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Common Name: zebra mussel
Taxonomy: available through 
Identification: Zebra mussels are small shellfish named for the striped pattern of their shells. Color patterns can vary to the point of having only dark or light colored shells and no stripes. They are typically found attached to objects, surfaces, or each other by threads underneath the shells. Although similar in appearance to the quagga mussel (Dreissena bugensis), the two species can be easily distinguished. When placed on a surface zebra mussels are stable on their flattened underside while quagga mussels, lacking a flat underside, will fall over. See Mackie and Schlosser (1996) for a key to adult Dreissenids.
Size: < 50 mm
Native Range:
Zebra mussels are native to the Black, Caspian, and Azov Seas. In 1769, Pallas first described populations of this species from the Caspian Sea and Ural River.
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Hawaii |
Caribbean |
Interactive maps: Continental US, Alaska, Hawaii, Caribbean
Means of Introduction: A release of larval mussels during the ballast exchange of a single commercial cargo ship traveling from the north shore of the Black Sea to the Great Lakes has been deduced as the likely vector of introduction to North America (McMahon 1996). Its rapid dispersal throughout the Great Lakes and major river systems was due to the passive drifting of the larval stage (the free-floating or "pelagic" veliger), and its ability to attach to boats navigating these lakes and rivers (see Remarks, below). Its rapid range expansion into connected waterways was probably due to barge traffic where it is theorized that attached mussels were scraped or fell off during routine navigation. Overland dispersal is also a possibility for aiding zebra mussel range expansion. Many small inland lakes near the Great Lakes unconnected by waterways but accessed by individuals trailering their boats from infested waters, have populations of zebra mussels living in them. At least nineteen trailered boats crossing into California had zebra mussels attached to their hulls or in motor compartments; all were found during inspections at agricultural inspection stations. Under cool, humid conditions, zebra mussels can stay alive for several days out of water.
Status: They are established in all the Great Lakes, most of the large navigable rivers in the eastern United States and in many inland lakes in the Great Lakes region.
Impact of Introduction: Zebra mussels are
notorious for their biofouling capabilities by colonizing water supply
pipes of hydroelectric and nuclear power plants, public water supply
plants, and industrial facilities. They colonize pipes constricting
flow, therefore reducing the intake in heat exchangers, condensers,
fire fighting equipment, and air conditioning and cooling systems.
Zebra mussel densities were as high as 700,000/m2
at one power plant in Michigan and the diameters of pipes have been
reduced by two-thirds at water treatment facilities. Although there is
little information on zebra mussels affecting irrigation, farms and
golf courses could be likely candidates for infestations. Navigational
and recreational boating can be affected by increased drag due to
attached mussels. Small mussels can get into engine cooling systems
causing overheating and damage. Navigational buoys have been sunk under
the weight of attached zebra mussels. Fishing gear can be fouled if
left in the water for long periods. Deterioration of dock pilings has
increased when they are encrusted with zebra mussels. Continued
attachment of zebra mussel can cause corrosion of steel and concrete
affecting its structural integrity.
Zebra mussels can have profound effects on the ecosystems they
invade. They primarily consume phytoplankton, but other suspended
material is filtered from the water column including bacteria,
protozoans, zebra mussel veligers, other microzooplankton and silt.
Large populations of zebra mussels in the Great Lakes and Hudson River
reduced the biomass of phytoplankton significantly following invasion.
Diatom abundance declined 82-91% and transparency as measured by Secchi
depth increased by 100% during the first years of the invasion in Lake
Erie (Holland 1993). As the invasion spread eastward during 1988 to
1990, successive sampling stations recorded declines in total algae
abundance from 90% at the most western station to 62% at the most
eastern (Nichols and Hopkins 1993). In Saginaw Bay, sampling stations
with high zebra mussel populations experienced a 60-70% drop in
chlorophyll-a and doubling of Secchi depth (Fahnenstiel et al. 1993). Phytoplankton biomass declined 85% following mussel invasion in the Hudson River (Caraco et al.
1997). The extent of change that zebra mussels can exert on species
composition of the phytoplankton community is unresolved. Increased
water clarity allows light to penetrate further, potentially promoting
macrophyte populations (Skubinna et al. 1995). As macrophytes
can be colonized by veligers, the macrophyte community may be altered
if such colonization proves detrimental. Increased light penetration
may also cause water temperatures to rise and thermoclines to become
deeper, but these effects have not yet been documented. As
phytoplankton are consumed, the dissolved organic carbon (DOC)
concentration may drop. Indeed, inland lakes with zebra mussels have
been found to have lower concentrations of DOC (Raikow 2002).
