vvEPA
United States
Environmental Protection
Agency
An Introduction to
Freshwater Mussels as
Biological Indicators
Including Accounts of Interior Basin,
Cumberlandian, and Atlantic Slope
Species
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EPA-260-R-08-015
November 2008
An Introduction to
Freshwater Mussels as
Biological Indicators
Including Accounts of Interior Basin,
Cumberlandian, and Atlantic Slope Species
Prepared by:
Jeffrey D. Grabarkiewicz1 and Wayne S. Davis2
1Ecological Survey and Design, LLC
1517 W. Temperance Rd.
Temperance, Ml 48182
2U.S. Environmental Protection Agency
Office of Environmental Information
Office of Information Analysis and Access
Washington, DC 20460
U.S. Environmental Protection Agency
Office of Environmental Information
Office of Information Analysis and Access
Washington, DC 20460
Printed on chlorine free 100% recycled paper with
100% post-consumer fiber using vegetable-based ink.
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An Introduction to Freshwater Mussels as Biological Indicators
NOTICE
This document has been reviewed and approved in accordance
with U.S. Environmental Protection Agency policy. Mention
of trade names, products, or services does not convey and
should not be interpreted as conveying official EPA approval,
endorsement, or recommendation for use.
Funding was provided by the U.S. Environmental Protection
Agency under Contract # 68-C-04-006, Work Assignment #4-79
with the Great Lakes Environmental Center, Inc.
The appropriate citation for this report is:
Grabarkiewicz, J. and W. Davis. 2008. An introduction to
freshwater mussels as biological indicators. EPA-260-R-
08-015. U.S. Environmental Protection Agency, Office of
Environmental Information, Washington, DC.
The entire document can be downloaded from:
http://www.epa.gov/bioindicators/html/publications.html
ACKNOWLEDGEMENTS
We would like to thank the many individuals who provided
manuscripts and papers for our review and reference. Thanks
also to the University of Michigan Museum of Zoology and the
Ohio State Museum of Biological Diversity for providing access
to their collections. Last, but certainly not least, thank you to
the reviewers who took the time to look over this document,
including Dr. Tom Augspurger, Dr. Chris Barnhart, Dr. Arthur
Bogan, Dr. Hans Gottgens, Edward Hammer, Tina Hendon, Dr.
Teresa Newton, Dr. Brenda Rashleigh, Brett Rodstrom, and John
Tetzloff.
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An Introduction to Freshwater Mussels as Biological Indicators
CONTENTS
Notice v
Acknowledgements v
Part One - Introduction and Indicator Use
Introduction 1
Distribution and Conservation Status 2
Freshwater Mussel Ecology 4
Freshwater Mussel Reproduction 6
Shell Morphology 8
Sampling Freshwater Mussels 10
Freshwater Mussels as Biological Indicators 15
Tolerance to Habitat Alteration 17
Indicators of Biological Integrity 18
Sensitivity to Toxic Contaminants 20
Heavy metals 21
Ammonia 25
Chlorine 27
Insecticides, Herbicides, and Fungicides 27
Shells as Indicators 29
Part Two - Genus and Species Accounts
Genus Accounts
Alasmidonta Say, 1818 31
Epioblasma Rafinesque, 1831 34
Fusconaia Simpson, 1900 37
Lampsilis Rafinesque, 1820 40
Lasmigona Rafinesque, 1831 44
Pleurobema Rafinesque, 1819 47
Quadrula Rafinesque, 1820 50
Species Accounts
Mucket (Actinonaias ligamentina) 53
Pheasantshell (Actinonaias pectorosa) 54
Dwarf Wedgemussel (Alasmidonta heterodon) 55
Elktoe (Alasmidonta marginata) 56
Threeridge (Amblema plicata) 57
VI
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An Introduction to Freshwater Mussels as Biological Indicators
CONTENTS (CON'T)
Species Accounts*
Purple Wartyback (Cyclonaias tuberculata) 58
Dromedary Pearlymussel (Dromus dromas) 59
Eastern Elliptic (Elliptic complanata) 60
Spike (Elliptic dilatata) 61
Oyster Mussel (Epioblasma capsaeformis) 62
Northern Riffleshell (Epioblasma tomlosa rangiana) 63
Tubercled Blossom (Epioblasma torulosa torulosa) 64
Wabash Pigtoe (Fusconaia flava) 65
Pink Mucket (Lampsilis abrupta) 66
Plain Pocketbook (Lampsilis cardium) 67
Wavyrayed Lampmussel (Lampsilis fasciola) 68
Fat Mucket (Lampsilis siliquoidea) 69
White Heelsplitter (Lasmigona complanata complanata) 70
Flutedshell (Lasmigona costata) 71
Black Sandshell (Ligumia recta) 72
Cumberland Moccasinshell (Medionidus conradicus) 73
Threehorn Wartyback (Obliquaria reflexa) 74
Sheepnose (Plethobasus cyphyus) 75
Clubshell (Pleurobema clava) 76
Ohio Pigtoe (Pleurobema cordatum) 77
Round Pigtoe (Pleurobema sintoxia) 78
Kidneyshell (Ptychobranchus fasciolaris) 79
Fluted Kidneyshell (Ptychobranchus subtentum) 80
Giant Floater (Pyganodon grandis) 81
Rabbitsfoot (Quadrula cylindrica cylindrical 82
Pimpleback (Quadrula pustulosa pustulosa) 83
Mapleleaf (Quadrula quadrula) 84
Creeper (Strophitus undulatus) 85
Pistolgrip (Tritogonia verrucosa) 86
Rain bow (Villosa iris) 87
Conservation Status Table 88
Glossary 89
Literature Cited 90
*Taxonomic Note: To simplify the taxonomy of this guide, all
names follow Turgeon et al. (1998). However, we recognize that
this may not represent the most current taxonomic scheme.
VII
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An Introduction to Freshwater Mussels as Biological Indicators
FIGURES
Figure 1. (Top) Total number of freshwater mussel species by state.
(Bottom) Percentage of imperiled freshwater mussel species by state 2
Figure 2. Freshwater mussel faunal provinces 3
Figure 3. Proportion of species at risk by plant and animal group 3
Figure 4. Various beak sculptures 8
Figure 5. Basic shell anatomy 8
Figure 6. Basic shell orientation 9
Figure 7. Inner soft tissue 9
Figure 8. Hypothetical qualitative bridge survey. 12
Figure 9. A systematic sampling design along 2 transects with 3 random starts 13
Figure 10. Hypothetical survey layout using the ORVET protocol 15
Figure 11. A listing of the most sensitive aquatic genera to copper
excluding freshwater mussels 23
Figure 12. A listing of the most sensitive aquatic genera to copper
including freshwater mussel taxa 23
Figure 13. A listing of the most sensitive aquatic genera to ammonia
excluding freshwater mussel taxa 25
Figure 14. A listing of the most sensitive aquatic genera to ammonia
including freshwater mussel taxa 26
Figure 15. External rings of the Ohio Pigtoe (Pleurobema cordatum) 29
VIM
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An Introduction to Freshwater Mussels as Biological Indicators
PHOTOGRAPHS
Photo 1. A Kidneyshell (Ptychobranchus fasciolaris) repositioning in the substrate 4
Photo 2. Small fishes, such as this Brindled Madtom (Noturus miurus), often take
shelter in the spent shells of freshwater mussels (Photo: Threeridge shell) 5
Photo 3. The Spike (Elliptic dilatata) blanketed in periphyton 5
Photo 4. Water pennies (Psephenidae sp.) grazing on a White Heelsplitter
shell (Lasmigona complanata complanata) 5
Photo 5. The mantle flap lure of the Plain Pocketbook (Lampsilis cardium) 6
Photo 6. The charged gills of the Plain Pocketbook (Lampsilis cardium) 6
Photo 7. A 12 mm Rayed Bean (Villosa fabalis) attached to a small piece
of gravel via byssal threads 7
Photo 8. Clinch River, TN 10
Photo 9. Sampling a small creek with a view-bucket and snorkeling gear 10
Photo 10. Typical diving gear used by biologists searching for mussels 11
Photo 11. Mussel researcher utilizing an underwater viewer 11
Photo 12. Researcher sorting mussels for identification and measurement 11
Photo 13. Unionids collected during a qualitative survey. 12
Photo 14. Excavating sediments within a 0.25 m2 quadrat during a
quantitative survey. 14
Photo 15. Excavating sediments within a 0.25 m2 quadrat during a
quantitative survey 14
Photo 16. The Green River, KY, home to 71 mussel species and 151
fish species 16
Photo 17. The Shelbyville dam, Duck River, TN 17
Photo 18. A researcher measures a mussel with a caliper to collect
demographic data 19
Photo 19. Swan Creek, OH, headwater habitat of the Slippershell
Mussel (Alasmidonta viridis) 31
Photo 20. Slippershell Mussel (Alasmidonta viridis), Swan Creek, OH 31
Photo 21. The Oyster Mussel (Epioblasma capsaeformis), Clinch River, TN 34
Photo 22. The Powell River (TN, VA) was historically home to several
species of Epioblasma, including the extinct Forkshell (Epioblasma lewisii)
and Acornshell (Epioblasma haysiana) 34
Photo 23. Roanoke River, VA, home to the Atlantic Pigtoe (Fusconaia masoni) 37
Photo 24. The Wabash Pigtoe (Fusconaia flava), Swan Creek, OH 38
IX
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An Introduction to Freshwater Mussels as Biological Indicators
PHOTOGRAPHS (CON'T)
Photo 25. Streamline chubs (Erimystax dissimilis) foraging above a
Longsolid (Fusconaia subrotunda), French Creek, PA 38
Photo 26. Cumberland River, KY, home to the Ebonyshell (Fusconaia ebena),
Longsolid (Fusconaia subrotunda), and Wabash Pigtoe (Fusconaia flava) 39
Photo 27. Lake Michigan, home to the Plain Pocketbook (Lampsilis cardium)
and Fatmucket (Lampsilis siliquoidea) 41
Photo 28. In-situ mantle flap lure of the Wavyrayed Lampmussel 43
Photo 29. In-situ apertures of the Plain Pocketbook 43
Photo 30. The Flutedshell (Lasmigona costata) in French Creek, PA 45
Photo 31. Green River, KY, home to the Flutedshell (Lasmigona costata) and
White Heelsplitter (Lasmigona complanata complanata) 46
Photo 32. The Green River (KY), home of the federally endangered
Rough Pigtoe (Pleurobema plenum) 48
Photo 33. The federally endangered Clubshell (Pleurobema clava),
French Creek, PA 48
Photo 34. East Fork West Branch St. Joseph River, Ml, habitat of the
Clubshell (Pleurobema clava) 49
Photo 35. Rabbitsfoot (Quadrula cylindrica cylindrical, French Creek, PA 51
Photo 36. French Creek, PA, habitat of the Rabbitsfoot (Quadrula cylindrica cylindrical 52
All photographs by Jeff Grabarkiewicz and Todd Crail
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An Introduction to Freshwater Mussels as Biological Indicators
INTRODUCTION
While marine environments harbor the delicate beauty of the seahorse, coral reef, and
anemone, North American freshwater streams and lakes support a splendor all their own.
Freshwater mussels, also called pearly mussels, naiads, or clams, are a diverse group of
creatures that are both unassuming and inconspicuous. Most will not confound passersby
with oddities or evoke awe-inspired gasps from onlookers. In fact, many spend a good
deal of their life partially or wholly buried in stream sediments, detectable to only the most
astute observer. Yet, despite their mild manner and cryptic nature, evolution has bestowed
a great variety of these creatures on the North American continent. And we should
consider ourselves fortunate! Not only do these mussels possess a unique elegance and
beauty, but they also exhibit a dizzying array of adaptations and life history strategies.
As a group, freshwater mussels are differentiated from other bivalves by their unique life
cycle. This life cycle includes a parasitic larval stage that requires, in most cases, a fish
host to complete. Adult mussels can measure up to ten inches in length and, under certain
conditions, live more than 100 years (Bauer 1987; Cummings and Mayer 1992).
From an economic perspective, mussels have been valued for their beauty, shell material,
and natural pearls for centuries. Unfortunately, this has also led to the overharvesting
of mussel resources. For example, during the mid-1800s, freshwater mussels were
commonly collected by people seeking fortune in the form of freshwater pearls. Following
a valuable discovery, successive collecting often led to the wholesale destruction of entire
mussel beds (Anthony and Downing 2001). The pearl button industry, founded during the
late 1800s, provides another example of overharvesting leading to resource depletion. This
industry created buttons from the durable shells of freshwater mussels. Thousands of tons
of shells were harvested in Eastern North America from the late 1800s to the mid 1900s to
fuel the button industry. Coker et al. (1921) eloquently observed "equal as they were to the
vicissitudes of natural conditions, they were unable to withstand the unchecked ravages
of commercial fishery". Freshwater mussels continue to be an important economic and
ecological resource (Anthony and Downing 2001, Pritchard 2001).
The primary purpose of this guide is to encourage the use of freshwater mussels for water
quality assessment and to briefly review their use as biological indicators and biomonitors.
This document is not intended as a "how to" manual or methods document, but as an
educational tool. It was designed and written with a wide audience in mind, including
academic institutions, natural resource managers, park naturalists, conservationists,
monitoring groups, environmental managers, and interested citizens. Several topics will
be examined in detail, including ecology, reproduction, indicator use, sensitivity to toxic
contaminants, survey methodologies. In addition, species records are included with
pictures to assist in identification and indicator usage.
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An Introduction to Freshwater Mussels as Biological Indicators
DISTRIBUTION AND CONSERVATION STATUS
Total Number
Freshwater mussels (Bivalvia: Unionoida) are distributed nearly worldwide, inhabiting
every continent on Earth except Antarctica. Approximately 780 species belonging to 140
genera have been identified, with species diversity maximized in the creeks, rivers, and
lakes of North America (Graf and Cummings 2007). Globally, six families of freshwater
mussels are known, although
only two occur in North
America - the Unionidae and
Margaritiferidae. The Unionidae
makeup the vast majority of the
North American fauna. Overall,
approximately 300 species
of mussels are found in the
U.S., with the highest species
richness found in the Southeast
(Fig. 1) (Neves et al. 1997).
Various unionoid faunal
zones have been identified by
malacologists during the past
century (e.g. Simpson 1900;
van der Schalie and van der
Schalie 1950; Parmalee and
Began 1998; Abell et al. 2000).
This guide borrows from the
interpretation introduced by
Parmalee and Bogan (1998)
(Fig. 2). Faunal regions are
useful when attempting to
describe the distribution
patterns and evolutionary
characteristics of freshwater
mussels. For example, some
faunal provinces represent
areas of considerable
endemism, such as the
Ozarkian, Cumberlandian, and
Mobile Basin. Within these
regions, faunal "hotspots" occur
that support unionoid species
found nowhere else on Earth.
Percentage Imperiled
(Source: LaRoe et al. 1995)
Figure 1: (Top) Total number of freshwater mussel species
by state. (Bottom) Percentage of imperiled freshwater mussel
species by state.
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An Introduction to Freshwater Mussels as Biological Indicators
(Source: Parmalee and Bogan (1998), The Freshwater Mussels of Tennessee.
University of Tennessee Press. Permission granted by the authors.)
Northern Atlantic Slope
Southern Atlantic Slope
Interior Basin or
Mississippian
Cumberlandlan
Mobil Basin
Eastern Gulf Coast or
Apalachlcolan
Peninsular Florida
Ozarkian
Sablne
Central Texas
Rio Grande
Pacific
Figure 2: Freshwater mussel faunal provinces.
While historically highly diverse
and abundant throughout a good
portion of the U.S., unionoids are
now one of the most imperiled
groups nationwide (Fig. 3).
Approximately 70% of the North
American fauna is in various
states of decline (Williams et
al. 1993; Master et al. 2000;
NatureServe 2008). Sadly, 37
species are now presumed
extinct (Master et al. 2000). This
decline is often attributed to
habitat destruction, water quality
degradation, damming, exotic
species, and hydrologic changes
(Williams et al. 1993; Strayer et
al. 2004).
Crayfishes
Stoneflies
Freshwater Fishes
Amphibians
Flowering Plants
1 II 51%
QB 43%
1 13
1 1 369
1 1 1 33%
Gymnosperms H
I 24%
Ferns/Fern Allies I •
|| 22%
Tiger Beetles I I
Butterflies/Skippers
Reptiles
Dragonflies/Damselflies I •
Mammals
Birds rZ^^B
0% 10%
(Sou
19% P-, D
I1 19% • Critic
, , .
|| 18% ° lmPE
CH Vuln
HI™
] 16%
14%
%
i
jmed/Poss
allylmper'
riled (G2)
arable (G3)
blyF-xtinct
ed(G1)
69%
(GX/GH)
20% 30% 40% 50% 60% 70%
Percent of Species
ce: Precious Heritage, ©TNC and NatureServe, 2000)
Figure 3: Proportion of species at risk by plant and animal
group.
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An Introduction to Freshwater Mussels as Biological Indicators
FRESHWATER MUSSEL ECOLOGY
Freshwater mussels play a number of important roles in aquatic ecosystems. As sedentary
suspension feeders, unionoids remove a variety of materials from the water column,
including sediment, organic matter, bacteria, and phytoplankton. Siphoned material is
either transferred to the mouth for digestion or sloughs off the gills and exits via the ventral
margin of the shell (pseudofeces). Digested material is either used as fuel for various
life processes or excreted as feces. The amount and rate of particulate matter removed
from the water column and subsequent deposition of waste is largely dependent on
temperature, particle concentration, flow regime, mussel size, and species (Vaughn and
Hakenkamp 2001). While the siphoning activities of mussels are often overlooked, they
provide an integral resource link between pelagic and benthic habitats (Nelepa et al. 1991;
Howard and Cuffey 2006).
Mussels also interact with stream sediments.
The burrowing behavior of unionids mixes
sediment pore water, releasing nutrients and
oxygenating substrates (Photo 1) (Vaughn
and Hakenkamp 2001). Particularly dense
assemblages of mussels may influence
substrate stability and provide nutrients and
microrefugia for benthic life (Vaughn and
Hakenkamp 2001; Zimmerman and de Szalay
2007).
Juvenile mussels have demonstrated the
ability to pedal feed by sweeping their foot
to collect food particles from sediments.
Studies conducted by Gatenby et al. (1996)
documented the importance of sediments
to the growth of juvenile Rainbow (Villosa
iris). Researchers reported increased shell
growth and survival rates when algal diets
were supplemented with a fine sediment
substratum.
Photo 1: A Kidneyshell (Ptychobranchus
fasciolahs) repositioning in the substrate.
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An Introduction to Freshwater Mussels as Biological Indicators
Freshwater mussels also provide food
for a number of terrestrial and aquatic
species. Raccoons, muskrats, otters,
fishes, turtles, and birds all feed on
mussels. The cracked valves and
weathered remains of unionids often litter
gravel bars and floodplains, a testament
to the efficiency of terrestrial predators.
The spent valves of freshwater mussels
play a role in aquatic ecosystems as well.
Shells provide habitat for a variety of
life, including fish (Photo 2), periphyton
(Photo 3), crustaceans, molluscs,
and macroinvertebrates (Photo 4).
Additionally, the weathering and eventual
erosion of shell material recycles calcium
carbonate back to aquatic ecosystems.
Photo 2: Small fishes, such as this Brindled
Madtom (Noturus miurus), often take shelter
in the spent shells of freshwater mussels
(Photo: Threeridge shell).
Photo 3: The Spike (Elliptic dilatata) blanketed
in periphyton.
Photo 4: Water pennies (Psephenidae sp.)
grazing on a White Heelsplitter shell (Lasmigona
complanata complanata).
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An Introduction to Freshwater Mussels as Biological Indicators
FRESHWATER MUSSEL REPRODUCTION
The reproductive characteristics and
processes found among the Unionoidea
are diverse, complex, and more than a
little intriguing. As a group, they exhibit
extraordinary variations in fecundity, brooding
tendencies, host specificity, and "luring"
techniques (Photo 5) (Walters 1995; Haag
and Warren 1999; Haag and Staton 2003;
Haag and Warren 2003).
While sexes are separate in most freshwater
mussels, hermaphroditic species have
been reported (van der Schalie 1970). The
reproductive process is initiated when an
upstream male releases sperm into the water
column and a downstream female collects it
via the incurrent aperture. Fertilization occurs
internally, with embryo development ensuing
within the marsupia (gill pouches) (Photo 6).
The resulting larvae, termed "glochidia," are
brooded by the female for a period of time
ranging from a few weeks to several months.
While some species use all four gills to brood
(e.g. Quadrula), others use only the outer gills
or specialized portions of the outer gills (e.g.
Lampsilis). Once released, glochidia are, for
the most part, obligate parasites that must
find a suitable host or perish.
With few exceptions, freshwater mussels
utilize freshwater and anadromous fishes
as hosts. While some mussel species have
proven capable of parasitizing a wide range
of fish hosts, others are seemingly more
specific. For example, laboratory host studies
Photos 5 and 6: (Top) The mantle flap lure of the
Plain Pocketbook (Lampsilis cardium). (Bottom)
The charged gills of the Plain Pocketbook
(Lampsilis cardium).
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An Introduction to Freshwater Mussels as Biological Indicators
suggest that the Giant Floater (Pyganodon
grandis) may be capable of successfully
transforming on nearly 40 species (Walters
1994; Walters 1995). Conversely, Layzer et al.
(2003) exposed 18 species of fish (from six
families) to the glochidia of the Cumberland
Pigtoe (Pleurobema gibberum) and reported
just two species as suitable hosts. Glochidia
that successfully locate a host species attach to
the fins, skin, or gills. Once attached, glochidia
feed on host tissue and develop anatomical
structures. After a period of time, the glochidia
excyst and drop off into the substrate or attach
to objects with byssal threads (Photo 7).
Photo 7: A 12 mm Rayed Bean (Villosa
fabalis) attached to a small piece of gravel via
byssal threads.
Freshwater mussels utilize a variety of structures and techniques to attract potential
host species. For example, several members of the genus Lampsilis display a mantle
flap lure (Photos 5 and 6) to entice various piscivorous hosts, including species such
the Largemouth Bass, Rock Bass, and Black Crappie. When the mantle lure is struck
by an unsuspecting fish, the female responds by expelling glochidia out of the excurrent
siphon and infecting the potential host. The Fusconaia utilize a markedly different
strategy, packaging glochidia in capsule-like cases termed "conglutinates." After being
ejected into the water column, the conglutinates are fed on by fishes, initiating glochidial
attachment to the gills of potential host fishes. A few species have the ability to produce
"superconglutinates," gelatinous masses that are attached to the female mussel via
a mucus cord. The minnow-like masses "swim" back and forth in the current, and are
presumably preyed upon by species such as the Smallmouth Bass (Haag and Warren
1997).
7
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An Introduction to Freshwater Mussels as Biological Indicators
SHELL MORPHOLOGY
In order to accurately identify freshwater
mussels, some basic knowledge of shell
morphology is required. While some species
have distinctive features that allow for instant
identification, others are much more cryptic. In
addition, young mussels often vary significantly
in shape, thickness, length, color, and inflation
when compared to older individuals. To
further complicate matters, the same species
may vary in appearance from watershed to
watershed, or even within the same watershed.
Perhaps the easiest (and most efficient) way
to become proficient with the identification of
mussels is to visit an established collection.
Many major universities maintain collections
of mollusks that are open to researchers
and interested naturalists by appointment.
Additionally, numerous regional and state
guides are now available at reasonable prices
(e.g. Oesch 1984; Cummings and Mayer 1992;
Parmalee and Bogan 1998; WDNR 2003).
elevated and
heavy ridges
double-looped
ridges
coarse ridges
numerous wavy
ridges •
fine ridges
Figure 4: Various beak sculptures.
Figure 5: Basic shell anatomy.
