Figure 190. Ptyehobranchus fasciolare.
33
-------
39 (38) Posterior mantle margins with long papillate
projections (Fig. Ill); shell rayed (Fig. 110):
Villosa iris and Villosa fabilis*
*The species of Villosa are difficult to distinguish
but the above two have been recorded from the Great
Lakes. Burch (1975b) gives outlines of other eastern
North American forms.
Posterior mantle margins with ribbon-like flaps (Fig. 114) 40
110
111
112
Figures 110-112. Villosa iris: 110- lateral aspect; 111- papillate mantle
margin. Villosa fabilis: 112- outline of lateral aspect.
40 (39) Shell elongate (L/H ratio of males greater than 1.6)
(Figs. 115-116) 41
Shell higher (L/H ratio of males less than 1.6);
beaks sculptured with small double-looped ridges;
color rays extend evenly over shell (Fig. 113):
Lampsilis fasoiola
113
114
Figures 113-114. Lampsilis fasciola: 113- lateral aspect; 114- flap-like
mantle margin.
41 (40) Color rays extend over entire shell and extend
to ventral margin without fading in color (Fig. 115):
Lampsilis radiata radiata
Color rays absent or limited to posterior slope
or fading before reaching ventral margin (Fig. 116):
Lampsilis radiata siliquoidea
34
-------
115
116
Figures 115-116. 115- Lampsilis vadiata radiata; 116- Lampsilis radiata
siliquoidea.
35
-------
SPHAERIIDAE
General hinge characters for Sphaeriidae are given in Figs. 117, 118.
Though a typical Pisidium is figured, the tooth placement remains the same
for both Musoulium and Sphaeriwn. In all other drawings in the key, the
position of cusps on the lateral teeth is indicated by an "X". Scale lines
indicate 1.0 mm. Photographs were taken with the aid of a scanning electron
microscope and are 10 to 150*.
Cusps
Anterior lateral (~A III
teeth |_A I
Anterior slope —
117
Posterior lateral
tooth
Pll
118
PMll Posterior lateral
PI J tooth
Cusps
Posterior sulcus
Figures 117-118. General hinge and shell characters for Sphaeriidae:
117- interior of right valve of Pisidium dubium; 118- interior of
left valve of Pisidium dubium.
36
-------
1 Beaks anterior, or if subcentral, on the anterior side
of center (Figs. 124, 132); both anal and branchial
siphons present (Figs. 119, 120) subfamily Sphaeriinae... 2
Beaks posterior, or if subcentral, on the posterior
side of center (Figs. 117, 118); only the anal
siphon present, the branchial siphon being
represented by the mantle cleft (Fig. 121)
subfamily Pisidiinae.... 13
2 (1) True calyculae (caps) present on adults born near the
end of summer (Figs. 124, 140); absent on those born
during summer; living adult shells thin, yellowish,
and semitransparent; siphons united only at basal
half (Fig. 119); sulcus of AI, AIII, and PI, PHI
not distinctly tuberculated Musculium 3
Caps absent, or if present, are not true calyculate;
living adult shells usually heavy and/or opaque,
periostracum usually brownish or blackish (juveniles
yellowish; sulcus of .AI, AIII and PI, PHI usually
with distinct tubercles (Fig. 160); siphons united
for entire length (Fig. 120) Sphaerium 8
119 120
Figures 119-121. Branchial (lower) and anal (upper) siphons of
Sphaeriidae: 119- Musoulium; 120- Sphaerium; 121- Pis-idiwn.
3 (2) H/L ratio less than 0.75; cusps of lateral teeth occur
before the dorso-anterior and dorso-posterior angles
(Figs. 122, 123, 127, 129); posterior end of C3
greatly enlarged and triangular, usually bifid; anterior
part narrow and of uniform thickness throughout
(Figs. 122, 126); shell shape as in Figs. 124, 125:
Musaulium transversum
H/L ratio greater than 0.76; cusps of lateral teeth
occur before or after the dorso-anterior and
dorso-posterior angles; posterior end of C3
usually not greatly swollen, but if so, not
trian'gular; anterior part thick or thin (Figs. 130-
133, 138-141) 4
37
-------
122
123
124
125
Figures 122-125. Musaulium transversim: 122- right hinge; 123- left hinge;
124- lateral aspect of left valve; 125-end view. Also see Figs. 126-
129, page 39.
4 (3) Cusps of lateral teeth arising before dorso-anterior
and dorso-posterior angles of shell (Figs. 130-135);
C3 usually enlarged posteriorly and anteriorly
(Figs. 130, 134) (also Figs. 131-133, 136, 137):
Musculiym partwneiwn
Cusps of lateral teeth arising on or after the
dorso-anterior and dorso-posterior angles of
shell (Fig. 138); C3 only slightly enlarged
posteriorly or of uniform thickness throughout
(Figs. 138, 142) 5
130
131
132
133
Figures 130-133. Musauliwn partumeiim: 130-right hinge; 131- left hinge;
132- lateral aspect of left valve; 133- end view. Also see Figs.
134-137, page 40.
38
-------
126
127
Figures 126-129. Musauliwn
transversum:
126- cardinal teeth of left valve;
127- outline of left valve;
128- cardinal teeth of left valve;
129- outline of left valve.
12S
129
39
-------
134
135
Figures 134-137. Musculium
paptumeium:
134- cardinal tooth of right valve;
135- outline of right valve;
136- cardinal teeth of left valve;
137- outline of left valve.
136
137
40
-------
5 (4) Ventral margin sloped steeply upward beginning near
posterior end, anterior margin rounded (Fig. 140)
(also Figs. 138-145): Musculiim seauifis
Ventral margin with only a slight upward slope or
none, anterior margin angled (Fig. 154) 6
Figures 138-141. Musauliwn sectafis: 138- right hinge; 139- left hinge;
140- lateral aspect of left valve; 141- end view. Also see Figs.
142-145, page 42.
6 (5) Shell more or less square in outline, dorsal margin
straight (Fig. 148) (also Figs. 146-151):
Musculium lacustve f. jayense
Shell more or less triangular in outline, dorsal
margin curved (Fig. 154) 7
146
147
148
149
Figures 146-149. Musauliwn laeustre f. jayense: 146- right hinge;
147- left hinge; 148- lateral aspect of left valve; 149- end view.
Also see Figs. 150, 151, page 43.
41
-------
142
143
Figures 142-145. Museulium securis:
142- cardinal tooth of right valve;
143- outline of right valve;
144- cardinal teeth of left valve;
145- outline of left valve.
144
145
42
-------
150
Figures 150-151. MusGuliivn laaustre
f. jayense:
150- outline of right valve;
151- outline of left valve.
151
43
-------
7 (6) Dorsal margin joined by posterior and anterior
margins with similar, usually steep, angles;
C2 straight (see Herrington, 1962; Burch, 1975a).
Dorso-posterior margin with less slope than dorso-
anterior margin (Fig. 154); C2 curved or bent
(Figs. 152-159): fJusculiwn laeustre f. lacustre
155
Figures 152-155. Musoulium laoustre f. laeustre: 152- right hinge;
153- left hinge; 154- lateral aspect of left valve; 155- end view.
Also see Figs. 156-159, page 45.
8 (2) Shell sculptured with coarse striae (8 or fewer
per mm in middle of shell); shell heavy; C2
stump-like (Fig. 167) or canine shaped (Fig. 161)
Shell sculptured with fine striae (12 or more per
mm in middle of shell); shell usually thin; C2
thin, straight and incisor-shaped or bent and
chevron-shaped (Figs. 179, 185)
10
44
-------
156
157
Figures 156-159. tfoisculiwn lacustre
f. lacustre:
156- cardinal tooth of right valve;
157- outline of right valve;
158- cardinal teeth of left valve;
159- outline of left valve
158
159
45
-------
Sphaerium. simile
9 (8) Striae evenly spaced; adults longer than 1.6 cm;
shell long in outline (Fig. 160-165):
Striae not evely spaced; adults shorter than
1.5 cm; shell usually short in outline
(Figs 166-177): Sphaerium striatinum*
*Two forms have been recorded from the Great
Lakes which differ in shell shape and position
of C2 and C4; the forms may not always be distinct.
(Figs. 166-169, 174-177): Sphaerium striatinum f. emarainatum
(Figs. 170-173): Sphaerium striatinum f. acuminatum
160
161
163
162
Figures 160-163. Sphaerium simile: 160- right hinge showing detail of
AI and AIII; 161- left hinge showing shape of C2 and C4 from medial
view; 162- lateral aspect of left valve; 163- end view. Also see
Figs. 164-165, page 47.
46
-------
164
Figures 164-165. Sphaeriwn simile:
164- right valve;
165- left valve.
165
47
-------
168
169
Figures 166-169. Sphaeriiffn striatimm f. emarginatwn: 166- right hinge;
167- left hinge showing shape of C2 and C4 rom medial view; 168-
lateral aspect of left valve; 169- end view. Also see Figs. 175-
177, page 49.
170
171
172
173
Figures 170-173. Sphaeriim striatimian f. aoiminaUcm: 170- right hinge;
171- left hinge showing shape of C2 and C4 from medial view; 172-
lateral aspect of left valve; 173- end view.
48
-------
174
175
Figures 174-177. Sphaerium
striatinwn f. emapginatum:
174- cardinal tooth of right valve;
175- right valve;
176- cardinal teeth of left valve;
177- left valve.
176
177
49
-------
10 (8) All I and usually PHI of right valve less than 1.5
times longer than C3, or cusp of All less than
1.5 times longer than combined length of C2 and
C4; adults usually less than 8 mm long (Figs. 188,
194, 200) 11
A1II and usually PHI of right valve more than
twice the length of C3, or cusp of AH more than
twice the combined length of C2 and C4; adults
longer than 8 mm; shell rectangular in outline
(Figs. 178-183): Sphaeriwn Thomboi-dewn
180
181
Figures 178-181. Sphaerium rhomboideum: 178- right hinge; 179- left
hinge showing shape of C2 and C4 from medial view; 180- lateral
aspect of left valve; 181- end view. Also see Figs. 182-183, page
51.
11 (10) Shell glossy; striae maintain spacing and height
in region of beaks; C3 of more or less uniform
thickness throughout, only slightly curved and
parallel with hinge plate (Figs. 184-189): Sphaerium nit-idum
Shell dull; striae fade in region of beaks; C3 bent
and swollen posteriorly (Figs. 190-195) .' 12
12 (11) Umbo slightly raised above dorsal margin; posterior
margin truncate, ventral margin sloped slightly
upward anteriorly; C4 almost completely overlapping
C2 and more or less parallel (Figs. 190-195): Sphaerium corneum
Umbo distinctly raised above dorsal margin; posterior
end rounded, ventral margin symmetrical; C2 and C4
slope in opposite directions with little overlapping;
in habitats wihich dry up part of the year (Figs.
196-201): Sphaeriwn oooidentale
50
-------
182
Figures 182-183. Sphaeriwn
rhomb o-ideum:
182- right valve;
183- left valve.
183
51
-------
184
Figures 184-187. Sphaerium nitidum: 184- right hinge; 185- left hinge
showing shape of C2 and C4 from medial view; 186- lateral aspect of
left valve; 187- end view. Also see Figs. 188-189, page 53.
190
191
192
193
Figures 190-193. Sphaerium oornewn: 190- right hinge; 191- left hinge;
192- lateral aspect of left valve; 193- end view. Aslo see Figs.
194-195, page 54.
52
-------
18S
Figures 188-189. Sphaerium nitidum:
188- right valve;
189- left valve.
189
53
-------
194
Figures 194-195. Sphaeriwn aornewn:
194- right valve;
195- left valve.
195
54
-------
196
197
198
199
Figures 196-199. Sphaeriwn occidentale: 196- right hinge; 197- left hinge;
198- lateral aspect of left valve; 199- end view. Also see Figs. 200-
201, page 56.
13
(1) C3 greatly recurved and an inverted J or U shape
(Figs. 202, 206, 208, 212); striae coarse, 10 or
fewer per mm; adults 6 mm long or more
only slightly curved, never an inverted J or
U shape (Figs. 214, 222); striae fine, 15 or
more per mm; except for Pisidi-tm adamsi, adults
less than 6 mm long
C3
14
15
Pis-idium dubium
14 (13) Striae fade on beaks; umbo positioned vary far
back, near posterior margin (Fig. 204); C3
and inverted U with both anterior and posterior
ends enlarged (Figs. 202, 206); cardinals
nearer posterior end than anterior end (Figs.
202, 203):
Striae prominent on beaks; umbo more central and
not as close to posterior margin (Fig. 210);
C3 an inverted J with only the posterior end
enlarged (Figs. 208, 212); cardinals nearer
anterior end than posterior end (Figs. 208, 209): Pisidiwn armieum
15 (13) Shell with one or more prominent dorsal ridges
(Figs. 217- 225, 231, 239) 16
Shell without prominent dorsal ridges (Fig. 249) 19
16 (15) Cardinals on anterior side of center of hinge
(Figs. 242, 250) 17
Cardinals central in position (Figs. 228, 229, 233,
235) 18
55
-------
200
Figures 200-201. Sphaerium
OGcidenta1e\
200- right valve;
201- left valve.
201
56
-------
202
205
Figures 202-205. Pisiditm dubiwn: 202- right hinge; 203- left hinge:
204- lateral aspect of left valve; 205- end view. Also see Figs.
206-207, page 58.
211
Figures 208-211. Pisidium cormiown: 208- right hinge; 209- left hinge;
210- lateral aspect of left valve; 211- end view. Also see Figs.
212-213, page 59.
57
-------
206
Figures 206-207. Pisidium dubium:
206- right valve;
207- left valve.
207
58
-------
212
Figures 212-213. Pisidiim armiawn:
212- right valve;
213- left valve.
213
59
-------
17 (16) Shell dull; striae distinct and evenly spaced;
shell longer in outline; C2 chevron-shaped
and longer than C4 (Figs. 214-221):
Shell silky to glossy, short in outline; striae
fine, indistinct, irregular; C2 stump-like and
shorter than C4 (Figs. 222-227): Pisidium lilljebovgi f.
Pisidium henslowanwn
214
215
216
217
Figures 214-217. Pisidium henslowanum: 214- right hinge; 215 left hinge;
216- lateral aspect of left valve; 217- end view showing umbonal
ridges at arrow. Also see Figs. 218-221, page 61.
222
223
224
225
Figures 222-225. Pisidium lilljeborgi f. eristatum: 222- right hinge;
223- left hinge: 224- lateral aspect of left valve; 225- end view
showing umbonal ridges. Also see Figs. 226-227, page 62.
60
-------
218
219
Figures 218-221. Pisidiwn
hens lowanum:
218- cardinal tooth of right valve;
219- right valve;
220- cardinal teeth of left valve;
221- left valve.
220
221
61
-------
226
Figures 226-227. Pisidium
Hlljeborgi f. eristatwn:
226- right valve;
227- left valve.
227
62
-------
18 (16) Intercuspal to shell length ratio greater than
0.50 (Fig. 229); C2 chevron-shaped (Figs.
229, 234, 235, also Figs. 228-235):
Intercuspal to shell length ratio less than
0.45 (Fig. 237); C2 stump-like (Figs. 237,
241, also Figs. 236-241):
Pisidium supinwn
Pisidiwn compression
223
229
230
231
Figures 228-231. Pisidium sup^n^cm: 228- right hinge; 229- left hinge
showing shape of C2 and C4 from medial view, and method of measuring
intercuspal length; 230- lateral aspect of left valve; 231- end view.
Also see Figs. 232-235, page 64.
236
237
23S
239
Figures 236-239. Pisidiwn compression: 236- right hinge; 237- left hinge
showing shape of C2 and C4 from medial view; 238- lateral aspect of
left valve; 239- end view. Also see Figs. 240-241, page 65.
63
-------
232
233
Figures 232-235. Pisidiim supinwn:
232- cardinal tooth of right valve;
233- right valve;
234- cardinal teeth of left valve;
235- left valve.
234
235
64
-------
240
Figures 240-241. Pisidiim
compression:
240- right valve;
241- left valve.