Macrophytes could eventually compensate for this since they are also a
source of DOC, but there may be a lag period between the time when
phytoplankton biomass is down and macrophytes proliferate. This could
produce a period of time when UV-B light penetrates deeper into the
water column, because DOC absorbs UV-B radiation. Zebra mussels have
also recently been shown to be able to directly assimilate DOC (Roditi et al.
2001). Zebra mussels are able to filter particles smaller than 1µm in
diameter, although they preferentially select larger particles (Sprung
and Rose 1988). Thus bacteria may represent an important food source
(Cotner et al. 1995, Silverman et al. 1996). At a
90% efficiency rate, zebra mussels are much more efficient at
filtration of such small particles than are unionids and Asiatic clams.
Filtering rate is highly variable, depending on temperature,
concentration of suspended matter, phytoplankton abundance, and mussel
size (reviewed by Noordhuis et al. 1992). Although European
zebra mussels are less active in winter, this seasonal pattern is
temperature driven. No diel patterns of filtration rate have been
found. During spring, filtration rates rise dramatically between 5 and
10oC, then level off with respect to temperature, and may be inhibited at temperatures over 20oC.
Increased suspended matter can reduce filtration activity to a minimum
required to maintain oxygen demand. A sigmoidal relationship exists
with filtration rate and size, but this may be an affect of aging.
Material filtered by zebra mussels is either ingested or expelled as
feces or mucus covered pseudofeces. True fecal pellets are chemically
altered, larger and more dense. Pseudofeces production increases with
increasing suspended solid concentration, as well as increasing
temperature, albeit to a much lesser extent (Noordhuis et al. 1992, MacIsaac and Rocha 1995). The rate of biosedimentation through pseudofeces production was very high (28mg/cm2 day at a density of 1180 individuals/m2)
under turbid conditions in Lake Erie, lending support to the hypothesis
that zebra mussels are responsible for increased water clarity observed
since mussel introduction (Klerks et al. 1996). Filtration rate was not related to seston composition (POC:TSS, chl:TSS) in Saginaw Bay (Fanslow et al.
1995). Veligers also filter material, but their impact is far less than
that of sessile adults. Settled mussels exerted 103 times the grazing
rate of veligers in western Lake Erie, for example (MacIsaac et al.
1992). Microzooplankton, e.g. rotifers and veligers, are ingested by
zebra mussels, but larger zooplankton are not eaten (MacIsaac et al. 1991, MacIsaac et al.
1995). It has been speculated that benthic deposition of feces and
pseudofeces may aid bacterial productivity, thus producing a source
culture that zebra mussels can feed upon (Silverman et al.
1996). It has also been speculated that biodeposition of feces and
pseudofeces might cause observed increases in benthic macroinvertebrate
populations (Stewart and Haynes 1994). Biomagnification of
Polychlorinated Biphenyls (PCBs) was observed in Gammarus associated
with zebra mussels, indicating concentration of pollutants in zebra
mussel feces or pseudofeces can transfer to other trophic levels
(Bruner et al. 1994). In an experimental study, however, Botts et al.
(1996) found greater abundances of macroinvertebrates associated with
both living and non-living (i.e. empty shell) zebra mussel druses
compared with their no-druse treatment. Thus the increased physical
habitat complexity of a mussel colony may benefit macroinvertebrates
rather than deposition of feces and pseudofeces. Zebra mussels can
reduce filtration rates (more frequent interruption of filtering or
slower pumping rates) and/or produce pseudofeces above an incipient
limiting concentration (ILC) of algae to maintain a constant
consumption rate (Sprung and Rose 1988, Fanslow et al. 1995
MacMahon 1996,). Feeding activity can be described by the clearance
rate (percentage of algal biomass removed from the water column over
time), biomass of cleared algae (BCA), feces production and pseudofeces production (µg F or P/BCA). For example, Berg et al.