8
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An Introduction to Freshwater Mussels as Biological Indicators
postero-
dorsal
dorsal
antero-
dorsal
posterior
anterior
postero-
ventral
ventral
antero-
ventral
Figure 6: Basic shell orientation.
Figure 7: Inner soft tissue.
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 8: Clinch River, TN.
SAMPLING FRESHWATER MUSSELS
General Overview
Freshwater mussel sampling designs and techniques are largely dependent on the
resources available, sampling conditions, survey objectives, and prior knowledge of the
target population(s) (Strayer and Smith 2003). Surveys range from informal or qualitative
timed searches to intensive, quantitative designs aimed at providing precise population
estimates.
Survey area size and water depth play an
important role in determining sampling
techniques. For example, small, shallow
streams can often be effectively sampled with
little more than a view-bucket or mask and
snorkel (Photos 9 and 11). However, large
streams, rivers, and lakes usually require
SCUBA gear (Photo 10) or surface-supplied
air (SSA) systems. Biologists and commercial
divers utilize a widerange of SSA systems,
some of which are intended for recreational
purposes while others are designed for
deepwater diving. Hookah compressors
(recreational) are popular when surveying
small streams where the water is clean and
relatively shallow. Conversely, navigational channels often require the use of SCUBA
gear or commercial SSA dive stations. Commercial dive stations usually permit divers to
communicate with a topside dive supervisor as well as each other; thereby adding an extra
degree of comfort and safety to dive operations. Commercial systems may also utilize a
"hardhat," a helmet that completely encapsulates the head, protecting the ears and oral-
Photo 9: Sampling a small creek with a view-
bucket and snorkeling gear.
10
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 10: Typical diving gear used
by biologists searching for mussels.
nasal passages
from contact
with the water
column. This may
be especially
important when
surveying in
contaminated
environments.
Sampling
conditions and
the habits of
freshwater
mussels can
make them difficult to detect. Detection is usually
related to substrate type, species, mussel length,
instream vegetation, and observer proficiency (Smith et
al. 2001 a). Mussel investigators often use their hands
to gently feel or disturb the substrate, a technique
informally termed "noodling." Noodling is effective when
searching for young mussels or species that burrow
deep into the substrate.
Photo 11: Mussel researcher utilizing an
underwater viewer.
Photo 12: Researcher sorting mussels for identification and
measurement.
11
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An Introduction to Freshwater Mussels as Biological Indicators
Qualitative Searches
Qualitative searches are generally performed in a bounded area (such as 30m x 20m cell)
for a finite amount of time. Time is often expressed in person-minutes or person-hours
when multiple investigators survey the same area. Qualitative timed searches usually are
designed to optimize species detection without expending the effort required to derive
population estimates, calculate relative abundances, or detect statistically significant
changes in mussel populations over time. As such, qualitative searches are generally more
efficient for species detection than quadrat-based surveys where sediments are excavated
(Obermeyer 1998; Smith et al. 2001 a;
2001 b; Smith 2006).
Example. Figure 8 provides a
plan view example of a typical
qualitative survey. In this study, a
small area near a bridge is being
surveyed qualitatively by two
malacologists. The survey design
partitions the study area into 15m
x 20m sampling units, which are
delineated onsite using a variety
of markers. The investigators
then survey each cell using
snorkeling gear for 40 person-
minutes, searching for mussels
visually and factually. Assuming a
search efficiency of 1 m2 /minute,
an "effective sampling fraction"
of 0.13 can be calculated (see
Smith et al. 2001 a). Essentially,
the effective sampling fraction is
defined as the percentage of the
cell that is thoroughly searched
for mussels, which is 13% in this
example. To increase the amount
of coverage in the study area,
the cell dimensions could be
altered or the survey time per cell
increased.
Figure 8: Hypothetical qualitative bridge survey.
Photo 13: Unionids collected during a qualitative
survey.
12
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An Introduction to Freshwater Mussels as Biological Indicators
Quantitative Studies
Quantitative studies are performed when population estimates are desired for a particular
area or target population. Unlike qualitative sampling, a "comprehensive" sampling effort
requiring excavation of each sampling unit (typically a 0.25 m2 quadrat) is needed to
ensure mussel detection. Such sampling efforts are usually time intensive and expensive
due to the number of samples required and the need to excavate sediments. Smith
(2006) found that excavation required 3 to 12x more time than surface counts. However,
guidelines have been developed by Smith et al. (2001 a; 2001 b) to limit the amount of
excavation required based on the traits of the target population.
A number of sampling strategies can be utilized independently or in combination to
assess a study area, including random sampling, systematic sampling, double sampling,
and adaptive cluster sampling. For a detailed explanation of probability-based sampling
strategies, refer to Strayer and Smith (2003). Essentially, the goal of the survey designer
is to take enough samples to achieve the desired amount of precision. Where the target
population is abundant, less effort is generally required. Unfortunately, due to the patchy
nature of unionoid populations (even within the limits of a mussel bed) and the low
densities at which imperiled species often occur, a large number of samples is usually
needed.
Example. The objective
of a survey may be to
estimate the population
size of federal
candidate Sheepnose
(Plethobasus cyphyus)
near a bridge site.
From prior knowledge
of the site, the
designer anticipates
finding Sheepnose at
densities of 0.5/m2.
With these data and
a few additional site
attributes (e.g. survey
area), the designer
uses quantitative, systematic sampling techniques (Photos 14 and 15;
Fig. 9) to collect an unbiased sample of the survey area. The number
of quadrats required to achieve the desired amount of precision can
be calculated using the methods presented in Smith et al. (2001 a) or
Strayer and Smith (2003).
i L
•>J
1
3
1 '
T1 T2
2
1
1
2
3
3
meter
1
1
2
2
3
3
Figure 9: A
systematic sampling
design along 2
transects with 3
random starts (see
Strayer and Smith
2003). Generally,
0.25 m2 quadrats
are used for
quantitative work; 1
m2 sampling units
are used here for
simplicity.
13
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 14 (left) and 15 (right): Excavating sediments within a 0.25 m2 quadrat during a quantitative
survey.
Big River Surveys - ORVET Protocol
Protocols have been developed by the Ohio River Valley Ecosystem Team (ORVET
- Mollusk Subgroup) with the intent of providing a consistent and reliable approach to
mussel survey activities in the Ohio River (ORVET 2004). This methodology has also been
applied to large rivers throughout the Midwest and eastern United States. It is a simple,
adaptable, qualitative method that that does not require quadrat-based sampling or
sediment excavation. The following is a summary of ORVET (2004):
The protocol calls for 100 meter transects laid perpendicular to
flow, spaced 100 meters apart. Buffer areas are added to the total
survey area based on the type of planned disturbance or presence of
federal species. Transects are divided into (10) 10 meter segments or
"samples." A diver searches a 1 meter wide path over each sample (10
meter segment) for a minimum search time of 5 minutes. Mussels are
bagged at the end of each 10 meter segment. This protocol assumes
that the diver detects only 50% of the actual mussels present,
therefore densities of 0.5/nf may actually equal 1.0/m2.
Example. Figure 10 provides a plan view example of a typical large
river survey using the ORVET protocol. In this example, transects
are laid according to protocol along the left bank of the river. Divers
would then proceed to search each 10 meter segment for a minimum
of 5 minutes. Mussels are usually identified on a boat by a qualified
malacologist.
14
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An Introduction to Freshwater Mussels as Biological Indicators
Figure 10: Hypothetical survey layout using the
ORVET protocol.
FRESHWATER MUSSELS AS BIOLOGICAL INDICATORS
Freshwater mussels possess several characteristics that make them suitable indicator
organisms (Ortmann 1909; Wurtz 1956; Bedford et al. 1968; Simmons and Reed 1973;
Imlay 1982; Neves 1993; Naimo 1995). These attributes allow individuals or assemblages
to function as "environmental logbooks," effectively recording changes in water and habitat
quality over time. North American unionoids have, in fact, been sounding the alarm for
over the past century. What follows is a short list of qualities that support the use of
mussels as bioindicators:
Unionoidean Attributes
1. Long-lived: some species may reach 70+ years.
2. Sedentary: juvenile and adult mussels move little during their entire
lifetime.
3. Burrowers: some species and juveniles burrow deep into the
streambed.
4. Filter feeders: mussels obtain food and oxygen from the water column
and via interstitial flow.
5. Fairly large: mussels contain ample soft tissue for chemical analysis.
6. Spent valves: dead mussels leave a historical record.
15
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 16: The Green River, KY, home to 71 mussel species and 151 fish species.
Sensitivity to Environmental
Perturbations
1. Unionoids demonstrate a gradient
of tolerances to both chemical
contaminants and physical alterations.
2. Exhibit sensitivity to habitat or
watershed changes that alter flow
regimes, reduce substrate stability, or
cause siltation.
3. Vulnerable to periods of low dissolved
oxygen.
4. Sensitive to exotic species invasions,
such as the zebra mussel (Dreissena
polymorpha).
Sampling and Monitoring
1. Unionoids are widespread throughout
the United States, and are particularly
speciose in the eastern United States.
2. Protocols have been developed to
survey mussels, although sampling
techniques have not been standardized.
3. Freshwater mussels are relatively easy
to tag and monitor.
Biological Indicator
A numerical value(s) derived from actual
measurements, has known statistical
properties, and conveys useful information
for environmental decision making. It can
be a measure, an index of measures, or
a model that characterizes an ecosystem
or one of its critical components (USEPA
2007).
Biological Integrity
The capability of supporting and
maintaining a balanced, integrated,
adapted community of organisms having
a species composition, diversity, and
functional organization comparable to the
natural habitats of the region (Karr and
Dudley 1981).
Biomonitor
An organism that is sensitive to, or is
capable of detecting, changes in the
surrounding environment.
Indicator Organism
An organism whose characteristics are
used to point out the presence or absence
of environmental conditions which cannot
be feasibly measured from other taxa
or the environment as a whole (slightly
modified from Landres et al. 1988).
16
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 17: The Shelbyville dam, Duck River, TN.
Tolerance to Habitat Alterations
From the sinking mud of turbid embayments to the sand and gravel of morainal streams,
the Unionidae and Margaritiferidae can be found in a wide range of habitats throughout
North America. Yet, despite the seemingly ubiquitous distribution of some species, the vast
majority of freshwater mussels thrive in clear, oxygenated streams and rivers where the
bed is comprised of sand, gravel, and cobble substrates.
Of the habitat alterations initiated by humans, the systematic damming of creeks and
rivers has likely had the most profound effect on freshwater mussels (USFWS 1985a;
Bogan 1993; Neves 1993; Yeager 1993). The alteration of shallow, flowing habitats to long,
linear pools has drastically altered the physical, chemical, and biological characteristics
of numerous North American rivers (Ellis 1942; Bates 1962; Coon et al. 1977; Yeager
1993; Hughes and Parmalee 1999). Impoundment not only reworks the depth and
hydraulics of a river reach, but also prevents the migration of host fishes and may
severely alter downstream water quality (Walters 1996c; Vaughn and Taylor 1999; Walters
2000). As a result, mussel species adapted to shallow, flowing rivers are now some
of the most imperiled animals in the United States. The destruction of the Epioblasma
(riffleshells), for example, has been largely attributed to the impoundment of small and
large rivers (USFWS 1983b; USFWS 1985a; USFWS 2004). Nonetheless, diverse
unionid assemblages have persisted under impounded conditions. For example, Haag
and Warren (2006) recently documented a fairly diverse, recruiting unionid community in
an impounded portion of the Little Tallahatchie River (MS) below Sardis Dam. However,
it should be noted that the assemblage consisted of many species known to tolerate
impoundment or lentic conditions.
Sedimentation is another process that may have harmful impacts on freshwater mussel
communities. With the exception of some anodontines, the majority of North American
unionoid mussels occur in coarse substrates and flowing water. Soft, cohesive substrates
17
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An Introduction to Freshwater Mussels as Biological Indicators
and suspended fine sediments are deleterious for most species and may affect respiration,
feeding, and growth (Marking and Bills 1979).
Waterway modifications such as channelization and dredging homogenize habitat,
alter flow regimes, and may increase streambed and bank shear stresses. The physical
straightening and deepening of streams has been associated with the decimation of
mussel communities and changes in faunal composition (Stansbery and Stein 1971;
Walters 1988). In addition, stream "maintenance" is often a reoccurring event, as unstable
streams "silt in" or develop various obstructions. Continuous channel maintenance may
destabilize streams, resulting in shifting, unstable substrates, excessive erosion, and soft
mid-channel bars.
While not frequently named as a primary contributor to the decline of mussels in North
America, landscape alterations such as urbanization have been documented to reduce
stream quality and alter macroinvertebrate communities (Stepenuck et al. 2002; Deacon
et al. 2005). Watershed modifications that increase the volume and change the timing of
stormwater runoff may initiate substrate instability, increase bank erosion, and promote
the siltation of downstream habitats. Strayer (1999) found mussels to be correlated closely
with areas of hydraulic stability, moreso than other habitat features such as depth and
substrate size. While American cities and suburbs continue to expand, stable habitat will
likely become increasingly rare to the detriment of freshwater mussel communities.
Indicators of Biological Integrity
Freshwater mussels are commonly labeled as "good" indicators of biological integrity
and water quality by scientists. Despite this, there remains little guidance available for
monitoring groups or agencies as to how to utilize unionoids in this capacity. A review
of published literature, technical reports, and research does provide some information,
however. For example, Kearns and Karr (1994) used mussels from the genus Epioblasma
and three snail genera as an intolerant metric when developing a B-IBI (Benthic Index of
Biotic Integrity) for the Tennessee Valley. Pip (2006) analyzed water quality and mollusk
communities in southern Lake Winnipeg, Manitoba, Canada. Freshwater mussel species
richness was positively correlated with total dissolved solids and negatively correlated with
lead. She also found a significant reduction in mussel species diversity and suggested the
change was possibly due to oxygen depletion, algal toxins, sewage and agricultural spills
and runoff, application of copper sulphate, and habitat changes. Hoggarth and Goodman
(2007) utilized a multimetric Mussel Index of Biotic Integrity developed by Goodman (2007)
to evaluate changes in the mussel fauna of the Little Miami River, OH. The Index included
metrics that assessed distribution and abundance, reproductive potential, and community
structure.
18
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An Introduction to Freshwater Mussels as Biological Indicators
While freshwater mussels do present unique challenges to monitoring (e.g. patchy
distributions, sampling conditions, etc.), the information gained from their study can
provide unique insights regarding biological integrity and water quality. Although
developing complex multimetric mussel indices is beyond the scope of this guide, below
are a few basic recommendations on monitoring techniques that may provide valuable
information when assessing local waterways.
Community Demographics
Characterizing community demographics over time may yield important data on
species viability, conditions conducive to reproductive success, water quality, and
ecosystem stability.
When monitoring community demographics, multiple stations with known mussel
populations should be setup within the desired study area. If possible, monitoring
should be done quantitatively to limit sample bias.
To evaluate community age class structure, a malacologist typically uses a metric
caliper and measures live individuals in one (anterior to posterior) or three dimensions.
Basic summary statistics can be generated to analyze shell length diversity, age class
heterogeneity, and recent recruitment. This may be especially interesting when the
collected data is contrasted with "ideal" study sites that maintain a "healthy" cross
section of age class diversity. For example, Grabarkiewicz and Crail (2008) used
simple summary statistics to compare age class diversity in three different waterways.
This method might be further developed as a metric in a full multimetric index.
Mark and Recapture
Mark and recapture is a method often
used when translocating mussels
from one area to another. With this
technique, mussels are first marked
or given a unique tag and moved to
a defined area up or downstream.
Mark and recapture may also prove
useful in water quality assessment. For
example, with a unique tag the health
and growth of multiple individuals may
be monitored over time by measuring
shell dimensions, weight, or biological
markers. This type of monitoring
may assist when assessing lethal or
sublethal impacts of nearby industrial
or sewage outfalls.
Photo 18: A researcher measures a mussel with a
caliper to collect demographic data.
19
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An Introduction to Freshwater Mussels as Biological Indicators
Long Term Quantitative Monitoring
Quantitative sampling generally involves the excavation of quadrats and the sieving
of sediments. Long-term quantitative monitoring is useful when assessing trends of
community diversity, abundance, and overall system integrity. For example, Ahlstedt
and Tuberville (1997) reported trends in species composition, abundance, and
recruitment between 1979 and 1994 in the Clinch and Powell River (TN, VA).The
researchers reported that a severe drought had a strong impact on mussel populations
during the mid-1980s and that mussel distribution and abundance patterns were
"influenced by proximity of mined land." They also noted that declines in tributary
streams were generally more severe for freshwater mussels than fish, suggesting that
mussels are more sensitive to environmental perturbations.
Sensitivity to Toxic Contaminants
Toxic contaminants have long been implicated in the reduction or extirpation of mussel
populations throughout the country (Lewis 1868; Ortmann 1909; Clark and Wilson 1912;
Baker 1928). Early 20th century accounts of industrial stream pollution tell of dyestuff
discharges from knitting mills "causing widespread destruction" (Clark and Wilson 1912)
and rivers "acting as the sole receptacle of sewage and manufacturing waste" (Howe
1900). More recently, regional and continent-wide assessments of unionoid populations
have cited toxic contaminants as a contributor to widespread faunal declines (Havlik and
Marking 1987; Bogan 1993; Neves et al. 1997).
Toxic contaminants are defined here as inorganic or organic contaminants that have the
potential to be lethal or elicit sublethal responses from a chosen study population. The
concentration and exposure required to elicit a response from a particular species may
vary greatly from pollutant to pollutant. Additionally, the toxicity of a particular pollutant
may be influenced by a number of variables, including concentration and exposure route,
frequency, and duration.
Freshwater mussels exhibit a variety of sensitivities to toxic contaminants based on
species, life stage (glochidium, juvenile, or adult), and environmental conditions. For
example, Wang et al. (2007a) reported that glochidial Oyster Mussel (Epioblasma
capsaeformis) and Scaleshell (Leptodea leptodon) were far more sensitive to copper than
glochidial Dwarf Wedgemussel (Alasmidonta heterodon). Interestingly, all three species
are listed as federally endangered by the U.S. Fish and Wildlife Service. Interspecific
disparities like these can be helpful to bioassessment programs in compiling and
diagnosing sources and causes of impairment.
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An Introduction to Freshwater Mussels as Biological Indicators
Heavy Metals
The American Society for Testing and Materials (ASTM 2006) issued new guidelines
for freshwater mussel toxicity testing. The studies included in this document meet
the requirements of the ASTM guidance. Heavy metals are a concern to freshwater
mussels due to their ability to cause mortality, disrupt enzyme efficiency, alter filtration
rate, reduce growth, and change behavior (Naimo 1995). Because freshwater mussels
exhibit suspension and deposit feeding behaviors (Gatenby et al. 1996; Raikow and
Hamilton 2001), heavy metals may be available to unionoids through both the water
column (dissolved or attached to suspended particles) and streambed sediments
(Naimo 1995). However, the overall bioavailability of metals is complicated and
influenced by a suite of factors, including metal concentration and speciation, water
column chemistry, redox potential, particulate matter, and flow regime characteristics
(Luoma 1989).
While it is well-known that freshwater mussels readily bioaccumulate metals, the rate
and location(s) of accrual may vary greatly by unionoid species, unionoid size, and
heavy metal species (Adams et al. 1981; Naimo et al. 1992b; Pip 1995). For example,
studies have suggested that metals such as zinc accumulate most readily in the gills
(Adams et al. 1981), while others such as cadmium become concentrated in the heart
(Pip 1995). In addition to the heart and gills, metal accumulation may occur in the
kidney, digestive gland, foot, and mantle.
Before discussing heavy metals in greater detail, some perspective on the background
levels at which they occur may be helpful. The following was modified from the
excellent review done by Naimo (1995):
Metal
Cd
Cu
Hg
Zn
Locality
Rhone River, France1
Mississippi River, USA2
Lake Vanda, Antartica3
Rhone River, France1
Mississippi River, USA2
Lake Vanda, Antartica3
Silver Lake, CA5
Clear Lake, CA5
Onondaga Lake, NY6
Ohio River, USA7
Lake Erie, USA8
Yangtze River, China7
Condition/Area
Industrial
At mouth
Pristine
Industrial
At mouth
Pristine
Pristine
Polluted
Polluted
Industrial
-
At mouth
Total*
20-117
-
10-70
405-1340
-
400-600
0.6
3.6-104
7-19
-
-
-
Dissolved*
17-80
8-16
-
119-1240
1810-1960
400-600
0.4
1.1-1.5
2-10
288-3203
26-55
39-78
*in units of ng/L
References: 1Huynh-Ngoe etal. 1988a 2Trefryetal. 1986 3Greenetal. 1986 4Huynh-Ngoe et al. 1988b
5Gill and Bruland 1990 6Bloom and Effler 1990 7Shiller and Boyle 1985 8Coale and Flegal 1989
21
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An Introduction to Freshwater Mussels as Biological Indicators
Cadmium. Cadmium is a common
pollutant found in mine drainage, industrial
discharges, insecticides, fungicides, and
urban runoff. Generally considered highly
toxic to aquatic life (Eisler 1985), the
effects of cadmium have been evaluated
by mussel toxicologists under both
laboratory and "semi-field" conditions
(Jenner et al. 1991; Lassee 1991;
Hansten et al. 1996). Jenner et al. (1991)
described the accumulation patterns in
adult Painter's Mussel (Unio pictorum) as
follows:
"....the process of concentration
(accumulation) to high levels
is rapid without direct toxic
(lethal) effects. Detoxification
by metal-binding proteins will
require energy and resources
which are drawn from the
energy and nitrogen (protein)
pool of the animal. A delayed
effect will occur in which the
population size diminishes due
to the constant drain of energy
and protein, causing energy
exhaustion."
AWQC Ambient Water Quality Criteria
Quantitative concentrations or
qualitative assessments of the levels
of pollutants in water which, if not
exceeded, will generally ensure
adequate water quality for a specified
water use (U.S. EPA 2000).
GMAV Genus Mean Acute Value
The geometric mean of the SMAVs for
the genus.
SMAV Species Mean Acute Value
The geometric mean of the results
of all acceptable flow-through acute
toxicity tests with the most sensitive
tested life stage of the species.
LC50 Lethal Concentration (50%)
The concentration of a toxicant within
a medium that causes the death of
50% of a population.
EC50 Effective Concentration (50%)
The dose at which a defined non-lethal
response occurs in at least 50% of the
study population.
Lasee (1991) and Naimo et al. (1992) also investigated the chronic effects of cadmium
exposure. Lasee (1991) documented the dissolution of the crystalline style, an
anatomical structure that assists in food digestion, when mussels were exposed
to mercury, cadmium, and copper. Naimo et al. (1992a) studied the physiological
responses of adult Plain Pocketbook (Lampsilis cardium) to cadmium concentrations
ranging from 22-305 ug Cd/L. While most physiological responses were widely variable,
respiration rates significantly decreased with increased cadmium exposure.
Copper. Elevated levels of copper in the aquatic environment may come from a variety
of sources, including mine drainage, coal ash effluent, industrial discharges, and urban
runoff. Recent studies and data compilations suggest that freshwater mussels are
among the most sensitive aquatic taxa to copper (Figs. 11 and 12). It should also be
noted that a significant amount of variability has been observed among the individual
responses unionids exhibit to copper (Milam et al. 2005).
22
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An Introduction to Freshwater Mussels as Biological Indicators
Ranked GMAVs for Copper (1996 AWQC Update)
10 most sensitive taxa in the dataset
Figure 1 1 : A listing of the most sensitive aquatic genera to copper excluding
freshwater mussels (Augspurger et al. 2006). Used with permission.
Ranked GMAVs for Copper (1996 AWQC Update)
adding data for mussel genera (shaded)
rdness
8 §
3
* *
* *
****
**
* * *
Figure 12: A listing of the most sensitive aquatic genera to copper including freshwater
mussel taxa (Augspurger et al. 2006). Used with permission.