241
65
-------
19 (15) Cardinals on anterior side of center of hinge
(Figs. 242, 243, 250, 251) 20
Cardinals central on hinge (Figs. 323, 324,
327, 328) 30
20 (19) Hinge short, less than 3/4 shell length (Figs.
242, 243, 250, 251) 21
Hinge long, more than 3/4 shell length (Figs.
315, 316, 320, 322) 29
21 (20) Anterior end of posterior sulcus in right valve
closed (Figs. 242, 250); length of hinge plate
very short, distance between All and PII only
1/3 of shell length (Fig. 243); shell inflated
(Figs. 245, 255)
Anterior end of posterior sulcus in right valve
open (Figs. 268, 272); distance between All
and PII greater than 1/3 of shell length; shell
inflated or not inflated
22
23
22 (21) Beaks subcentral; space between C2 and cusp of PII
less than 1.5 times length of C2 (Figs. 243,
251, also Figs. 242-251): Pisidium ventrioosurn f. rotundatwn
Beaks terminal; space between C2 and cusp of PII
more than 1.5 times the length of C2 Figs. 253,
257, also Figs. 252-257): Pisidium ventriaosum f. ventviaoswri
242
243
245
244
Figures 242-245. Pisidium ventrioosum f. rotundatum: 242- right hinge;
243- left hinge showing method of determining hinge length (distance
between junction of posterior lateral tooth with shell and junction
of anterior lateral tooth with shell) and center of hinge (1/2 hinge
length): 244- lateral aspect of left valve showing method of
determining shell length; 245- end view. Also see Figs. 246-249, page
67 and Figs. 250-251, page 68.
66
-------
248 249
Figures 246-249. Variant of P-isidium ventricoswn f. potundatum: 246- right
hinge; 247- left hinge showing shape of C2, C4, and All from medial
view; 248- lateral aspect of left valve; 249- end view. Also see Figs.
242-245, page 66 and 250-251, page 68.
252
254 255
253
Figures 252-255. Pisidium ventrieoswn f. ventricosum: 252- right hinge;
253- left hinge showing shape of C2, C4 and All from medial view;
254- lateral aspect of left valve. Also see Figs. 256-257, page 69.
67
-------
250
Figures 250-251. Pisidium
ventrioosum f. rotundatwn:
250- right valve;
251- left valve.
251
68
-------
256
Figures 256-257. Pisidium
ventrieosum f. ventriaosum:
250- right valve;
251- left valve.
257
69
-------
23 (21) Posterior end of C3 enlarged 3 to 5 times more
than anterior end (Figs. 268, 272), often
toothbrush-shaped (Fig. 258): shell dull or
glossy 24
Posterior end of C3 less than twice the thickness
of anterior end (Figs. 280, 284); shell glossy 26
24 (23) Shell dull; umbo clearly posterior; dorso-
anterior margin rounded (Fig. 270) 25
Shell glossy; umbo more central; dorso-anterior
margin angulate (Figs. 258, 260, also Figs.
258-267): Pisidium lilljeborgi f. lilljeborgi
258
259
260
262
265
Figures 258-265. Pisidium lilljeborgi f. lilljeborgi: 258- right hinge;
259- left hinge; 260- lateral aspect of left valve; 261- end view.
Variant of Pisidium lilljeborgi f. lilljeborgi: 262- right hinge;
263- left hinge; 264- lateral aspect of left valve; 265- end view.
Also see Figs. 266-267, page 71.
70
-------
266
Figures 266-267. Pisidium
lilljeborgi f.
266- right valve;
267- left valve.
267
71
-------
25 (24) C2 low and canine-shaped (Figs. 269,273); shell
pointed anteriorly (Fig. 270, also Figs. 268-
273): Pisidiwn walkeri f. walkeri
C2 high and incisor shaped (Figs. 275, 279); shell
rounded anteriorly (Fig 275, also Figs. 274-
279): Pisidiwn walkeri f. mainense
270
271
Figures 268-271. Pisidiwn walkeri f. walkeri: 268- right hinge; 269- left
hinge showing shape of C2 and C4 from medial view; 270 lateral aspect
of left valve; 271- end view. Also see Figs. 272-273, page 73.
277
Figures 274-277. Pisidiwn walkeri f. mainense: 274- right hinge; 275- left
hinge showing shape of C2 and C4 from medial view; 276- lateral aspect
of left valve; 277- end view. Also see Figs. 278-279, page 74.
72
-------
272
Figures 272-273. Pisidiwn walkeri
f. walkeri:
272- right valve;
273- left valve.
273
73
-------
278
Figures 278-279. Pisidium walkeri
f. mainense:
278- right valve;
279- left valve.
279
74
-------
26 (23) Shell inflated, ventral margin more or less straight
in side view and truncated in end view (Figs. 282,
283, also Figs. 280-287): Pisidium miliwrn
Shell not inflated, or if so, without a truncated
or straight ventral margin (Figs. 290, 304) 27
2S2 283
Figures 280-283. Pisidium miliwn: 280- right hinge; 281- left hinge;
282- lateral view of left valve; 283- end view. Also see Figs. 284-
287, page 76.
27 (26) Space between cardinals and All small, less
than 1.5 times length of C3 (Figs. 288, 289,
295); beaks often tuberculate (Figs. 288-295):
Pisidiwn fevruginevm
Space between cardinals and All large, usually
more than twice the length of C3 (Figs. 296,
297); beaks never tuberculate (Figs. 289-304) 28
283
289
290 291
Figures 288-291. Pisidium ferrugineum: 288- right hinge; 289- left hinge;
290- lateral aspect of left valve; 291- end view. Also see Figs. 292-
295, page 77.
75
-------
284
2S5
Figures 284-287. Pisidiim miliwn:
284- cardinal tooth of right valve;
285- right valve;
286- cardinal teeth of left valve;
287- left valve.
2S6
287
76
-------
292
293
Figures 292-295. Pisidium
ferrugineiffn:
292- cardinal tooth of right valve;
293- right valve;
294- cardinal teeth of left valve;
295- left valve.
294
295
77
-------
28 (27) Anterior slope long and gradual, the angle at
dorso-anterior margin beginning above the
cusps of anterior laterals; dorsal and
ventral margins not parallel; dorso-anterior
margin without angle (Fig. 298, also Figs. 296-301):
Pisidium subtruncatum
Anterior slope short and steep, the angle at
dorso-anterior margin beginning below the
cusps of anterior laterals (Figs. 302-303);
dorsal and ventral margins nearly parallel
and dorso-anterior margin distinctly angulate
(Fig. 304, also Figs. 302-307): Pisidium nitidum f. nitidum
296
297
29S
299
Figures 296-299. Pisidium subtrunoatum: 296- right hinge; 297- left hinge;
298- lateral aspect of left valve; 299- end view. Also see Figs. 300-
301, page 79.
303
304 305
Figures 302-305. Pisidium nitidum f. nitidum: 302- right hinge; 303- left
hinge; 304- lateral aspect of left valve; 305- end view. Also see
Figs. 306-307, page 80.
78
-------
300
Figures 300-301. Pisidivm
subtrunaatum:
300- right valve;
301- left valve.
301
79
-------
306
Figures 306-307.
f. nitidwn:
306- right valve;
307- left valve.
Pi-si-di-un ni,tidum
307
80
-------
29 (20) C3 club-shaped, C2 and C4 short and of similar
shape and parallel with each other (Figs. 308,
309, 312-314); shell shaped like a parallel-
ogram (Fig. 310); shell thin; found only in
cold waters, northern or deep lakes (Figs. 308-
314): Pisidiim eonventus
C3 bent and usually enlarged posteriorly, C2 and
C4 of different shapes and usually not parallel
(Figs. 315, 316); anterior and posterior margins
sloped at different angles, not parallel; shell
thin to heavy (Figs. 315-322); umbones often
flattened (Fig. 318): Pisidiwn fallax
310 311
Figures 308-311. Pisidiwn aonventus: 308- right hinge; 309- left hinge;
310- lateral aspect of left valve; 311- end view. Also see Figs. 312-
314, page 82.
315
316
317 318
Figures 315-318. Pisidiwn fallax: 315- right hinge; 316- left hinge;
317- lateral aspect of left valve; 318- end view. Also see Figs. 319-
322, page 83.
81
-------
312
Figures 312-314. Pisidium
conventus:
312- left valve;
313- right valve;
314- cardinal teeth of left valve
313
314
82
-------
319
320
Figures 319-322. Pisidium fallax:
319- cardinal tooth of right valve;
320- right valve;
321- cardinal teeth of left valve;
322- left valve.
321
322
83
-------
30 (19) Hinge short, less than 3/4 shell length (see
Figs. 243, 244) 31
Hinge long, more than 3/4 shell length (see
Figs. 243, 244) 33
31 (30) Cusp of All proximal (nearest utnbo) or on
proximal side of center (Fig. 324); hinge
heavy; shell very glossy (Figs. 323-328):
Cusp of All distal (farthest from umbo) or on
distal side of center (Fig. 330); hinge light;
shell glossy to dull 32
Pisidiwn var-Labile
323
324
325 326
Figures 323-326. Pisidium variabile: 323- right hinge; 324- left hinge
showing proximal position of cusp of All; 325- lateral aspect of left
valve; 326- end view. Also see Figs. 327-328, page 85.
32 (31) H:L ratio more than 0.86; shell glossy and
laterally compressed (Fig. 332); adults
longer than 6 mm (Figs. 329-334):
H:L ratio less than 0.85; shell silky to dull,
not laterally compressed (Fig. 338); adults
less than 6 mm long Figs. 335-344):
Pisidium idahoense
Pisidium aasertanwn
33 (30) C3 long and thin and usually of similar thickness
throughout (Figs. 245, 249); C2 and C4 long, thin,
of similar shape but may be of different lengths,
more or less parallel with each other and with
hinge (Figs. 345-350): Pisidium nitidum f. pauperculum
C3 enlarged posteriorly (Figs. 351, 355); C2 and C4
short and parallel, or if of different shapes,
not parallel with each other or with hinge
(Figs. 352-356) 34
84
-------
327
Figures 327-328. Pisidiwn
varidb-ile:
327- right valve;
328- left valve.
32S
85
-------
329
330
331 332
Figures 329-332. Pisidiwn iddhoense: 329- right hinge; 330- left hinge
showing distal position of cusp of All; 331- lateral aspect of left
valve; 332- end view. Also see Figs. 333-334, page 88.
337 338
Figures 335-338. Pisidiwn casertamm: 335- right hinge; 336- left hinge;
337- lateral aspect of left valve; 338- end view. Also see Figs. 339-
342, page 87, Figs. 343-344, page 89, and Figs. 396-374, page 96.
86
-------
341 342
Figures 339-342. Variant of Pisidiim casertanwn: 339- right hinge; 340- left
hinge; 341- lateral aspect of left valve; 342- end view. Also see Figs.
335-338, page 86, Figs. 343-344, page 89, and Figs. 369-374, page 96.
347
348
Figures 345-348. Pisidium nitidum f. pauperculum: 345- right hinge;
346- left hinge; 347- lateral aspect of left valve; 348- end view.
Also see Figs. 349-350, page 90.
87
-------
333
Figures 333-334. Pisidiwn
'Ldahoense:
333- right valve;
334- left valve.
334
-------
343
Figures 343-344. P-isidium
easertcmwn:
343- right valve;
344- left valve.
344
89
-------
349
Figures 349-350. Pisidiim nitidwn
f. pauperculum:
349- right valve;
350- left valve.
350
90
-------
34 (33) Shell glossy; hinge very heavy (Figs. 351-356): Pisidittm equilatevale
Shell dull to glossy; hinge heavy or light 35
351
352
353
354
Figures. 351-354. Pisidiwn equilaterale: 351- right hinge; 352- left hinge;
353- lateral aspect of left valve; 354- end view. Also see Figs. 355-
356, page 92.
35 (34) C2 and C4 short and parallel with hinge and
with each other (Figs. 358, 362): C3 club-
shaped (Figs. 357, 361); striae distinct, evenly
spaced; adult shell less than 2 mm long
(Figs. 357-362): Pisidiwn punotatum
C2 and C4 of different shapes (Figs. 364, 368);
C3 enlarged posteriorly but not club-
shaped (Figs. 363, 367); striae usually not
distinct and evenly spaced; adult shell more
than 3 mm long 36
36 (35) H:L ratio more than 0.86; hinge heavy (Figs. 237,
241, 324, 327: adults always less than 6 mm
long 37
H:L ratio less than 0.85; shell silky to dull;
hinge light adults may be more than 6 mm long
(Figs. 363-368) 38
37 (36) Shell dull; umbo narrow and prominent; ridge on
umbo present or absent (Figs. 236-241):
Shell glossy; umbo broad and prominent; ridge
never present on umbo (Figs. 323-328):
Pisiditon compression
Pisidiwn variabile
91
-------
355
Figures 355-356. Pisidittm
equilaterale:
355- right valve;
356- left valve.
356
92
-------
357
359 360
358
Figures 357-360. Pisidium punetatum: 357- right hinge; 358- left hinge;
359- lateral aspect of left valve; 360- end view. Also see Figs.
362, page 94.
365
366
Figures 363-366. Pisidiwn adamsi: 363- right hinge; 364- left hinge;
365- lateral aspect of left valve showing trapezoidal outline; 366- end
view. Also see Figs. 367-368, page 95.
93
-------
361
Figures 361-362. P-isidium
punotatum:
361- right valve;
362- left valve.
362
94
-------
367
Figures 367-368. Pisidiwn adconsi:
367- right valve;
368- left valve.
368
95
-------
38 (36) Anterior slope distinctly angled at dorsal margin
or shell trapezoidal in shape (dorsal and
ventral margins are parallel but anterior and
posterior slopes diverge at nearly equal
angles from dorsal margin, Fig. 365); adults
usually longer than 6 mm (Figs. 363-368):
Anterior slope evenly rounded (Fig. 371); shell
not trapezoidal in shape; adults less than
6 mm long (Figs. 335-344, 369-374):
Pisidiion adamsi-
Pisidium aasertamon
369
370
371
373
374
372
Figures 369-374. Pisidiwn easertanum: 369- C3 of right hinge; 370- C2 and
C4 of left hinge; 371- right hinge; 372- left hinge; 373- lateral
aspect of left valve; 374- end view. Also see Figs. 335-338, page 86,
Figs. 339-342, page 87, Figs.343-344, page 89.
96
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SECTION IV
ECOLOGY AND DISTRIBUTION OF
GASTROPODA
VIVIPARIDAE
Campeloma
C. decision has been reported from Superior, Huron, and Ontario where it
may be common in quiet waters with soft substrates containing some organic
matter (Bovbjerg, 1952; Chamberlain, 1958; Mackie, 1971; Mackie and Ouadri,
1971a; Clarke, 1973). C. rufttm apparently is quite rare and recorded only
by Baker (1928a); Goodrich (1932) and Van Cleave and Altringer (1937) list
it as a species of smaller lakes.
Viviparus
The Great Lake's species of Viviparus are most abundant in moderately
eutrophic areas. V. georgianus has been collected in Huron. V. japoniaus
and its apparent synonym V. malleatus is an Asian introduction particularly
abundant in Sandusky Bay, other areas of western Lake Erie, and Green Bay of
Michigan (Wolfert and Hiltunen, 1968).
VALVATIDAE
Valvata
Most species and forms of this genus occur in limited numbers throughout
the lakes. V. bicarinata (possibly a subspecies of V. triaarinata")is known
from Lake Michigan (LaRocque, 1953), V. lewisi from Superior, V. pisoinalis
from Ontario^ V. triearinata (including the morphs basalis and perconfusa)
from eutrophic bays of Ontario and Superior, and V. sincera from Erie,
Ontario and Superior. V. perdepressa is abundant in many areas of Lake
Michigan (Shelford, 1913; Baker, 1928a, 1930) and common in the open lake
down to 18 m near Waukegan (Industrial Bio-Test, 1973).
HYDROBIIDAE
Armicola
A. limosa and A. walkevi are most common in dense masses of aquatic
macrophytes and also are present in open areas protected from strong wave
action and currents (Berry, 1943; Harman and Berg, 1971). A. limosa is
recorded from Ontario, Michigan and Superior; A. walkeri is from Michigan.