(1996) examined the effects of zebra mussel size and algae species and
concentration on zebra mussel feeding activity. Clearance rates were
constant over varying concentrations of pure cultures of Chlamydomonas reinhardtii,
a spherical unicellular species of 7.42 µm (± 0.13µm) in diameter. This
indicates that the concentrations used in experiments were below the
ILC. However, clearance rates decreased, with increasing concentrations
of Pandorina morum, a species made up of colonies with varying numbers of cells that are individually as large as C. reinhardtii.
This indicates that the concentrations used in experiments were above
the ILC. Large zebra mussels (20-25 mm in length) displayed a higher
clearance rate across all concentrations of C. reinhardtii
than did small mussels (10-15 mm). Incipient limiting concentration
differed in this study from previous studies done with European
populations. Thus zebra mussel size, phytoplankton species, and
regional population differences affect clearance rates, ILC and
feces/pseudofeces production. Zebra mussels produce pseudofeces to
avoid ingesting non-food material (e.g. clay), as a mechanism to deal
with overabundance of food (e.g. algal concentrations above the ILC),
and possibly as a way to reject unpalatable algae. Zebra mussels
readily reject blue-green algae, such as Microcystis, as pseudofeces
(Vanderploeg et al. 2001). The presence of this cyanobacterium does not inhibit filtering, except in mass abundances such as a bloom (Noordhuis et al. 1992, Lavrentyev et al.
1995). Zebra mussels can select material for rejection through
pseudofeces production internally, perhaps identifying cyanobacteria by
chemical cues (Ten Winkel and Davids 1982). Inland lakes with lower
nutrient levels have been observed to be more frequently dominated by
Microcystis when invaded by zebra mussels (Raikow et al.
2004). Understanding of the fate of pseudofeces once it expelled is
poor. Zebra mussels removed metals from the water column of Lake Erie
and deposited it to the bottom at high rates (Klerks et al. 1997). Roditi et al.
(1997) found that the biodeposits of zebra mussel were organically
enriched, including 3.9% live algae by weight. Resuspension of this
material occurred in their system, a tidal estuary, reducing the
potential impact of biodeposition to the benthos. Less well known is
the fate of live algae bound into pseudofeces. Bastviken et al.
(1998) speculate that phytoplankton which survives the pseudofeces
process must be resuspended in order for long term survival, a process
less likely to occur in inland lakes than in tidal estuaries. If
survivorship following filtration is equal between phytoplankton
species, then community species composition can remain unchanged. Other
factors may affect the phytoplankton community, however, including
increased light. The zooplankton community has also been affected by
the invasion of zebra mussels. Zooplankton abundance dropped 55-71%
following mussel invasion in Lake Erie, with microzooplankton more
heavily impacted (MacIsaac et al. 1995). Mean summer biomass of zooplankton decreased from 130 to 78 mg dry wt. m-3 between
1991 and 1992 in the inner portion of Saginaw Bay. The total biomass of
zooplankton in the Hudson River declined 70% following mussel invasion,
due both to a reduction in large zooplankton body size and reduction in
microzooplankton abundance. These effects can be attributed to
reduction of available food (phytoplankton) and direct predation on
microzooplankton. Increased competition in the zooplankton community
for newly limited food should result from zebra mussel infestation. The
size of individual zooplankters might decrease. Hypotheses can be
formulated specifying which species will prevail based on knowledge of
competitive ability.
Effects may continue through the food web to fish. Reductions in
zooplankton biomass may cause increased competition, decreased survival
and decreased biomass of planktivorous fish. Alternatively, because
microzooplankton are more heavily impacted by zebra mussels the larval
fish population may be more greatly affected than later life stages.
This may be especially important to inland lakes with populations of
pelagic larval fish such as bluegills. Benthic feeding fish may benefit
as opposed to planktivorous fish, or behavioral shifts from pelagic to
benthic-feeding may occur. In addition, proliferation of macrophytes
may alter fish habitat. Experimental evidence exists that zebra mussels
can reduce the growth rate of larval fish through food web interactions
(Raikow 2004). Conclusive negative impacts on natural populations of
fish, however, have yet to observed (see Raikow 2004). Other effects
include the extirpation of native unionid clams through epizootic
colonization (Schloesser et al. 1996, Baker and Hornbach
1997). Zebra mussels restrict valve operation, cause shell deformity,
smother siphons, compete for food, impair movement and deposit
metabolic waste onto unionid clams. Survival rates of native unionid
mussels in the Mississippi River, Minnesota have been shown to decline
significantly with the increase in zebra mussel colonization (Hart et
al. 2001).To date, unionids have been extirpated from Lake St. Clair
and nearly so in western Lake Erie. Many species of birds known to be
predators of zebra mussels occur in the Great Lakes region. While a new
food source may benefit such predators, biomagnification of toxins into
both fish and birds is possible. Some effects have been hypothesized as
worst-case scenarios. For example, zebra mussels may cause a shift from
pelagically to benthically-based food webs in inland lakes. Zebra
mussels may also shift lakes from a turbid and phytoplankton-dominated
state to clear and macrophyte-dominated state, i.e. between alternative
stable equilibria (Scheffer et al. 1993).