The toxicity of copper to freshwater mussels has been investigated by several
researchers (e.g. Jacobson et al. 1993; 1997; Cherry et al. 2002; Milam et al. 2005;
Wang et al. 2007a; Wang et al. 2007b).
During 24-hour exposures, Jacobson et al. (1993) calculated EC50s (valve closure) as
low as 33 ug Cu/L and 27 ug Cu/L for juvenile Giant Floater (Pyganodon grandis) and
Rainbow (Villosa iris), respectively. Reported LC50s ranged from 44 ug Cu/L (hardness
= 70 mg/L) to 83 ug Cu/L (hardness = 190 mg/L) for Giant Floater (P. grandis) and
Rainbow (V. iris), respectively.
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An Introduction to Freshwater Mussels as Biological Indicators
Jacobson et al. (1997) further investigated the toxicity of copper to the early life
stages of freshwater mussels by assessing exposures to brooded, released, and
encapsulated glochidia. Brooded and encapsulated glochidia exhibited little sensitivity
to copper exposures, while calculated LC50s for released glochidia ranged from 26 to
347 ug Cu/L (hardness = 55-190 mg/l).
Wang et al. (2007a; 2007b) evaluated the acute and chronic toxicity of copper to the
early life stages (glochidia and juveniles) of 11 unionid species. Reported EC50s
for glochidia varied widely between species, ranging from 10 to >100 ug Cu/L
during 24-hour exposures. The most sensitive species to acute exposures of copper
included Wavyrayed Lampmussel (Lampsilis fasciola), Oyster Mussel (Epioblasma
capsaeformis), Ellipse (Venustaconcha ellipsiformis), Scaleshell (Leptodea leptodon),
and Pink Papershell (Potamilus ohiensis). As a result of their study, Wang et al. (2007a)
suggested that the U.S. EPA 1996 acute WQC for copper may not be protective of the
early life stages of freshwater mussels.
March et al. (2007) evaluated toxicity data in the derivation of water quality guidance
and standards for copper. Freshwater mussel SMAVs were generally similar to
the more sensitive species in the U.S. EPA database. The Ellipse (Venustaconcha
ellipsiformis), Oyster Mussel (E. capsaeformis), and Pink Papershell (Potamilus
ohiensis) were among the most sensitive aquatic species to copper. On the basis
of established ASTM standards, in addition to historical and ongoing research, the
researchers advocated that state and federal agencies consider using freshwater
mussel toxicity data in determining water quality standards.
Zinc. Zinc is commonly discharged into surface waters as a result of mining activities,
industrial processes, and urban runoff. While studies suggest zinc is not as acutely
toxic to freshwater mussels as copper or cadmium (McCann 1993), it may accumulate
to high concentrations in surface waters impacted by mining waste or industrialization
(Adams et al. 1981). At high concentrations, zinc may elicit sublethal responses or
cause mortality.
Results reported by McCann (1993) for a pair of Cumberlandian species and the
Rainbow (Villosa iris), with LC50s ranging from 274 to 1230 ug Zn/L during 48-hour
exposures (hardness = 40-160). Likewise, Cherry et al. (1991) documented LC50s
ranging from 212 to 656 ug Zn/L during 48-hour exposures to four species of unionids
(hardness = 170).
Mercury. Mercury contamination in surface waters may result from pesticides,
hazardous waste disposal, waste incineration, and fossil fuel combustion. Mercury is a
concern in freshwater ecosystems due to its ability to biomagnify up the food chain and
cause various health problems in humans and wildlife.
24
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An Introduction to Freshwater Mussels as Biological Indicators
Valenti et al. (2005) evaluated the acute and chronic toxicity of mercury to the early
life stages V. iris. A glochidial LC50 of >107 ug Hg/L was reported during 24-hour
exposures. A juvenile LC50 of 99 ug Hg/L was reported during 96-hour exposures.
Glochidial Rainbow were determined to be more acutely sensitive to mercury than two
month old juveniles.
Valenti et al. (2006b) also assessed the acute toxicity of inorganic and organic mercury
salts to the glochidia of four freshwater mussel species. Included as test subjects were
two federally endangered species, the oyster mussel (Epioblasma capsaeformis) and
Cumberlandian combshell (Epioblasma brevidens). Reported LC50s ranged from 25 to
54 ug HgCI/L during 24-hour exposures to mercuric chloride (HgCI2). Methylmercuric
chloride (CH3CIHg) was more acutely toxic to E. capsaeformis and E. brevidens, with
reported LC50s of 21 to 26 ug/L during 24-hour exposures. The Rainbow (V. iris) was
found to be far more tolerant to methylmercuric chloride than both E. capsaeformis and
E. brevidens, with less than 50% mortality observed at 120 ug Hg/L after 24 hours.
Ammonia
Anthropogenic sources of ammonia include livestock waste, sewage treatment plants,
faulty septic systems, and industrial wastewater. Like copper, recent toxicity studies
have suggested that freshwater mussels are particularly sensitive to ammonia (e.g.
Goudreau et al. 1993; Bartsch et al. 2003; Augspurger et al. 2003; 2006; Newton et al.
2003; Wang et al. 2007a; 2007b) (Figs. 13 and 14).
Ranked GMAVs for Total Ammonia
(10 most sensitive taxa in the 1985 water quality criteria dataset)
30
25
20
15
(9
**
Figure 13: A listing of the most sensitive aquatic genera to ammonia excluding freshwater mussel
taxa (Augspurger et al. 2006). Used with permission.
25
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An Introduction to Freshwater Mussels as Biological Indicators
Ranked GMAVs for Total Ammonia
[10 most sensitive taxa in the 1985 water quality criteria
dataset adding data for freshwater mussels (shaded)]
s- 30
S 25
* 20
I 15
I 10
_j**_
* *
///S'///////////'/
Figure 14: A listing of the most sensitive aquatic genera to ammonia including freshwater mussel taxa
(Augspurger et al. 2006). Used with permission.
Goudreau et al. (1993) assessed the toxicity of ammonia and monochloroamine
(MCH) to Rainbow glochidia and investigated the effects of wastewater treatment plant
(WWTP) effluents on freshwater mussels. In laboratory toxicity studies, V. iris glochidia
were exposed to ammonia and MCH for 24 hours. Reported EC50s were 0.237 and
0.042 mg/L for unionized ammonia and MCH, respectively. Reported LC50s were
0.284 and 0.084 mg/L for unionized ammonia and MCH, respectively. In field studies of
treatment plant effluents, researchers examined unionid communities above and below
a pair of WWTPs in the upper Clinch River, VA. River segments directly downstream
(up to 3.7 km) of both WWTPs were devoid of mussels, while live unionids were found
above each plant. Researchers suggested that even if glochidia were not killed outright
by MCH or ammonia, the sublethal impacts may reduce glochidial viability and prevent
the colonization of river segments near WWTPs.
Newton et al. (2003) and Newton and Bartsch (2007) studied the effects of ammonia
in sediments and water-only exposures to juvenile Plain Pocketbook (L cardium)
and Higgins' Eye (Lampsilis higginsii). Newton et al. (2003) documented mortality at
concentrations as low as 93 ug NH3-N/L and growth reduction at 31 ug NH3-N/L. These
results are at or below acute national water quality criteria. Researchers also noted
that control survival was much higher when compared with assays where sediments
were not part of the study. Because juveniles may collect food particles from sediments
by pedal feeding (Yaeger and Cherry 1994), this observation further accentuated the
need to examine sediments and pore water as an important exposure route.
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An Introduction to Freshwater Mussels as Biological Indicators
Wang et al. (2007a) examined the acute toxicity of ammonia to the early life stages
of several unionid species. Glochidial mussels exhibited an array of tolerances, with
reported EC50s ranging from 5 to >16 mg N /L during 24-hour exposures. Glochidia
of the Oyster Mussel (E. capsaeformis) and Ellipse (V. ellipsiformis) were among the
most sensitive species tested. During 4-day and 10-day exposures to juvenile mussels,
reported EC50s ranged from 5.7 to 11 mg N/L and 1.7 to 4.5 mg N/L, respectively. The
authors concluded that the 1999 acute WQC for ammonia many not be protective of
the freshwater mussels species tested.
Chlorine
Anthropogenic sources of chlorine include wastewater treatment plants and industrial
facilities (Valenti 2006a). Chlorine is generally considered highly toxic to most forms of
life and is commonly used as a disinfectant.
Valenti et al. (2006a) evaluated the toxicity of total residual chlorine (TRC) to the early
life stages of five unionid species, including three federally endangered taxa: the
Dwarf Wedgemussel (Alasmidonta heterodon), Cumberland Combshell (E. brevidens),
and Oyster Mussel (E. capsaeformis). Mean LC50s ranged from 70 to 220 ug TRC/L
during 24-hour exposures. Federally endangered mussels were found to be slightly
to far more sensitive than Wavyrayed Lampmussel (L. fasciola) and Rainbow (V. iris),
respectively. The study also reported reduced growth in juvenile E. capsaeformis at
concentrations as low as 20 ug TRC/L during 21 -day chronic exposures. The authors
suggested that while endpoints were above U.S. EPA WQC for TRC, potential sublethal
effects to federally endangered juvenile mussels were still a concern.
Wang et al. (2007a) evaluated the acute toxicity of chlorine to the early life stages
of several unionid species. Reported EC50s for mussel glochidia during 24-hour
exposures ranged from 58 to >100 ug TRC/L. Reported EC50s for juvenile mussels
during 4-day and 10-day exposures ranged from 68 to >100 ug TRC/L and 16 to >100
ug TRC/L, respectively. Researchers suggested that the early life stages of mussels
were relatively tolerant of chlorine.
Insecticides, Herbicides, and Fungicides
Insecticides, herbicides, and fungicides are common contaminants in both rural
and urban settings. In fact, scientists estimate that approximately 1.1 billion pounds
of pesticides are spread in the United States annually (Aspelin 1994). Freshwater
mussels exhibit an array of tolerances to current-use pesticides, largely dependent on
the species of mussel, identity of the contaminant, and length of exposure.
Recent research has provided new insights into the acute effects of various
insecticides, herbicides, and fungicides (e.g. Keller 1993; Moulton et al. 1996; Keller
and Ruessler 1997; Milam et al. 2005; Bringolf et al. 2007a). Keller (1993) evaluated
27
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An Introduction to Freshwater Mussels as Biological Indicators
the toxicity of several organic compounds to Paper Pondshell (U. imbecillis) and
contrasted the results with Daphnia (Daphnia magna) and Bluegill Sunfish (Lepomis
macrochirus}. U. imbecillis was generally less sensitive to most contaminants when
compared with other test organisms, including the pesticides toxaphene, chlordane,
and PCP (pentachlorophenol).
Keller and Ruessler (1997) evaluated the toxicity of malathion, a commonly used
mosquito and fruit fly insecticide, to the early life stages of several unionid species.
Glochidia trials yielded LC50s ranging from 7 to 374 mg/L during exposures of 4 to 48
hours. Glochidial Paper Pondshell were by far the most tolerant species, with reported
LC50s of 324 to 374 mg/L. Reported LC50s for juvenile mussels ranged from 74 to
129 mg/L during exposures of 96 hours in hard water. Researchers concluded that
"expected environmental concentrations should not be lethal to unionids."
Bringolf et al. (2007a) assessed the toxicities of various current-use pesticides to the
glochidia and juveniles of several freshwater mussel species. Fatmucket (Lampsilis
siliquoidea) glochidia and juveniles were found to be highly sensitive to chlorothalonil,
propiconazole, and pyraclostrobin, with reported glochidial EC50s ranging from 0.09
to 20.75 mg/L during 24-hour exposures. Juvenile 96-hour EC50s ranged from 0.03 to
10.01 mg/L. Technical grade atrazine, permethrin, fipronil, and pendimethalin were not
acutely toxic to the unionids tested. However, chronic studies found juvenile Fatmucket
to be sensitive to atrazine at low concentrations, with reported EC50s of 15.8 mg/L and
4.3 mg/L during 14-day and 21-day exposures, respectively.
Glyphosate is one of the most widely used herbicides today, yet few studies have
analyzed its effects on freshwater mussels. Bringolf et al. (2007b) investigated the
toxicity of several forms of glyphosate, its formulations, and a surfactant (MON 0818) to
juvenile and glochidial Fatmucket (L. siliquoidea). Reported 24-hour EC50s were as low
as 3.0 mg/L and 0.6 mg/L for Roundup and MON 0818, respectively. Reported 96-hour
EC50s for juvenile L. siliquoidea were 5.9 mg/L and 3.8 mg/L for Roundup and MON
0818, respectively. Researchers concluded that the early life stages of the Fatmucket
are among the most sensitive aquatic organisms to glysphosate-based chemicals and
MON 0818 tested to date.
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An Introduction to Freshwater Mussels as Biological Indicators
Shells as Indicators
The use of freshwater mussel shells as indicators of ecological integrity and environmental
stress has been informally exercised by scientists since the early 1900s (Ortman
1909; Coker et al. 1921). However, only recently have researchers started to collect
quantitative information from shell material (Imlay 1982; Ravera et al. 2005; Brown et al.
2005). Because spent valves persist in aquatic ecosystems for decades or more, shell-
based studies often offer information inaccessible to investigators through traditional
bioassessment strategies.
Mussel shells are comprised of five primary
layers: the periostracum, prismatic layer,
peripheral layer, laminar layer, and inner nacreous
layer (Imlay 1982). The periostracum is mainly
proteinaceous in nature, while the other four
layers are comprised of calcium carbonate, in the
form of calcite or aragonite. Periods of rest are
delineated by internal and external rings (Fig. 15)
laid down during periods of latency. These rings
are often used to age mussels, with each well
defined line constituting a rest period during cold
weather. In addition to growth rings, disturbance
rings may also be present, possibly reflecting
periods of pollution, drought, displacement,
or handling. While aging studies have utilized
both internal and external annuli, some debate
remains in regards to which is more accurate
(Metcalfe-Smith and Green 1992). Uncertainty
is often due to shell weathering, the presence
of disturbance rings, and ring crowding resulting
from old age (Ray 1977; Strayer 1981; Anthony
et al. 2001). Although aging remains a precarious
endeavor with certain specimens, the information
gleaned from growth and disturbance rings can
provide useful insights into historical growth rates,
disturbance, and water quality (Imlay 1982; Haag
2007}. Figure 15: External rings of the Ohio Pigtoe
(Pleurobema cordatum).
Metals may be present in shell material as a result
of surface adsorption or as metabolic analogues of calcium. The metal content of shell
material often varies greatly from what is found in soft tissues. For example, Anderson
(1977) found overall metal concentrations to be higher in soft tissues than shell material
during a study of the Fox River (IL, Wl). Zinc, in particular, was reportedly accumulated
29
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An Introduction to Freshwater Mussels as Biological Indicators
to levels 10-40 times the concentration found in shell material. Ravera et al. (2003) found
shells to contain higher concentrations of Ca, Cr, Mn, Ni, and Mo than soft tissues,
while concentrations of As, Cd, Cu, Ni, and Pb were lower in shells than soft tissues.
Considerable variation was also observed in heavy metal concentrations between different
species.
Chatters et al. (1995) utilized the valves of freshwater mussels to recreate ancient stream
environments in the Columbia River basin (western North America). By analyzing the
archaeological presence of Western Ridged Pearlshell (Gonidea angulata) and Western
Pearlshell (Margaritifera falcata), two species with markedly different habitat requirements,
scientists inferred the crude substrate composition and suspended sediments of historical
stream systems. In addition, researchers analyzed growth increments of Western
Pearlshell to determine historical temperature patterns. The report concluded that the
study area was likely poor for salmon 6,000-7,000 B.P, due mainly to higher stream
temperatures, greater quantities of fine sediments, and lower flows.
Perhaps one of the most interesting application of shell-based strategies is the
examination of heavy metal trends over long periods of time. For example, Brown et
al. (2005) found freshwater mussel shells of the North Fork Holston River to provide
an otherwise unavailable record of mercury contamination at five sites near Saltville,
VA. Through analysis of over 350 shells, researchers verified significant differences in
mercury concentrations between shell assemblages above, within, and below an area of
contamination.
Similarly, Ravera et al. (2005) analyzed shell material from a pair of Italian lakes to
document changes in metal concentrations over two distinct time periods. Using recently
collected shells and preserved valves from a museum, researchers were able to analyze
metal concentrations from 1928-1934 and 1995-2000. Several metals significantly differed
in concentration between the two periods, which also varied greatly between the two lakes.
30
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An Introduction to Freshwater Mussels as Biological Indicators
GENUS PROFILE - ALASMIDONTA
Photo 19: Swan Creek, OH, headwater habitat of the Slippershell Mussel (Alasmidonta viridis).
Species Diversity and Conservation: A total of 12 species have been assigned to the
genus Alasmidonta (Turgeon et al. 1998). Williams et al. (1993) reviewed the conservation
status of unionoids in North America and reported three Alasmidonta as special concern,
two as threatened, and six species as endangered. Currently, three species are listed as
endangered by the U.S. Fish and Wildlife Service (USFWS 2007), while three species are
considered extinct (Turgeon et al. 1998).
Shell Characteristics: Shell rhomboidal,
elongate, or subquadrate; often quite thin when
young, becoming moderately stout with age.
Maximum shell length 2.0 to 4.5 inches (50-
115 mm). Periostracum light yellowish-green
to yellowish tan, often with green rays. Some
species with green spots. Beak sculpture of
elevated and heavy double-looped ridges,
concentric loops, or double-looped bars.
Pseduocardinal teeth well developed but not
massive; generally small. Lateral teeth poorly
developed. Nacre white, often with salmon
tinge.
Photo 20: Slippershell Mussel (A. viridis), Swan
Creek, OH.
Habitat: The Alasmidonta are a wideranging group, inhabiting the lower and upper
Mississippi River basin, Great Lakes-St. Lawrence basin, Atlantic Slope, and Gulf
drainages. Several species are endemics and have limited distributions (e.g. Cumberland
River drainage, Altamaha River drainage, etc.). This group may be found in both
headwaters and large rivers in sluggish or swift current. The Alasmidonta may occur in
a variety of substrates, from mud to sand, gravel, and cobble (Gordon and Layzer 1989;
Parmalee and Bogan 1998).
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An Introduction to Freshwater Mussels as Biological Indicators
Reproduction: Researchers have identified suitable host fishes for many of the
Alasmidonta. Confirmed hosts belong to one of six families: Cyprinidae (minnows),
Catostomidae (suckers), Ictaluridae (catfishes), Cottidae (sculpins), Centrarchidae
(sunfishes), and Percidae (perches) (Howard and Anson 1923; Gordon and Layzer 1993;
Moorman and Gordon 1993; Michaelson and Neves 1995; Schulz and Marbain 1998;
Walters et al. 1998c). Many of the Alasmidonta have been documented as bradytictic
(Ortmann 1921; Baker 1928).
Tolerance to Habitat Alteration: In
general, the Alasmidonta are sensitive
to alterations that change habitat
dynamics. Walters (1995) noted that the
Elktoe (A. marginata) and Slippershell
Mussel (A. viridis) do not tolerate
impoundment. Several members of the
genus have experienced population
declines rangewide due to dams and
associated hypolimnetic discharges,
coal mine runoff, sedimentation, and
pollutants (Oesch 1984; Master 1986;
USFWS 1993b; Parmalee and Bogan
1998; USFWS 2004; Natureserve).
Taxa List (Turgeon et al. 1998)
Altamaha Arcmussel (Alasmidonta arcula)
Cumberland Elktoe (Alasmidonta atropurpurea)
Dwarf Wedgemussel (Alasmidonta heterodon)
Elktoe (Alasmidonta marginata)
Coosa Elktoe (Alasmidonta mccordi)
Appalachian Elktoe (Alasmidonta raveneliana)
Carolina Elktoe (Alasmidonta robusta)
Southern Elktoe (Alasmidonta triangulata)
Triangle Floater (Alasmidonta undulata)
Brook Floater (Alasmidonta varicosa)
Slippershell Mussel (Alasmidonta viridis)
Ochlockonee Arcmussel (Alasmidonta wrightiana)
Sensitivity to Toxic Contaminants: Recently, Wang et al. (2007a) evaluated the acute
toxicity of copper, ammonia, and chlorine to the federally endangered Dwarf Wedgemussel
(Alasmidonta heterodon) and 10 other species. The glochidia of A. heterodon were among
the most tolerant species tested to copper and ammonia. Reported 24-hour EC50s were
>100 ug/L and >16 mg N/L for copper and ammonia, respectively.
Keller and Augspurger (2005) evaluated the toxicity of fluoride to the federally endangered
Appalachian Elktoe (Alasmidonta raveneliana) in response to concerns regarding feldspar
mining operations. The reported 24-hour LC50 for glochidial A. raveneliana was 288 mg F/
L. The reported 96-hour LC50 for juvenile Appalachian Elktoe was 303 mg F/L. The study
concluded that acute fluoride toxicity was unlikely although longer tests would be helpful to
determine sublethal effects.
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An Introduction to Freshwater Mussels as Biological Indicators
Indicator Use: Bedford et al. (1968) utilized
A. marginata as a biomonitor of pesticides
in the Red Cedar River (Ml). Reported
concentrations ranged from 0.0153-0.198
ppm total DDT and metabolites. The study
concluded that mussels were excellent
indicators of pesticide contamination; mainly
because of their ability to concentrate
contaminants.
The Alasmidonta are valuable indicators
of habitat and water quality, as they
generally inhabit clear, good quality, flowing
habitats with stable substrates (Walters
1995; Nedeau et al. 2000; Bogan 2002;
N at u reserve).
Turgeon et al. 1 998 Taxa
Presumed Extinct Taxa
Federally Listed Taxa
Williams et al. 1993
Special Concern
Threatened
Endangered
Total
Natu reserve
G5
G4-G5
G4
G3
G2
G1-G2
G1
GH
GX
12
3 (25.0%)
3 (25.0%)
3 (25.0%)
2(16.7%)
6 (50.0%)
11 (91.7%)
0 (0.0%)
1 (8.3%)
2(16.7%)
1 (8.3%)
1 (8.3%)
2(16.7%)
3 (25.0%)
1 (8.3%)
1 (8.3%)
33
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An Introduction to Freshwater Mussels as Biological Indicators
GENUS PROFILE - EPIOBLASMA
Species Diversity and Conservation:
A total of 17 species and eight
subspecies comprise the genus
Epioblasma (Turgeon et al. 1998), 14 of
which are considered extinct (Hoggarth
et al. 1995; Turgeon et al. 1998). The
remaining 11 Epioblasma are classified
as vulnerable, highly imperiled, or
critically imperiled (Natureserve). A total
of 14 taxa are listed as endangered
by the U.S. Fish and Wildlife Service
(USFWS 2007).
Photo 21: The Oyster Mussel (Epioblasma
capsaeformis), Clinch River, TN.
Shell Characteristics: Shell shape
highly variable; adults usually solid and small (< 70 mm). Most taxa exhibit strong
sexual dimorphism; which varies in pronouncement and morphology. Females generally
possess an expanded and rounded posterior, elongated posterior, or a pronounced
marsupial swelling along the posterior ridge. Males are often shallow to moderately
sulcate; although some taxa lack a sulcus altogether (e.g Epioblasma triquetra).
The periostracum is yellowish-tan to brown with green rays or chevron markings.
Pseudocardinal teeth and lateral teeth are generally well developed; often heavy in
large specimens. Nacre is usually white or purple.
Photo 22: The Powell River (TN, VA) was historically home to several species of Epioblasma, including the
extinct Forkshell (Epioblasma lewisii) and Acornshell (Epioblasma haysiana).
34
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An Introduction to Freshwater Mussels as Biological Indicators
Habitat: The Epioblasma were
formerly widespread throughout the
eastern United States, inhabiting
the Mississippi River basin, Great
Lakes basin, and Mobile basin.