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Bithyn-ia
B. tentaculata is listed from Ontario and Erie but probably will
continue to spread throughout the Great Lakes (Berry, 1943) where there are
submerged macrophytes and eutrophic conditions (Harman, 1968).
Cinoinnat-la
C. cineinnatiensis is recorded only from Michigan where it occurs on a
variety of substrates from soft ooze to sand with or without aquatic
vegetation (Berry, 1943; Harman and Berg, 1971; Clarke, 1973) but "is
largely a species of rivers" (Baker, 1928a).
Marstonia
Af. deoepta is often associated with A. limosa in Ontario and Superior
(Berry, 1943). Clarke (1973) found it in mesotrophic to eutrophic waters
where vegetation was moderately thick, but Harman and Berg (1971) collected
the species only from inorganic substrates.
Probyth-inella
Berry (1943) states that P. laaustris is common in the Great Lakes.
Clarke (1973) and Harman and Berg (1971) suggest that it may require cold
water particularly in the southern part of the range as it is found only in
large, deep lakes.
Somatogyrus
A few specimens of S. subglobosus have been reported from western Lake
Erie and Lake Michigan (Baker, 1928a, 1930).
PLEUROCERIDAE
Goniobasis
G. livescens is common and recorded from all the Great Lakes from
"unprotected, rocky lettoral habitats ... on limestone rock ... and on piers
of wharves, etc.", (Lake Michigan, Baker, 1911), "in sand runs or along
rocky shores or on rocky shoals" (Lake Huron, Robertson, 1915a, 1915b),'on
pebbles and small stones of a bar "subject to almost continuous wave action"
(Lake Erie, Wiebe, 1926), and "on marly clay bottom in water 0.3-1.3 m deep,
among Chara, on sand bottom" (Lake Michigan, Baker, 1928a) (see also
Goodrich, 1945, for a list of other habitats).
Pleurooera
P. aauta is known from Lake Erie where it has a limited and localized
distribution and can be found in quiet, sheltered areas (Dazo, 1965) as well
as shores exposed to wave action (Baker, 1928a; Harman and Berg, 1971).
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LYMNAEIDAE
(The genera listed below are often regarded as subgenera under Lymnaea.)
Bulirmea
B. megasoma is reported from Lake Superior (LaRoque, 1962), Georgian
Bay (Rush, in Baker, 1928a), and western Lake Erie (Wolfert and Hiltunen,
1968). It occurs mainly in muddy, heavily vegetated habitats (Clarke, 1973)
but has been collected from rocky shores exposed to constant wave action
(Baker, 1928a).
Fossaria
F. decampi usually is considered a subspecies of F. obrussa, but Clarke
(1973) considers both as distinct species with the Great Lakes and St.
Lawrence River as the dividing point for their distributions; F. deoampi
occurs in the northwest in cold-water lakes and rivers, and F. dbrussa
extends south to the Gulf of Mexico in smaller bodies of water. Both have
been recorded for Lake Superior (Clarke, 1973). Baker (1911) and LaRoque
(1953) list F. himilis from Lake Superior.
Lymnaea
L. stagnalis appressa occurs in all the Great Lakes and is
characteristic of all permanent bodies of water which support macrophytes
(Baker, 1911; Mozley, 1938). Lake Superior is cited as the type locality
for Lymneus appressus Say by Clarke (1973).
Pseudosuccinea
P. columella is rare but may be found on aquatic macrophytes (Baker,
1911).
Radix
R. auricularia is rare in the Great Lakes and recorded only from Lake
Ontario. The habitat is quite variable (Harman and Berg, 1971).
Stagnieola
Species of Stagn-ioola occur on a wide variety of substrates from silt
in open waters to aquatic vegetation in protected bays. S. oaperata is
recorded from southwestern Lake Michigan (Limnetics, 1973), S. catascopiwn
f. catascop-ium from Lake Ontario, 5. catascopium f. nasoni from exposed
shores of Lake Huron and drift along Lake Michigan, S. emavginata from
shallow areas of Lake Superior, and 5". veflexa from areas of Erie and
Superior (Baker, 1911, 1928a; Harman and Berg, 1971; Clarke, 1973). Lakes
Erie and Superior are cited as type localities for Lymnaea reflexa Say and
the synonym L. lanoeata Gould by Clarke (1973).
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PHYSIDAE
Physella
Following the recent revision by Te (1978) it is difficult to describe
the distributions and habitats of P. gyrina sayi, P. -Integra and P.
vinosa. Species of Physella have been recorded from all the Great Lakes and
are most abundant on hard surfaces and aquatic vegetation.
PLANORBIDAE
Gyraulus
The species of Gyraulus are most abundant on aquatic vegetation and are
tolerant of polluted and harsh environments. G. defleetus probably is
present in all the Great Lakes but reported only from Ontario but is more
characteristic of temporary waters (Taylor, 1960); G. parvus is a
cosmopolitan species occurring in great abundance on aquatic vegetation
(Baker, 1928a; Clarke, 1973).
Helisoma
H. anoeps and H. trivolvis are widespread and often occur together.
Both occur in shallow waters with or without vegetation; however, anceps is
found at the upper edge of the profundal zone while trivolvis remains in
shallow waters (Baker, 1928a; Harman and Berg, 1971). H. campanulatum is
not common but may be found on bare stones and rock faces and is recorded
from Ontario and Superior. H. oorpulentum has been collected from Lake
Erie but is very uncommon (Kalas, pers. comm.).
Promenetus
P. exaouaus is widespread in shallows and among decaying and living
vegetation.
ANCYLIDAE
Ferrissia
F. parallela is fairly common on aquatic vegetation and in substrates
ranging from mud to sand (Basch, 1963). F. tarda usually does not occur
directly in the lakes but in the mouths of rivers entering the lakes where
the substrate is cobble (Clarke, 1973).
Laevapex
L. fuscus is very rare but may be found in backwaters and densely
vegetated portions of the lakes (Ward, 1896; Basch, 1963).
100
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SECTION V
ECOLOGY AND DISTRIBUTION OF
UNIONIDAE
The Unionidae are not uniformly distributed throughout the Laurentian
Great Lakes nor within any one of the lakes. In general, it can be said
that areas which once had the most developed unionid populations are those
which have come under the greatest biological disturbance due to pollution
(e.g., western Lake Erie, Saginaw Bay in Lake Huron, Green Bay in Lake
Michigan). In this light, it is most practical to discuss the ecology and
distribution by habitat and by lake rather than at the generic or specific
level. For indepth notes on the ecology and biology of the Great Lakes'
taxa, one should consult the references listed in the introduction in
addition to the more general works by Baker (1928b), van der Schalie (1938),
Goodrich and van der Schalie (1944), Mirray and Leonard (1962), Parmalee
(1967), Clarke (1973), Taylor (1975) and Burch (1975b).
Of the 39 taxa recorded here from the Laurentian Great Lakes, 36 are
known from Lake Erie, primarily western Lake Erie. The exceptions are
Anodonta grandis simpsoniana known only from Lake Ontario, Lampsilis radiata
Tad-lota from Lakes Ontario and Superior, and Quadrula nodulata from Lake
Michigan. The taxonomy and status of the two former species in the Great
Lakes has been questioned by some, and the latter species has not been
recorded since before the turn of this century (Ward, 1896). The following
list of 19 species records those unionids collected only from Lake Erie.
None are particularly common but may occur in isolated patches along
protected shorelines and shallow areas of the western basin. Some may now
be extinct from the lake or their populations eliminated from or greatly
reduced in open waters. Included after each name is the most probable
substrate type.
Anodonta imbecillis - soft substrates, particularly soft, silty mud.
Carunculina parva - soft substrates, particularly soft, silty mud.
Cyctonaias tuberculata - soft substrates.
Dysnomia suleata - substrate variable.
Lampsilis fasoiola - substrate variable.
Lasmigona costata - substrates of hard gravel and rock to soft mud.
Obliquaria reflexa - firm substrates, hard sand to rocky shoals.
Obovavict olivaria - substrate variable, usually sand and gravel.
Obovaria subrotunda - substrate variable, usually sand and gravel.
Pleurobema oordatum f. oooeineum - substrate variable.
Pr>optera laevissima - firm to silty sand.
Ptyehobvan.ohus fasaiolare - firm substrates with wave action.
Qu.adur'la pustulosa - more common on firm substrates, particularly clay.
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Qy.adru~ia quadpula - generally firm substrates.
Simpsoniconcha cffribigua - substrate variable.
Truncilla donacifovmis - quiet waters with firm, sandy substrates, clay.
TTuncilla truncata - moving waters with firm, sandy substrates, gravel.
Villosa fabilis - substrate variable, usually sandy.
Villosa iris - substrate variable, usually sandy.
The following 5 species have been recorded for both Lake Erie and Lake
Huron. None are wide spread but may be locally common in protected, °hallow
areas of the lakes which are relatively free from organic enrichment.
Arriblema plicate. - firm substrates (Lake Erie specimens tend to be
stunted by their environment, Brown et al., 1938).
Dysnomia triquetva - substrate variable, usually firm.
Eliptio dilatata - substrate variable.
Fusconaia flava - usually firm substrates, hard mud.
Ligwnia nasuta - soft to fairly firm substrates, hard sand.
The remaining unionid species have been recorded from Lake Erie and one
or more of the other Great Lakes.
Leptodea fragilis - (Erie and Ontario) - firm, sandy substrates.
PvopteTa alata - (Erie and Ontario) - substrate variable, often in rocky
shoals with some wave action.
Alasmidonta calceolus - (Erie and Michigan) - substrate variable.
Strophitus undulatus - (Erie and Superior) - substrate variable, fine
but firm sediments.
Anodonta cataracta - (Erie, Michigan, and Ontario) - soft substrates,
particularly silt and mud.
Lasmigona complanata - (Erie, Huron and Superior) - substrate varies
from soft silt in still waters to rocky, wave swept areas.
Ligwnia recta - (Erie, Huron and Ontario) - usually firm but sandy
substrates, mud.
Anodontoides ferussacianus - (Erie, Huron, Michigan, and Superior) -
soft substrates.
Lcanpsilis ovata - (Erie, Huron, Michigan, and Superior) - substrate
quite variable.
Anodonta grandis grandis - (Erie, Michigan, Ontario, and Superior) -
soft substrates.
Eliptio complanata - (Erie, Huron, Ontario, and Superior) - substrate
variable.
Lampsilis radiata siliquoidea - (present in all the Great Lakes) -
substrate quite variable. This species possibly is the only
freshwater mussel still present in the deeper waters of the western
basin of Lake Erie (Wood, 1953, per. comm.). Where it occurs in
soft sediments, it may be of great importance in the reworking of
bottom materials (McCall et al., 1979).
Most of the unionids listed for Lake Michigan have been recorded for
Green Bay; those recorded for Lake Huron are primarily from Saginaw Bay. It
is doubtful if populations are still present in these areas but if found,
most likely will be away from sources of pollution.
102
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SECTION VI
ECOLOGY, DISTRIBUTION, AND VARIATION OF
SPHAERIIDAE
MUSCULIUM
Musculiwn laoustre f. lacustre and Museuliwn laeustre f. jayense
Of the two forms, jayense appears to be the most common and has been
recorded from all except Lake Superior. The preferred habitat is hard mud.
Form lasustre has a distribution similar to jayense but is uncommon. Its
habitats range from shallow, temporary pools and streams to deep, permanent
lakes and rivers (Herrington, 1962). Both have been recorded from depths
down to 20 m.
Musauliwn partwne'iwn
Uncommon but reported from Lakes Michigan and Ontario. It usually is
found on silty or muddy bottoms of large lakes or among plant debris where
oxygen content often falls to near anoxic levels. Usually in less than
10 m of water.
Musaulium seauris
Reported from Lakes Michigan, Ontario, and Superior. As with M.
paTtumeium, this species is common in mud and plant debris. It is unlikely
that the distribution is limited by oxygen supply although it is not common
in organically rich waters. Usually in less than 10 m of water.
Museulium transversim
This is the most common MuscuHwn in Lakes Michigan, Erie, and
Ontario. Unlike other Musaulium, transverswn inhabits permanent bodies of
water on a variety of substrates from mud to stones. Soft mud in less than
10 m of water has been shown to be the preferred habitat (Gale, 1971).
PISIDIUM
Pisidiwn adamsi
Uncommon and localized in all the Great Lakes (yet to be recorded from
Superior). Herrington (1962) and Henson and Herrington (1965) give the
preferred habitat as mud and decaying vegetation in small lakes, ponds,
rivers, and (rarely) creeks, also from "black sand caly" and shallow
103
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pockets of debris in the Great Lakes. Generally found in very shallow water
less than 5 m deep.
P. adamsi typically has a long trapezoidal outline with the dorsal and
ventral margins parallel (Fig. 365). The relative length of the shell is
variable, and shorter individuals may resemble either P. aompressum or
P. casertanum. The umbo of P. adamsi barely protrudes over the dorsal margin
(Fig. 366) and does not exhibit the flattening or ridge formation found on
most P. compression (Figs. 238, 239). In contrast to P. oasertanum, the
periostracum of P. adamsi, is quite dull, striae are coarse, and the lateral
teeth are broad and blunt on top. P. oasertanum is moderately glossy, finely
striated, and the lateral teeth are well-develloped and pointed (Fig. 344) .
Young P. adamsi. are proportionally shorter than adults making the trapezoidal
shape less obvious. Separation of young specimens may be based on the
parallel dorsal and ventral margins, coarse striae, and the dull-iridescent
periostracum.
Pisidium amniewn
Recorded from Lakes Ontario, Erie, and Michigan, this species generally
is not abundant and too few collections have been make to determine substrate
and depth relationships. It may prefer sandy substrates (Herrington, 1962)
but has been collected on mud in waters less than 10 m deep.
P. armiawn is distinct in its longish-oval outline (Fig. 210), laterally
compressed shell, strong and widely spaced striae, and relatively large size.
The striations are uniform but may fade in the area of the umbo thereby
resembling P, dubium. The umbo of P. armicum usually is positioned 2/3 back
along the length of the shell (Fig. 210). Occasionally, placement closer to
the dorsal margin may create further confusion with P. dubium. Doubts may be
resolved by examining the cardinal teeth. C3 of P. arrmicum is "J" shaped
with the anterior arm much thinner than the posterior (Figs. 208, 212). In
P. dubium, C3 is "U" shaped and both arms are of equal thickness or neraly
so (Figs. 212, 206). C2 of P. amniaum is a peg-like structure overlapped by
the thinner, gently arched C4 (Figs. 209, 213). The corresponding teeth in
P. dubium are each approximately as thick (Figs. 203-207). Young P. amnicum
are separable from P. dubium on the same basis.
Pisidium oasertanum
Occurring in all the Great Lakes on clay, mud, sand, and gravel
substrates, this cosmopolitan species is very common in water down to 25 m
deep but has been collected as deep as 100 m.
P. oasertanum is the most variable and widely distributed of the
Pisidium. The average shell is long in appearance with an extended, fully
rounded anterior margin. There are two general morphotypes; a short, high,
heavy shelled form, and a long, low, thin to moderately thick form. The
short, high form (Figs. 335-338). is most commonly confused with P.
subtrunoatum, P. walkeri f. mainense, and P. variabile. P. subtruncatum is
distinguished by cardinal teeth C2 and C4 which take the appearance of thin,
parallel plates (Fig. 297). In P. oasertanum, C2 is either a "D" shaped peg
104
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or a bent, thickened plate which is overlapped by a curving C4 pointing at or
interior of PII (Figs. 336, 3AO, 344). Additionally, the umbo of P.
subtrunoatwn is narrow and prominent, not broad as P. casertanwn; and the
dorsal and ventral margins are skewed (Fig. 289), not parallel as in P.
oasertanwn Figs. 337, 341). Finally, the anterior and posterior margins of
P. subtrunoatwn are nearly straight or gently curved (Figs. 337, 341). Long,
low specimens of P. variabile have low, blunt lateral teeth, highly glossed
periostracum, and often a prominent umbo. The lateral teeth of P. aasertanwn
are long and sharp, the periostracum moderately dull to slightly glossy, an.d
the umbo barely projects over the dorsal margin. P. waVkeri f. mainense is
distinguished by its shorter hinge (Fig. 279) which is less than 3/4 of the
total shell length. P. casertanum exhibits a hinge which is 3/4 or greater
of the total shell length.