Remarks: Zebra mussels represent
one of the most important biological invasions into North America,
having profoundly affected the science of Invasion Biology, public
perception, and policy. In the 1980's Invasion Biology began to emerge
as a true sub-discipline of ecology as evidenced by an exponential
increase in scientific output on the subject (Raikow, unpubl. data).
Most work on the subject was terrestrial. Invasions were not a large
component of the popular environmental movement, and no serious
legislation existed concerning invasions beyond agricultural pests.
After the discovery of zebra mussels in 1988 the exponential rate of
scientific output on invasions itself increased (Raikow, unpubl. data),
the Nonindigenous Aquatic Nuisance Prevention and Control Act was
written and passed, and invasions became a topic discussed in the
media. Today biological invasions are described as the second leading
cause of extinction behind habitat destruction. Aquatic invasions are a
topic of much research. For these reasons the zebra mussel is often
described as the "poster child" of biological invasions.
A long tradition of zebra mussel study exists in Europe and the
former Soviet Union, where the zebra mussel has been present for 150
years (see Mackie et al. 1989 for an annotated bibliography
of European references). Work includes spatial distribution patterns,
demography, tolerance limits for physical and chemical parameters, and
physiology. Extensive ecological work in the United States began soon
as the zebra mussel was discovered and peaked in the early 1990's. The
literature on ecosystem and community-level effects of zebra mussels
has been dominated by work investigating Lake Erie, Saginaw Bay, the
Hudson River, and Oneida Lake (e.g. Fahnenstiel 1993, Holland 1993,
Pace et al. 1998, Idrisi et al. 2001.
The life history of zebra mussels differs greatly from most endemic
Great-Lakes region bivalves (Pennak 1989, Mackie and Schlosser 1996).
Exotic dreissenids are dioecious, with fertilization occurring in the
water column. Endemic bivalves are monoecious, dioecious or
hermaphroditic, and some internally fertilized by filtering sperm from
the water column. Under natural thermal regimes, zebra mussel oogenesis
occurs in autumn, with eggs developing until release and fertilization
in spring. In thermally polluted areas, reproduction can occur
continually through the year. Females generally reproduce in their
second year. Eggs are expelled by the females and fertilized outside
the body by the males; this process usually occurs in the spring or
summer, depending on water temperature. Optimal temperature for
spawning is 14-16oC. Over 40,000 eggs can be laid in a
reproductive cycle and up to one million in a spawning season. Spawning
may last longer in waters that are warm throughout the year. After the
eggs are fertilized, the larvae (veligers) emerge within 3 to 5 days
and are free-swimming for up to a month. Optimal temperature for larval
development is 20-22oC. Dispersal of larvae is normally
passive by being carried downstream with the flow. The larvae begin
their juvenile stage by settling to the bottom where they crawl about
on the bottom by means of a foot, searching for suitable substratum.
They then attach themselves to it by means of a byssus, an "organ"
outside the body near the foot consisting of many threads. Although the
juveniles prefer a hard or rocky substrate, they have been known to
attach to vegetation. As adults, they have a difficult time staying
attached when water velocities exceed two meters per second.
Zebra mussels are filter feeders having both inhalant and exhalant
siphons. They are capable of filtering about one liter of water per day
while feeding primarily on algae. Once the veliger undergoes
morphological changes including development of the siphon, foot, organ
systems and blood, it is known as a postveliger. Further subdivision of
the larval stage has been delineated: (veliger) preshell,
straight-hinged, umbonal, (postveliger) pediveliger, plantigrade, and
(juvenile) settling stage (ZMIS 1996). The settling stage attaches to a
substrate via protienaceous threads secreted from the byssal gland. The
vast majority of veliger mortality (99%) occurs at this stage due to
settlement onto unsuitable substrates. Sensitivity to changes in
temperature and oxygen are also greatest at this stage. Once attached,
the life span of D. polymorphais variable, but can range from 3-9 years. Maximum growth rates can reach 0.5 mm/day and 1.5-2.0 cm/year1. Adults are sexually mature at 8-9 mm in shell length (i.e. within one year).