Today, their ranges have been
dramatically reduced, and for the
most part, are small and disjunct
(USFWS1983b;1985a; 1990;
1994; 2004). As a group, this
genus inhabits shallow riffles,
runs, or shoals of moderate-sized
creeks and rivers (Stansberry
1970; Walters 1995; Parmalee and
Bogan 1998). They are often most
abundant in clear water and clean
substrates comprised of sand,
gravel, and cobble.
Reproduction: Because the
Epioblasma have experienced such
a precipitous decline, very little
information is available regarding
the reproductive habits of most
taxa. However, observations made
by several researchers suggest that
some Epioblasma employ brutal
tactics to entrap host fishes. Much
of the following is paraphrased from
Barnhart (2007):
Taxa List (Turgeon et al. 1998)
Altamaha Arcmussel (Alasmidonta arcula)
Angled Riffleshell (Epioblasma biemarginata)
Cumberland Combshell (Epioblasma brevidens)
Oyster Mussel (Epioblasma capsaeformis)
Leafshell (Epioblasma flexuosa)
Curtis Pearlymussel (Epioblasma florentina curtisii)
Yellow Blossom (Epioblasma florentina florentina)
Tan Riffleshell (Epioblasma florentina walkeri)
Acornshell (Epioblasma haysiana)
Narrow Catspaw (Epioblasma lenoir)
Forkshell (Epioblasma lewisii)
Upland Combshell (Epioblasma metastriata)
Catspaw (Epioblasma obliquata obliquata)
White Catspaw (Epioblasma obliquata perobliqua)
Southern Acornshell (Epioblasma othcaloogensis)
Southern Combshell (Epioblasma penita)
Round Combshell (Epioblasma personata)
Tennessee Rifflshell (Epioblasma propinqua)
Wabash Riffleshell (Epioblasma sampsonii)
Cumberland Leafshell (Epioblasma stewardsonii)
Green Blossom (Epioblasma torulosa gubernaculum)
Northern Riffleshell (Epioblasma torulosa rangiana)
Tubercled Blossom (Epioblasma torulosa torulosa)
Snuffbox (Epioblasma triquetra)
Turgid Blossom (Epioblasma turgidula)
Dubbed "fish snappers" by Jess Jones (Virginia Tech University), this
moniker aptly describes a technique utilized by gravid females to trap
unwary fishes. To initiate glochidia-host interaction when brooding,
females migrate to the surface and lie-in-wait. While resting (or slightly
burrowed) on the substrate some of the Epioblasma gape widely, exposing
their spongy "mantle pads" (termed cymapallia by Barnhart). When contact
is initiated with the shell or mantle pads, the female quickly snaps her shell
shut. This action essentially traps unsuspecting fish, allowing the female to
directly infect her captive with glochidia. This technique seems especially
brutal when fishes are observed ensnared by the snout or head.
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An Introduction to Freshwater Mussels as Biological Indicators
Identified hosts include fishes from families Percidae (perches), Cottidae (sculpins), and
Salmonidae (trout) (Buchanan 1987; Sherman 1993; Yaeger and Saylor 1995; O'Dee and
Walters 2000; Rogers et al. 2001). More specifically, the most commonly reported hosts
are darters from two genera, Etheostoma and Percina, and sculpin belonging to the genus
Cottus.
Turgeon et al. 1 998 Taxa
Presumed Extinct Taxa
Federally Listed Taxa
Williams et al. 1993
Special Concern
Threatened
Endangered
Total
Natu reserve
G5
G4
G3
G2
G1
G2TX
G2T2
G1
G1TX
G1T1
GHQ
GH
GX
25
14(56.0%)
14(56.0%)
0 (0.0%)
1 (4.0%)
24 (96.0%)
1 1 (91 .7%)
0 (0.0%)
0 (0.0%)
1 (4.0%)
0 (0.0%)
2 (8.0%)
1 (4.0%)
3(12.0%)
1 (4.0%)
1 (4.0%)
4(16.0%)
1 (4.0%)
1 (4.0%)
1 1 (44.0%)
Tolerance to Habitat Alteration: The
Epioblasma are, with few exceptions, habitat
specialists adapted to shallow segments of large
creeks and rivers. The ubiquitous damming and
modification of river habitats has likely impacted
this specialized group more than any other
genera of unionids (USFWS 1983b; 1985a).
Many of the now extinct Epioblasma formerly
occured in riffles of large rivers, a habitat that
has been virtually eliminated by impoundment
(Stansberry 1970).
Sensitivity to Toxic Contaminants: Recently
Wang et al. (2007a) assessed the acute toxicity
of copper, ammonia, and chlorine to the early life
stages of 11 species, including the oyster mussel
(E. capsaeformis). Newly-transformed Oyster
Mussel juveniles were among the more sensitive
species tested during 96-hour exposures.
Reported EC50s were 17 ug/L and 5.7 mg N/L
for copper and ammonia, respectively.
Indicator Use: While generally not used for contaminant biomonitoring studies, the
Epioblasma are certainly among the most sensitive genera to anthropogenic disturbance
and have great value for measuring the biological integrity of surface waters (Ahlstedt
1991; Neves et al. 1997). As such, this group has been utilized as a sensitive indicator. For
example, Kearns and Karr (1994) used Epioblasma as an intolerant mussel genera when
developing a benthic index of biotic integrity (B-IBI) for rivers of the Tennessee Valley.
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An Introduction to Freshwater Mussels as Biological Indicators
GENUS PROFILE - FUSCONAIA
Species Diversity and Conservation: A total of 13 species belong to the genus
Fusconaia (Turgeon et al. 1998). Williams et al. (1993) reviewed the conservation status
of unionoids in North America and reported six Fusconaia as special concern, two
as threatened, and two species as endangered. At present, two species are listed as
endangered by the U.S. Fish and Wildlife Service (USFWS 2007).
Taxa List (Turgeon et al. 1998)
Texas Pigtoe (Fusconaia askewi)
Tennessee Pigtoe (Fusconaia barnesiana)
Southern Pigtoe (Fusoconaia cerina)
Shiny Pigtoe (Fusconaia cor)
Finerayed Pigtoe (Fusconaia cuneolus)
Ebonyshell (Fusconaia ebena)
Narrow Pigtoe (Fusconaia escambia)
Wabash Pigtoe (Fusconaia flava)
Triangle Pigtoe (Fusconaia lanaensis)
Atlantic Pigtoe (Fusconaia masoni)
Ozark Pigtoe (Fusconaia ozarkensis)
Longsolid (Fusconaia subrotunda)
Purple Pigtoe (Fusconaia succissa)
Shell Characteristics: Shell shape highly
variable; often subtriangular, subovate, or
oval. Valves usually moderately thick and
more often inflated than not (especially big
river forms). Beaks raised above hinge-line.
Shell surface smooth and devoid of sculpture.
Maximum length ranges from approximately
2.5 to 4.5 inches (65-110 mm). Shell
sexual dimorphism absent or very weakly
pronounced. Periostracum yellowish-tan,
reddish-brown, brown, or dark brown; often
with faint green rays when young. Teeth well
developed. Nacre white, rarely with a salmon
tinge.
Habitat: The Fusconaia are widely
distributed throughout the lower and upper
Mississippi River basin, Great Lakes-St.
Lawrence basin, several Gulf drainages, and the southern Atlantic drainage. They occur in
small to large creeks, rivers, lakes, and reservoirs in both fine and coarse substrates.
Reproduction: Researchers have
identified suitable host fishes for several
Fusconaia. Confirmed hosts belong
to one of five families: Clupeidae
(herrings), Cyprinidae (minnows),
Centrarchidae (sunfishes), Percidae
(darters), and Cottidae (sculpins) (Coker
et al. 1921; Wilson 1916; Bruenderman
and Neves 1993; O'Dee and Walters
2000; Haag and Warren 2003). The
Fusconaia are tachytictic, with the
reproductive period extending from May
to August (Baker 1928).
Photo 23: Roanoke River, VA, home to the Atlantic
Pigtoe (Fusconaia masoni).
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 24: The Wabash Pigtoe (Fusconaia flava),
Swan Creek, OH.
Photo 25: Streamline chubs (Erimystax
dissimilis) foraging above a Longsolid
(Fusconaia subrotunda), French Creek, PA.
Female Fusconaia package glochidia in semi-cohesive capsules (termed "conglutinates")
that are expelled into the water column (Utterback 1915: 1916). Conglutinates are fed on
by fishes, thereby infecting potential hosts.
Tolerance to Habitat Alteration: The Fusconaia exhibit varying degrees of tolerance
to habitat alteration. The Wabash Pigtoe (Fusconaia flava), for example, is a habitat
generalist capable of tolerating turbid streams, impounded habitats, and a variety of
substrates (Gordon and Layzer 1989; Walters 1995; Strayer and Jirka 1997). Likewise,
the Ebonyshell (Fusconaia ebena) has exhibited the ability to adapt to impoundment
in reaches of the lower Ohio River (Payne and Miller 2001). Other species, such as the
Cumberlandian endemic Finerayed Pigtoe (Fusconaia cuneolus), are adapted to shallow,
swiftly flowing waters of creeks and rivers. Habitat alterations such as impoundment and
siltation have contributed to the decline of this species (USFWS 1984). The Finerayed
Pigtoe is currently listed as endangered by the U.S. Fish and Wildlife Service.
Sensitivity to Toxic Contaminants: Relatively little laboratory toxicity testing has been
performed with the Fusconaia.
Waller et al. (1998b) evaluated the acute toxicity of 3-trifluoromethyl-4-nitrophenol (TFM),
a common lampricide, to the Wabash Pigtoe and Threehorn Wartyback (Obliquaria
reflexa). The Wabash Pigtoe was less sensitive than the Threehorn Wartyback, with
reported LC50s of 3.81 and 1.80 mg/L, respectively. Researchers concluded that sea
lampricide treatments would likely have negligible effects on the species tested.
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 26: Cumberland River, KY, home to the Ebonyshell (Fusconaia ebena), Longsolid (Fusconaia
subrotunda), and Wabash Pigtoe (Fusconaia flava).
Indicator Use: Pip (1995) evaluated heavy
metal contamination in the Assiniboine River
(Manitoba, Canada) using several species of
unionids as biomonitors. The Wabash Pigtoe
accumulated levels of cadmium, lead, and
copper comparable to other species. However,
the Mapleleaf (Quadrula quadrula) accumulated
significantly more copper in the gill, while
the Fatmucket (Lampsilis siliquoidea) and
Black Sandshell (Ligumia recta) accumulated
substantially more cadmium in the heart.
Turgeon et al. 1 998 Taxa
Presumed Extinct Taxa
Federally Listed Taxa
Williams etal. 1993
Special Concern
Threatened
Endangered
Total
Natu reserve
G5
G4-G5
G4
G3-G4
G3
G2-G3
G2
G1Q
G1
13
0 (0.0%)
2(15.4%)
6 (46.2%)
2(15.4%)
2(15.4%)
10(76.9%)
2(15.4%)
1 (7.7%)
0 (0.0%)
2(15.4%)
1 (7.7%)
2(15.4%)
2(15.4%)
1 (7.7%)
2(15.4%
Imlay (1982) advocated the use of several
Fusconaia as biomonitors of heavy metals
based on annual shell growth, including the
Longsolid (Fusconaia subrotunda), Finerayed
Pigtoe, Wabash Pigtoe, Tennesse Pigtoe
(Fusconaia barnesiana), and Shiny Pigtoe
(Fusconaia cor).
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An Introduction to Freshwater Mussels as Biological Indicators
GENUS PROFILE - LAMPSILIS
Taxa List (Turgeon et al. 1998)
Pink Mucket (Lampsilis abrupta)
Finelinded Pocketbook (Lampsilis altilis)
Southern Sandshell (Lampsilis australis)
Lined Pocketbook (Lampsilis binominata)
Texas Fatmucket (Lampsilis bracteata)
Plain Pocketbook (Lampsilis cardium)
Yellow Lampmussel (Lampsilis cariosa)
Atlamaha Pocketbook (Lampsilis dolabraeformis)
Wavyrayed Lampmussel (Lampsilis fasciola)
Waccamaw Fatmucket (Lampsilis fullerkati)
Haddleton Lampmussel (Lasmpsilis haddletoni)
Higgins Eye (Lampsilis higginsii)
Louisiana Fatmucket (Lampsilis hydiana)
Southern Pocketbook (Lampsilis ornate)
Pocketbook (Lampsilis ovata)
Orangenacre Mucket (Lampsilis perovalis)
Arkansas Fatmucket (Lampsilis powellli)
Carolina Fatmucket (Lampsilis radiata conspicua)
Eastern Lampmussel (Lampsilis radiata radiata)
Neosho Mucket (Lampsilis rafinesqueana)
Ozark Brokenray (Lampsilis reeveiana brevicula)
Northern Brokenray (Lampsils reeveiana brittsi)
Arkansas Brokenray (Lampsilis reeveiana reeveiana)
Sandbank Pocketbook (Lampsilis satura)
Fatmucket (Lampsilis siliquoidea)
Rayed Pink Fatmucket (Lampsilis splendida)
Southern Fatmucket (Lampsilis straminea claibornensis)
Rough Fatmucket (Lampsilis straminea straminea)
Speckled Pocketbook (Lampsilis streckeri)
Shinyrayed Pocketbook (Lampsilis subangulata)
Yellow Sandshell (Lampsilis teres)
Alabama Lampmussel (Lampsilis virescens)
Species Diversity and
Conservation: A total of 25
species and seven subspecies
have been assigned to the
genus Lampsilis (Turgeon et
al. 1998). A single species, the
Lined Pocketbook (Lampsilis
binominata), is presumed extinct
(Turgeon et al. 1998). Williams
et al. (1993) reviewed the
conservation status of unionoids
in North America and reported
nine Lampsilis as special
concern, 10 as threatened, and
six species as endangered.
Currently, eight Lampsilis are
listed as endangered by the
U.S. Fish and Wildlife Service
(USFWS 2007).
Shell Characteristics: Shell
elliptical, subovate, elongate,
or subquadrate; moderately
thin to heavy and moderately
compressed to inflated. Most
Lampsilis exhibit strong
sexual dimorphism. Females
are expanded and rounded
posteriorly or obliquely flared
posteriorly. Males are usually
bluntly pointed. Maximum shell
length variable; usually ranging
from 2.5 to 7.0 inches (65-170
mm). Periostracum often yellow,
yellowish-green, yellowish-tan or
brown; thin to wide green rays
common but not always present.
Beak nearly even or raised well
above hinge-line; sculpture often
40
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 27: Lake Michigan, home to the Plain Pocketbook (Lampsilis cardium) and Fatmucket (Lampsilis
siliquoidea).
double-looped. Teeth well developed; may
be large and heavy in old individuals. Nacre
usually white, bluish-white, pink, or salmon.
Habitat: The Lampsilis are a widely
distributed group, ranging geographically
from the Central United States to the
East Coast and north into Canada. They
are known from the upper and lower
Mississippi River basin, Great Lakes-St.
Lawrence basin, Atlantic slope, several Gulf
drainages, and a few Canadian drainages.
Lampsilis are found in small to large creeks,
rivers, and lakes; occurring in both fine and
coarse substrates.
Reproduction: Researchers have
identified suitable host fishes for several
Lampsilis. Confirmed hosts belong to one of
seven families: Acipenseridae (sturgeons),
Lepisosteidae (gars), Cyprinidae (minnows),
Fundulidae (topminnows), Centrarchidae
(sunfishes), Percidae (perches), or
Sciaenidae (drums) (Surber 1913; Coker et
al. 1921; Fuller 1974;Zaleand Neves 1982;
Waller and Holland-Bartels 1988; O'Dee
and Walters 2000).
Turgeon et al. 1 998 Taxa
Presumed Extinct Taxa
Federally Listed Taxa
Williams et al. 1993
Special Concern
Threatened
Endangered
Total
Natureserve
G5
G5T5
G5T3
G5T2Q
G5T1Q
G4
G4T3
G3G4
G3
G2
G1Q
G1
GXQ
GX
32
1 (3.1%)
8 (25.0%)
9(28.1%)
10(31.2%)
6(18.8%)
25 (84.0%)
7(24.1%)
2 (6.9%)
1 (3.4%)
1 (3.4%)
1 (3.4%)
1(3.4%)
1 (3.4%)
2 (6.9%)
1 (7.7%)
3(10.3%)
2 (6.9%)
3(10.3%)
1 (7.7%)
1 (7.7%)
41
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An Introduction to Freshwater Mussels as Biological Indicators
The Lampsilis have evolved some of the most
extraordinary reproductive adaptations found
in freshwater bivalves. To enhance glochidia-
host interaction, many Lampsilis possess a
conspicuous mantle flap "lure" that appears,
in form and pigmentation, similar to a small
fish (photo 28). Mantle lures may rapidly or
occasionally "pulse" or "swim," distinctive
movements that expose the charged gills. This
motion can be quite variable among species,
season, and time of day, ranging from 180
movements per minute to virtually no movement
at all (Kraemer 1970). Gravid females use this
amazing tool to lure potential host fishes into
striking the mantle flap, thereby rupturing the
marsupia and liberating hundreds to thousands
of glochidia.
Tolerance to Habitat Aleration: The Lampsilis
vary widely in their tolerance to habitat
alteration. Widespread species such as the
Plain Pocketbook (Lampsilis cardium) and
Fatmucket (Lampsilis siliquoidea) may tolerate
impoundment, turbidity, and sedimentation
(Parmalee and Bogan 1998). Conversely, several
federally listed taxa, many endemic to southern
drainages, have been severely reduced in
number by siltation, dredging, channelization,
impoundment, and hypolimnetic releases
(USFWS 1985b; USFWS 1991; USFWS 1992).
Sensitivity to Toxic Contaminants: The Lampsilis have been widely used to evaluate
the toxicity of various contaminants to unionids (e.g. Jacobson et al.1993; Jacobson et
al. 1997; Wang et al. 2007a; Wang et al. 2007b). Refer to the Wavyrayed Lampmussel,
Fatmucket, and Plain Pocketbook species accounts for more information.
42
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An Introduction to Freshwater Mussels as Biological Indicators
Indicator Use: Several of the Lampsilis have been used to successfully biomonitor heavy
metals, including Plain Pocketbook (Lampsilis cardium), Eastern Lampmussel (Lampsilis
radiata radiata), and Fatmucket (Lampsilis siliquoidea). Refer to the species accounts for
more information.
Photo 28: In-situ mantle flap lure of the
Wavyrayed Lampmussel.
Photo 29: In-situ apertures of the Plain Pocketbook.
43
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An Introduction to Freshwater Mussels as Biological Indicators
GENUS PROFILE - LASMIGONA
Taxa List (Turgeon et al. 1998)
Alabama Heelsplitter (Lasmigona complanata
alabamensis)
White Heelsplitter (Lasmigona complanata
complanata)
Creek Heelsplitter (Lasmigona compressa)
Flutedshell (Lasmigona costata)
Carolina Heelsplitter (Lasmigona decorata)
Tennessee Heelsplitter (Lasmigona holstonia)
Green Floater (Lasmigona subviridis)
Species Diversity and Conservation: A total
of five species and two subspecies have been
assigned to the genus Lasmigona (Turgeon et
al. 1998). Williams et al. (1993) reviewed the
conservation status of unionoids in North America
and reported two Lasmigona as special concern,
one as threatened, and one species as endangered.
Currently, the Carolina Heelsplitter (Lasmigona
decorata) is listed as endangered by the U.S. Fish
and Wildlife Service (USFWS 2007).
Shell Characteristics: Shell variable; generally
rhomboidal, trapezoidal, or subovate; often quite thin
when young. A few members of Lasmigona possess
distinctive undulations or "flutes" posteriorly (e.g.
Lasmigona c. complanata and Lasmigona costata).
Maximum shell length from 2.5 to 7.0 inches (70-
180 mm). Periostracum usually yellowish-brown or
green when young; some species may have fine
rays. Shell becoming dark with age. Beak sculpture
often of well formed double-loops or moderately
heavy bars. Pseduocardinal teeth well developed
although variable in form; may be low and serrated
or elevated. Nacre white, often with a salmon tinge.
44
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 30: The Flutedshell (Lasmigona costata) in French Creek,
PA.
Habitat: The Lasmigona are a
wideranging group, inhabiting
the lower and upper Mississippi
River basin, Great Lakes-St.
Lawrence basin, Lake Winnipeg
drainage, Atlantic Slope, and
Gulf drainages. This group may
be found in both headwaters
and large rivers, as well as
lakes and reservoirs.
Reproduction: Researchers
have identified suitable
host fishes for many of the
Lasmigona. Confirmed hosts
belong to one of nine families:
Lepisosteidae (gars), Clupeidae (herrings and shad), Cyprinidae (minnows), Catostomidae
(suckers), Ictaluridae (catfishes), Fundulidae (topminnnows), Cottidae (sculpins),
Centrarchidae (sunfishes), and Percidae (perches) (Lefevre and Curtis 1910; Young 1911;
Lefevre and Curtis 1912; Hove et al. 1995; Steg and Neves 1997; Walters 1998a; Walters
et al. 1998b; McGill et al. 2002; Walters et al. 2005).
While the reproductive traits of each species belonging to Lasmigona have not been
thoroughly investigated, most members have been confirmed as bradytictic.
Tolerance to Habitat Alteration: The
Lasmigona exhibit a wide array of responses to
habitat alteration. The White Heelpslitter (L.c.
complanata), for example, has been reported
to tolerate impoundment, eutrophication, and
disturbed or silty substrates (Walters 1995;
Strayer and Jirka 1997; Parmalee and Bogan
1998; Metcalfe-Smith et al. 2003; Sietman
2003). Furthermore, Walters (1995) noted
that L.c. complanata may be abundant below
sewage outfalls. Conversely, the federally
endangered Carolina Heelsplitter (L. decorata)
has unfortunately exhibited sensitivity to a
suite of disturbances and stressors, including
stream impoundment, unstable stream banks,
channelization, siltation, road construction and
maintenance, and mining runoff (Keferl 1991;
USFWS 1996).
Turgeon et al. 1998 Taxa
Presumed Extinct Taxa
Federally Listed Taxa
Williams etal. 1993
Special Concern
Threatened
Endangered
Total
Natureserve
G5
G4
G3
G2
G1
7
0 (0.0%)
1 (14.3%)
2 (28.6%)
1 (14.3%)
1 (14.3%)
4 (57.2%)
4 (57.2%)
0 (0.0%)
2 (28.6%)
0 (0.0%)
1 (14.3%)
45
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 31: Green River, KY, home to the Flutedshell (L costata) and White Heelsplitter (L.c.
complanata).
Sensitivity to Toxic Contaminants: The Lasmigona are rarely used in laboratory toxicity
testing. However, Black (2001) did utilize the Green Floater (L. subviridis) as a surrogate
to derive water quality standards for the protection of North Carolina's endangered
mussels.
Indicator Use: The Lasmigona are valuable indicators of habitat and water quality.
For example, Metcalfe-Smith et al. (2003) considered the range expansion of the White
Heelsplitter and Flutedshell in the Sydenham River basin a biological indicator of
environmental deterioration.
Bedford et al. (1968) utilized the Flutedshell as a biomonitor of pesticides in the Red
Cedar River (Ml). Reported concentrations ranged from 0.0153-0.198 ppm total DDT
and metabolites. The study concluded that mussels were excellent indicators of pesticide
contamination; mainly because of their ability to concentrate contaminants.
Pip (1995) assessed the bioaccumulation and biomonitoring potential of the White
Heelsplitter and several other species in the Assiniboine River, Manitoba (Canada). Unlike
the other test unionids, cadmium, lead, and copper were concentrated to higher levels in
the mantle tissue of L.c. complanata than the heart, gills, gonad, muscle, or foot. It should
be noted, however, that only a single White Heelsplitter was analyzed.