The long, low form of P. oaseTtanum (Figs. 338-342) is most often
confused with small P. adamsi or P. nitidwn f. nitidwn, P. adamsi, however,
has coarse striae, a long dorsal margin (Fig. 365) and the lateral teeth are
low and blunt as in P. variabile above. P. casertanwn has fine striae and
moderately long but an evenly rounded dorsal margin (Fig. 341). The
separation of P. easertanum from P. nitidwn f. nitidwn may be difficult
without a large series of each for reference. Cardinal teeth are the primary
diagnostic trait. C3 of P. oasertccnum typically is club-shaped, the
posterior end greater than twice the width of the anterior (Figs. 335, 339),
and C2 and C4 are as described above. The cardinal teeth C2 and C4 of P.
nitidwn f. nitidwn are similar to P. subtTunaatwn above (Fig. 303); however,
C2 varies somewhat in thickness (Fig. 307). The posterior arm of C3 usually
is less than twice the thickness of the anterior arm (Figs. 302, 306). In
specimens of P. nitidwn f. nitidwn with a thickened C2 and C3, the cardinal
teeth are less diagnostic. In such a case, an evaluation of the ligament,
periostracum, and dorsal margin should be made. The ligament of P. nitidwn
f. nitidwn is short and wide, often not much longer than C3 (Fig. 306); the
periostracum is glossy; and the dorsal margin generally intersects the
anterior and posterior margins with an angle or rounded corner (Fig. 304).
In contrast, the ligament of P. easertanwn is long and wide, usually twice
the length of C3 (Fig. 343); the periostracum ranges from dull to only a
slight gloss; and the dorsal margin is evenly rounded, the intercepting
anterior and posterior margins smooth without an angle (Figs. 337, 341).
Pisidiwn oompresswn
Rarely abundant but occurs in all the Great Lakes on clay, mud, sand,
and gravel in shallow water usually less than 25 m deep. P. eornpvesswn may
prefer sandy bottoms with some vegetation (Herrington, 1962) or fine sand
(Henson and Herrington, 1965).
The shell outline of P. aompressum varies from short and high to
moderately long. The shorter form closely resembles P. supinwn: however,
the hinge plate of P. supinwn is shorter, the striae are more widely spaced,
and the cusp of All is proximal (Fig. 234), not distal as in P. compresswn
(Fig. 241). The low, long from of P. oompresswn exhibits a broader umbo and
may be mistaken for P. Gasertanum. Such individuals may be distinguished
from P. oasertanwn by their short, blunt lateral teeth (Fig. 241). The
105
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laterals are well-developed and pointed in P. easertanum (Fig. 344). Further
distinctions can be made by comparing the generally coarse striae and
dull-iridescent periostracum of P. compression with the finely striate,
moderately glossy shell of P. easertanum. The umbo of P. compression usually
shows some degree of flattening on the dorsal face similar to P. fallax
(Fig. 318) and may exhibit a ridge or plate not unlike P. henslowanum
(Fig. 217). P. compression is distinguishable from P. fallax by the
well-inflated shell (Fig. 239) and the position of All which is parallel to
the anterior margin (Fig. 241). In contrast, P. fallax is laterally
compressed (Fig. 318) and All twists inwardly creating a thickening of the
hingeplate at the anterior laterals in both valves (Figs. 319, 321). The
hinge characters and outlines of most P. compressum are the same as P.
varlabile (Figs. 323-328), but the periostracum of P. compression is dull,
not glossy, the umbo is ridged or flattened, and the striae of P. compressum
are coarse while those of P. variabile are quite fine. Young compression
have a short dorsal margin, dull periostracum, and a well developed
hingeplate; however, the characteristic flattening of ridge on the umbo may
not be present. See also P. adamsi.
Pisidium conventus
Abundant and widely distributed throughout the Great Lakes, P. conventus
has been recovered from all types of substrates including stones, gravel,
sand and mud. In Lake Michigan, it seems to be most abundant in sand to
coarse silt (Phi 4-5 on the Wentworth, 1922, grade scale). A review of the
biology of P. conventus was given by Heard (1963), and additional notes on
habitat and distribution for the Great Lakes can be found in Herrington
(1950, 1962), Heard (1962), Henson and Herrington (1965), Brinkhurst (1969),
and Clarke (1973). P. conventus is considered an oligothermal relect of the
primarily tropical subgenus Neopisidium and, with few exceptions, is limited
to high altitudes or to cooler, boreal lakes where it rarely is found above
the limits of the thermocline. Maximum densities in the Great Lakes occur
between 35 and 40 m; living specimens may be collected in less than 7 m to
nearly 200 m of water. Herrington (1950) recorded a few P. conventus from
219 m in Great Slave Lake, and it is the only mollusk to be collected
deeper than 150 m in the Great Lakes.
Shell morphology and diagnostic traits of P. conventus are quite
constant throughout the Great Lakes. The periostracum is very dull or
iridescent in deep waters (greater than 25 m) but may exhibit a glossy
quality at shallower depths. Glossy P. conventus may be mistaken for young
P. easertanum but are distinguished by the nearly parallel and flattened
anterior and posterior margins (Fig. 310). P. easertanum differs in having
well-rounded margins (Figs. 337-341). The degree of prominence of the umbo
of P. conventus varies from no overlap of the dorsal margin to a sizeable
protrusion which may be similar to young P. illjeborgi. f. lilljeborgi and
P. variabile. Shell outline of the three species will help to separate them;
P. conventus has a long dorsal margin approximately 2/3 of the total shell
length (Fig. 310); P. variabile has an extremely short dorsal margin
beginning and ending at the umbo (Fig. 325); and P. Hlljeborgi f.
lilljeborg-i has a well-rounded, symmetrical outline (Figs. 260,264) which
contrasts to the long rectangular shape of P. conventus. If the external
106
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traits are not definitive, hinge characters (Figs. 308, 309) should be the
final basis for diagnosis as they seem to be less variable.
Pisidium dubium
Not a common species but recorded for all the Great Lakes with the
exception of Erie. Distributions appear to be localized and patchy, usually
associated with areas of fine sand in less than 5 m of water (Henson and
Herrington, 1965).
With few exceptions, P. dubium rarely is confused with other Pisidium.
The shell is large, laterally compressed, and heavily striated except on
the umbo where the striations fade to near smoothness: P. omnioum is
similar, but the striations continue onto the umbo. Occasionally, P. dubium
may possess striations on the umbo, and the cardinal teeth should be used as
the final indicator. P. dubium has a "U" shaped C3 (Figs. 202, 203) with
C2 and C4 of equal thickness (Figs. 203, 207). C3 of P. comioum is "J"
shaped with the anterior arm much thinner than the posterior (Figs. 208,
212), and C2 is a peg which is overlapped by a thin, gently arching C4
(Figs. 209, 213). Young P. dubium may be recognized by their larger size
(rarely less than 3 mm long), their laterally compressed shell, and the
heavy striae divided by widely spaced furrows.
Pisidiwn equilaterale
This species is quite rare in the Great Lakes with most specimens
having been collected from Lake Ontario in the Bay of Quinte where it
inhabits the fine, sandy substrates (Herrington, 1962).
There is little significant variation of the diagnostic traits within
P. equilaterale. The .short, high form of P. variabile resembles this
species quite closely; however, the umbo of P. equilateyale is central
(Fig. 353) while that of P. variabile is further back (Fig. 325). Also,
the space between C2 and C4 is a slit directed toward the cusp of PII in
P. equilaterale (Fig. 352) while this space in P. variabile usually is much
greater owing to the enlargement of the posterior end of C3 (Fig. 324).
Immature P. equilaterale may be confused with heavy-shelled P. nitidum
f. nitidum but there are differences in the umbo, striae and periostracum.
The umbo of P. equilatevale is swollen and prominent (Figs. 353-356), the
striae are coarse, and the periostracum dull to moderately glossy. P.
nitidum f. nitidum exhibits a broad umbo which barely projects over the
dorsal margin (Figs. 304-307). The striae are quite fine, and the
periostracum of P. nitidum f. nitidum is very glossy, often mirror-like.
Pisidium fallax
Recording P. fallax from all the Great Lakes except Superior,
Herrington (1962) states that it "has a preference for coarse sand or
gravel, even sandy gravel in cracks on a flat rock bottom ... and appears
to like water in motion ... i.e. lakes, or bays where there is considerable
wave action". Though most abundant at less than 10 m, it is common at
depths down to 20 m.
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P. fallax ranks among the least protean of the Pisidium species. The
extent of flattening of the dorsal surface of the umbo is the only trait
subject to variation in Great Lakes specimens. Individuals with a high
degree of flattening often exhibit low, incipient ridges or ripplings at the
apex of the umbo. These specimens are similar to P, henslowanwn, but the
shell of P. fallax is laterally compressed (Fig. 318) and the umbo rarely
protrudes over the dorsal margin owing to the flattening on the dorsal face
of the beak (Fig. 317). P. henslowanwn, in contrast, is not compressed but
moderately inflated (Fig. 217) and its narrow umbo generally protrudes quite
significantly over the dorsal edge (Fig. 216). The thickening of the
hingeplate at the anterior lateral cusps AI and AIII due to the inward twist
of All which is present in P. fallax (Figs. 315, 319) is not present in
P. henslowanum (Figs. 215, 218). Shells in which the flattening of the umbo
is slight or not apparent may be mistaken for P. compression. These shells
are distinguished on the basis of the position of the anterior laterals or
the prominence of the umbo for the reasons discussed above. Additionally,
the dorsal margin of P. fallax generally extends beyond the umbo to join
the anterior and posterior margins (Fig. 317) whereas the dorsal edge of
P. compression is very short with the anterior and posterior margins often
beginning at the umbo (Fig. 238). Young P. fallax are separated from their
closest morphotypes, P. henslowanum and P. compression on the same bases
stated above. In distinguishing P. fallax from all other Pisidium, the
diagnostic traits are the silky-iridescent periostracum, the flattened umbo,
and the laterally compressed shell.
Pisidium ferrugineum
This species is common in all the Great Lakes. The habitat ranges
from clay, mud, sand, to gravel; maximum abundance is reached at about 10 m,
but it commonly is taken at depths down to 30 m.
There are two general forms of P. ferrugineum: one is short with a
prominent umbo, the other is long with the umbo barely protruding over the
dorsal margin. The shorter form of this species is similar to individuals
of both P. lilljeborgi f. lilljeborgi and P. nitidum f. paupereulum.
P. ferrugineum has a glossy periostracum (often concealed beneath brownish
accretions) and a gently rounded dorsal margin which blends easily into the
anterior and posterior edges (Figs. 290, 293, 295). P. lilljeborgi
f. lilljeborgi usually has a silky periostracum and a straighter dorsal
margin which creates an elbow or corner with the anterior and posterior
margins (Figs. 260, 264). In distinguishing P. nitidum f. paupereulum one
must examine hinge characters. The lateral teeth of P. ferrugineum are
short and pointed and cardinal teeth are situated close to the anterior
lateral cusps (Figs. 288, 293). The laterals of P. nitidum f. paupereulum
are large and long with blunt cusps and the cardinals are central (Figs. 345,
349).
The longer form of this species resembles specimens of P. miliwn and
glossy P. conventus. In end view, the outline of P. milium is truncate at
the ventral margin (Figs. 282, 283) whereas P. ferrugineum exhibits an even
taper from the apex of the umbo to the ventral edge (Figs. 290, 291). As
stated above, the dorsal margin of P. ferrugineum is evenly rounded and
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blends into the anterior and posterior margins without an angle. P.
oonoentus, in contrast, has a long dorsal margin .which is 2/3 or more of the
total shell length and intercepts the anterior and posterior margins with
distinct angles or corners (Fig. 310). The characters above will separate
young P. fevrugineum from closely related Pisidium.
Pisidium henslowanum
This species is common in Lakes Michigan, Erie, and Ontario.
Herrington (1962) lists P. henslowanum mainly from shore debris; however,
most Lake Michigan collections are from sandy substrates down to 20 m deep.
This species is an introduction from Europe.
The shell features of P. henslowanum are quite consistant. The most
significant variation occurs in the development of the umbonal ridge which
towers from the apex of the umbo and diverges from opposite shell halves
(Fig. 217). In some stunted specimens, the ridge may develop only to the
point of a slight rippling of the calcified shell at the umbo. These
stunted forms may resemble P, vaviabile\ however, the longer dorsal margin,
the angular intersection with the anterior and posterior margins, and the
silky-iridescent periostracum of P. henslowanum serve to distinguish it from
the glossy, more triangular shell of P. variabile. Additional variability
is found in both the striae and umbo of P. henslowanum. While the striae
are generally uniform, coarse, and concentric, the degree of uniformity
and coarseness is subject to variation. Specimens often exhibit moderately
fine striae, and the uniformity of the striae may be interrupted by the
presence of two or three deeper growth rings in mature specimens.
Individuals with finer striae appear to have a glossier periostracum, but
the degree of gloss in P. henslowanum never reaches that of P. nitidwn or
P. varidbile. The umbo of P. henslowanum is quite narrow and prominent in
adult specimens; however, in some instances a broader and less prominent
umbo similar to P. oasertanum or P. nitidwn f. nitidwn is observed. P.
henslowanum is distinguished from P. aasertanum by the lamella-like ridge
on the umbo and the angular intersection of the dorsal edge with the
anterior and posterior margins in P. henslowanum (Figs. 216, 217). P.
oasertanum exhibits no plate on the umbo and the dorsal margin flows in a
continuous curve to the anterior and posterior margins (Figs. 337, 338, 341,
342). In contrast to P. nitidwn f. nitidwn, the striae of P. henslowanum
never reach the degree of fineness exhibited by P. nitidwn f. nitidwn.
Additionally, the periostracum of P. henslowanum does not attain the usual
mirror-like glossiness of P. nitidwn f. nitidwn^ and the cardinal tooth C3
is much thickened posteriorly (Figs. 214, 218) whereas C3 of P. nitidwn
f. nitidwn is a narrow, evenly tapering plate (Figs. 302, 306).
The character traits used to distinguish adult specimens (lamella-like
ridge on the umbo, angular intersection of the dorsal margin, and coarse,
evenly spaced striae) are present in the nepionic shell and may be used to
distinguish immature P. henslowanum from other Pisidium.
Pisidium iddhoense
With the exception of Lake Erie, this species is common in all the
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the Great Lakes in muddy and sandy substrates. It often is found as deep as
40 m, but greatest abundance occurs at 20 m or shallower. Herrington (1962)
suggests that P. idahoense may be restricted to cold waters of northern
lakes; however, many recent collections have come from warmer waters of
temperate climates.
P. idahoense varies significantly only in shell length. The large
(adults 5 mm or longer), heavy, high shell usually is quite short with a
height to length ratio greater than 0.86. Some specimens will appear longer
and may be confused with P. amnicwn. The striae of P. armiaim are very
coarse, widely spaced, and uniform over the shell; those of P. idahoense
generally are fine and uniform, broken only by deeper growth rings
sculptured into the calcified surface. Cardinal teeth of P. amnicwn are
larger and heavier than P. idahoense, C3 is tightly arched and well-
thickened posteriorly (Figs. 208, 212), and C4 follows the curvature quite
closely so that it points to the interior of the shell (Figs. 209, 213).
In contrast, C3 of P. idahoense is gradually curved and less thickened
posteriorly (Figs. 329, 333) and C4 is directed to the posterior lateral
cusp of All. Young individuals of this species may strongly resemble P.
vaviabile; however, the anterior margin of P. vafiabile is straight or
nearly so (Fig. 325) while that of P. idahoense is well rounded.
Pisidiwn lilljeborgi f. lilljeborgi
This species is very common in all Great Lakes in clay, mud, sand, and
gravel substrates, although Meier-Brook (1969) demonstrated that it prefers
an endopelic mode of life in fine-grained organic sediment. Herrington
(1957) found empty shells in depths exceeding 45 m in Lake Huron.