The rapid invasion of North American waterways has been facilitated
by the zebra mussel's ability to disperse during all life stages.
Passive drift of large numbers of pelagic larval veligers allows
invasion downstream. Yearlings are able to detach and drift for short
distances. Adults routinely attach to boat hulls and floating objects
and are thus anthropogenically transported to new locations.
Transporting recreational boats disperses zebra mussels between inland
lakes. In addition, speculation exists that waterfowl can disperse
zebra mussels, but this has yet to be conclusively demonstrated. While
byssal threads develop in the larvae of some non-dresissenid endemic
bivalves and are used to attach to fish gills, there are no endemic
freshwater bivalves with byssal adult stages. This adaptation has been
important to the zebra mussel's success in invading North America.
Zebra mussels attach to any stable substrate in the water column or
benthos: rock, macrophytes, artificial surfaces (cement, steel, rope,
etc.), crayfish, unionid clams, and each other, forming dense colonies
called druses. Long-term stability of substrate affects population
density and age distributions on those substrates. Within Polish lakes,
perennial plants maintained larger populations than did annuals
(Stanczykowska and Lewandowski 1993). Populations on plants also were
dominated by mussels less than a year old, as compared with benthic
populations. These populations of small individuals allow higher
densities on plants. In areas where hard substrates are lacking, such
as a mud or sand, zebra mussels cluster on any hard surface available.
Given a choice of hard substrates, zebra mussels do not show a
preference, indicating that veligers cannot discriminate between
substrates (with the exception of substrate rejection due to
contaminants). Research on danish lakes shows that factors exist,
however, that cause substrate to be unsuitable for both initial and
long term colonization: extensive siltation, some sessile benthic
macroinvertebrates, macroalgae, and fluctuating water levels exposing
mussels to desiccation (Smit et al. 1993). The dispersion of
zebra mussels within a lake is controlled by physical conditions
including wind strength, lake/shore morphometry, and current patterns
(Stanczykowska and Lewandowski 1993). These conditions affect both
spatial patterns of pelagic veliger density and benthic adult
dispersion. Population density of benthic adults has been observed to
vary as widely as two orders of magnitude (e.g. <100 to >1500
individuals/m-2) within individual Polish lakes due to these
physical conditions. Tolerance limits of physical and chemical
parameters are well known (Sprung 1993, Vinogradov et al. 1993, McMahon 1996).
Although discrepancy exists when comparing temperature tolerance
limits of North American and European populations, this is probably due
to the American population being founded by mussels from the southern
limit of the European population's range. Most work in Europe has been
done in the northern range. North American populations are generally
adapted to warmer temperature regimes than their European counterparts.
Although shell growth has been reported to occur at temperatures as low
as 3¢ª C, Lake St. Clair populations and some European populations
display shell growth at 6-8¢ª C. Eggs are released when the
environmental temperature reaches 13¢ª C and release rate is maximized
over 17¢ª C. The optimal temperature range for adults extends to
20-25¢ª C, but D. polymorpha can persist in temperatures up
to 30¢ª C. Short term tolerance of temperatures up to 35¢ª C is
possible if the mussels were previously acclimated to high
temperatures. Rapid warming of shallow lakes has been hypothesized to
detrimentally affect reproductive rates in Danish populations (Smit et al.
1993). Oxygen demands are similar to those of other freshwater bivalves
including unionids. Tolerance of "anaerobic" conditions has been
reported for short time periods under certain temperatures and sizes,
but zebra mussels cannot persist in hypoxic conditions. The lower limit
of pO2 tolerance is 32-40 Torr at 25¢ª C. Zebra mussels have
been found in the hypolimnetic zone of lakes with oxygen levels of
0.1-11.2 mg/l, and in the epilimnetic zone with oxygen levels of
4.2-13.3 mg/l. Zebra mussels are described as poor O2
regulators, possibly explaining their low success rate in colonizing
eutrophic lakes and the hypolimnion. Zebra mussels can tolerate only
slight salinity.