46
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An Introduction to Freshwater Mussels as Biological Indicators
GENUS PROFILE - PLEUROBEMA
Species Diversity and Conservation:
A total of 32 species belong to the
genus Pleurobema (Turgeon et al.
1998). Williams et al. (1993) reviewed
the conservation status of unionoids
in North America and reported four
Pleurobema as special concern, one
as threatened, and 22 species as
endangered. At present, 12 species
are listed as endangered by the U.S.
Fish and Wildlife Service (USFWS
2007) while a total of 13 species are
presumed extinct (Turgeon et al. 1998).
Shell Characteristics: Shell shape
highly variable; often subtriangular or
subelliptical; less often subrhomboidal
or subquadrate. Shells usually
moderately thick to thick and more often
inflated than not (especially big river
forms). With rare exception (e.g. James
spinymussel), shell surface devoid of
sculpture. Maximum lengths range from
approximately 1.5 to 5.0 inches (35-120
mm). Shell sexual dimorphism absent or
weakly pronounced. Periostracum often
yellowish-tan, brown, or dark brown;
clubshells and some pigtoes with
rays extending from umbo. Teeth well
developed and heavy. Nacre usually
white or pink.
Habitat: The Pleurobema are a
wideranging genus, with species
richness maximized in the large creeks
and rivers of the Cumberland Plateau
and Mobile River basin. As a group,
they inhabit the lower and upper
Mississippi River basin, Great Lakes
basin, James River drainage, Roanoke
Taxa List (Turgeon et al. 1998)
Highnut (Pleurobema altum)
Hazel Pigtoe (Pleurobema avellanum)
Mississippi Pigtoe (Pleurobema beadleianum)
Scioto Pigtoe (Pleurobema bournianum)
Painted Clubshell (Pleurobema chattanoogaense)
Clubshell (Pleurobema clava)
James Spinymussel (Pleurobema collina)
Ohio Pigtoe (Pleurobema cordatum)
Black Clubshell (Pleurobema curtum)
Southern Clubshell (Pleurobema decisum)
Yellow Pigtoe (Pleurobema flavidulum)
Dark Pigtoe (Pleurobem furvum)
Southern Pigtoe (Pleurobema georgianum)
Cumberland Pigtoe (Pleurobema gibberum)
Brown Pigtoe (Pleurobema hagleri)
Georgia Pigtoe (Pleurobema hanleyianum)
Alabama Pigtoe (Pleurobema johannis)
Flat Pigtoe (Pleurobema marshal!!)
Coosa Pigtoe (Pleurobema murrayense)
Longnut (Pleurobema nucleopsis)
Tennessee Clubshell (Pleurobema oviforme)
Ovate Clubshell (Pleurobema perovatum)
Rough Pigtoe (Pleurobema plenum)
Oval Pigtoe (Pleurobema pyriforme)
Louisiana Pigtoe (Pleurobema riddellii)
Warrior Pigtoe (Pleurobema rubellum)
Pyramid Pigtoe (Pleurobema rubrum)
Round Pigtoe (Pleurobema sintoxia)
Fuzzy Pigtoe (Pleurobema strodeanum)
Heavy Pigtoe (Pleurobema taitianum)
Alabama Clubshell (Pleurobema troschelianum)
True Pigtoe (Pleurobema verum)
47
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 32: The Green River (KY), home of the federally endangered Rough Pigtoe (Pleurobema plenum).
River drainage, and several Gulf drainages. The Pleurobema are often most abundant in
small and large rivers, where substrates are comprised of sand, gravel, and cobble.
Reproduction: Researchers have identified suitable host fishes for many of the
Pleurobema. Confirmed hosts belong to one of four families - Cyprinidae (minnows),
Fundulidae (topminnows), Percidae (perches), or Centrarchidae (sunfishes) (Surber 1913;
Yokley 1972; Weaver et al. 1991; Hove 1995a; O'Dee and Walters 2000; Haag and Warren
2003). Although the reproductive traits of numerous species remain undocumented,
studies have reported several Pleurobema as tachytictic, with the reproductive period
extending from May to August (Ortmann 1909; Baker 1928; Haag and Warren 2003).
Tolerance to Habitat Alteration: The Pleurbema
have been severely impacted by habitat alterations
that have changed free-flowing, riffle-pool river
segments to impounded, lentic habitats. For example,
widespread modification of the Upper Tombigbee River
(Mobile River basin) occurred when construction of
the Tombigbee-Tennessee Waterway was completed
in 1984. The Waterway shortened the river by 48
miles, destroyed and fragmented suitable streambed
habitat, profoundly altered the flow regime, and initiated
geomorphic instability (ARA et al. 1999). Once home
to 50 species of freshwater mussels, including several
pleurobemids, approximately 30% of the fauna is now
imperiled, extirpated, or extinct as a direct result of river
impoundment and related impacts (ARA et al. 1999).
Four federally endangered species, the Black Clubshell
(Pleurobema curtum), Southern Clubshell (Pleurobema decisum), Ovate Clubshell
(Pleurobema perovatum), and Heavy Pigtoe (Pleurobema taitianum) once occurred in the
Tombigbee River (USFWS 2000). They are now considered extirpated or limited to small
segments where suitable habitat still exists (Hartfield and Jones 1989; USFWS 2000).
Photo 33: The federally
endangered Clubshell (Pleurobema
clava), French Creek, PA.
48
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 34: East Fork West Branch St. Joseph River, Ml, habitat of the Clubshell (P. clava).
Sensitivity to Toxic Contaminants: The
Pleurobema have not been used in laboratory
toxicity testing.
Indicator Use: The Pleurobema are among
the most imperiled mollusks in North America,
with 78% of the genus either listed as federally
endangered or extinct (Turgeon et al. 1998;
USFWS 2007). Many are endemic to Mobile
Basin, including over a dozen extinct species
(Neves et al. 1997; Turgeon et al. 1998). They
are sensitive to habitat alterations such as river
impoundment and channelization, in addition to
stressors such as water quality degradation and
sedimentation (USFWS 2000).
The use of Pleurobema in bioaccumulation
studies has not been reported on. However, Imlay
(1982) recommended the use of the Ohio Pigtoe
(Pleurobema cordatum), Tennessee Clubshell
(Pleurobema oviforme), and Round Pigtoe
(Pleurobema sintoxia) as biomonitors of heavy
metals on the basis of annual shell growth. This
recommendation was based on "widespread
distribution, age, pollution tolerance, and/or
conchological reflection of stream location."
Turgeon et al. 1 998 Taxa
Presumed Extinct Taxa
Federally Listed Taxa
Williams et al. 1993
Special Concern
Threatened
Endangered
Total
Natureserve
G4G5
G4
G3
G2G3
G2
G1G2Q
G1G2
G1Q
G1
GHQ
GH
GX
32
13(41.0%)
12(38.0%)
4(13.0%)
1 (3.0%)
22 (69.0%)
27 (84.0%)
1 (3.0%)
1 (3.0%)
1 (3.0%)
3(9.1%)
3(9.1%)
1 (3.0%)
1 (3.0%)
3(9.1%)
10(30.3%)
1 (3.0%)
1 (3.0%)
7(21.2%)
49
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An Introduction to Freshwater Mussels as Biological Indicators
GENUS PROFILE - QUADRULA
Species Diversity and Conservation: A total
of 16 species and four subspecies have been
assigned to the genus Quadrula (Turgeon et
al. 1998). A single species, the rough rockshell
(Quadrula tuberosa), is presumed extinct
(Turgeon et al. 1998). Williams et al. (1993)
reviewed the conservation status of unionoids
in North America and reported five Quadrula
as special concern, three as threatened, six
species as endangered, and one as possibly
extinct. Currently, five taxa are listed as
endangered by the U.S. Fish and
Wildlife Service (USFWS 2007).
Shell Characteristics: Shell
quadrate or round; often heavy
and moderately inflated to inflated.|
Maximum shell length variable;
usually ranging from 2.5 to 6.0
inches (64-153 mm). Shell sexual
dimorphism weakly pronounced
or absent. Periostracum often
yellowish-tan or brown; with green
smudges, chevrons, or faint rays
commonly found in several taxa. If present, shell sculpture consists of few to numerous
bumps, pustules, or knobs. Teeth well developed and heavy. Nacre usually white.
Habitat: The Quadrula are a wideranging group, inhabiting the lower and upper
Mississippi River basin, Great Lakes basin, and several Gulf drainages. In general, this
group is most abundant in small and large rivers, where substrates are comprised of
packed sand, gravel, and cobble. Several Quadrula may also occur in lakes and reservoirs.
Reproduction: Researchers have identified suitable host fishes for several Quadrula.
Confirmed hosts belong to one of three families - Cyprinidae (minnows), Ictaluridae
(catfishes), or Centrarchidae (sunfishes) (Surber 1913; Coker 1921; Yeager and Neves
1986; Barnhart 2000; Hove et al. 2001; Haag and Warren 2003). However, host studies
have not been performed on a fair number of taxa (many endemic to southern drainages).
The genus is apparently tachytictic, with the reproductive period extending from May to
August (Utterback 1915; Baker 1928).
50
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An Introduction to Freshwater Mussels as Biological Indicators
Taxa List (Turgeon et al. 1998)
Southern Mapleleaf (Quadrula apiculata)
Alabama Orb (Quadrula asperata)
Golden Orb (Quadrula aurea)
Rio Grande Monkeyface (Quadrula couchiana)
Rabbitsfoot (Quadrula cylindrica cylindrica)
Rough Rabbitsfoot (Quadrula cylindrica strigillata)
Winged Mapleleaf (Quadrula fragosa)
Smooth Pimpleback (Quadrula houstonensis)
Cumberland Monkeyface (Quadrula Intermedia}
Monkeyface (Quadrula metanevra)
Wartyback (Quadrula nodulata)
Texas Pimpleback (Quadrula petrina)
Western Pimpleback (Quadrula pustulosa
mortoni)
Pimpleback (Quadrula pustulosa pustulosa)
Mapleleaf (Quadrula quadrula)
Purple Pimpleback (Quadrula refulgens)
Ridged Mapleleaf (Quadrula rumphiana)
Appalachian Monkeyface (Quadrula sparsa)
Stirrupshell (Quadrula stapes)
Rough Rockshell (Quadrula tuberosa)
Photo 35: Rabbitsfoot (Quadrula cylindrica
cylindrica), French Creek, PA.
Tolerance to Habitat Alteration:
Members of genus Quadrula have
demonstrated a wide array of
sensitivities to habitat alteration.
For example, species such as the
Wartyback (Quadrula nodulata),
Pimpleback (Quadrula pustulosa
pustulosa), and Mapleleaf (Quadrula
quadrula) appear capable of tolerating
river impoundment (Oesch 1984;
Parmalee and Bogan 1998; ESI
2001; WDNR 2003). Conversely,
unionids such as the Rabbitsfoot
(Q.c. cylindrica) have experienced
widespread population reductions due
to river impoundment and associated
hypolimnetic releases (Bates 1962;
Butler 2005b). The Rabbitsfoot is
currently under assessment for
federal status, while the Wartyback,
Pimpleback, and Mapleleaf remain
secure throughout their respective
ranges (Williams et al. 1993; Butler
2005b).
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An Introduction to Freshwater Mussels as Biological Indicators
Photo 36: French Creek, PA, habitat of the Rabbitsfoot (Q.c. cylindrica).
Sensitivity to Toxic Contaminants: The
Quadrula are not commonly used in laboratory
toxicity testing.
Indicator Use: Foster and Bates (1978) reported
the use of Q. quadrula as a biomonitor of copper
electroplating wastes in the Muskingum River, OH.
Caged mussels were placed at various intervals
downstream of the study outfall for 14, 30, and 45
days. Mortality of caged mussels was observed
after 11 days and associated with body burdens
of 20 ug Cu/g wet tissue weight. Researchers
concluded that elevated levels of copper were
likely responsible for mass mortalities of freshwater
mussels.
In addition to the study above, Q. quadrula has
been used as a biomonitor of iron-dominated
mine discharges (Milam and Farris 1998) and
ambient cadmium, lead, and copper levels in
the Assiniboine River, Canada (Pip 1995). The
Monkeyface (Quadrula metanevra) and Pimpleback
were utilized by Allen et al. (2001) to assess
potential heavy metal contamination within the
habitat range of the federally threatened Neosho Madtom (Noturus placidus).
Imlay (1982) advocated the use of several species as biomonitors of heavy metals based
on annual shell growth, including the Rabbitsfoot, Pimpleback, and Monkeyface. This
recommendation was based on "widespread distribution, age, pollution tolerance, and/or
conchological reflection of stream location."
Turgeon et al. 1 998 Taxa
Presumed Extinct Taxa
Federally Listed Taxa
Williams etal. 1993
Special Concern
Threatened
Endangered
Total
G5
G5T5
G5T3Q
G4
G3G4
G3G4T3
G3G4T2
G2
G1
GH
GXQ
20
1 (5.0%)
5 (25.0%)
4 (20.0%)
3(15.0%)
7 (35.0%)
14(70.0%)
3(13.6%)
1 (4.5%)
1 (4.5%)
4(18.1%)
2(9.1%)
1 (4.5%)
1 (4.5%)
2(9.1%)
4(18.1%)
2(9.1%)
1 (4.5%)
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An Introduction to Freshwater Mussels as Biological Indicators
Mucket
Actinonaias ligamentina
(Lamarck, 1819)
Identification tips: Shell oval to elliptical,
thick, and somewhat compressed when young,
becoming inflated with age. Periostracum light
tannish-green to brown; young individuals often
with wide green rays (A) Beak slightly raised above
hinge-line; sculpture of a few concentric ridges
that are often eroded (B) Teeth well developed;
especially heavy in older individuals. Nacre white.
Shell up to 7 inches in length.
Indicator use: A widespread and often abundant
species, the Mucket is currently stable throughout
its range (Williams et al. 1993; Natureserve).
Spooner et al. (2005) studied the physiological
effects of high water temperatures on eight species
of unionids, including A. ligamentina.The mucket
was reported as the most thermally sensitive
species studied, with reduced respiration rates
at 32°C. The authors suggested the use of A.
ligamentina as an indicator species for mussel
bed health in the Kiamachi River, OK. It was
also advised that managing to protect Mucket
populations should translate into thermal protection
for federally listed species.
GMAVs compiled and ranked by Augspurger et
al. (2003), Augspurger et al. (2006), and March et
al. (2007) placed Actinonaias as "intermediately
tolerant" of ammonia and copper when compared
to other unionid genera.
Habitat: Widespread in the Mississippi River basin
and Great Lakes-St. Lawrence basin. The Mucket
is found in creeks and rivers where substrates are
comprised of stable sand, gravel, and cobble.
Reproduction: Identified host species include
the American Eel, Central Stoneroller, Silverjaw Minnow, Common Carp, Tadpole Madtom,
Banded Killifish, White Bass, Rock Bass, Green Sunfish, Orangespotted Sunfish, Bluegill
Sunfish, Black and White Crappie, Smallmouth Bass, Largemouth Bass, Tippecanoe
Darter, and Yellow Perch (Young 1911; Lefevre and Curtis 1912; Coker et al. 1921; Walters
et al. 1998b). The Mucket is bradytictic, with the reproductive period extending from May to
August (Surber 1912).
53
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An Introduction to Freshwater Mussels as Biological Indicators
Pheasantshell
Actinonaias pectorosa
(Conrad, 1834)
Identification tips: Shell elliptical, moderately
thick, and moderately inflated. Posterior ridge
usually well defined. Periostracum greenish-yellow,
yellowish-tan, or brown; often with distinctive,
broken green rays. Beak sculpture of indistinct lines
(B); usually eroded. Teeth well developed. Nacre
white. Shell up to 7 inches in length.
Indicator use: A Cumberlandian endemic that has
experienced declines in portions of its range, the
Pheasantshell is considered a species of "special
concern" (Williams et al. 1993; Parmalee and
Bogan 1998).
GMAVs compiled and ranked by Augspurger
et al. (2003), Augspurger et al. (2006), and March
et al. (2007) placed Actinonaias among the
"intermediately tolerant" unionid genera to ammonia
and copper.
Keller and Augspurger (2005) evaluated the
toxicity of fluoride from feldspar mining operations
as a limiting factor in the recovery of the federally
endangered Appalachian Elktoe (Alasmidonta
raveneliana). Four unionid species were tested, with
96-hour LC50s of 178 ug/L and 303 ug/L observed
for juvenile A. pectorosa and A. raveneliana,
respectively. The researchers concluded that acute
toxicity from fluoride was unlikely, although further
sublethal tests at lower concentrations and longer
durations would be worthwhile.
Habitat: Endemic to the Cumberland River system
and Tennessee River system. The Pheasantshell
is a species of large creeks and rivers where the
water is shallow and flow is swift (Gordon and
Layzer 1989; Parmalee and Bogan 1998). Occurs in
sand, gravel, and cobble substrates.
Reproduction: Layzer and Khym (2005) identified several suitable hosts species, including
the Rock Bass, Smallmouth Bass, Largemouth Bass, Spotted Bass, Banded Sculpin, and
Sauger. Successful transformation was most often observed on Microptems spp. with
reported percentages of 35-43%. Several species (e.g. Etheostoma and cyprinids) sloughed
glochidia and were found to be unsuitable hosts (Layzer and Khym 2005).
54
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An Introduction to Freshwater Mussels as Biological Indicators
Dwarf Wedgemussel
Alasmidonta heterodon (I. Lea, 1830)
Identification tips: Shell subovate to
subtrapezoidal, somewhat elongate, moderately
thin, and moderately inflated. Periostracum
yellowish-tan to brown; green rays may be
present in young individuals (A). Beak elevated
slightly above hinge-line; sculpture of a few
conspicuous ridges (B), may be eroded. Teeth
well developed, although small (C); two lateral
teeth in right valve. Nacre bluish-white or white.
Shell up to 2 inches in length.
Indicator use: Widely distributed but rare
throughout its range, the Dwarf Wedgemussel is
currently listed as endangered by the U.S. Fish
and Wildlife Service. Like many other freshwater
mussel species, the Dwarf Wedgemussel is
considered relatively intolerant of impoundment
and chemical pollutants (Master 1986; USFWS
1993c).
GMAVs compiled and ranked March et al.
(2007) placed Alasmidonta heterodon among the
15 most sensitive taxa to copper.
Habitat: Distributed throughout the Atlantic
Slope, from North Carolina to New Brunswick,
Canada. Populations are often very patchily
distributed (Nedeau 2008). Occurs in headwaters
to rivers where the current is slow to moderate.
Generally found in stable muddy sand, sand,
or sand and gravel substrates (USFWS 1993c;
Nedeau 2008).
Reproduction: Confirmed host fishes include
the Atlantic Salmon, Mottled Sculpin, Tessellated Darter, and
Johnny Darter (Michaelson and Neves 1995; Wicklow 1999).
Michaelson and Neves (1995) reported glochidial metamorphosis
on 3 of 15 fish species.
The Dwarf Wedgemussel is bradytictic, with glochidia released
during April and May (Michaelson and Neves 1995; McLain and
Ross 2005).
55
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An Introduction to Freshwater Mussels as Biological Indicators
"
Elktoe
Alasmidonta marginata (Say, 1818)
Identification tips: Shell elongate to elliptical,
somewhat thin and inflated, with an abruptly angled
posterior (A). Periostracum yellowish-tan to brownish;
with wide green rays and flecks (A). Beak sculpture
of distinctive, heavy ridges (B); usually conspicuous
in both young and old individuals. Pseudocardinal
teeth weakly developed, lateral teeth present as a
thickened hinge (C). Nacre white. Shell up to 4 inches
in length. Notes: Orange foot. Live Elktoe are often
covered with periphyton (see upper right photo).
Indicator use: While widely distributed, the Elktoe
occurs sporadically throughout its range and is
considered a species of "special concern" (Williams
et al. 1993; USFWS 2007c). It is characteristic of
flowing streams of good quality and does not appear
to tolerate impoundment (Walters 1995; Parmalee
and Bogan 1998). Oesch (1984) reported that wastes
from mining activities reduced its abundance in the
Big River, MO.
Bedford et al. (1968) harvested Elktoe and
several other unionid species from the Red
Cedar River (Ml) for pesticide analysis. Reported
concentrations ranged from 0.0153-0.198 ppm
total DDT and metabolites. Concentrations did
not significantly vary between species. The study
concluded that mussels were excellent indicators
of pesticide contamination; mainly because of their
ability to concentrate contaminants.
Habitat: Distributed throughout the Mississippi
River basin, Great Lakes-St. Lawrence basin, and
Susquehanna drainage. Occurs in moderate-sized
streams to small rivers where the current is swift and streambed comprised
of sand, gravel, and small cobble substrates.
Reproduction: Identified host species include the White Sucker, Northern
Hog Sucker, Shorthead Redhorse, Rock Bass, and Warmouth Sunfish
(Howard and Anson 1922; Howard and Anson 1923).
The Elktoe is bradytictic, with the reproductive period extending from
June to July (Baker 1928).
56
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An Introduction to Freshwater Mussels as Biological Indicators
Threeridge
Amblema plicata
(Say, 1817)
Identification tips: Shell subquadrate or
subrhomboidal, thick, and somewhat compressed to
inflated. Periostracum light tannish-green (A), brown,
or dark brown; rayless. Beak nearly even to raised
above hinge-line; sculpture of a few coarse ridges
(B). Teeth well-developed (C); especially heavy in
older individuals. Nacre white. Shell up to 7 inches in
length.
Indicator use: A widely distributed species,
the Threeridge has demonstrated tolerance to
impoundment, pollutants, and regular harvest
(Starrett 1971; Oesch 1984; WDNR 2003). ESI
(2000) reported A. plicata as common to dominant
in several of the navigational pools of the upper
Ohio River. The Threeridge is currently considered
secure throughout its range (Williams et al. 1993;
Natu reserve).
Spooner et al. (2005) studied the physiological
effects of high water temperatures to eight species
of unionids, including A. plicata. The Threeridge
was the among the more tolerant unionid species
to high temperatures, exhibiting stress at 34-35°C
and maximal stress at 42°C. The authors suggested
the use of A. ligamentina as an indicator species in
lieu of more thermally tolerant species such as the
Threeridge.
The Threeridge has been used extensively
to biomonitor pollutants, for more information see
Adams et al. (1980; 1981), Naimo et al. (1992), and
Pip (1995).
Habitat: Widespread in the Mississippi basin, Great
Lakes-St. Lawrence basin, north into Canada, and
several Gulf drainages. The Threeridge occurs in small creeks to rivers, embayments,
and lakes. It may be found at depths ranging from 0.5-30 feet, in both fine and coarse
substrates.
Reproduction: Over 20 species of fish belonging to eight taxonomical families have
been identified as suitable hosts for A. plicata (Howard 1914; Wilson 1916; Coker et al.
1921; Stein 1968; Weiss and Layzer 1995).
57
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An Introduction to Freshwater Mussels as Biological Indicators
Purple Wartyback
Cyclonaias tuberculata
(Rafinesque, 1820)
Identification tips: Shell round or subquadrate,
thick, and moderately compressed. Shell surface
with numerous pustules, usually most abundant
medially and posteriorly (A). Periostracum brown to
dark brown, with well defined rest lines (A). Beak
even or slightly raised above hinge-line; sculpture of
numerous ridges (B). Teeth well developed (C); may
be particularly heavy in old individuals. Nacre purple
(C) or white. Shell up to 6 inches in length.
Indicator use: Widely distributed but sporadically
occurring, the Purple Wartyback is an inhabitant of
good quality creeks and rivers where the water is
clear (Walters 1995; Badra and Goforth 2002). C.
tuberculata has also been recorded from lentic or
impounded habitats, although the long-term viability
of these populations remains unclear (Bates 1962;
Parmalee and Bogan 1998). Williams et al. (1993)
considered it a species of special concern.