Collections of P. lilljeborgi f. lilljebovgi from Lake Michigan show that
maximum abundance is reached at about 20 m; however, this taxon is not
uncommon at 70 m with only P. oonventus being recorded from greater depths.
The form lilljeborgi differs from form cristatwn in having the shell
much longer and ridges absent from the umbo. Glossy specimens of P.
lilljeborgi f. lilljeborgi often resemble P. nitidwn f. pauperculwn, but
the umbo of the form pauper'cu'lwn is broad and not very prominent (Figs. 348,
350) while P. lilljeboTgi f. lil'Ljeborgi has a narrow umbo which overlaps
the dorsal margin to a considerable degree (Figs. 265, 267). The ligament
of P. lilljeborgi f- lilljeborgi is long and thin while that of P. nitidwn
f. pauperaulwn is short and wide. P. lilljebovgi f. 1i 11jebovgi with a dull
periostracum may resemble P. compression, but there are significant
differences. In P. compression the dorsal margin is very short, often
yielding a three-cornered appearance, there usually is a flattening or
incipient ridge on the umbo (Fig. 239); and, the lateral cusps All and PII
are short and blunt (Fig. 241). P. lilljebovgi f. lilljeborgi exhibits
no ridge or flattening (Figs. 261, 265), the dorsal margin extends beyond
the umbo, and the cusps of All and PII tend to be rather sharp on top (Fig.
267).
When the dorsal margin of P. lilljeborgi f. lilljeborgi is short,
confusion with P. subtrunctwn may occur. The umbo of P. subtrunotum lies in
the last two-thirds of the shell and the posterior margin slopes nearly
110
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vertically from the posterior lateral teeth, becoming undercut at the ventral
margin (Fig. 298). In contrast, the umbo of P. lilljeborgi f. lilljeborgi
is centrally situated and the curvature of the posterior margin is evenly
rounded, not undercut at the ventral margin (Figs. 260, 264). Specimens
with longer dorsal margins are the predominant type of P, lilljeborgi
f. lilljeborgi. These may resemble P. nitidum f. nitidum with both having
the hinge area with the dorsal edge forming corners at the intersection
with the anterior and posterior margins; however, the umbo of P, nitidum
f. nitidum is broad and and barely protrudes over the dorsal margin (Figs.
306, 307), and the degree of inflation exhibited by the shell of P. nitidum
f. nitidum never approaches that of P. lilljeborgi £. lilljeborgi. Young
P. lilljeborgi f. lilljeborgi may be mistaken for P. ventrioosum
f. ventrioosum or P. ventrioosum f. rotundatum but the'dorsal margins of
the latter two do not extend out from the umbo as in P. lilljeborgi f.
lilljeborgi (Figs. 260, 264), rather this edge follows the curvature of the
umbo (Figs. 244, 248, 254).
Pisidium lilljeborgi f. oristatum
This taxon is most likely a valid species, but until further analysis
of the soft anatomy is done, it will be considered a form of P. lilljeborgi
Often having been lumped with P. lilljeborgi f. lilljeborgi, we are unsure
of its actual distribution; however, specimens from Lake Michigan rarely are
collected in more than 10 m of water and are most abundant on sandy
substrates.
As with P. lilljeborgi f. lilljeborgi there is little variation among
populations of P. lilljeborgi f. oristatum. The latter differs from the
former by the presence of a ridge on the umbo and the shorter shell (Figs.
224, 225). P. lilljeborgi f. oristatum may be confused with P. henslowanum
and P. supinum both of which have umbonal ridges. P. lilljeborgi f.
oristatum (Fig. 224) is bulbous and rotund while P. henslowanum and P.
supinum are laterally compressed (Figs. 217, 231). The anterior margin of
the form oristatum is fully and evenly rounded (Fig. 224) whereas those of
P. henslowanum and P. supinum often are straight along the middle or mildly
curved and long (Figs. 216, 230). Finally, the umbo of P. lilljeborgi f.
oristatum is very narrow and prominent (Fig. 224) while that of both P.
henslowanum and P. supinum is broad and projects only slightly over the
dorsal margin (Figs. 216, 230). The form oristatum, including young
specimens, is distinguished from the remaining Pisidium by its swollen shell,
narrow and prominently ridged umbo, and by the angles created at the
intersection of the dorsal margin with the anterior and posterior margins.
Pisidium milium
P. milium is rare in the Great Lakes reported only for the Bay of
Quinte, Lake Ontario. More characteristically, it is a species of mud or
ooze in creeks, rivers, small lakes, and ponds (Herrington, 1962). A few
additional specimens have since been taken from Lake Michigan.
Intraspecific variation of P. milium is confined to inflation of the
shell and the degree of truncation. Individuals of this species ordinarily
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exhibit a well-inflated, bulbous shell, but specimens in which the inflation
is less pronounced often resemble the glossy form of P. oonoentus. The
parallel anterior and posterior margins give P. conventus a rectangular
outline (Fig. 310) whereas the rounded anterior and posterior edges of
P. m-ilium yield an overal appearance (Figs. 282, 285, 287). In end view,
the well-inflated shell of P. miliwn cuts off abruptly at the ventral margin
(Fig. 283). This truncate outline contrasts to the gentle, evenly-tapering
slope from the apex of the umbo to the ventral margin found in P. oonoentus
(Fig. 311).
The degree of truncation is also somewhat variable within this species.
Specimens from Lake Michigan are generally less inflated and truncate than
their counterparts in smaller bodies of water. Individuals with a lesser
degree of truncation are similar to P. ventvioosum f. ventriaosum, but the
hinge length of this form is less than one-half of the shell length (Figs.
252, 254) while the hinge length of P. miliwn is greater than one-half the
total shell length (Fig. 287).
In overall outline and degree of inflation, P. miliwn resembles P.
punotatum but is separable by the fine striae and highly glossed periostracum
of P. miliwn which contrasts to the very coarse striae and flat-dull
periostracum of P. punotatum. Young P. miliwn may be confused with P.
conventus or P. aasevtanum. From P. conventus, they may be separated as
described above. Young P. miliwn are lower and longer than nepionic P.
oasevtanwn and exhibit a greater degree of inflation.
Pisidium nitidum f. nitidum
Common in all the Great Lakes in sandy substrates in water up to 40 m
deep. Meier-Brook (1969) states, " P. nitidum is strongly restricted to
biotopes below the sediment surface ... but prefers coarse organic sediment
with large-pored interstitial spaces which enable the animal to provide
itself with water sufficiently rich in oxygen".
The ecological morph P. nitidum f. nitidum is distinct from P. nitidum
f. paupevoulum in several characters. The shell of form paupevoulum is
short, high, and heavy with a short, tightly-curved dorsal margin and a
prominent umbo (Fig. 347). In contrast, P. nitidum f. nitidum is longer,
less inflated with a straight or gently curved dorsal margin, and the umbo
barely projects above the dorsal edge (Fig. 304).
P. nitidum f. nitidum exhibits much variation in structure and
appearance. Of particular importance are the gradations in (1) the
sharpness of the angle formed at the intersection of the dorsal margin with
the anterior and posterior margins, (2) the thickness of the posterior arm
of C3, and (3) the degree of glossiness of the periostracum. Where the
angle of the intersection of the dorsal margin is rounded, specimens closely
resemble P. casertanum and P. subtrunoatum. The posterior arm of C3 in P.
casertanum generally is greater than two times the thickness of the anterior
arm (Figs. 335, 339), the ligament is long and wide, and the dorsal margin
blends easily into the anterior margin (Fig. 341). P. nitidum f. nitidum
contrasts in the posterior arm of C3 being two times or less the thickness
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of the anterior arm (Fig. 302), the ligament is short and wide, and the
dorsal margin creates angles or corners with the anterior and posterior edges
(Fig. 304). An additional point of distinction is the anterior margin which
is generally well-rounded and evenly curved in P. casertanum (Fig. 341)
while P. nitidum f. nitidum is straight or abruptly curved (Fig. 304) and
does not flow into the dorsal and ventral margins in a continuous curve. P.
subtvunoatum is distinguished by its long and wide ligament, nearly vertical
and undercut posterior margin, and shorter hinge. The hingeplate length of
P. subtruneatum is less than half the total shell length, where that of P.
nitidum f. nitidum is half or greater. Also, the posterior margin of P.
nitidum f. nitidum is evenly rounded (Fig. 301) in contrast to the very
steep slope of P. subtvunoatum (Fig. 298). While the thickness of C3 of this
species varies, the posterior arm never exceeds twice the thickness of the
anterior arum. In its thickest form, C3 may resemble thin-toothed specimens
of P. casertanum, but these species may be distinguished on the alternative
bases noted above.
The periostracum of P. nitidum f. nitidum varies from a moderate degree
of gloss to a mirror-like finish, often resembling broad-beaked specimens of
P. varidbile. Distinctions between these two species are made by an
examination of C3 and the dorsal margin of the specimen. The posterior arm
of C3 in P. vavidbile is consistantly greater than twice the thickness of
the anterior arm (Figs. 323, 327), while that of P. nitidum f. nitidum is
twice or less as previously mentioned (Figs. 302, 306). Additionally, the
dorsal margin of P. varidbile is tightly covered, barely extending much
beyond the umbo (Fig. 326). The margin in P. nitidum f. nitidum invariably
extends beyond the curve of the umbo. Young P. nitidum f. nitidum may be
separated from other Pisidium on the shell shape characters given above.
Pisidium nitidum f. pauperaulum
The form pauperoulum most likely is a species distinct from form
nitidum. The distribution within the Great Lakes is incomplete and overlaps
with P. nitidum f. nitidum with which it is often included without
reference. Where the two have been distinguished, paupepeulum is less common
but habitat characteristics are similar with both occurring on sandy
substrates to a depth of 40 m.
There is little variation among specimens of P. nitidum f. paupeTGuium.
This form differs from P. nitidum f. nitidum by its short, high, heavy
shell; shorter and much curved dorsal margin; and narrow, often prominent
umbo (Fig. 348). P. nitidum f. nitidum is longer and less inflated, the
dorsal margin is long and nearly straight, and the umbo barely projects
above the dorsal margin if at all (Fig. 305).
P. variabile and P. lilljeborgi f. lilljeborgi have some similarities
to P. nitidum f. paupevoulum. The cardinal tooth C3 of P. varidbile is
well-thickened posteriorly (Figs. 323, 327), the posterior arm often three
times the thickness of the anterior arm, and the anterior margin is a
straight slope or nearly so (Fig. 325). In contrast, P. nitidum f.
pauperoulum has a broad and prominent umbo, and the dorsal margin is very
short, often blending into the anterior and posterior margins imperceptively
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(Fig. 347). In some specimens? the hinge characters of P. variabile are
similar to those of P. nitidum f. pauperculum. The cardinal tooth C3 may
not exhibit the degree of thickening in nepionic P, variabile that is
observed in the adults. Thus, any distinction of these immature specimens
must be based on the slope of the anterior margin as above.
Pisidium punctatum
This species has been recorded from all areas of the Great Lakes with
the exception of Superior. It is not common but may be found in relatively
small and localized populations in sandy substrates.
Variation of P. punctatum occurs in shell outline and prominence and
position of the umbo. Specimens in which the umbo is central and prominent
may be confused with P. ferrungineum, P. lilljeborgi f. lilljeborgi, or P.
compress-urn. The invariably coarse, uniformly spaced striae and dull
periostracum of P, punctatum serve to distinguish it from P. ferrugineum and
most specimens of P. lilljeborgi f. lilljeborgi, and P, compression,
Additionally, P. lilljeborgi f, lilljeborgi has a longer dorsal margin which
creates an elbow or angle where it intercepts the anterior and posterior
margins (Figs. 260, 264) while this transition in P, punctatum is a gradual,
continuous curve, and the dorsal margin is short (Fig. 359). P. compressum
is often separable on the basis of a flattening or incipient ridge on the
umbo (Figs. 238, 239), and the striae are generally fine and closely spaced
rendering them less distinct than in P, punctatum. Individuals on which the
umbo is prominent and posteriorly situated are similar to P. ventricosum f.
ventricosum; however, differences in periostracum, striae, and the degree of
inflation distinguish the two. The periostracum of P, ventricosum f.
ventricosum is glossy, the striae quite fine and the shell is very bulbous,
often resembling a hazel or hickory nut (Fig. 255). In contrast, P.
punctatum exhibits a dull periostracum with coarse, uniform striae and the
degree of inflation does not approach the bulb-like appearance of P.
ventricosum f. ventricosum but rather tapers evenly and gradually from the
apex of the umbo to the ventral margin (Fig. 360). Young P. punctatum are
separable on the same characters.
Pisidium subtruncatum
Records exist from all the Great Lakes where it is relatively common in
clay, mud, and sand substrates. Maximum abundance is reached at about 12 m,
but live specimens often are taken in as much as 40 m of water.
Major sources of variability within P, subtruncatum are the prominence
and breadth of the umbo and the relative thickness of the posterior arm of
cardinal tooth C3. Specimens exhibiting a prominent umbo are similar to
P. lilljeborgi f. lilljeborgi, but there are differences in the posterior
end of the umbo and the dorsal margin. In P. lilljeborgi f. lilljeborgi the
umbo is situated centrally on the shell, yielding a symmetrical outline, and
the dorsal margin extends beyond the umbo, creating elbows or corners with
the anterior and posterior margins (Figs. 260, 264). In P. subtruncatum the
umbo is positioned in the last two-thirds of the shell and the dorsal margin
is short, blending with the anterior and posterior margins (Fig. 298).
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Specimens of P. subtrunoatum with a broader, less prominent umbo are
morphologically close to P, oasertaunwn and P. nitidum f. nitidum. From P.
casertanum, this species is distinguished by the general anterio-ventral
extension of the shell, the thin and evenly tapering C3, and the posterior
margin which slopes nearly vertically from the posterior-lateral cusps and
cuts under to the ventral margin (Fig. 289). In contrast, C3 of P.
easertanum is club-shaped rather than evenly tapering (Figs. 355, 339), and
the posterior margin extends in a gradual but even curve from the posterior-
lateral teeth to the ventral margin (Figs. 337, 341).
P. subtrunoatum differs from P, nitidum f. nitidum in the dorsal margin,
relative hingeplate length and the ligament. The dorsal margin of P. nitidum
f. nitidum extends beyond the umbo to form corners or elbows with the lateral
margins (Fig. 304) and the distance between the cusps of All and PII is half
or greater of the total shell length. The dorsal margin of P. subtruncatum
is short, rarely protruding much beyond the curve of the umbo (Fig. 298),
and the distance between All and PII is always less than half the total
shell length. The ligament of P. subtvuneatum generally is long and wide
(Fig. 301) in contrast to the short, wide ligament of P. nitidum f. nitidum
(Fig. 307), but this trait is less consistent in P. subtvuneatum than the
hinge and dorsal margin traits discussed above.
Immature P. subtruneatum may be separated from morphologically similar
Pisidium species on the basis of characters discussed above.
Pisidium supinim
This species is rare in the Great Lakes and recorded only from shore-
line debris of Lake Ontario (Herrington, 1962). It most likely is an
introduction from Europe.
There is little apparent variation in the species P. supinim.
Herrington (1962) recognized P, supinum only as an ecological form of P.
henslowanum. The two are remarkably similar but most likely represent
separate species. P. supinum, however, is shorter in length rendering it
nearly symmetrical (Fig. 230) whereas P. henslowanaum is clearly
asymmetrical in outline (Fig. 216). Additionally, the striae of P. supinum
are coarse and widely spaced, and the cardinal teeth are situated near the
center of the hingeplate (Figs. 228, 229). P. henslowanum has coarse by
closely spaced striae and the cardinal teeth are located near the anterior
cusps (Figs. 214, 215). Another possible source of confusion is with the
morphotype P. lilljeborgi f. cristatum. The dorsal margin of P. cpistatum
is gradually curved, joining the lateral margins with an angle (Figs. 224,
226). P. supinum contrasts by the greater curve of its dorsal margin and
lack of angles or elbows where it joins the anterior and posterior margins
(Figs. 230, 233, 235). Alternatively, the anterior margin of P, supinum is
long and not much curved while that of P, lilljeborgi f, cristatum is fully
rounded. This may be used as a basis for distinction, even among the
immatures, where the characters of the dorsal margin are ambigous.