Although some populations of European zebra mussels can be found in
estuaries, their persistence has been speculatively attributed to
reduced tidal fluctuation. Upper limits of freshwater bivalve salinity
tolerance reach 8-10 ‰, and populations of European zebra mussels have
been found to tolerate and range of salinities, from 0.6 ‰ (Rhine
River) to 10.2 ‰ (Caspian Sea). North American populations generally
tolerate salinity up to 4 ‰. In European populations, calcium
concentrations of 24 mg Ca2+/l allow only 10% larval survival due to inhibition of shell development. Optimal calcium concentrations ranges from 40-55 mg Ca2+/l,
but North American populations have been found in lakes with lower
concentrations. North American populations require 10 mg Ca2+/l to initiate shell growth and 25 mg Ca2+/l
to maintain shell growth. Larval development is inhibited at pH of 7.4.
Higher rates of adult survival occur at a pH of 7.0-7.5, but
populations have been found in the hypolimnetic zone of lakes with a pH
of 6.6-8.0, and in the epilimnetic zone with a pH of 7.7-8.5. Optimal
larval survival occurs at a pH of 8.4, and optimal adult growth occurs
at pH 7.4-8.0. There are many methods that have been investigated to
help control zebra mussels. They are listed below in no particular
order. Some methods will work better than others in a particular
situation.
References
Baker, S. M., and D. J. Hornbach, 1997, Acute physiological effects of zebra mussel (Dreissena polymorpha) infestation on two unionid mussels, Actinonaias ligmentina and Amblema plicata, Canadian Journal of Fisheries and Aquatic Sciences 54:512-519.
Bastviken, D. T. E., N. F. Caraco, and J. J. Cole, 1998, Experimental measurements of zebra mussel (Dreissena polymorpha) impacts on phytoplankton community composition, Freshwater Biology 39:375-386.
Berg, D. J., S. W. Fisher, and P. F. Landrum, 1996, Clearance and processing of algal particles by zebra mussels (Dreissena polymorpha), Journal of Great Lakes Research 22:779-788
Botts, P., B. Silver, A Patterson, and D. W. Schloesser, 1996, Zebra mussel effects on benthic invertebrates: physical or biotic?, Journal of the North American Benthological Society 15:179-184.
Bruner, K. A., S. W. Fisher, and P. F. Landrum, 1994, The role of the zebra mussel, Dreissena polymorpha, in contaminant cycling: II. Zebra mussel contaminant accumulation from algae and suspended particles, and transfer to the benthic invertebrate, Gammarus fasiatus, Journal of Great Lakes Research 20:735-750.
Caraco, N. F., J. J. Cole, P. A. Raymond, D. L. Strayer, M. L. Pace, S. E. G. Findlay, and D. T. Fischer, 1997, Zebra mussel invasion in a large, turbid river: phytoplankton response to increased grazing. Ecology 78:588-602.
Cotner, J. B, W. S. Gardner, J. R. Johnson, R. H. Sada, J. F. Cavaletto, and R. T. Heath, 1995, Effects of zebra mussels (Dreissena polymorpha) on bacterioplankton: evidence for both size-selective consumption and growth stimulation, Journal of Great Lakes Research 21:517-528.
Fahnenstiel, G. L., T. B. Bridgeman, G. A. Lang, M. J. McCormik, and T. F. Nalepa, 1993, Phytoplankton productivity in Saginaw Bay, Lake Huron: Effects of zebra mussel (Dreissena polymorpha) colonization, Journal of Great Lakes Research 21:465-475
Fanslow, D.L., Nalepa, T. F., and Lang, G. A., 1995, Filtration rates of zebra mussels (Dreissena polymorpha) on natural seston from Saginaw Bay, Lake Huron, Journal of Great Lakes research 21:489-500.
Hart, R. A., A. C. Miller and M. Davis. 2001. Empirically Derived Survival Rates of a Native Mussel, Amblema plicata, in the Mississippi and Otter Tail Rivers, Minnesota. American Midland Naturalist. 146: 254-263.
Holland, R. E., 1993, Changes in planktonic diatoms and water transparency in Hatchery Bay, Bass Island Area, Western Lake Erie since the establishment of the zebra mussel, Journal of Great Lakes Research 19:617-624.