Imlay (1982) suggested of the use of C.
tuberculata as a biomonitor of heavy metals on the
basis of annual shell growth. This recommendation
was based on "widespread distribution, age, pollution
tolerance, and/or conchological reflection of stream
location."
Habitat: Fairly widespread throughout the Mississippi
River basin and Great Lakes basin. Occurs in both
creeks and rivers where the current is slow to swift.
May also be found in lentic habitats. Often most
abundant in a mixture of mud, sand, and gravel.
Reproduction: Confirmed host fishes include the
Black Bullhead, Yellow Bullhead, Channel Catfish,
and Flathead Catfish (Hove 1994b; Hove et al. 1997).
The Purple Wartyback is tachytictic, with the
reproductive period extending from June to August
(Ortmann 1919).
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An Introduction to Freshwater Mussels as Biological Indicators
Dromedary Pearlymussel
Dromus dramas (Lea, 1834)
Identification tips: A round, deep, thick, and slightly
inflated shell. Periostracum yellowish-tan with very
fine, broken green rays or lines; often more prominent
anteriorly (A). Beak sculpture of indistinct ridges; often
eroded (B). Teeth well developed and large (C). Nacre
white or pink. Shell up to 4 inches in length.
Indicator use: This once widespread and abundant
Cumberlandian species has declined dramatically during
the 20th century. A study by Hughes and Parmalee
(1999) reported the notable abundance of D. dromas at
prehistoric aboriginal sites along the Tennessee River.
Of the 159,450 freshwater mussel shells recovered
from 15 sites, the Dromedary Pearlymussel constituted
32% of the identified specimens. The widespread
impoundment of the Tennessee and Cumberland
drainages has probably contributed more to the decline
of D. dromas than any other factor, although dredging,
sand and gravel mining, coal mining, and sewage
wastes have also been implicated (USFWS 1983a;
Hughes and Parmalee 1999; Jones et al. 2004). This
species is now listed as endangered by the U.S. Fish
and Wildlife Service.
Habitat: Endemic to the Cumberland River and
Tennessee River drainages. Viable populations of D.
dromas occur in the Clinch and Powell rivers (Jones
et al. 2004). Historically, this species occurred in
moderate sized streams to rivers in shallow water (<3
ft.). Most often found in packed sand, gravel, and cobble
substrates.
Reproduction: Identified host fishes include numerous
darter and sculpin species, including the Black Sculpin,
Greenside Darter, Fantail Darter, Snubnose Darter, Tangerine
Darter, Blotchside Logperch, Logperch, Channel Darter, Gilt
Darter, and Roanoke Darter (Jones et al. 2004). Glochidia are
packed into 20-40 mm conglutinates that resemble leeches
or flatworms (Jones et al. 2004). D. dromas is bradytictic,
although the species shares some traits of tachytictic taxa
(Ortmann 1912; Jones et al. 2004).
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An Introduction to Freshwater Mussels as Biological Indicators
Eastern Elliptic
Elliptic complanata
(Lightfoot, 1786)
Identification tips: Shell variable; subtrapezoidal,
elongate, moderately thin to thick, and compressed
to slightly inflated. Periostracum yellowish-tan to
dark brown; green rays may be present in young
individuals. Beak roughly even to slightly elevated
above hinge-line; sculpture of several ridges, usually
eroded (C). Teeth well developed (D). Nacre white,
salmon, or purple. Shell up to 5 inches in length.
Indicator use: Widely distributed in eastern North
America, the Eastern Elliptic has been reported as a
habitat generalist that tolerates disturbance (Ortmann
1919; Johnson 1970; Nedeau 2008; Natureserve).
Of the freshwater mussels found in North
America, no other species has been so extensively
used as a biomonitor of pollutants (e.g. Campbell and
Evans 1991; Renaud et al. 1995; Beckvar et al. 2000;
Gagne et al. 2001; Mierzykowski and Carr 2001;
Gewurtz et al. 2002; Gewurtz et al. 2003; Martel et al.
2003). In addition, E. complanata has been frequently
employed to better understand the bioavailability,
bioaccumulation, and biotransformation of pollutants
(e.g. Day et al. 1990; Metcalfe-Smith 1994; Tessier et
al. 1994; Muncaster et al. 2002; Gewurtz et al. 2002;
O'Rourke et al. 2004; Thorsen et. al 2004).
Habitat: Distributed throughout the northern and
southern Atlantic Slope, Hudson Bay basin, and
parts of the Great Lakes-St. Lawrence basin. Occurs
in a wide variety of habitats, including creeks, rivers,
embayments, and lakes. May be found in both coarse
and fine substrates.
Reproduction: Confirmed host fishes include the Banded
Killifish, Green Sunfish, RedearSunfish, Orangespotted Sunfish,
Largemouth Bass, White Crappie, and Yellow Perch (Young 1911;
Walters 1994; Walters et al. 2005).
The Easlern Elliplio is lachyliclic, wilh Ihe reproduclive
period extending from May lo July. Orlmann (1919) reported lhal
Ihe glochidia of E. complanata are packaged in conglulinales.
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An Introduction to Freshwater Mussels as Biological Indicators
Spike
Elliptic dilatata
(Rafinesque, 1820)
Identification tips: Shell elongate, moderately thick,
and somewhat compressed. Periostracum yellowish-
tan to brown; young often with faint green rays (A).
Older individuals brown to dark brown and rayless.
Beak with distinctive, coarse ridges (C). Teeth well
developed. Nacre white, salmon, or purple (B). Shell
up to 6 inches in length.
Indicator use: A wideranging unionid that may be
locally abundant, the Spike is secure throughout its
range (Williams et al. 1993; Natureserve). Stansberry
(1965) considered the spike intolerant of water
pollution and Oesch (1984) reported a reduction in its
range where severe channelization and siltation had
occurred.
Wren and MacCrimmon (1983) analyzed the
bioaccumulation and biomagnification of metals in an
undisturbed lake in Ontario, Canada. Tissue samples
were collected from a number of fish, birds, mammals,
and the freshwater mussel - E. dilatata. The following
wet weight metal concentrations (ug/g) were reported:
Cd 5.8; Cu 2.0; Pb 4.5; Hg 0.17; Zn 78.5. In general,
E. dilatata bioaccumulated higher concentrations of
cadmium, copper, lead, and zinc than fish species.
However, mercury was found to biomagnify up the food
chain, with concentrations of 1.0 and 1.7 ug/g found
in the Northern Pike (Esox lucius) and Herring Gull
(Larus argentatus), respectively.
Habitat: Widespread in the Mississippi River basin
and Great Lakes-St. Lawrence basin. The Spike
occurs in small to large creeks, rivers, lakes, and
impoundments. Often most abundant in flowing
habitats where the streambed is comprised of stable
sand and gravel substrates.
Reproduction: Identified host fishes include the Flathead Catfish, Gizzard Shad,
Rock Bass, White and Black Crappie, Banded Sculpin, Rainbow Darter, Yellow Perch,
and Sauger (Howard 1914; Wilson 1916; Clark 1981; Luo 1993).
The Spike is tachytictic, with the reproductive period extending from mid-May to
August (Baker 1928).
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An Introduction to Freshwater Mussels as Biological Indicators
Oyster Mussel
Epioblasma capsaeformis (Lea, 1834)
Identification tips: General: Shell elliptical,
somewhat thin, and moderately inflated. Sexually
dimorphic: Females (A) expanded and rounded
posteriorly, males (B) elliptical and without expansion.
General: Periostracum yellowish-tan to light brown
with numerous green rays of varying widths. Beak
sculpture of indistinctive loops; often eroded. Teeth well
developed. Nacre white. Shell up to 3 inches in length.
Indicator use: The Oyster Mussel is listed as
federally endangered, occupying just a minute fraction
of its former range (USFWS 2004). Historically, it
was one of the most widely distributed and common
Cumberlandian species (Ortmann 1918; 1924;
1925). The decline of E. capsaeformis is most often
associated with impoundment, channelization, and
mineral extraction (Bates 1962; USFWS 2004).
Recent toxicological studies have evaluated
the sensitivity of the Oyster Mussel to a variety of
pollutants. For example, Valenti et al. (2006a) evaluated
the toxicity of chlorine to five unionid species, including
three federally listed taxa. Federally listed species
were reportedly more sensitive to TRC (total residual
chlorine) than two non-listed species. Mean 24-hr
LC50s for non-listed species ranged from 145 to 220
TRC ug/L, while federally listed LC50s ranged from 70
to 107 TRC ug/L. The reported 24-hr mean LC50 for E.
capsaeformis was 107 ug/L.
Habitat: Endemic to the Cumberland River system
and Tennessee River system. Occurs in moderate-
sized streams to rivers where the current is swift.
Generally most abundant at shallow depths (< 3 ft.)
where substrates are comprised of stable sand, gravel,
and cobble.
Reproduction: Identified host fishes include the Banded Sculpin,
Wounded Darter, Redline Darter, Spotted Darter and Dusky Darter
(Hill 1986; Yaeger and Saylor 1995).
The Oyster Mussel has been documented with mature glochidia
during May, June, and July (Gordon and Layzer 1989; Yaeger and
Saylor 1995).
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An Introduction to Freshwater Mussels as Biological Indicators
Northern Riffleshell
Epioblasma torulosa rangiana (Lea, 1838)
Identification tips: General: Shell moderately thick
and moderately inflated. Sexually dimorphic: Females
(A) expanded, rounded, and compressed posteriorly,
males (B) somewhat triangular and sulcate. General:
Periostracum yellowish-tan to light brown with fine green
rays. Beak sculpture of double-loops; often eroded or
indistinct (C). Teeth well developed (D). Nacre white.
Shell up to 3 inches in length.
Indicator use: This once widespread subspecies
has declined sharply and is now listed as federally
endangered, occupying less than 5% of its former
range (USFWS 2007). E.t. torulosa is thought to be
sensitive to siltation, impoundment, and water pollution
(USFWS 1994; USFWS 1997).Toxicity testing of a
congener (Epioblasma capsaeformis) suggests that the
Epioblasma may be among the most sensitive aquatic
genera to copper and ammonia.
Habitat: Occurs in the Mississippi River basin and
Great Lakes basin. Found in flowing reaches of large
creeks and rivers where substrates are comprised of
stable sand, gravel, and cobble. The largest remaining
populations occur in the Allegheny River and French
Creek, PA (USFWS 1994).
Reproduction: Identified host fishes include the Brown
Trout, Banded Darter, Bluebreast Darter, and Banded
Sculpin (Walters 1996a). Anecdotal evidence and recent
investigations suggest that E.t. rangiana may capture
host fishes to facilitate glochidia transfer (Barnhart 2006).
The Northern Riffleshell is likely bradytictic, with gravid
females having been observed in September (Ortmann
1919).
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An Introduction to Freshwater Mussels as Biological Indicators
Tubercled Blossom
Epioblasma torulosa torulosa
(Rafinesque, 1820)
Identification tips: General: Shell moderately thick
and moderately inflated; with a row of irregular medial
knobs often extending from the beak to ventral margin.
Sexually dimporphic: Females (A) greatly expanded
and rounded posteriorly, males (B) sulcate and with
a distinctive posterior-ventral indentation. General:
Periostracum yellowish-tan with fine green rays. Beak
sculpture of indistinct ridges, often absent in adults (C).
Teeth well developed (D). Nacre white. Shell up to 3
inches in length.
Indicator use: The Tubercled Blossom is a federally
endangered species and may now be extinct (Parmalee
and Bogan 1998; USFWS 2007a; USFWS 2007b). E.t.
torulosa formerly occurred throughout the Tennessee
River drainage, Cumberland River, and upper Ohio
River drainage (USFWS 1985a). Investigations of
archaeological shell middens along the Tennessee
River suggest that this species was once fairly common
in the Tennessee mainstem (Hughes and Parmalee
1999). It was last found fresh dead below Kanawha
Falls (WV) in 1969 (USFWS 2007a).
The decline of E.t torulosa (and many of its
congeners) has been attributed to ubiquitous river
impoundment, increased turbidity and siltation
associated with agriculture and deforestation, and
pollution (USFWS 1985a; USFWS 2007a).
Habitat: Formerly occurred in the Tennessee River
drainage, Cumberland drainage, and upper Ohio
drainage. E.t. torulosa has been found in shallow
reaches of rivers with swift current and substrates
comprised of stable sand and gravel.
Reproduction: Unknown.
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An Introduction to Freshwater Mussels as Biological Indicators
Wabash Pigtoe
Fusconaia flava (Rafinesque, 1820)
Identification tips: Shell highly variable; usually sub-
triangular, moderately thick, and moderately inflated.
Ventral margin of shell may have a velvet-like texture.
Big river forms usually with greater inflation and umbo
often more anterior. Periostracum yellowish-tan to brown,
rest lines distinctive; young often with fine green rays.
Beak sculpture of indistinct ridges as adult (B). Teeth well
developed (C). Nacre white, may have a salmon tinge.
Shell up to 4 inches in length.
Indicator use: A unionid that occurs in a broad range of
habitats, the Wabash Pigtoe is one of the most ubiquitous
mussels in North America. It reportedly adapts to a
variety of substrate types and habitat conditions (Walters
1995; Parmalee and Bogan 1998). Strayer (1983)
commented that the Wabash Pigtoe "seems to do well in
muddy, hydrologically unstable, low-gradient streams."
Spooner et al. (2005) studied the physiological
effects of high water temperatures on eight species of
unionids, including F. flava. The Wabash Pigtoe was
reported as among the more thermally tolerant species,
exhibiting stress at 34-35°C and maximal stress at 38°C.
The authors suggested the use of A. ligamentina as
an indicator species in lieu of more thermally tolerant
species, such as the Wabash Pigtoe.
Waller and Fisher (1998a) exposed F. flava and a
number of other unionids to various chloride salts. The
Wabash Pigtoe was among the most tolerant unionids
to the various salt treatments, while the Threehorn
Wartyback (Obliquaria reflexa) and Fragile Papershell
(Leptodea fragilis) were among the more sensitive.
Habitat: Occurs throughout the Mississippi River basin,
Great Lakes-St. Lawrence basin, and Mobile basin. Found
in creeks, rivers, embayments, and lakes. Often most
abundant where substrates are comprised of stable sand,
gravel and cobble, although F. flava may also occur in fine
substrates.
Reproduction: Identified host fishes include the Creek
Chub, Silver Shiner, Bluegill Sunfish, Black Crappie, and
White Crappie (Howard 1914; Wilson 1916; Walters and
O'Dee 1997). The Wabash Pigtoe is lachyliclic, wilh Ihe
reproduclive period extending from May lo August
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An Introduction to Freshwater Mussels as Biological Indicators
Pink Mucket
Lasmpsilis abrupta (=orbiculata) (Say, 1831)
Identification tips: General: Shell elliptical or
subquadrate, thick, and slightly inflated to inflated.
Sexually dimorphic: Females expanded and inflated
posteriorly (A), males bluntly pointed (B). General:
Periostracum yellowish-tan or brown; often rayless,
although faint rays may be seen near the beak (A) and
in young individuals. Beak sculpture of indistinct double-
loops; often eroded (C). Teeth well developed (D);
pseudocardinal teeth large and heavy in older individuals.
Nacre white or pink. Shell up to 5 inches in length.
Indicator use: Although widely distributed, the Pink
Mucket remains imperiled or critically imperiled throughout
its range (Natureserve). Collections made by early
naturalists suggest that L. abrupta may have also been
uncommon in the recent past (USFWS 1985c).The Pink
Mucket is currently listed as endangered by the U.S. Fish
and Wildlife Service (USFWS 1985c; USFWS 2007).
Wang et al. (2007a) evaluated the toxicity of copper,
and ammonia to the early life stages of L. abrupta and
several other unionid species. Short-term exposures of
copper to L. abrupta glochidia yielded a 24-hr EC50 of
34 ug Cu/L, a test result comparable to other species.
Juvenile (2-month old) L abrupta were slightly more
sensitive than other test species during 96-hour exposures
to ammonia, with a reported EC50s of 2.3 mg N/L.
Habitat: Widespread throughout the Mississippi Basin,
Tennessee River, Cumberland River, and Ohio River
drainage (USFWS 1985c).The Pink Mucket is found in
small to large rivers where substrates are comprised
of stable sand, gravel, and cobble. L. abrupta typically
inhabits shallow, swiftly flowing areas, which may include
the tailwaters and upper reaches of navigational pools
where adequate habitat is present (USFWS 1985).
Reproduction: Identified host fishes include the
Smallmouth Bass, Largemouth Bass, Spotted Bass, and
Walleye (Barnhart et al.1997). The confirmed hosts of a
relative, Lampsilis higginsi, include the Sauger, Freshwater
Drum, and Yellow Perch, in addition to some of the fishes
named above (Surber 1913; Waller and Holland-Bartels
1988).
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An Introduction to Freshwater Mussels as Biological Indicators
Plain Pocketbook
Lampsilis cardium (Rafinesque, 1820)
Identification tips: General: Shell moderately thick
to thick and inflated. Sexually dimorphic: Females (A)
inflated and rounded posteriorly, males (B) pointed
posteriorly. General: Periostracum yellowish-tan to
brown; with few to numerous green rays (A-B). Beak
elevated above hingeline; sculpture double-looped (C),
often more distinctive in younger individuals. Teeth well
developed. Nacre white. Shell up to 7 inches in length.
Indicator use: The Plain Pocketbook is a widespread
species that occurs in a variety of habitats. In
Tennessee, it has been reported to tolerate
impoundment, poor water quality, and substrates
comprised of mud and silt (Parmalee and Bogan 1998).
In an effort to assess lead and cadmium
contamination from tailing ponds and chat piles in the
Big River watershed (MO), Czarnezki (1987) used
L. cardium to biomonitor contaminant levels at five
sites. The mussels were found to accumulate metals
quickly and assisted in identifying the main sources of
contamination.
Under laboratory conditions, juvenile L. cardium
exhibited sensitivity to ammonia during acute pore
water exposures (Newton et al. 2003). Fifty percent
mortality was observed at concentrations as low as 93
ug NH3-N/L and sublethal effects (50% reduced growth)
occurred at 31 ug NH3-N/L. An in-situ companion
study performed by Bartsch et al. (2003) deployed
caged juveniles below the substrate surface in the St.
Croix River, Wl. Survival and growth were reported as
highly variable and generally unrelated to ammonia
concentrations found at the study sites.
Habitat: Widely distributed throughout the Mississippi
River basin, Great Lakes-St. Lawrence Basin, and
north into Canada. Occurs in moderate-sized streams
to rivers, lakes, reservoirs, and coastal marshes. Found
in a variety of substrates, from sand, gravel, and cobble
to mud and silt.
Reproduction: Identified host fishes include the Pumpkinseed Sunfish, Bluegill
Sunfish, Green Sunfish, Largemouth Bass, Smallmouth Bass, White Crappie, Black
crappie, Yellow Perch, Sauger and Walleye (Coker 1921; Waller et al. 1985; Walters
1996b; Draxler et al. 2006). The Plain Pocketbook utilizes a minnow-like lure to attract
host fishes (partially visible in upper right photo).
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An Introduction to Freshwater Mussels as Biological Indicators
Wavyrayed Lampmussel
Lasmpsilis fasciola (Rafinesque, 1820)
Identification tips: General: Shell oval, moderately
thick, and moderately inflated to inflated. Sexually
dimorphic: Females (A) expanded and rounded
posteriorly, males (B) bluntly pointed. General: Shell
periostracum with distinctive, wavy green rays (A-
B). Beak slightly raised above hinge-line; sculpture
of somewhat irregular double-loops (C). Teeth well
developed. Nacre white. Shell up to 5 inches in length.
Indicator use: The Wavyrayed Lampmussel is a
species of hydrologically stable and good quality
streams (Strayer 1983; Walters 1995). It may be
locally abundant when found and is considered stable
throughout its range (Williams et al. 1993; Natureserve).
Several researchers have used L fasciola as a test
organism when evaluating the sensitivity of unionids
to contaminants (e.g. Jacobson et al. 1997; Cherry
et al. 2002; Keller and Augspurger 2005; Valenti et al.
2006; Bringolf et al. 2007; Wang et al. 2007a; 2007b).
Wang et al. (2007) conducted a series of ammonia
toxicity trials using L. fasciola glochidia and juveniles,
in addition to the early life stages of several other
unionid species. Reported EC50s for Wavyrayed
glochidia ranged from 6.2 to 8.7 mg N/L during 24-hour
exposures. These results were generally comparable to
the values reported for other species, with the exception
of two particularly sensitive species - the Ellipse
(Venustaconcha ellipsiformis) and Oyster Mussel (E.
capsaeformis). GMAVs compiled and by Augpsurger
et al. (2003), Augspurger et al. (2003), and March et al.
(2007) rank L. fasciola among the most sensitive aquatic
species to ammonia and copper.
Habitat: Occurs disjunctly throughout the Mississippi
River basin and Great Lakes basin. Generally found in
creeks to small rivers at shallow depths in stable sand
and gravel. This species has also been documented
to occur at depths of 15 feet or more in the Tennessee
River and Allegheny River.
Reproduction: Identified host fishes include the
Smallmouth Bass (Zale and Neves 1982). The
Wavyrayed Lampmussel utilizes a mantle flap lure (upper right photo) to entice
host fishes. The striking action of predators triggers the expulsion of hundreds to
thousands of glochidia, thereby facilitating the parasitization of the host fish.
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An Introduction to Freshwater Mussels as Biological Indicators
Fatmucket
Lampsilis siliquoidea
(Barnes, 1823)
Identification tips: General: Shell moderately thick
and somewhat compressed to moderately inflated.
Sexually dimorphic: Females (A) inflated and rounded
posteriorly, males (B) bluntly pointed posteriorly.
General: Periostracum yellowish-tan to brown; with
few to numerous green rays (A-B). Beak elevated
slightly above hinge-line; sculpture double-looped (C),
often more prominent in younger individuals. Teeth well
developed. Nacre white. Shell up to 5 inches in length.
Indicator use: Widely distributed and often abundant,
the Fatmucket has been described as tolerant of silt
and disturbance (Dawley 1947; Walters 1995; McRae
etal.2004).
Manny and Kenaga (1991) reported the use of
L. siliquoidea (= L. radiata siliquoidea, see Turgeon
et al. 1998) as a biomonitor of heavy metals, PCBs,
and octachlorostyrene in the Detroit River, a heavily
urbanized watershed. Researchers found toxic
contaminants concentrated by L. siliquoidea to levels
59 times nearby sediments (GLI 1984; Pugsley et al.
1985).
Bringolf et al. (2007b) investigated the toxicity
of several forms of glyphosate, its formulations, and
a surfactant (MON 0818) to juvenile and glochidial
Fatmucket (L. siliquoidea). Reported 24-hour EC50s
were as low as 3.0 mg/L and 0.6 mg/L for Roundup
and MON 0818, respectively. Reported 96-hour
EC50s for juvenile L. siliquoidea were 5.9 mg/L and
3.8 mg/L for Roundup and MON 0818, respectively.
Researchers concluded that the early life stages of
the Fatmucket are among the most sensitive aquatic
organisms to glysphosate-based chemicals and MON
0818 tested to date.
Habitat: Widely distributed throughout the Mississippi
River basin, Great Lakes-St. Lawrence drainage, and
north into Canada. Occurs in a wide variety of habitats,
including creeks, rivers, lakes, and backwaters. Generally most abundant in sluggish to
moderate currents, in both fine and coarse substrates.
Reproduction: A total of 20 fishes have been identified as suitable host species for
L. siliquoidea (Evermann and Clark 1918; Coker et al. 1921; Fuller 1978; Trdan 1981;
Walters and O'Dee 1997; O'Dee and Wallers 2000; Wallers el al. 2005).
The Falmuckel is bradyliclic, wilh Ihe reproduclive period extending from Augusl lo
July (Baker 1928).