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Pisidiwn variabile
This species is relatively common in all the Great Lakes, particularly
on mud and fine sand substrates where it may be locally abundant. Specimens
have been collected from 1-30 m but are most common at depths less than 10 m.
The most significant diagnostic traits in P. variabile are the degree of
gloss of the periostracum, the extent of prominence of the umbo, and the
shape of cardinal tooth C2. Individuals of this species vary from a short,
high, heavy, triangular shell to a long, low, thinner shell which is not
triangular but exhibits a short dorsal margin. Short, high shells tend to
have prominent umbos and may generate confusion with both P. lilljeborgi f.
lilljeborgi and P. sub truncation. In P. lilljeborgi f. lilljeborgi, however,
the dorsal margin extends well beyond the umbo creating corners or elbows
with the anterior and posterior margins (Figs. 260, 264), the periostracum is
slightly dull to moderately glossy, and the lateral cusps All and PII tend to
be sharp on top. These traits contrast to the shorter margin of P. variabile
which blends smoothly into the side margins (Fig. 325), the glossy
periostracum which is often mirror-like in its luster, and the blunt cusps
of All and PII. P. subtruncatum is distinct in that the umbo is positioned
in the last two-thirds of the shell (Fig. 298) and cardinal tooth C3 is not
much thickened posteriorly; the posterior end is never more than twice the
thickness of the anterior end (Figs. 296, 300). The umbo of P. variabile is
situated centrally on the shell (Fig. 325), the tooth C3 is club-shaped, and
the posterior end is invariably greater than twice the thickness of the
anterior end (Figs. 323, 327).
Long, lower shells have broad, less prominent umbos and may be confused
with specimens of P. casertanwn and P. nitidum f. nitidum. However, P.
casertanum usually has sharp cusps at All and PII and the periostracum never
reaches the mirror-like quality found in many P. variabile. Additionally,
the anterior margin of P. casertanum is evenly and continously rounded (Figs.
337, 341) while this edge of P. variabile is straight or only mildly curved
(Fig. 325). With respect to P. nitidwn f. nitidum, C3 exhibits the same
qualities as P. subtrunaatum above (Figs. 302, 306). The dorsal margin
creates corners or elbows as in P. lillg'eborgi f. lilljeborgi (Fig. 304) and
C2 is never peg-like (Figs. 303, 307) as is often the case with P. variabile
(Figs. 321, 328). P. nitidum f. pauperculum also resembles P. variabile in
hinge characters, as discussed earlier. The degree of gloss exhibited by
the periostracum ranges from a mirror-like luster to a moderate glossiness.
The degree of gloss in P. variabile is similar to P. nitidum spp., but these
two species are separable on the bases discussed above. Finally, C2 varies
from a plate-like structure to a stout D-shaped peg. Where C2 is plate-like,
the hinge characters largely resemble those of P. casertanum,, but these
species are distinguished as previously described. Where C2 is peg-like, the
specimen will be close in form to P. compression, but the periostracum of P.
compression is dull or silky-luster and the shell usually shows a flattening
or incipent ridge on the umbo (Figs. 238, 239).
All of the above characters should hold for immature specimens of P.
variabile.
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Pisidium ventriaosum f. ventriaosum
This form is very common in the Great Lakes with the exception of Lake
Superior. It occurs on mud and sandy substrates but prefers soft sediments
in quiet shallow waters. Live specimens have been taken in up to 20 m of
water, but greater densities are reached at depths less than 5 m.
P. ventriaosum f. ventriaosum is quite distinct, and variation is
minimal. It differs from P. ventriaosum f. rotundatum in hingeplate
characters and umbo placement. The umbo of P. ventriaosum f. ventriaosum is
positioned well posterior yielding an asymmetrical appearance (Fig. 254)
while the umbo of P. ventriaosum f. rotundatum is central thus giving a
symmetrical outline (Figs. 244, 248). Additionally, the hingeplate of P.
ventriaosum f. rotundatum is very narrow (Figs. 242, 246, 250, 251) while
that of P. ventriaosum f. ventrieosum is relatively wide (Figs. 252, 256,
257). The above coupled with other anatomical and ecological characters
have lead some workers to propose elevating rotundatum to species status.
The shell of P. ventriaosum f. ventriaosum is the most bulbous of the
Pisidium, resembling a hazel or hickory nut (Fig. 255). The other species
which may approach this degree of inflation are P. lilljeborgi f.
lillj'eborgi3 P. nitidum f. pauperaulum, P. milium, P. ventrieosum f.
rotundatwn and P. ferrugineum. P. lillje'borgi f. lilljeborgi is symmetrical
and the dorsal margin intercepts the anterior and posterior edges with a
distinct angle (Figs. 260, 264); P. ventrieosum f. ventrioosum is
asymmetrical because of the posterior placement of the umbo and exhibits a
very short, tightly curved dorsal margin (Fig. 254). From P. ferrugineum
this species is separable on the basis of hingeplate length and umbo
placement. As in P. tilljeborgi f. 1i11jeborgi, the umbo of P. ferrugineum
is central yielding a symmetrical shell (Fig. 290). The hinge length of P.
ventriaosum f. ventriaosum is only one-third of the shell length (Figs. 252,
254); in P. ferrugineum, the hingeplate length is one-half or more of the
total shell length (Figs. 293, 295). P. nitidum f. pauperaulum has a longer
dorsal margin (Fig. 347), its hinge plate length is one-half the total shell
length, it is symmetrical or nearly so, and it exhibits a short and wide
ligament (Fig. 349). As noted above, P. ventriaosum f. ventriaosum has a
short dorsal margin, it is asymmetrical, and the ligament is generally quite
long and thin (Fig. 257).
The young of P. ventriaosum f. ventriaosum are separable from the
remaining Pisidium species by their rotund, nut-like end view and very short
hingeplate length.
Pisidium ventriaosum f. rotundatum
As with the nominal form, this taxon is common in most of the Great
Lakes, but its distribution may be obscured by reports only at the species
level. P. ventriaosum f. rotundatum occurs most frequently in shallow, calm
waters over soft substrates but may be locally abundant on firm, coarse sand.
P. ventriasoum f. rotundatum is separable from P. ventriaosum f.
ventriaosum in having the umbo centrally placed giving the shell a
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symmetrical appearance (Figs. 244, 248)? and the hingeplate is quite thin
(Figs. 242, 246, 250, 251) in contrast to the moderately wide hinge of
P. ventrioosum f. ventrioosum (Figs. 252, 256, 257). This and other
characters may indicate that rotundatum is a distinct species.
The umbo of P. ventrioosum f. rotundatum overlaps the dorsal margin
to varying degress. Specimens which exhibit little overlap may resemble P.
nitidum f. pauperoulum, but the two are distinguishable by the hinge plate
length, degree of inflation, and ligament. The distance between All and PII
in P. nitidum f. pauperoulum is one-half or more of the shell length while
that of P. ventrioosum f. rotundatum is invariably less than one half (Figs.
250, 251). In end view, P. ventrioosum f. rotundatum is very bulbous and
similar to a hazel nut (Figs. 245, 249); whereas P, nitid.um f. pauperaulwn
exhibits an evenly tapering slope from the apex of the umbo to the ventral
margin (Fig. 348). Finally, the ligament of P. ventrioosum f. rotundatum is
generally long and thin while that of P, nitidum f. pauperoulum is short and
wide. Specimens in which the amount of umbo overlap is great may be confused
with P. lilljeborgi f. lilljeborgi or P, ferrugineum, The dorsal margin of
the form rotundatum is very short and tightly curved (Figs. 244, 248) while
this edge in both P. ferrugineum and P. lilljeborgi f. lilljeborgi extends
well beyond the umbo (Figs. 260, 264, 290). The dorsal margin of P.
lilljeborgi f. lilljeborgi forms distinct angles or corners with the anterior
and posterior margins while in P. ventrioosum f. rotundatum the anterior and
posterior margins begin at the umbo.
Nepionic P. ventricoswn f. rotundatum are distinguished from the
remaining Pisidium by the characteristic, rotund, nut-like end view and
short hingeplate length which extends only one-third of the total shell
length.
Pisidium walkeri f. walkeri and Pisidium walkeri f. mainense
These forms are considered together because they differ only slightly
in ecological and morphological characteristics. Both are common and occur
together on soft, muddy to firm, sandy substrates in water less than 10 m
deep where wave action and current are not too pronounced. They have been
recorded from all the Great Lakes with the exception of Lake Superior,
Pisidium walkeri f, walkeri varies only slightly from P. walkeri f,
mainense. The latter is generally smaller with finer striae and a short
dorsal margin. The anterior margin of P. walkeri f. mainense begins at or
very near the umbo; whereas, that of P, walkeri f. walkeri is more distant
from the umbo owing to the longer, more extended dorsal margin, C2 of P.
Walkeri f. walkeri is short, bent plate and C4 is curved and directed toward
the interior of the shell (Figs. 269, 273), while C2 of P, walkeri f,
mainense is nearly as long as C4 and parallel to it as in P. subtruncatum
(Fig. 275). Specimens of P. walkeri f, walkeri in which the umbo is not
prominent are similar to P. oasertanum; however, the umbo of P, walkeri f.
walkeri is generally narrower (cf. Figs. 270 with 337, 341), the ligament
is long and thin rather than long and wide, and the anterior margin is long
and barely curved (Fig. 270); the anterior margin of P. oasertanum is fully
rounded anteriorly (Figs. 337, 341, 373). Where the umbo of P, walkeri
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f. walkeri is prominent it may resemble P. subtruncatum, P. lilljeborgi f.
lilljebovgi or P. vavi.abi.1e. In P. waUkevi f. walkeri, C3 is enlarged
posteriorly (Figs. 268, 272) and C2 is short and bent with C4 overlapping C2
as it curves toward the interior of the shell (Figs. 269, 273). In contrast,
C3 of P. subtvunaatwn is not thickened posteriorly (Figs. 296, 300), and C2
and C4 are parallel and often nearly the same length (Figs. 297, 301). P.
lilljebovgi f . Ulljebovgi differs from P. walkefi f . walkevi in that the
dorsal margin forms corners or elbows with the anterior and posterior
margins, and the umbo is positioned centrally yielding a nearly symmetrical
outline (Figs. 260, 264). Even where extended significantly, the dorsal
margin of P. walkeri f. walkeri does not intersect the lateral margins with
an angle, and the umbo is situated at or near the last two-thirds of the
shell giving an asymmetrical appearance (Fig. 270). P. variabile is
distinct on the basis of periostracal features and the lateral cusp. The
periostracum of P. vaviabile is glossy, often mirror-like in appearance, and
the lateral cusps are short and blunt. P. walkeri f. walkeri generally
exhibits a dull to silky periostracum and the lateral cusps are rather long
and sharp on top .
There is little variability among specimens of P. waUkevi f.
however, they may be confused with P. nitidum f. pauperculwn3 P. lilljebovgi
f. lilljeborgi, and P. variabile. C2 and C4 are of the same length and
parallel in P. walkeri f. mainense. In both P. lillj eborgi f. lillgebovgi
and P. variabile C2 is short and thick or a bent plate and C4 is curved and
directed interior of PII (Figs. 263, 328). The umbo of P. nitidum f.
paupepculwn is broad while that of P. walkeri f. mainense is narrow and
generally projects above the dorsal margin. The periostracum of P. nitidum
f. paupereulum is very glossy, almost mirror-like; the ligament is very
short and thick, covering more than half of the hingeplate width. P. walkeri
f. mainense exhibits a dull to silky periostracum; the ligament is long and
thin, covering less than half the hingeplate width.
The above characters will hold for immature P. waVkeT-l forms.
SPHAERIUM
Sphaeriwn aorneum
This species is locally abundant in Lake Erie and common in localized
populations in Lake Ontario and Lake Michigan but uncommon in Lake Huron
or Lake Superior. The usual habitat is fairly quiet water with little or
no wave action, and there is an association with organic enrichment (Wurtz,
1956) . Live specimens have been taken from Lake Michigan in water down to
10 m deep on substrates ranging from soft mud to hard gravel.
Sphaerium nitidwn
Clarke (1973) and Herrington (1962) consider S. nitidum to be a cold
water species typically occurring only in deep or unusually cold waters,
particularly in the southern part of its range. The species has been
recorded from all the Great Lakes with the exception of Lake Erie which
would tend to confirm its cold stenothermic nature. However, S. nitidum is
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common in warm, shallow areas of southern Lake Michigan. Greatest abundances
are found at about 20 m where the substrate is sandy. Live specimens are not
uncommon at 40-50 m in Lake Michigan.
Sphaeriion oooidentale
Heard (1962) records this species for all the Great Lakes with the
exception of Lake Michigan. These records seem anomolous since the species
is characteristic of vernal pools and "has a preference for, or requires, a
habitat that dries up for part of the year" (Herrington, 1962). The
explanation for its presence in the Great Lakes may be that it occurs only
in shore zones that partly dry up during the year. This has yet to be
confirmed. There is little ecological data with museum records or in Heard
(1962).
Sphaerium rhomboideum
Records exist only for Lake Ontario. Its habitat preference in the
Great Lakes is unknown, but Herrington (1962) associates it with muddy
bottoms in creeks and small rivers and quiet, sheltered places in small
lakes.
Sphaeriwn simile
S. simile is locally common in sandy substrates less than 10 m deep in
Lake Ontario. It has not been recorded for the other Great Lakes.
Sphaeviwn striatinum
This is the most common species of Sphaerium and occurs in all the
Great Lakes. It may be collected in a variety of substrates from clay and
mud to sand and gravel at depths from 0-30 m. Gale (1973) showed that
adult stream specimens exhibit no real substrate preferences, although
small clams did select mud over sand and sandy-mud.
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SYNONYMY
The synonyms listed below are for the convenience of those working with
the fauna of the Laurentian Great Lakes and reflect names frequently
encountered in museum catalogues and the literature. The list is not
complete for many species as it includes only synonyms used in a Great Lakes
publication or report. More comprehensive synonymies can be found in
Baker (1928a, 1928b), Berry (1943), Robertson and Blakeslee (1948),
Herrington (1962), Clarke (1973), Burch (1975a, 1975b), and Te (1978).