Idrisi, N., E. L. Mills, L. G. Rudstam, and D. J. Stewart, 2001, Impact of zebra mussels (Dreissena polymorpha) on the pelagic lower trophic levels of Oneida Lake, New York, Canadian Journal of Fisheries and Aquatic Sciences 58:1430-1441.
Klerks, P. L., P. C. Fraleigh, and J. E. Lawniczak, 1996, Effects of zebra mussels (Dreissena polymorpha) on seston levels and sediment deposition in western Lake Erie, Canadian Journal of Aquatic Sciences 53:2284-2291.
Lavrentyev, P. J., W. S. Gardner, J. F. Cavaletto, and J. R. Beaver, 1995, Effects of the zebra mussel (Dreissena polymorpha Pallas) on protozoa and the phytoplankton from Saginaw Bay, Lake Huron, Journal of Great Lakes Research 21:545-557.
MacIsaac, H. J., W. G. Sprules, and J. H. Leach, 1991, Ingestion of small-bodied zooplankton by zebra mussels (Dreissena polymorpha): can cannibalism on larvae influence population dynamics, Canadian Journal of Fisheries and Aquatic Sciences 48:2051-2060.
MacIsaac, H. J., R. Rocha, 1995, Effects of suspended clay on zebra mussel (Dreissena polymorpha) faeces and pseudofaeces production, Archiv fur Hydrobiologie 135:53-64.
MacIsaac, H. J., W. G. Sprules, O. E. Johannsson, and J. H. Leach, 1992, Filtering impacts of larval and sessile zebra mussels (Dreissena polymorpha) in Western Lake Erie, Oecologia 92:30-39.
MacIsaac, H. J., C. J. Lonnee, and J. H. Leach, 1995, Suppression of microzooplankton by zebra mussels: importance of mussel size, Freshwater Biology 34:379-387.
Mackie, G. L., and D. W. Schlosser, 1996, Comparative biology of zebra mussels in Europe and North America: an overview, American Zoologist 36:244-258.
Mackie, G. L, W. N. Gibbons, B. W. Muncaster, and I. M. Gray, 1989, The zebra mussel, Dreissena polymorpha, a synthesis of European experiences and a preview for North America, Queen's Printer for Ontario.
McMahon, R. F., 1996, The physiological ecology of the zebra mussel, Dreissena polymorpha, in North America and Europe, American Zoologist 36:339-363.
Nichols, K. H., and G. J. Hopkins, 1993, Recent changes in Lake Erie (north shore) phytoplankton: cumulative impacts of phosphorus loading reductions and the zebra mussel introduction, Journal of Great Lakes Research 19:637-647
Pace, M. L., S. E. G. Findlay, and D. Fischer, 1998, Effects of an invasive bivalve on the zooplankton community of the Hudson River, Freshwater Biology 39:103-116
Pennak, R. W., 1989, Fresh-water invertebrates of the United States, Third Edition, John Wiley & Sons Inc., New York, NY, 628 p.
Raikow, D. F., 2002, How the feeding ecology of native and exotic mussels affects freshwater ecosystems, Doctoral Dissertation, Michigan State University.
Raikow, D.F., 2004, Food web interactions between larval bluegill (Lepomis macrochirus) and exotic zebra mussels (Dreissena polymorpha), Can. J. Fish. Aquat. Sci. 61:497-504.
Raikow, D. F., O. Sarnelle, A E. Wilson, and S. K. Hamilton. 2004. Dominance of the noxious cyanobacterium Microcystis aeruginosa in low-nutrient lakes is associated with exotic zebra mussels. Limnol. Oceanogr. 49: 482-487.
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Other Resources:
Distinguishing D. polymorpha from D. bugensis
Dreissena species (ANS Clearninghouse Bibliography)
Dreissena polymorpha (Global Invasive Species Database)
Pennsylvania Sea Grant Factsheet
Great Lakes Indian Fish and Wildlife Commission Maps
NOAA Sea Grant Nonindigenous Species Site (SGNIS)
Author: Benson, A. J. and D. Raikow
Contributing Agencies: 
NOAA - GLERL
Revision Date: 8/28/2009 Citation for this information:
Benson, A. J. and D. Raikow. 2010. Dreissena polymorpha. USGS Nonindigenous Aquatic Species Database, Gainesville, FL.
<http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=5> Revision Date: 8/28/2009
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