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An Introduction to Freshwater Mussels as Biological Indicators
White Heelsplitter
Lasmigona complanata complanata (Barnes 1823)
Identification tips: Shell large, compressed, and
moderately thin to moderately thick; with a distinctive,
often ribbed dorsal wing (A-B). Periostracum light to dark
brown; usually with fine green rays in young individuals.
Beak sculpture of conspicuous double-loops (C).
Pseudocardinal teeth well developed although low and
irregularly serrated; lateral teeth present as an under
developed raised and striated ridge. Nacre white. Shell
up to 8 inches in length.
Indicator use: This wideranging unionid has been
reported to be tolerant of silt, habitat disturbance, and
impoundment by numerous authors (Stansberry 1965;
Walters 1995; Dean et al. 2002; Metcalfe-Smith et al.
2003; Sietman 2003). Stayer and Jirka (1997) noted
that L.c. complanata was "one of the few unionoids that
seemed to do well in disturbed substrates."
The White Heelsplitter may potentially exploit areas
that other unionids find unfavorable for colonization.
For example, in the early 1900s, Goodrich and van der
Schalie (1932) observed the colonization of Michigan's
Saginaw Bay by L.c. complanata during a period of
eutrophication resulting from increased sewage wastes.
Metcalfe-Smith et al. (2003) recently noted the range
expansion of L.c. complanata in the Sydenham River,
Ontario. The researchers speculated that the rapid
expansion of such an opportunistic unionid indicated
deteriorating conditions within the study area.
Habitat: Distributed throughout the Mississippi River
basin, Great Lakes-St. Lawrence basin, and north
into Canada. Occurs in creeks, rivers, reservoirs, lakes, and
embayments. Adapts well to turbid waters. Tolerates a wide range
of substrates, including unstable sand and gravel, silt, and mud.
Reproduction: Identified host fishes include the Common
Carp, Banded Killifish, Green Sunfish, Orangespotted Sunfish,
Largemouth Bass, and White Crappie (Lefevre and Curtis 1910;
Young 1911).
The White Heelsplitter is bradytictic, with the reproductive
period extending from September to May (Baker 1928).
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An Introduction to Freshwater Mussels as Biological Indicators
Flutedshell
Lasmigona costata
(Rafinesque, 1820)
Identification tips: Shell elongate, moderately
thick, and somewhat compressed; with several
undulations or "flutes" present on the posterior slope
(A-B). Periostracum brownish with striking green
rays in young individuals; adults often dark brown
and rayless. Beak even with hinge-line; sculpture of
a few concentric ridges that are often eroded (C).
Pseudocardinal teeth distinctive, low, and heavy (D).
Lateral teeth weakly developed (D). Nacre white.
Shell up to 7 inches in length.
Indicator use: While not often locally abundant,
this widespread species is secure throughout its
range (Williams 1993; Natureserve). Kidd (1973)
documented an increase in L costata in the the
polluted waters of the Grand River, indicating that
this species may be capable of colonizing disturbed
or polluted habitats.
Tetzloff (2001) investigated the survival rates
of unionid species following a low dissolved oxygen
(DO) event in Big Darby Creek, OH. The event
was initiated by an agribusiness spill of molasses,
fermented grain, and other organic substances.
The Flutedshell, a common species within the
study area, experienced lower rates of mortality
than several other species at the site (-10%).
The Kidneyshell (P. fasciolaris) and Wavyrayed
Lampmussel (L. fasciola) were among the most
sensitive species to the event.
Habitat: Widespread throughout the Mississippi
River basin and Great Lakes-St. Lawrence basin.
The Flutedshell is found in small creeks to rivers
where substrates are comprised of stable sand,
gravel, and cobble. While usually most abundant in
shallow water, L. costata has been documented from
depths of 15-20 feet.
Reproduction: Identified host species include the Creek
Chub, Central Stoneroller, Longnose Dace, Common Carp,
Northern Hog Sucker, Bluegill Sunfish, Pumpkinseed
Sunfish, Largemouth Bass, and Banded Darter (Lefevre
and Curtis 1910; Walters et al. 1998c; Walters et al. 2005).
The Fluledshell is lachyliclic, wilh Ihe reproduclive
period extending from September unlil May (Baker 1928).
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An Introduction to Freshwater Mussels as Biological Indicators
Black Sandshell
Ligumia recta (Lamarck, 1819)
Identification tips: General: Shell elongate,
thick, and moderately inflated. Sexually dimorphic:
Females obliquely flared posteriorly (A), males
bluntly pointed (B). General: Periostracum dark
brown to black, smooth, and lustrous; young
individuals often with brillant green rays. Beak
raised slightly above hinge-line; sculpture of a few
indistinct ridges that are often eroded. Teeth well
developed; lateral teeth distinctive, long, narrow,
and straight (C). Nacre white; often with purplish
tinge near beak cavity and pseudocardinal teeth.
Shell up to 9 inches in length.
Indicator Use: Although widespread, L. recta
is relatively uncommon throughout its range and
has significantly declined in some states (e.g.
Kansas, Illinois). Khym and Layzer (2000) observed
that the "decline and low abundance of L. recta
is concordant with a decline in sauger (Sander
canadensis) runs," a suitable host species. Williams
et al. (1993) considered the Black Sandshell a
species of "special concern".
Pip (1995) evaluated the use of L. recta and
eight other unionids as biomonitors of heavy metals
in the Assiniboine River, Manitoba, Canada. Of the
nine unionids studied, the Black Sandshell and
Mapleleaf (Quadrula quadrula) were determined
to have sequestered the highest concentrations of
copper.
Habitat: Known from the Mississippi River basin,
Great Lakes-St. Lawrence River basin, a few Gulf
Coast drainages, and north into Canada. The Black
Sandshell is found in large creeks, rivers, and lakes
where substrates consist of mud, sand, gravel, or
cobble. In riverine habitats, it occurs in both current and slackwater areas.
Reproduction: Identified host species include the Banded Killifish, Green Sunfish,
Bluegill Sunfish, Orangespotted Sunfish, White Crappie, Largemouth Bass, and Sauger
(Young 1911; Surber 1913; Pearse 1924; Walters 1994; Hove 1994; Barnhart 2000; Khym
et al. 2000). Khym and Layzer (2000) reported poor glochidia metamorphosis in Black
and White Crappie, Bluegill Sunfish, and Largemouth Bass (<5%). Conversely, Sauger
proved an excellent host, with 53% of glochidia transforming into juveniles.
The Black Sandshell is bradytictic, with the reproductive period extending from
August to July (Ortmann 1919).
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An Introduction to Freshwater Mussels as Biological Indicators
Cumberland Moccasinshell
Medionidus conradicus (Lea, 1834)
Identification tips: Shell small, elongate,
moderately thin, and compressed. Periostracum
glossy and yellowish-tan with numerous green rays
(A). Beak sculpture of fine, broken double-looped
ridges (B); usually eroded in adults. Teeth well
developed although delicate (C). Nacre bluish-
white, often iridescent. Shell up to 3 inches in
length.
Indicator use: A Cumberlandian endemic
that has experienced declines in portions of
its range (Williams et al. 1993; Natureserve),
the Cumberland Moccasinshell occurs in
unimpounded, high quality stream and river
reaches. Williams et al. (1993) considered M.
conradicus a species of "special concern".
Jacobson et al. 1997 evaluated the toxicity of
copper to the glochidial stages of M. conradicus
and four additional unionid species. Reported
LC50s for released glochidia ranged from 37 to
81 ug Cu/L during 24-hour exposures. Similar
results were documented by Cherry et al. (2002),
who reported an acute mean LC50 of 41 ug Cu/L.
GMAVs compiled and ranked by Augspurger (2006)
and March et al. (2007) placed Medionidus among
the most sensitive aquatic genera to copper.
Cherry et al. (1991) and McCann (1993)
assessed the acute toxicity of zinc to the early life
stages of M. conradicus and several other unionid
species. Reported LC50s ranged from 492 to 570
ug Zn/L during 48-hour exposures (hardness = 60
to 170).
Habitat: Endemic to the Cumberland River
system and Tennessee River system. Occurs in small streams to small rivers where
the current is swift. Generally most abundant at shallow depths (< 3 ft.), where
substrates are comprised of stable sand, gravel, cobble, and slab rock (Gordon and
Layzer 1989; Parmalee and Bogan 1998).
Reproduction: Identified host fishes include the Warmouth Sunfish, Redline
Darter, Fantail Darter, Striped Darter, and Rainbow Darter (Stern and Felder 1978;
Zale and Neves 1982; Luo and Layzer 1993).
Cumberland Moccasinshell glochidia have been documented in Big Moccasin
Creek (VA) drift from January to June, September to October, and infrequently
during November (Zale and Neves 1982).
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An Introduction to Freshwater Mussels as Biological Indicators
Threehorn Wartyback
Obliquaria reflexa (Rafinesque, 1820)
Identification tips: Shell round, moderately thick,
and moderately inflated; anterior of shell noticeably
thicker than posterior. A single row of large medial
knobs extends from the umbo to dorsal margin (A).
Knobs alternate in position on left and right valve (B).
Posterior edge with fine ribs or small undulations (C).
Shell periostracum variable, usually tan or green with
fine green rays. Beak sculpture of fine, indistinctive
lines often with small knob near hinge-line. Teeth
well developed (D). Nacre white. Shell up to 3 inches
in length.
Indicator use: This wideranging unionid has
exhibited tolerance to river impoundment and a wide
variety of substrates. It often sustains or increases
in relative abundance where impoundment has
occurred (Fuller 1985; Parmalee and Bogan 1998;
WDNR 2003). ESI (2000) consistently reported O.
reflexa as a dominant species in the impounded
pools of the upper Ohio River.
Levengood et al. (2004) evaluated contaminant
concentrations in freshwater mussels at the
confluence of the Mississippi River and Illinois
River. Five regionally abundant mussels, including
O. reflexa, were used to evaluate contaminant
burdens of local unionids. Cadmium and arsenic
were below detection limits in O. reflexa, while most
EPA priority metals were accumulated to comparable
concentrations by Amblema plicata, Obovaria
olivaria, and O. reflexa upstream of the Mississippi-
Illinois confluence. Threehorn wartyback were not
collected below the confluence.
Habitat: Widespread in the lower and upper
Mississippi basin, Great Lakes basin, and Gulf
drainages. Found in small to large rivers, lakes,
reservoirs, and embayments. Often most abundant
in substrates comprised of mud, sand, and gravel
where the current is slow to moderate.
Reproduction: Identified host fishes include the Common Shiner, Longnose Dace, and
Silverjaw Minnow (Walters et al. 1998). However, only a few juveniles were recovered from
the above species, giving credence to the possibility that the true host(s) has yet to be
found.
The Threehorn Wartyback is tachytictic, with the reproductive period extending from
June to August (Utterback 1915).
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An Introduction to Freshwater Mussels as Biological Indicators
Sheepnose
Plethobasus cyphyus (Rafinesque, 1820)
Identification tips: Shell subovate, thick, and
moderately inflated; often with a row of medial
tubercles extending from below the umbo to the
ventral margin. The tubercles may vary in number,
size, and shape. Periostracum yellowish-tan and
rayless (A) usually brown to dark brown in old
individuals. Beak slightly raised above hinge-line;
sculpture of indistinct ridges, often eroded in adults
(B). Teeth well developed (C). Nacre white. Shell up to
5 inches in length.
Indicator use: Once widespread in the small
and large rivers of the Mississippi drainage, the
Sheepnose now occupies just 34% of its former
range (Butler 2002). It is currently a candidate
for federal listing and is considered threatened,
imperiled, or critically imperiled throughout its range
(Cummings and Mayer 1992; Williams et al. 1993;
Natureserve). Interestingly, archaeological evidence
suggests that this species was rare in some systems
historically (Parmalee et al. 1980; Parmalee and
Hughes 1994; Hughes and Parmalee 1999; Butler
2002).
Perhaps the largest contributing factor to the
decline of P. cyphyus has been habitat fragmentation
and associated hypolimnetic releases resulting
from the construction of dams (Butler 2002). Other
suspected contributors include large channel
maintenance, mine drainage, in-stream gravel mining,
sedimentation, turbidity, and chemical contaminants
(Hartfield and Hartfield 1996; Butler 2002).
Habitat: Widespread in the upper and lower
Mississippi River, Ohio River, Tennessee River, and
Cumberland River drainages. Occurs in small to large
rivers where the current is moderate to swift. Often
found at shallow depths (<3 ft.) where the streambed
is comprised of sand and gravel substrates
(Parmalee and Bogan 1998).
Reproduction: Identified host fishes include the
Central Stoneroller Minnow and Sauger (Wilson
1916; Walters et al. 2005). Glochidia are packaged in "narrow, lanceolate shapes which
are solid, red and discharge in unbroken form" (Oesch 1984).
The Sheepnose is tachytictic, with the reproductive period occurring in early
summer (Parmalee and Bogan 1998).
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An Introduction to Freshwater Mussels as Biological Indicators
Clubshell
Pleurobema clava
(Lamarck, 1819)
Identification tips: Shell triangular, elongate, and
moderately inflated; noticeably thicker anteriorly
than posteriorly. The shell is perhaps best described
as "wedge-shaped" (Walters 1995). Periostracum
yellowish-tan; umbo with prominent, broken green
rays (upper right photo) (A). Rays generally less
conspicuous in older individuals. Beak slightly
raised above hinge-line and set anteriorly; sculpture
of indistinct ridges, usually eroded (B). Teeth well
developed (C). Nacre white. Shell up to 3 inches in
length.
Indicator use: The Clubshell was formerly
widespread and abundant throughout the upper
Ohio River drainage and western Lake Erie
tributaries. It now occupies less than 5% of its
former range (USFWS 1993; Natureserve). The
dramatic decline of P. clava has been attributed to
siltation, impoundment, in-stream sand and gravel
mining, pollutants, and exotic species (USFWS
1993a; USFWS 1994). While there is little doubt the
Clubshell is a sensitive species, the causal factors
of decline have not been evaluated, quantitatively
or otherwise, for their individual impacts. It is now
listed as an endangered species by the U.S. Fish and
Wildlife Service.
Habitat: Formerly widespread in the upper Ohio
River basin and the western tributaries of Lake Erie.
Occurs in creeks and rivers where the streambed
is comprised of clean sand, gravel, and small
cobble substrates. Many authors have reported on
its propensity to burrow deep into the streambed
(Ortmann 1919; Walter 1995; USFWS 1997),
although il may also be found in ihe more lypical life
posilion assumed by mosl unionids.
Reproduction: Idenlified hosl fishes include Ihe
Cenlral Sloneroller Minnnow, Slriped Shiner, Blackside
Darler, and Logperch (Wallers and O'Dee 1997; O'Dee and
Wallers 2000).
The Clubshell is apparenlly lachyliclic, wilh Ihe
reproduclive season extending from May lo mid-June
(Orlmann 1919).
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An Introduction to Freshwater Mussels as Biological Indicators
Ohio Pigtoe
Pleurobema cordatum
(Rafinesque, 1820)
Identification tips: Shell subtriangular, deep, heavy, and
inflated; usually with a shallow sulcus. Periostracum light
brown to reddish-brown; young individuals may have faint
green rays. Rest lines prominent (A). Beak raised and
turned forward (A-B); sculpture generally indistinct or of
elevated ridges (Cummings and Mayer 1992). Teeth well
developed (C). Nacre white. Shell up to 4 inches in length.
Indicator use: The Ohio Pigtoe is a large river species
that is of conservation concern throughout its range
(Williams 1993; Roe 2002; Natureserve). Parmalee
and Bogan (1998) reported P. cordatum to be intolerant
of impounded reservoirs in Tennessee. Conversely,
Gordon and Layzer (1989) commented that it may be
somewhat tolerant of lentic habitats. Investigations into
the impounded pools of the upper Ohio River found P.
cordatum to be "fairly abundant" at Greenup and Meldahl
(ESI 2000). Thus, the tolerance of this unionid (and
hosts) to impoundment may depend on local habitat
conditions or undetermined factors. It is also suspected
to be sensitive to poor water quality, siltation, and zebra
mussels (Roe 2002).
Chen (1998) analyzed the ability of P. cordatum
and several other unionid species to regulate oxygen
consumption (OC) with declining dissolved oxygen levels.
The study results suggested that P. cordatum was likely
vulnerable to hypoxia due to its poor ability to regulate OC.
Habitat: Distributed throughout the upper Mississippi
River, Ohio River, Cumberland River, and Tennessee
River basins. Generally occurs in flowing sections of
medium to large rivers in shallow or deep water (up to
25 ft.). Often found in stable sand, gravel, and cobble
substrates.
Reproduction: Identified host fishes include the Guppy,
Brook Stickleback, Creek Chub, Rosefin Shiner, and
Bluegill Sunfish (Fuller 1974; Walters and Kuehnl 2004).
The Ohio Pigtoe is apparently tachytictic, with the
reproductive period occurring in late spring to summer.
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An Introduction to Freshwater Mussels as Biological Indicators
Round Pigtoe
Pleurobema sintoxia
(Rafinesque, 1820)
Identification tips: Shell highly variable; generally
round to subquadrate and moderately thick. Shell
inflation variable; often compressed in smaller
systems and inflated in large rivers. Periostracum
yellowish-tan, reddish-brown, or dark brown; young
individuals may have faint rays. Rays often absent in
older individuals. Beak sculpture somewhat indistinct,
usually of irregular ridges. Beaks often low in creek
and small river forms; higher in large river forms.
Teeth well developed (B). Nacre white or pink (B).
Shell up to 5 inches in length.
Indicator use: The Round Pigtoe is widely
distributed although rarely abundant throughout its
range (Cummings and Mayer 1992; Sietman 2003;
WDNR 2003; Natureserve). It most commonly
occurs in areas of current and good water quality
(Walters 1995; Parmalee and Bogan 1998; Zanatta
and Metcalfe-Smith 2004). P. sintoxia has apparently
tolerated impoundment in some rivers, although
this may be dependent on localized conditions. For
example, while historically reported from the Upper
Ohio River, recent sampling has found it to be rare
with no evidence of recent recruitment (Taylor 1989;
ESI 2000). Populations of P. sintoxia may also be
vulnerable to zebra mussel infestations, as recent
reports suggest from the Detroit River (tributary
between Lake St. Clair and Lake Erie) (Schloesser et
al.2006).
Habitat: Widespread in the Mississippi River basin
and lower Great Lakes. Often most abundant in
large creeks or rivers with moderate current (Walters
1995; Parmalee and Bogan 1998). However, it is also
known from Lake Erie and Lake St. Clair. Occurs in
packed mud, sand, and gravel subslrales (Oesch
1984; Parmalee and Bogan 1998).
Reproduction: Idenlified hosl fishes include Ihe
Cenlral Sloneroller Minnow, Spolfin Shiner, Norlhern
Redbelly Dace, Soulhern Redbelly Dace, Blunlnose Minnow, and Bluegill Sunfish
(Surber 1913; Coker el al. 1921; Hove 1995a; Hove el al. 1997; Wallers el al. 2005).
The Round Pigloe is apparenlly lachyliclic, wilh Ihe reproductive season
extending from May lo late July (Baker 1928).
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An Introduction to Freshwater Mussels as Biological Indicators
Kidneyshell
Ptychobranchus fasciolaris
(Rafinesque, 1820)
Identification tips: Shell somewhat elongate
and sub-elliptical, moderately thick to thick, and
compressed to slightly inflated. Periostracum yellow,
yellowish-tan, or light brown; often with numerous
broken green rays (A). Beak sculpture of indistinct
ridges; usually eroded (B). Beak roughly even
to slightly elevated above hinge-line. Teeth well
developed (C). Nacre white. Shell up to 6 inches in
length.
Indicator use: Widely distributed but sporadically
occurring, the Kidneyshell may be locally abundant
in good habitat. It has reportedly tolerated
impoundment in Tennessee (Parmalee and Bogan
1998), but is more typical of shallow, flowing creeks
and rivers (van der Schalie 1938; Gordon and
Layzer 1989; Walters 1995). Williams et al. (1993)
considered it secure throughout its range.
Positive associations have been documented
between Kidneyshell populations and other species
of unionids, including the Spike (Elliptic dilatata),
Wavyrayed Lampmussel (Lampsilis fasciola),
Clubshell (Pleurobema clava), and Rainbow (Villosa
iris) (van der Schalie 1938; Metcalfe-Smith et al.
1999; Badra and Goforth 2001). These reports
suggest the use of the Kidneyshell as an indicator of
suitable conditions for the presence or reintroduction
of the species listed above.
Tetzloff (2001) investigated the survival rates
of unionid species following a low dissolved oxygen
(DO) event in Big Darby Creek, OH. The Kidneyshell,
an abundant species within the study area, was
reported with the Wayrayed Lampmussel (L. fasciola)
as experiencing the highest mortality rates (-95%).
Habitat: Fairly widespread throughout the Mississippi River basin and Great
Lakes basin. Occurs in creeks and rivers where the current is moderate to
swift. Often most abundant in sand and gravel substrates.
Reproduction: Confirmed host fishes include the Brook Stickleback (Walters
et al. 2005). Hosls observed infecled in Ihe field include Ihe Johnny Darler,
Fanlail Darler, Greenside Darler, and Banded Darler (While el al. 1996).
The Kidneyshell is bradyliclic, wilh glochidia overwinlering in Ihe marsupia
(Orlmann 1919).
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An Introduction to Freshwater Mussels as Biological Indicators
Fluted Kidneyshell
Ptychobranchus subtentum (Say, 1825)
Identification tips: Shell elongate, moderately thick,
with a fluted or undulating posterior dorsal slope (A-
B). Periostracum usually yellowish-tan with few to
several wide green rays (A). Rays generally fade with
age. Beak sculpture of indistinct loops; often eroded
(C). Teeth well developed (D). Nacre white. Shell up
to 5 inches in length.
Indicator use: The Fluted Kidneyshell has
undergone a drastic range reduction in the past 100
years, a result of stream and river impoundment,
toxic contaminants, and siltation (Butler 2005a). This
sensitive species is currently a candidate for federal
listing.
Habitat: Endemic to the Cumberland River system
and Tennessee River system. Occurs in moderate-
sized streams to small rivers where the current is
swift. Generally most abundant at shallow depths
(< 2 ft.) where substrates are comprised of packed
sand and gravel. This species requires flowing, well-
oxygenated waters to thrive (Butler 2005a).
Reproduction: Identified host fishes include
many riffle-dwelling species, including the Banded
Sculpin, Redline Darter, Rainbow Darter, Barcheek
Darter, and Fantail Darter (Luo and Layzer 1993).
The Fluted Kidneyshell utilizes a fascinating tool for
reproduction: a gelatinous capsule shaped like insect.
These "conglutinates" are packed with glochidia and
liberated into the stream current where they adhere
to stones. Unsuspecting host fishes then strike the
conglutinates, infecting their gill tissue with glochidia.
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An Introduction to Freshwater Mussels as Biological Indicators
Giant Floater
Pyganodon grandis
(Say, 1829)
Identification tips: Shell elongate, thin, and inflated
(A) Periostracum yellow, yellowish-tan, or green when
young; often with faint green rays. Older individuals light
brown to dark brown; usually rayless. Beak elevated
above hinge-line; sculpture of pronounced double-loops
(B). Teeth poorly developed; hinge thickened. Nacre
white; beak cavity may be tinged with salmon (C). Shell
up to 9 inches in length.
Indicator use: This wideranging unionid adapts well
to impoundments and tolerates moderate levels of silt
(Walters 1995; WDNR 2003). Pip (2006) described the
Giant Floater as one of the most tolerant unionids in
Manitoba, Canada. It may also persist in soft substrates
where heavier shelled species sink or suffocate (Coker
1921).
GMAVs compiled and ranked by Augspurger et
al. (2003) and Augspurger (2006) placed Pyganodon
among the more sensitive aquatic genera to copper and
ammonia.