Synonym
GASTROPODA
altissimus Baker
antrosus Conrad
appressus Say
arotious Beck
aurioularia Linnaeus (Lymnaed)
bioarinatus Say
binneyana Hannibal (Armicola)
oaperata Say (Lymnaed)
oatasoopiim Say (Lymnaed)
oolumella Say (Lymnaed)
aonteotoides Binney
deoampi Streng (Lymnaed)
desidiosa Say (Lymnaed)
emarginata Kuster (Lymnaed)
emarginata Say (Lymnaed)
exilis Lea (Lymnaed)
hirsutus Gould
hwnilis Say (Lymnaed)
hypnorum Linnaeus
•Integra Say (Amnioola)
kirklandi (Ferrissia)
laoustre Pilsbry (Amnioola)
lanoeata Gould (Lymnaed)
lustrioa Pilsbry (Amnioola)
malleatus Martens
megasoma Say (Lymnaed)
palustris Mtlller (Lymnaed)
pilsbryana Walker (Lymnaed)
porata Say
Present Status
status unknown (=lGyraulus parvus
(Say))
Eelisoma anoeps (Menke)
Lymnaea stagnalis appressa (Say)
status unknown (=lGyraulus
ciroimstri-atus Tryon)
Radix auricularia (Linnaeus)
Helisoma anoeps (Menke)
Probyth-inella laoustris (Baker)
Stagnicola oaperata (Say)
Stagnicola oatascopiwn (Say)
Pseudosuooinea eolwnella (Say)
Viviparus georgianus (Lea)
Fossaria deoampi (Streng)
Fossaria deoampi (Streng)
Probyfhinella laaustris (Baker)
Stagnioola emarginata (Say)
Stagnioola reflexa (Say)
Gyraulus defleatus (Say)
Fossaria hwnilis (Say)
Aplexa elongata Say
Cinoinnatia oincinnatiensis Anthony
status unknown
Marstonia deaepta (Baker)
Stagnioola reflexa (Say)
Marstonia deaepta (Baker)
Viviparus japonicus Martens
Bulimnea megasoma (Say)
Stagnioola elodes (Say)
Stagnioola aatasaopiim (Say)
Armicola limosa (Say)
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Synonym
Present Status
reflexa Say (Lymnaea)
sheldoni Pilsbry (Hoyia)
tentaculata Linnaeus (Bulimus)
truncation Miles
walkeriana Baker (.Lyrmaed)
woodruffi Baker (Lyrnnaea)
SPHAERIIDAE
abditum Haldeman
acuminatum Prime
aequilaterale Prime
altile Prime
cinereum Alder
emarginata Prime
kirklandi Sterki
medianwn Sterki
obtusale Herrington
ryckholti Normand
stamineum Conrad
subtransversum Prime
sulcatum Lamarck
UNIONIDAE
attenuata Rafinesque
borealis Gray
buchanensis Lea
canadensis Lea
corpulenta Cooper
costata Rafinesque
footiana Lea
gibbosus Barnes
graaHis Barnes
hippopaeus Lea
•iris Lea (M-iaromya")
katherinae Lea
kenniaottii Lea
latissima Rafinesque
marginata Say
parvula Grier
recta Lamarck (Unio)
rugosus Swainson
siliquoidea Barnes
simpsoniana Lea
sterkii Grier
Stagnicola reflexa (Say)
status unknown
Bithynia tentaculata (Linnaeus)
status unknown (,=?Helisoma tr-ivolv-is
(Say))
Stagn-icola catascop-ium (Say)
Stagnicola catascopium nason-L (Baker)
Pisidium casertanum (Poli)
form of Sphaerium striatinum (Lamarck)
not considered here
Pisidi-um equilaterale Prime
Pisidium compression Prime
Pisidium casertanum (Poli)
form of Sphaerium striati-num (Lamarck)
not considered here
Pisid-Lum fallax Sterki
Pisidium ferrugineum Prime
Pisidiwn ventricosum Prime
form of Musculium lacustre (MUller)
not considered here
Sphaerium striatinum (Lamarck)
Musculium transversum (Say)
Sphaerium simile (Say)
Ligumia nasuta Say
Lampsilis radiata Gmelin
Anodontoides ferussacianus (Lea)
Lampsilis ovata (Say)
Anodonta grandis Say »
Amblema plicata (Say)
Anodonta grandis Say
Elliptic dilatata (Rafinesque)
Leptodea fragilis (Rafinesque)
Amblema plicata (Say)
Villosa iris (Lea)
Lasmigona complanata (Barnes)
Anodonta grandis simpsoniana Lea
Ligumia recta (Lamarck)
Anodonta cataracta Say
Fusconaia flava (Rafinesque)
Ligumia recta (Lamarck)
Strophitus undulata (Say)
Lampsilis radiata siliquoidea Barnes
Anodonta grandis simpsoniana Lea
Elliptic dilatata (Rafinesque)
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Synonym
Present Status
superiorensis Marsh Lampsilis radiata siliquoidea Barnes
triquetra Rafinesque (Trunc-illa') Dysnomia triquetra (Rafinesque)
undata Barnes Fusconaia flava (Rafinesque)
ventrioosus Barnes Lampsilis ovata (Say)
wiridis Rafinesque Alasmidonta caloeolus (Lea)
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GLOSSARY
abaxially: Situated outside of or facing away from the axis.
adapically: Towards or facing the apex of the shell.
anal siphon: The dorsal exhalant tube derived from mantle-margin epithelium
and located near the anus at the posterior end of the animal.
aperture: The opening at the larger end of the shell of a gastropod.
apex: The tip or oldest part of a shell.
apical: At or belonging to the apex or oldest portion of a gastropod shell.
attenuate: Tapering gradually.
basal: Relating to or situated at the base.
beak: The raised part on the dorsal margin of a valve. It is formed by the
embryonic shell around which the later shell develops. Also called the
"umbo."
beak cavity: The cavity on the inside of each valve of a mussel shell going
into the beak. In some species the cavity is quite deep, in others it
is shallow so as to be little more than a weak depression.
bifid: Incised; cut in so as to be divided into two parts.
body whorl: The last whorl of the shell, it supports the aperture. The
term "body whorl" is misleading since the organs of the animal are
distributed through all the whorls.
branchial opening: The ventral posterior opening or siphon through which
water enters the mantle cavity of a mollusc such as a freshwater
mussel. Also called the "incurrent opening" or "siphon."
branchial siphon: The ventral inhalant tube derived from the mantle-margin
epithelium located at the posterior end of the animal.
calycle (calyculae): A growth variation of the beak or umbo, typical of
Musaulium born at or near the end of the summer, yielding a cap-like
appearance.
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canaliculate: Grooved or channeled longitudinally.
cardinal teeth: Lamellae on the center of the hinge of both valves of a
bivalve which serve to stabilize the two valves against shearing
forces. The opposing cardinal teeth of the two valves fit together in
a complimentary fashion. There are usually two cardinal teeth in the
left valve (C2, C4) and one in the right (C3).
carinae: Small, sharp, raised ridges or keels.
caruncle: A fleshy elevation or outgrowth; a characteristic protuberance on
the inner edge of each side of the mantle in front of the branchial
opening of members of the genus Carunculina.
chevron-shaped: Shaped like a wide-angled V, normally positioned or
inverted.
color ray: A more or less straight band of color, continuous or
discontinuous, contrasted to the ground color of the shell and
radiating from the umbonal area distally towards or to the peripheral
margins of the valve.
columella: The central axis or "column" of a gastropod shell; seen only as
the inner lip of the aperture.
compressed: Flattened or pressed together laterally, such as the appearance
of some freshwater molluscs in end view.
corrugations: Alternating ridges and furrows on the surface of a unionid
shell.
cusps: The highest elevations on the lateral teeth.
demibranch: One-half of one of the paired gills of a lamellibranch
pelecypod; i.e. the two opposing rows of gill filaments on one side of
the gill; a half-gill.
denticle: A toothlike projection usually at or just interior of the
aperture in some gastropod shells.
dextral: Coiled to the right. When held with the apex up and aperture
facing viewer, the aperture of a dextral shell is on the right side.
disc: The middle, central, or main portion of the exterior of a mussel
valve as distinct from the posterior slope and other areas immediately
adjacent to the marginal peripheries.
distal: Farthest from the beak or umbo of the shell in an anterior-
posterior axis.
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gill (branchia): The platelike or filamentous growth, usually located
within the mantle cavity, serving as the respiratory organ of aquatic
molluscs. In lamellibranch molluscs the gills are greatly enlarged,
serving also as a food gathering organ through filter-feeding.
gravid female: A female with a marsupium containing young embryos.
ground color: The basic or background color of a shell against which any
additional color markings are contrasted.
growth lines: Minute lines on the outer shell surface indicating a minor
rest period during growth. Not to be confused with the major "rest
marks" caused by prolonged growth arrest (as during winter).
gyrate: Circular, rounded; winding or coiled round.
hinge: (Unionidae) The portion of the dorsal margin between and including
the pseudocardinal teeth and lateral teeth.
(Sphaeriidae) The part of the dorsal margin of the shell between
and including the anterior and posterior lateral teeth of each valve.
hinge teeth: The opposing lamellae on the hinge plates of bivalved molluscs
which serve to stabilize the two valves against shearing forces.
hirsute: Covered with hairs.
imperforate: Not open; used in reference to the umbilicus or columella of a
gastropod shell.
inflated: Swollen or bulbous.
interdentum: The space on the hinge plate between the pseudocardinal and
lateral teeth in unionid clams.
lateral teeth: The elongated lamellae at each end of the hinge plate.
malleated: With flattened areas, appearing like beaten metal.
mantle: An extension of the dorsal body wall of molluscs as one or a pair
of folds which usually secretes a shell and encloses a mantle cavity,
typically containing gills.
marsupium: The pouchlike structure used to hold the young. In unionids,
internal spaces in the gills perform this function.
multicarinate: Having small, sharp, raised ridges or keels.
nacre: The white or iridescent inner layer of the shell in many molluscs
lying next to the mantle and often characteristically colored in many
unionacean clams.
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nodule: A small knot, lump, or irregularly shaped mass such as the
projections occurring on the shell surface of some freshwater mussels.
nodulous: Having small knobs, nodules, or projections.
nuclear whorl: The whorl located at the apex or oldest part of a gastropod
shell, also called a "protoconch."
operculum: A stiff corneous or calcareous plate attached to the odrsal part
of the foot of certain gastropods, arranged so that when the animal
withdraws into the shell the operculum seals the aperture.
pallial line: The line of attachment of the mantle to the shell on the
inside surface of a bivalved shell, often marked by a depression or
scar.
papillate: Having many small papillae or bumps on the surface.
perforate: With an opening; used in reference to the umbilicus or columella
of a gastropod shell.
periostracum: The pigmented outside layer of a molluscan shell.
peritreme: The lips of the aperture in limpets.
plaited: Having flattened folds, pleated.
planospiral: Spiraling in a single plane.
posterior ridge: A ridge on the external surface of many mussel shells
extending from the umbo posterioventrally to the shell margin.
posterior slope: The area on the external surface of a mussel shell between
the posterior ridge and the dorsal margin of the shell.
protoconch: The whorl located at the apex or oldest part of a gastropod
shell.
proximal: Nearest the beak or umbo of the shell in an anterior-posterior
axis.
pseudocardinal teeth: The anterior, opposing lamellae located on the hinge
plates of bivalved molluscs which serve to stabilize the two valves
against shearing forces.
pustule: A blister-like prominence found on the shell surface of some
freshwater mussels.
ray: A streak or linear mark. It may be continuous or interrupted at
intervals.
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ridge: A wrinkle or raised part on the beaks of some species of Pisidium.
septa: Partitions formed by interlamellar connections separating spaces
occurring between the two lamellae.
sinistral: Coiled to the left. When held with the apex up and aperture
facing viewer, the aperture of a sinistral shell is on the left side.
siphon: A tubular structure formed by the opposing posterior mantle margins
in mussels; a pair are commonly present on bivalves, providing
restricted incurrent and excurrent opening to the mantle cavity.
spire: All whorls of a gastropod shell except the last, or body, whorl.
striae: (Bivalvia) Concentric raised striations or lines on the exterior
surface of a shell. These may vary from very fine to coarse, sometimes
resembling ribs.
(Gastropoda) Small ridges running perpendicular to the growth
lines on the exterior surface of the shell.
subcentral: Not quite central; off-center.
sulcus: A groove, furrow, or channel.
suture: The line of junction between two contiguous whorls of the shell of
a gastropod.
terminal: Located at or near the posterior end.
truncate: Having the end cut off more or less squarely.
tubercle: A nodule or small eminence, such as a solid elevation, occurring
on the shell surface of some freshwater mussels.
tuberculated: Containing many small nodules or eminences.
umbilical: Of or pertaining to the umbilicus of a gastropod shell.
umbilicus: The depression in the base of the shell (opposite the spire)
around which the body whorl is coiled; it is hollow in tube-like
columella.
umbo: The oldest part of the bivalve shell identified as the raised
portion at or overlapping the dorsal margin of a valve. Also called
the "beak."
undulation: A wavy form, resembling that of a wave or waves.
vestigial: A small, degenerate or imperfectly developed bodily part or organ
that remains from one more fully developed at an earlier stage of the
individual or in past generations.
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whorl: A single volution of the spiral shell of a gastropod.
wing: The dorsal, thin, flat, plate-like extension of the posterior slope
of some freshwater mussels.
129
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REFERENCES
References marked with an asterisk (*) have not been cited in the text;
however, they are included here because they directly refer to benthic or
mollusk studies of the Laurentian Great Lakes or are pertinent to the study
of a species occurring in the Great Lakes.
Adams, C. E. and R. D. Kregar. 1969. Sedimentary and faunal environments
of eastern Lake Superior. Proc. 12th Conf. Great Lakes Res. 1969:
1-20.
*Adamstone, F. B. 1924. The distribution and economic importance of the
bottom fauna of Lake Ontario. Univ. Toronto Stud., Biol. Ser. 25:
33-100.
*Ahlstrom, E. H. 1930. Mollusk collected in Bass Island region, Lake Erie.
Nautilus 44:44-48.
Baker, F. C. 1902. The Mollusca of the Chicago area, the Gastropoda.
Bull. Chicago Acad. Sci. Nat. Hist. Surv. 3:131-418.
Baker, F. C. 1911. Lymnaeidae of North America. Chicago Acad. Sci. Spec.
Publ. No. 3. 539 pp.
Baker, F. C. 1928a. The fresh-water Mollusca of Wisconsin Pt. 1. (Bull.
No. 70) Gastropoda. Wisconsin Acad. Sci. Arts Let. 507 pp.
Baker, F. C. 1928b. The fresh-water Mollusca of Wisconsin Pt. 2.
Pelecypoda. Bull. Univ. Wisconsin No. 1527. 495 pp.
Baker, F. C. 1930. The molluscan fauna of the southern part of Lake
Michigan and its relationship to old glacial Lake Chicago. Trans.
Illinois Acad. Sci. 22:186-194.
Ball, G. H. 1922. Variation in fresh-water mussels. Ecology 3:93-121.
Basch, P. F. 1963. A review of the recent fresh-water limpet snails of
North America. Bull. Mus. Comp. Zool. 129:401-461.
*Bickel, D. 1965. The role of aquatic plants and submerged structures in
the ecology of a freshwater pulmonate snail, Physa integra Hald.
Sterkiana 18:17-20.
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Berry, E. G. 1943. The Amnicolidae of Michigan: distribution, ecology,
and taxonomy. Misc. Publ. Mus. Zool., Univ. Michigan 57:1-68.