Black et al. (1995) exposed the Giant Floater
to lead in the laboratory and field and found that
DMA strand breakage occurred only at the lowest
exposure level (50 ug/L). They attributed this to higher
lead concentrations inducing DMA repair processes,
whereas lower concentrations may have not initiated or
maximized repair processes. A secondary hypothesis
was localized cellular necrosis (cell death and
breakage) of the foot.
The Giant Floater has been used extensively as
a surrogate in contaminant uptake and loss studies by
several researchers (e.g. Bedford and Zabik 1973; Heit et al. 1980; Perry et al. 1980; Malley
et al. 1995; Stewart 1999; Wang et al. 1999; Perceval et al. 2002). Additionally, P. grandis has
been used to evaluate potential biomarkers to assess heavy metal exposure (Chamberland
et al. 1995; Couillard et al. 1995; Perceval 2002).
Habitat: Distributed throughout the Mississippi River basin, Great Lakes-St. Lawrence
basin, several Gulf drainages, and north into Canada. Typically found in low-gradient stream
reaches, backwaters, lakes, and impoundments. Often most abundant in fine substrates.
Reproduction: Over 35 species of fish (including some exotic species) have been identified
as hosts for the Giant Floater. Jacobson et al. (1997) reported that glochidia of P. grandis
survived longer in natural river water than did the glochidia of three lampsiline species.
Converted to mg/L total ammonia as N, normalized to pH 8.
81
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An Introduction to Freshwater Mussels as Biological Indicators
Rabbitsfoot
Quadrula cylindrica cylindrical (Say, 1817)
Identification tips: Shell elongate, rectangular,
moderately thick, and somewhat compressed
to inflated; sculpturing often present in the form
of knobs, bumps, or ribs. Shell periostracum
and sculpturing often variable intraspecifically.
Periostracum usually yellowish-tan or green, with
numerous dark green chevrons; streaking may also
be present (A). Beak slightly raised above hinge-
line; sculpture often eroded or indistinct (B). Teeth
well developed; lateral teeth very long and narrow
(C). Nacre white. Shell up to 5 inches in length.
Indicator use: A rare taxon throughout its range
(Williams et al. 1993; Natureserve), the Rabbitsfoot
is a unionid of clear, high quality streams (Oesch
1984). It is listed as endangered by several
midwestern states (IL, IN, KS, OH). A close relative
and Tennessee River endemic, Quadrula cylindrica
strigillata, is currently listed as federally endangered
(USFWS 2004).
Butler (2005b) noted the elimination of Q.c.
cylindrica from 66% of the waterways where
it formerly occurred, although realistically he
remarked that total range reductions and population
losses were closer to 90%. This dramatic decline
was primarily attributed to the inability of the
Rabbitsfoot to persist in impounded habitats and
tolerate cold hypolimnetic tailwater releases.
Other causal factors included channelization,
mining activities, chemical contaminants, and
sedimentation (Butler 2005b).
Habitat: Distributed throughout the upper and
lower Mississippi River basin and western Lake Erie basin. Occurs in both moderate-sized
streams and rivers in swift flow at depths up to 12 feet (Parmalee and Bogan 1998). Most
abundant in sand, gravel, and cobble substrates. This interesting mussel does not exhibit the
burrowing tendencies of its congeners, preferring instead to rest on the substrate surface
(photo - upper right).
Reproduction: Identified hosts for Q.c. cylindrica include the Blacktail Shiner, Spotfin
Shiner, and Rosyface Shiner (Barnhart and Baird 2000; Butler 2005b). Yeager and Neves
(1986) identified the Bigeye Chub, Spotfin Shiner, and Whitetail Shiner as suitable hosts for
Q.c. strigillata, a Tennessee River basin endemic.
The Rabbitsfoot packages its glochidia into lanceolate conglutinates, generally orange
or yellowish-brown in color (Ortmann 1919).
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An Introduction to Freshwater Mussels as Biological Indicators
Pimpleback
Quadrula pustulosa pustulosa (Lea, 1831)
Identification tips: Shell round or quadrate, thick, and
usually inflated. Periostracum yellowish-tan to brown
with few to numerous pustules (A). A conspicuous green
smudge or broken, wide green ray(s) present in young
individuals (B) less apparent or absent in adults (A) Beak
sculpture of indistinct lines; often eroded (C). Teeth well
developed and heavy (D). Nacre white. Shell up to 4
inches in length.
Indicator use: The Pimpleback tolerates a wide range
of substrates and adapts well to impounded conditions
(Oesch 1984; Parmalee and Bogan 1998; WDNR 2003).
ESI (2000) consistently reported Q. pustulosa from the
navigational pools of the Upper Ohio River, where it
comprised 1.4% -18.4% of the mussel community.
In Kansas, researchers utilized tissue samples
from Q. pustulosa to evaluate the bioavailability of
heavy metals in the Neosho River, home of the federally
threatened Neosho Madtom (Noturus placidus) (Allen et
al. 2001). Most of the metals of concern were found in low
concentrations.
Habitat: A wideranging species, the Pimpleback occurs
in large streams, rivers, lakes, and reservoirs. It may be
found in both shallow and deep water (-20 ft.). Generally
most abundant in packed sand and gravel.
Reproduction: Identified host fishes include the
Shovelnose Sturgeon, Black Bullhead, Brown Bullhead,
Channel Catfish, Flathead Catfish, and White Crappie
(Surber 1916; Coker 1921). The reproductive period is
from June through August (Howard 1914).
83
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An Introduction to Freshwater Mussels as Biological Indicators
Mapleleaf
Quadrula quadrula
(Rafinesque, 1820)
Identification tips: Shell subquadrate, thick, and
slightly inflated. Two rows of pustules present on shell;
separated by a shallow to well-defined sulcus (A).
Small flute-like projections or bumps often present
on posterior slope and posterior-dorsal margin (B).
Periostracum yellowish-tan, brown, or dark brown;
green rays often present in younger individuals. Beak
sculpture of double-looped ridges; often eroded or
indistinct (C). Teeth well-developed (D); pseudocardinal
teeth may be large and heavy in older individuals.
Nacre white. Shell up to 6 inches in length.
Indicator use: The Mapleleaf is an adaptable species,
tolerant of (and perhaps exploiting) impoundment,
moderate levels of turbidity, and a variety of
substrates (Oesch 1984; Parmalee and Bogan 1998;
Metcalfe-Smith et al. 2003; WDNR 2003). ESI (2000)
documented Q. quadrula as one of the most abundant
species in the navigational pools of the upper Ohio
River.
The Mapleleaf has been utilized extensively
as a biomonitor and surrogate to evaluate the
potential impacts of industrial effluents and coal mine
discharges. In the early 1970s, Foster and Bates
(1978) employed juvenile Q. quadrula as in-stream
biomonitors of copper electroplating wastes in the
Muskingum River, OH. Body burdens as high as
20.64 ug Cu/g (10x background levels) were reported
for caged Q. quadrula 0.1 km below electroplating
discharges. Ultimately, the study concluded that
electroplating wastes were responsible for the
decimation of 60% and 39% of all mussels up to 5 km
and 21 km downstream, respectively.
Habitat: Distributed throughout the Mississippi
River basin, Great Lakes-St. Lawrence drainage, and
north in to Canada. Occurs in small to large rivers,
embayments, and lakes. Found in firm of substrates of mud, sand, gravel, and
cobble at depths ranging from several inches to 30 ft.
Reproduction: Identified host fishes include the Flathead Catfish and
Channel Catfish (Howard and Anson 1923; Schwebach et al. 2002). The
Mapleaf is tachytictic, with the reproductive period extending from May and
August (Baker 1928).
84
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An Introduction to Freshwater Mussels as Biological Indicators
Creeper
Strophitus undulatus
(Say, 1817)
Identification tips: Shell highly variable; generally
subelliptical to subtrapezoidal, thin to moderately thick,
and compressed to inflated. Periostracum usually
light brown to dark brown (A); young individuals often
with bold green rays. Beak raised slightly above
hinge-line; sculpture of concentric, oblique ridges (B).
Pseudocardinal teeth present as a thickened, indented
hinge (C). Lateral teeth also present as a thickened
hinge. Shell up to 4 inches in length.
Indicator use: Widely distributed and relatively
common, the Creeper is frequently described as a
tolerant species (Gordon and Layzer 1989; Walters
1995; Parmalee and Bogan 1998). Williams et al.
(1993) reported the conservation status of S. undulatus
as "currently stable".
Harman (1997) utilized molluscan and
macroinvertebrate communities as indicators of water
quality and habitat degradation in Otsego Lake, NY.
Over a period of 25 years, he reported a decline in
molluscan and intolerant Ephemeroptera, Plecoptera
and Trichoptera (EPT) richness of 52.9% and 56.1%,
respectively. The Creeper was among the declining
bivalve species, having been eliminated from two sites
where it formerly occurred.
Pip (2002) utilized bivalves and gastropods in
a water quality study of southern Lake Winnipeg,
Canada. Occurrences of S. undulatus were correlated
with lower lead values, while the Giant Floater
(Pyganodon grandis) was linked to higher dissolved
solids and the Fatmucket (Lampsilis siliquoidea) to
higher dissolved organic matter
Habitat: Widespread in the Mississippi River basin,
Great Lakes-St. Lawrence basin, and northern Atlantic
drainages. Occurs in creeks, rivers, and lakes in both
still waters and moderate current. Found in both coarse
and fine substrates.
Reproduction: Well over 30 fishes from 9 taxonomical families have been
confirmed hosts for the Creeper (Baker 1928; Hove 1995b; Hillegrass and Hove
1997; Hove 1997; Walters et al. 1998c; Wicklow and Beisheim 1998; Van Snik
Gray el al. 1999; 2002; Cliff 2001).
The Creeper is bradyliclic, wilh Ihe reproduclive period extending from July
lo May (Baker 1928).
85
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An Introduction to Freshwater Mussels as Biological Indicators
Pistolgrip
Tritogonia verrucosa
(Rafinesque, 1820)
Identification tips: Shell subquadrate,
somewhat compressed to moderately inflated,
and thick. Shell covered with numerous tubercles
anterior of the posterior ridge (A). Posterior with
wavy undulations or irregular flutes (B). Shell
periostracum dark green or yellowish-brown,
sometimes with V-shaped markings. Beak with
indistinct ridges and tubercles (C). Teeth well
developed and heavy (D). Nacre white (D). Shell
up to 8 inches in length.
Indicator use: While intolerant of pollution, the
Pistolgrip has demonstrated the ability to adapt
to a variety of habitat conditions (Walters 1995;
Parmalee and Bogan 1998). In the state of Ohio,
T. verrucosa strongly repopulated a decimated
stretch of the Scioto River following major sewage
treatment plant upgrades (Tetzloff and Akison
1999).
Habitat: Well distributed throughout the
Mississippi River basin and Alabama River
system. Generally found in large streams where
substrates are comprised of stable sandy mud,
gravel, or cobble. May become abundant in
oxygen-rich riffles and runs (Stansberry 1965).
Often found resting on the substrate surface.
Reproduction:
Brown Bullhead,
Catfish (Howells
andKurth 1998)
period, Pepi and
mantle dorsal to
with a blue-gray
Identified host fishes include the
Yellow Bullhead, and Flathead
1997; Pepi and Hove 1997; Hove
, During the glochidia release
Hove (1997) observed an inflated
the excurrent siphon, crenulated
edge (top right).
86
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An Introduction to Freshwater Mussels as Biological Indicators
Rainbow
Villosairis (I. Lea, 1829)
Identification tips: General: Shell elliptical and
elongate, somewhat thin to moderately thick, and
compressed to moderately inflated. Sexually dimorphic:
Females (A) inflated and obliquely flared or rounded
posteriorly, males (B) pointed posteriorly. Sexual
dimorphism may be cryptic in some specimens. General:
Periostracum yellowish-tan to light brown with broken or
solid bright green rays. Beak relatively even to slightly
raised above hinge-line; sculpture of erratic double-loops
(C). Teeth well developed, although somewhat small and
delicate. Nacre white. Shell up to 4 inches in length.
Indicator use: Although fairly widespread and abundant
(Williams et al. 1993), the Rainbow is reportedly
declining in some portions of its range (Metcalfe-Smith
et al. 2003; COSEWIC 2006). It generally inhabits
clean, well oxygenated stream reaches (Walters 1995;
Parmalee and Bogan 1998).
The Rainbow has been used extensively as
a surrogate to evaluate the effects of various toxic
contaminants to freshwater mussels (e.g. Goudreau et
al. 1993; Jacobson et al. 1993; Jacobson et al. 1997;
Cherry et al. 2002; Mummert et al. 2003; Valenti et al.
2005; Valenti et al. 2006b; Wang et al. 2007a; 2007b).
GMAVs compiled and ranked by Augspurger et al.
(2003), Augspurger (2006), and March et al. (2007)
placed V. iris among the 15 most sensitive aquatic taxa
to copper and ammonia.
Habitat: Widespread in the upper Mississippi River
basin and Great Lakes basin. Occurs in creeks to small
rivers where the water is shallow and current sluggish to
moderately strong. May also be found in lakes (Gordon
and Layzer 1989; Walters 1995). Most abundanl in
mixlures of sand, gravel, and cobble.
Reproduction: Confirmed hosl fishes include Ihe Weslern
Mosquilofish, Rock Bass, Largemoulh Bass, Smallmoulh
Bass, Suwanee Bass, and Spoiled Bass (Zale and Neves
1982; Neves el al. 1985).
The Rainbow is bradyliclic, wilh Ihe reproduclive period
extending from July lo May (Orlmann 1919).
87
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An Introduction to Freshwater Mussels as Biological Indicators
CONSERVATION STATUS
Common Name
Mucket
Pheasantshell
Dwarf Wedaemussel
Elktoe
Threeridae
Purple Wartvback
Dromedary Pearlvmussel
Eastern Elliptic
Spike
Oyster Mussel
Northern Riffleshell
Tubercled Blossom
Wabash Piatoe
Pink Mucket
Plain Pocketbook
Wavvraved lampmussel
Fatmucket
White Heelsplitter
Flutedshell
Black Sandshell
Cumberland Moccasinshell
Threehorn Wartvback
Sheepnose
Clubshell
Ohio Piatoe
Round Piatoe
Kidnevshell
Fluted Kidnevshell
Giant Floater
Rabbitsfoot
Pimpleback
Mapleleaf
Creeper
Pistolarip
Rainbow
Scientific Name
Actinonaias liaamentina
Actinonaias pectorosa
Alasmidonta heterodon
Alasmidonta marainata
Amblema plicata
Cvclonaias tuberculata
Dromus dromas
Elliptic complanata
Elliptic dilatata
Epioblasma capsaeformis
Epioblasma torulosa ranaiana
Epioblasma torulosa torulosa
Fusconaia flava
Lampsilis abrupta
Lampsilis cardium
Lampsilis fasciola
Lampsilis siliauoidea
Lasmiaona complanata complanata
Lasmiaona costata
Liaumia recta
Medionidus conradicus
Obliauaria reflexa
Plethobasus cvphvus
Pleurobema clava
Pleurobema cordatum
Pleurobema sintoxia
Ptvchobranchus fasciolaris
Ptvchobranchus subtentum
Pvaanodon arandis
Quadrula cvlindrica cvlindrica
Quadrula pustulosa pustulosa
Quadrula Quadrula
Strophitus undulatus
Tritoaonia verrucosa
Villosa iris
Federal (1)
-
-
E
SC
-
-
E
-
-
E
E
E
-
E
-
-
-
-
-
-
-
-
C
E
SC
-
-
C
-
SC
-
-
-
-
-
Indiana (2)
-
-
-
-
-
-
-
-
-
-
E
-
-
E
-
SC
-
-
-
-
-
-
E
E
SC
-
SC
-
-
E
-
-
-
-
-
o"
.0
IS
O
-
-
-
SC
-
-
-
E
-
-
E
-
SC
-
-
-
T
-
T
E
E
E
SC
SC
-
-
E
-
-
-
-
-
Michigan (4)
-
-
-
SC
-
-
-
E
-
-
-
-
T
-
-
-
-
-
-
-
E
-
SC
-
-
-
-
-
-
-
-
SC
N. Carolina (5)
-
-
-
-
-
E
-
-
SC
-
-
-
-
-
-
SC
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
T
-
SC
S
re
'E
'5>
>
-
-
E
-
-
-
E
-
-
E
-
-
-
E
-
-
-
-
-
T
-
-
T
-
E
-
-
-
-
-
T
-
-
-
-
Natureserve (7)
G5
G4
G1G2
G4
G5
G5
G1
G5
G5
G1
G2T2
G2TX
G5
G2
G5
G5
G5
G5T5
G5
G5
G3G4
G5
G3
G2
G4
G4G5
G4G5
G2
G5
G3G4T3
G5
G5
G5
G4G5
G5Q
Williams et al. 1993
CS
SC
E
SC
CS
SC
E
CS
CS
E
E
E*
CS
E
SC
CS
CS
CS
CS
SC
SC
CS
T
E
SC
CS
CS
SC
CS
T
CS
CS
CS
CS
CS
CS = Currently stable SC = Special concern C = Candidate species T = Threatened E = Endangered
G5 = Secure G4 = Apparently secure G3 = Vulnerable G2 = Imperiled G1 = Critically imperiled
T = Infraspecific taxon X = Presumed extinct Q = Questionable taxonomy
(1) U.S. Fish and Wildlife Service, report generated 12/04/2007
(2) Indiana Department of Natural Resources, last updated 9/2004
(3) Ohio Department of Natural Resources, last updated 9/2007
(4) Michigan Natural Features Inventory, last updated 7/2002
(5) North Carolina Wildlife Resources Commission
(6) Virginia Department of Inland Game and Fishes, last updated 7/24/06
(7) Natureserve, Accessed 12/04/2007
88
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An Introduction to Freshwater Mussels as Biological Indicators
GLOSSARY
Adductor muscles: Muscles anterior and posterior that close the mussel shell.
Anterior: Forward portion of the shell.
Beak: Raised area on the dorsal margin near hinge.
Beak sculpture: Small, textured patterns on the beak.
Chevron: A V-shaped marking on the periostracum.
Cumberlandian: An ecoregion that encompasses the Cumberland River and Tennessee
River basins. This guide follows the interpretation of Parmalee and Bogan (1998) that
extends the region down to the mouth of both the Cumberland and Tennessee rivers.
Dorsal wing: An extension of a shell from the dorsal margin, posterior of the beak.
Extirpation: The elimination of a population from a certain geographic area.
Lateral teeth: The narrow, thin teeth located near the hinge-line.
Muscle scar: Impressions in the nacre where a muscle was previously attached.
Nacre: The inside of the shell, often pearly white, salmon, or purple.
Pallial line: A line present on the inner nacre where muscles previously attached the
mantle to the shell.
Periostracum: The outside layer of the shell or epidermis.
Posterior: Rear portion of the shell.
Posterior ridge: A ridge extending from the beak to the posterior margin.
Pseudocardinal teeth: Located anterior of the lateral teeth near the hinge, often raised
and triangular.
Unionid: A freshwater mussel belonging to the family Unionidae.
Unionoid: A freshwater mussel belonging to the order Unionoida.
89
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An Introduction to Freshwater Mussels as Biological Indicators
LITERATURE CITED
Abell, R.A., Olson, D.M., Dinerstein, E., and RT. Hurley. (2000). Freshwater Ecoregions of North
America: A Conservation Assessment. Island Press, Washington, DC. 368 pp.
Adams, T.G., Atchison, G.J., and R.J. Vetter. (1980). The impact of an industrially contaminated
lake on heavy metal levels in its effluent stream. Hydrobiologia 69(1-2):187-193.
Adams, T.G., Atchison, G.J., and R.J. Vetter. (1981). The use of the three-ridge clam (Amblema
perplicata) to monitor trace metal contamination. Hydrobiologia 83:67-72.
Ahlstedt, S.A. and J.D. Tuberville. (1997). Quantitative reassessment of the freshwater mussel
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Alderman, J.A. (1990). Fusconaia masoni (Conrad, 1834). pp 63-65. In: A report on the
conservation status of North Carolina's freshwater and terrestrial molluscan fauna. Scientific
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Allen, G.T., Blackford, S.H., Tabor, V.M., and M.S. Cringan. (2001). Metals, boron, selenium
in Neosho Madtom in the Neosho River in Kansas, U.S.A. Environmental Monitoring and
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ASTM (American Society for Testing and Materials). (2006). Standard guide for conducting
laboratory toxicity tests with freshwater mussels. In: Annual book of ASTM standards, Vol.
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Anderson, R.V. (1977). Concentration of cadmium, copper, lead, and zinc in six species of
freshwater clams. Bulletin of Environmental Contamination and Toxicology 18(4):492-496.
Anthony, J.J. and J.A. Downing. (2001). Exploitation trajectory of a declining fauna: a century of
freshwater mussel fisheries in North America. Canadian Journal of Fisheries and Aquatic
Sciences 58(10): 2071 -2090.
Anthony, J.L., Kesler, D.H., Downing, W.L., and J.A. Downing. (2001). Length-specific growth
rates in freshwater mussels (Bivalvia: Unionidae): extreme longevity or generalized growth
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Aspelin, A.L. (1994). Pesticide Industry Sales and Usage, 1992 and 1993 Market Estimates. U.S.
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Augspurger, T, Keller, A.E., Black, M.C., Cope, W.G., and F.J. Dwyer. (2003). Water quality
guidance for protection of freshwater mussels (Unionidae) from ammonia exposure.
Environmental Toxicology and Chemistry 22(11 ):2569-2575.
Augspurger, T, Dwyer, J., and C. Ingersoll. (2006). Implications of including freshwater mussel
toxicity data in water quality criteria derivation datasets. Presentation # 724 at the 27th annual
meeting of the Society of Environmental Toxicology and Chemistry, Montreal, Quebec,
November 2006.
Badra, RJ. and R.R. Goforth. (2001). Surveys for the clubshell (Pleurobema clava) and other rare
clams in Michigan: Final Report 2000. Report to U.S. Fish and Wildlife Region 3 Endangered
Species Office. 52pp. + appendices.
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An Introduction to Freshwater Mussels as Biological Indicators
Badra, RJ. and R.R. Goforth. (2002). Surveys of Native Freshwater Mussels in the Lower Reaches
of Great Lakes Tributary Rivers in Michigan. Report number MNFI 2002-03. Report to Michigan
Dept. of Environmental Quality, Coastal Zone Management Unit, Lansing, Ml. 39pp.
Baker, F.C. (1928). The fresh water Mollusca of Wisconsin. Part II. Pelecypoda. Bulletin of the
Wisconsin Geological and Natural History Survey. University of Wisconsin. 70(2):vi 495 + 76
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Barfield, M.L. and G.T. Walters. (1998). Non-parasitic life cycle in the green floater, Lasmigona
subviridis (Conrad, 1835).Triannual Unionid Report 16:22.
Barnhart, M.C., Riusech, F.A., and A.D. Roberts. (1997). Fish hosts of the federally endangered
pink mucket, Lampsilis abrupta. Triannual Unionid Report 13:35.
Barnhart, M.C. and M.S. Baird. (2000). Fish hosts and culture of mussel species of special
concern. Annual Report to U.S. Fish and Wildlife Service and Missouri Department of
Conservation. 39 pp.
Barnhart, M.C. (2006). Unio Gallery: http://unionid.missouristate.edu. Accessed 06/06/2007.
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Environmental Toxicology and Chemistry 22(1 1):2561 -2568.
Bates, J.M. (1962). The impact of impoundment on the mussel fauna of Kentucky Reservoir,
Tennessee River. American Midland Naturalist 68:232-236.
Bauer, G. (1987). Reproductive strategy of the freshwater pearl mussel Margaritifera margaritifera.
Journal of Animal Ecology 56:691-704.
Beckvar, N., Salazar, S., Salazar, M., and K. Finkelstein. (2000). An in-situ assessment of mercury
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