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138
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INDEX TO SPECIES, SUBSPECIES, AND FORMS
Page numbers in italics refer to figures.
aowninatwn (see Spha.er-i.ion striatinum') cataracta, Anodonta, 6, 26, 102
acuta, Pleuroeera, 6, 19, 98
adamsi, Pisidium, 7, 55, 93, 95, 96,
103, 104, 105, 106
Alasmidonta aaloeolus , 6, 27, 28,
102
alata, Proptera, 7, S3, 102
ambigua, Simpsonioonoha, 7, 27, 102
Amblema plioata,- 6, 24, 102
limosa, 6, 5, 20, 22, 97
walkeri, 6, 20, 22, 97
armiaum, Pisidium, 7, 55, 57, 59,
104, 107, 110
anoeps , Helisoma, 6, 11, 100
Anodonta
cataracta, 6, 26, 102
grandis grandis, 6, 25, 102
grandis simpsoniana, 6, 26, 101
imbeaillis, 6, 25, 101
Anodont aides ferussaoianus , 6, 2£,
102
appressa (see Lymnaea stagnalis)
auriaularia, Radix, 6, 25, 99
basalis (see 7aZ-vata tricarinata")
bioarinata, Valvata, 6, 17, 25, 97
Bithynia tentaoulata, 6, 2{7, 98
Bulirmea megasoma, 6, 16, 27, 99
aalaeolus , Alasmidonta, 6, 27, 28,
102
campanula-turn, Helisoma, 6, 11, 12,
100
Campe loma
deciswn, 6, 5, 18, 19, 97
rufum, 6, 18, 19, 97
aaperata, Stagnicola, 6, 16, 17, 99
Carunculina parva, 6, 30, 101
oasertamm, Pisidium, 1, 7, 84, 55,
57, 89, 96, 104, 105, 106, 109,
112, 113, 115, 116, 118
catasoopiwn f. catascopiwn,
Stagnieola, 6, 25, 17, 99
catascopiwn f. nasoni, Stagnicola, 6,
25, 99
Cinainnatia oinainnatiensis, 6, 20,
22, 98
cinoinnatiensis, Cinainnatia, 6, 20,
22, 98
circumstriatus, Gyraulus, 6, 14, 100
oooaineum (see Pleurobema cordatum)
colwnella, Pseudosucainea, 6, 15,
16, 99
complanata, Elliptic, 7, 29, 102
oomplanata, Lasmigona, 7, 27, 28,
102
compressum, Pisiditm, 7, 63, 65, 91,
104, 105, 106, 108, 110, 114, 116
conventus, Pisidium, 1, 3, 7, 52, 52,
106, 108, 109, 110, 112
cordatwn f. coccineum, Pleurdbema, 7,
29, 101
covnewn, Sphaeriwn, 7, 50, 52, 54,
119
eorpulentum, Helisoma, 6, 12, 13, 100
oostata, Lasmigona, 7, 27, 28, 101
cristatwn (see Pisidium lilljeborgi)
Cyolonaias tuberculata, 6, 29, 101
deoampi, Fossaria, 6, 16, 99
deaiswn, Campeloma, 6, 9t 18, 19, 97
defleatus, Gyraulus, 6, 13, 100
dilatata, Elliptic, 7, 29, 102
donaaiformis, Truneilla, 7, 32, 102
dubium, Pisidium, 7, 36, 55, 57, 58,
104, 107
Dysnomia
suloata, 6, 31, 101
triquetra, 6, 31, 102
139
-------
Elliptic
aomplanata, 7, 29, 102
dilatata, 7, 29, 102
elodes, Stagnicola (used for
illustrative purposes only), 8
emarginata, Stagnioola, 6, 16, 27,
99
emarginatum (see Sphaerium
striatinwn)
equilaterale, Pisidiim, 7, 91, 92,
107
exacuous, Promenetus, 6, 8, 13, 100
fdbilis, Villosa, 7, 34, 102
fallax, Pisidiim, 7, 81, 83, 106,
107, 108
fasoiola, Lcanpsilis, 7, 34, 101
fasoiolare, Ptyohobranohus, 7, 33,
101
Ferrissia
paralella, 6, 8, 10, 11, 100
tarda, 6, 10, 11, 100
ferrugineum, Pisidiwn, 7, 75, 77,
108, 109, 114, 117, 118
ferrussaaianus, Anodontoides, 6, 26,
102
flava, Fusoonaia, 7, 24, 25, 102
Fossaria
decampi, 6, 16, 99
humilis, 6, 16, 99
o&rwssa, 6, iff, 99
fragilis, Leptodea, 7, 33, 102
Fusoonaia flava, 7, 24, 25, 102
fuscus, Laevapex, 6, 10, 100
georgianus, Viviparus, 6, 18, IS,
97
Goniobasis livesoens, 6, S, 15, 98
grandis grandis, Anodonta, 6, 25,
102
grandis simpsoniana, Anodonta, 6,
25, 101
Gyraulus
drownstriatus, 6, 14, 100
defleotus, 6, 13, 100
parvus, 6, 14, 100
gyrina sayi, Physella, 6, 14, 100
Helisoma
anceps, 6, II, 100
oampanulatum, 6, 11, 12, 100
corpulentum, 6, 12, 13, 100
trivolvis, 6, 22, 100
henslowanim, Pisidiim, 7, 60, 61,
106, 108, 109, 111, 115
humilis, Fossaria, 6, 16, 99
idahoense, Pisidiim, 7, 84, #£, 55,
110
imbeaillis, Anodonta, 6, 25, 101
integra, Physella, 6, 14, 15, 100
iris, Villosa, 7, 34, 102
japoniaus, Viviparus, 6, 18, 13, 97
jayense (see Musoulium laeustre)
laoustre f. jayense, Musoulium, 7,
41, 43, 103
laoustre f. laoustre, Musoulium, 7,
44, 45, 103
laoustris, Probyfhinella, 6, 20, 21,
98
Laevapex fuscus, 6, 1(7, 100
laevissima, Proptera, 33, 101
LampsiZ-is
fasoiola, 7, 34, 101
ouata, 7, 32, 102
radiata radiata, 7, 34, 35, 101
radiata siliquoidea, 4, 7, 34,
35, 102
Lasmigona
oomplanata, 7, 27, 28, 102
oostata, 7, 27, 28, 101
Leptodea fragilis, 7, 33, 102
lewisi, Valvata, 6, 17, 15, 97
Ligumia
nasuta, 7, 30, 102
recta, 7, 30, 102
lilljeborgi, Pisidiim, 1
lilljeborgi f. oristatim, Pisidiim,
7, 60, 62, 110, 111, 115
lilljeborgi f. lilljeborgi, Pisidiim,
7, 70, 71, 106, 108, 110, 111, 113,
114, 116, 117, 118, 119
limosa, Amnioola, 6, 9, 20, 21, 97
livesoens, Goniobasis, 6, 9, 19, 98
Lymnaea stagnalis appressa, 6, 16,
17, 99
mainense (see Pisidiim walkeri)
Marstonia deoepta, 6, 20, 98
megasoma, Bulirmea, 6, 16, 17, 99
milium, Pisidiim, 7, 75, 7£, 108,
111, 112, 117
Musouliwn
laoustre f. jayense, 7, 41, 43,
103
140
-------
locustre f. lacustre, 7, 44, 45,
103
partumeium, 7, 38, 40, 103
seeuris, 7, 41, 42, 103
transversum, 7, 37, 35, 35, 103
nasoni (see Stagnicola catascopium)
nasuta, Ligumia, 7, 30, 102
nitidum, Pisidium, 109
nitidum f. nitidum, Pisidium, 7, 78,
80, 105, 107, 109, 111, 112, 113,
115, 116
nitidum f. pauperculum, Pisidium, 7,
84, 87, 90, 108, 110, 112, 113,
114, 117, 118, 119
nitidum, Sphaerium, 7, 50, 52, 53,
119
nodulata, Quadrula, 7, 24, 25, 101
Obliquaria reflexa, 7, 30, 101
Obovaria
olivaria, 7, 31, 101
subrotunda, 7, 31, 101
obrussa, Fossaria, 6, iff, 99
oeoidentale, Sphaerium, 7, 50, 55,
50, 120
olivaria, Obovaria, 7, 31, 101
ovata, Lampsilis, 7, 32, 102
paralella, Ferrissia, 6, fl, 10, 11,
100
partumeium, Musculium, 7, 38,40, 103
parva, Carunaulina, 6, 30, 101
paryue, Gyraulus, 6, 24, 100
paupereulton (see Pisidium nitidum)
peroonfusa (see Valvata tricarinata)
perdepressa, Valvata, 6, 18, 97
Fhysella
gyrina sayi, 6, 14, 100
integra, 6, 24, 15, 100
uinosa, 6, 24, 15, 100
pisoinalis, Valvata, 6, 25, 97
Pisidiwm
adamsi, 7, 55, 53, 55, 96, 103,
104, 105, 106
armiaum, 7, 55, 57, 55, 104, 107,
110
casertaman, 1, 7, 84, 86, 87, 55,
50, 104, 105, 106, 109, 112,
113, 115, 116, 118
compression, 7, 63, 65, 91, 104,
105, 106, 108, 110, 114, 116
conventus, 1, 3, 7, 52, 52, 106,
108, 109, 110, 112
dubiwn, 7, 36, 55, 57, 58, 104,
107
equilaterale, 7, 91, 92, 107
fallax, 7, 81, 83, 106, 107, 108
ferrugineum, 7, 75, 77, 108, 109,
114, 117, 118
henslowanum, 7, 60, 61, 106, 108,
109, 111, 115
idahoense, 7, 84, 86, 88, 110
lilljeborgi, 1
lilljeborgi f. cristatum, 7, 60,
62, 110, 111, 115
lilljeborgi f. lilljeborgi, 7,
70, 71, 106, 108, 110, 111, 113,
114, 116, 117, 118, 119
milium, 7, 75, 76, 108, 111, 112,
117
nitidum, 109
nitidum f. nitidum, 1, 78, 80,
105, 107, 109, 111, 112, 113,
115, 116
nitidum f. pauperculum, 7, 84,
87, 90, 108, 110, 112, 113,
114, 117, 118, 119
punctatum, 7, 91, 53, 94, 112, 114
subtruncatum, 7, 78, 79, 104, 105,
110, 112, 113, 114, 115, 116,
118, 119
supinum, 7, 63, 64, 105, 111, 115
variabile, 7, 84, 85, 91, 104,
105, 106, 107, 109, 110, 113,
114, 116, 119
ventricosum f. rotundatim, 7, 66,
67, 68, 111, 117, 118
ventricosum f. ventricosum, 7,
67, 69, 111, 112, 114, 117, 118
walkeri f. mainense, 7, 72, 74,
104, 105, 118, 119
walkeri f. walkeri, 7, 72, 73,
118, 119
Pleurobema cordatum f. coccineum, 7,
29, 101
Pleurocera acuta, 6, 19, 98
plicata, Amblema, 6, 24, 102
Probythinella lacustris, 6, 20, 22,
98
Promenetus exaeuous, 6, 5, 23, 100
Proptera
alata, 7, 33, 102
laevissima, 33, 101
141
-------
Pseudosuooinea oolwnella, 6, 15, 16,
99
Ptyehobranohus fasaiolare, 7, 33, 101
punatatwn , Pisidiwn, 7, 91, 93, 84,
112, 114
pustulosa, Quadrula, 7, 24, 25, 101
Quadrula
nodulata, 7, 24, 25, 101
pwetulosa, 7, 24, 25, 101
quadrula, 7, 24, 102
quadrula, Quadrula, 7, 24, 102
radiata radiata, Lampsilis, 7, 34,
35, 101
radiata siliquoidea, LampsiHs, 4, 7,
34, 35, 102
Radix aurioularia, 6, 25, 99
recta, Ligwnia, 7, 30, 102
reflexa, Obliquaria, 7, 30, 101
reflexa, Stagnioola, 6, 15, 99
rhomboideum , Sphaerium, 7, 50, 51,
120
rotundatwn (see Pisidiwn ventriooswn)
rufwn, Campeloma, 6, 18, 23, 97
sayi, gyrina, Physella, 6, 24, 100
seouris, Musouliwn, 7, 42, 42, 103
eiliquoidea (see Lampsilis radiatd)
simile , Sphaeriim, 7, 45, 47, 120
simpsoniana, grandis, Anodonta, 6,
26, 101
Simps oniconcha ambigua, 7, 27, 102
sinaera, Valvata, 6, 18, 97
Somatogyrus subglobosus, 6, 20, 98
corneum, 7, 50, 52, 54, 119
nitidum, 7, 50, 52, S3, 119
oeoidentale, 7, 50, 55, 55, 120
rhomboidewn, 7, 50, 51, 120
striatinum, 7, 46, 120
striatinum f. acwninatum , 46, 48
striatinim f. emapginatwn, 46, 48
stagnalis appressa, Lyrrmaea, 6, 16,
27, 99
oaperata, 6, 16, 27, 99
catascopium f. oatasoopiwn, 6,
25, 17, 99
oatasoopiwn f. nasoni, 6, 25, 99
elodes (used for illustrative
purposes only) , 8
emarginata, 6, 16, 27, 99
veflexa, 6, 25, 99
striatimon , Sphaerium, 7, 46, 120
striatimm f. aowninatum, 46, 48
striatinum f. emarginatum , 46, 48,
40
Strophitus undulatus, 7, 26, 102
subglobosus, Somatogyrus, 6, 2(9, 98
subrotunda, Obovaria, 7, 31, 101
subtrunoatw, Pisidiwn, 7, 78, 79,
104, 105, 110, ,,2, 113, 114, 115,
116, 118, 119
suloata, Dysnomia, 6, 32, 101
supinwn, Pisidiwn, 7, 63, 64, 105,
111, 115
tarda, Fevvissia, 6, 10, 11, 100
tentaoulata, Bithynia, 6, 20, 98
transverswn , Musouliwn, 7, 37, 38,
39, 103
tricarinata f. basalis, Valvata, 6,
17, 97
trioarinata f. peroonfusa, Valvata,
6, 17, 97
tricarinata f. tricarinata, Valvata,
6, 3, 17, 28, 97
triquetra, Dysnomia, 6, 32, 102
trivolvis, Helisoma, 6, 22, 100
truncata, Truncilla, 7, 32, 102
Trunoi lla
donaoiformis, 7, 32, 102
trunoata, 7, 32, 102
tuberculata , Cyolonaias, 6, 29, 101
undulatus, Strophitus, 7, 26, 102
bioarinata, 6, 17, 28, 97
lewisi, 6, 17, 28, 97
perdepressa, 6, 28, 97
pisainalis, 6, 28, 97
sincera, 6, 28, 97
tpicarinata f. basalis, 6, 17, 97
trioarinata f. peroonfusa, 6, 17,
57
tricarinata f. triearinata , 6, 3,
17, 28, 97
variabile, Pisidiwn, 7, 84, 85, 91,
104, 105, 106, 107, 109, 110, 113,
114, 116, 119
ventricoswn f. rotundatum, Pisidiwn,
7, 66, 67, 68, 111, 117, 118
ventrieoswn f. ventricosum, Pisidiwn,
7, 66, 57, 83, 111, 112, 114, 117,
118
142
-------
Villosa
fdbilis, 7, 34, 102
iris, 7, 34, 102
vinosa, Physella, 6, 14, 15, 100
Vivipams
georgianus, 6, 18, 19, 97
japonicus, 6, 18, 19, 97
malleatus, 97
walkeri, Arnnioola, 6, 20, 2i, 97
walkeri f. mainense, Pisidiwn, 7, 72,
74, 104, 105, 118, 119
walkeri f. walkevi,, Pi-sid-ium, 7, 72,
73, 118, 119
143
-------
TECHNICAL REPORT DATA
(I'ti-asc read lHsifueiu>>i\ on the reverse before completin/t}
I. FUPoru NO.
EPA-600/3-80-068
3. RECIPIENT'S ACCESSION-NO.
A. TITLE ANOSU8TITLE
A Guide to Freshwater Mollusks of the Laurentian
Great Lakes with Special Emphasis on the Genus
5. REPORT DATE
JULY 1980 ISSUING DATE
6. PERFORMING ORGANIZATION CODE
7. AUTHOR(S)
8. PERFORMING ORGANIZATION REPORT NO.
Gerry L. Mackie, David S. White, Thomas W. Zdeba
9. PERFORMING ORG \NIZATION NAME AND ADDRESS
10. PROGRAM ELEMENT NO.
Department of Zoology
University of Guelph
Guelph, Ontario NIG 2W1
11. CONTRACT/GRANT NO.
805333
17. SPONSORING AGENCY NAME AND ADDRESS
Environmental Research Laboratory
Office of Research and Development
U.S. Environmental Protection Agency
Duluth, Minnesota 55804
13. TYPE OF REPORT AND PERIOD COVERED
14. SPONSORING AGENCY CODE
EPA/600/03
10. SUPPLEMENTARY NOTES
Large Lakes Research Station, 9311 Groh Road, Grosse lie* Michigan 48138
16. ABSTRACT
Presented here are keys and notes on distribution and ecology for the freshwater
snails (Gastropoda), mussels (Unionidae), and fingernail clams (Sphaeriidae) which
have been collected from the Laurentian Great Lakes. Including subspecies and forms,
121 taxa are discussed: 47 Gastropoda, 39 Unionidae, and 35 Sphaeriidae. Relations
to substrate preferences and pollution are summarized where known. Special emphasis
is given to the sphaeriid genus Px^xuiauTi in discussions of morphological variability
and characters which will separate the species and forms both in the adult and
photomicrographs are presented for all sphaeriid taxa. A limited synonymy has
been compiled from publications and reports on the Laurentian Great Lakes and the
expanded reference section includes not only citations used in the text but also
publications and reports which may aid researchers in more indepth studies of the
fauna.
7.
KEY WORDS AND DOCUMENT ANALYSIS
DESCRIPTORS
b-IDENTIFIERS/OPEN ENDED TERMS C. COSATI Field/Croup
Mollusca, snails
Great Lakes
06C
B. UlSTmHUTION STATtMENT
Release to public
19. SECURITY CLASS (Tim Report)
Unclassified
21. NO. OF PAGES
150
20. SECURITY CLASS (Thitpage)
Unclassified
22. PRICE
EPA Form 2Z20-1 (9-73)
144
U S GOVERNMENT PRINTING OFFICE 1980-657-165/0067
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