Dreissena polymorpha

Invasion History

First Non-native North American Tidal Record: 1991
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1991

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General Invasion History:

Dreissena polymorpha was first collected from the Caspian Sea by Pallas in 1769. Its native region is believed to be the Caspian and Aral Seas, and low-salinity lagoons of the Black Sea and adjacent rivers (Son 2007). Early in the 19th century, D. polymorpha invaded canals connecting to the Danube and Dnieper basins, and from there spread rapidly through Europe, reaching England and Prussia by 1825. This mussel is still invading new lakes in Europe (Karatayev et al. 1997; Ludyanskiy et al. 1993), and is abundant in low-salinity (0-6 ppt) estuaries from Finland and Baltic Russia to Ireland, Britain, France, Spain and the lagoons bordering the Black Sea (Strayer and Smith 1993).

In North America, Zebra Mussels were transported to many lakes and river systems through the 1990s and 2000s (Ludyanskiy et al. 1993; USGS Nonindigenous Aquatic Species 2012; Ram et al. 2011). In 2008, D. polymorpha found in the upper Colorado and Arkansas drainages in Colorado (Grand Lake and Pueblo Reservoir), a lake in Utah, and a reservoir in the Monterey Bay watershed, California (Center for Aquatic Resource Studies 2008). Canals and barges permitted zebra mussels to spread between many eastern and midwestern river basins, but many isolated rivers and lakes were infested through the transport of mussels on trailered boats or with fishing gear (Carlton 1993; Johnson et al. 2001; Karatayev et al. 2011; Kelly et al. 2012).

North American Invasion History:

Invasion History on the East Coast:

In North America, D. polymorpha was first reported in 1986, in Lake Erie, Ontario, fouling natural gas wellheads. Dead shells of D. polymorpha were collected in Lake Michigan, off East Chicago, Indiana in 1988, and in the same year the mussels were collected at several locations in Lake Ontario (Carlton 2008). A later discovery, in 1989, in Lake St. Clair, Ontario, between Lakes Erie and Michigan, was widely publicized (Mills et al. 1993). It was probably introduced to the Great Lakes system in ballast water of cargo ships from Europe. By 1990, it was found in all five Great Lakes, and the upper St. Lawrence River, covering a range of ~600 km north-south and 1400 km east-west, with populations being most continuous downstream of Lake St. Clair, and by 1992, it reached the estuary of the St. Lawrence River at Quebec City (freshwater) (Mellina and Rasmussen 1994; Ram et al. 2012; USGS Nonindigenous Species Program 2012). By 1993, it also colonized Lake Champlain (NY-VT) (Marsden and Hauser 2009).

In the freshwater St. Lawrence River, D. polymorpha was seen in August-September 1992, in the vicinity of Quebec City, in the rocky freshwater intertidal zone, at densities averaging 25 m-2. Mussels on the tops of boulders did not survive winter ice scour, but mussels survived in rock crevices (Mellina and Rasmussen 1994). By 2000, zebra mussel veligers were the most abundant zooplankton in the fresh-oligohaline region of the estuary, below Quebec City and the Ile d'Orleans, with densities as high as 260 L-1. Their abundance decreases sharply below 2 PSU, though a few veligers were found at 8 PSU (Barnard et al. 2003; Barnard 2006).

In the Hudson River estuary, Zebra Mussels were first seen in May 1991, at Catskill, New York (NY) about 200 km upriver (River Km 112) from the mouth of the Hudson in New York Harbor (River Km 0), and about 40 km downriver from the head of tide at Troy, NY (River Km 243) (Strayer et al. 1996). By the end of 1992, they were abundant in much of the tidal river, and found downriver as far as Tarrytown, NY~135 km downriver (River Km 65) where salinities reached 5-6 PSU (Walton 1996). A second invasion occurred in the lower Mohawk, a major tributary joining the Hudson at Troy in 1991. The two populations joined in 1992. By 1994, mean densities in the fresh tidal river were 10-1000 M-3. Possible vectors include boats or boat trailers from the Great Lakes, or barges from the Erie Canals (Strayer et al. 1996). While the eastern end of the canal was colonized by 1990, Zebra Mussels reached the Hudson before mussels were reported from the eastern parts of the canal or the lower Mohawk River (Strayer et al. 1996; USGS Nonindigenous Aquatic Species Program 2012). In the tidal fresh Hudson River, from Troy to Newburgh, the population showed dramatic fluctuations, with several peaks and crashes in abundance from 1992 to 2009, but with an overall trend of decreasing mussel size, biomass, filtration rate, and survivorship. One cause of the decline may be predation (Carlsson et al. 2011), but major predators such as Blue Crabs (Callinectes sapidus) have not increased in abundance. The mechanism of the decline and associated changes in Hudson River benthos are not clear (Strayer et al. 2011).

The invasion of Chesapeake Bay and its watershed had been long anticipated. In 1991, D. polymorpha veligers were reported from the Susquehanna River near Binghamton, NY (Lange and Cap 1991). Veligers reportedly occurred in low densities in 1991-1993, but then disappeared. In 2001, an established population of D. polymorpha was found in Eaton Brook Reservoir, on the Chenango River, in Madison County, New York, in the northern reaches of the Susquehanna Watershed. Based on the size of the mussels, the invasion probably began in 1999. They spread to other lakes in the region, and by 2004, they had reached the upper mainstem of the Susquehanna in Colliersville, NY (Otsego County). By 2007, they had reached Binghamton, NY and Finger Island, Pennsylvania (PA), in the Susquehanna (USGS Nonindigenous Aquatic Species Program 2012). In November 2008, a single Zebra mussel was found alive, inside an intake at the Conowingo Dam, just above the head of tide of Chesapeake Bay (Thomson 2008, USGS Nonindigenous Aquatic Species Programs 2009). In November 2008, D. polymorpha was also found further upstream in Pennsylvania, in Muddy Run, a tributary near the Maryland border. In December 2008 and May 2009, clumps of mussels were found above Conowingo Dam (Thomson 2008; Halsey 2009). In October 2011, one dead mussel was found attached to a jet-ski mooring in the Sassafras River near Betterton, Maryland (MD) (USGS Nonindigenous Aquatic Species Program 2012). A recent report (Klauda and Ashton 2013) lists additional collections in the lower Susquehanna River and Susquehanna Flats, at the head of the Bay. In the summer of 2015, settling juveniles were collected from the Bush, Gunpowder and Middle River estuaries, tributaries of the upper Bay (Wheeler 2015). Veligers have also been collected at power plants and drinking water intakes in the lower Susquehanna. This mussel appears to be established in the lower Susquehanna and is colonizing tidal fresh regions of upper Chesapeake Bay (Klauda and Ashton 2013; Ashton and Klauda 2015).

Scattered occurrences have occurred in other East Coast watersheds, but have not yet reached tidal waters. In 2001, D. polymorpha was found in East Twin Lake, CT, in the Berkshire Mountains and the watershed of the Housatonic River, ultimately draining into Long Island Sound. In 2009, it was found in another Berkshire lake, Laurel Lake in Lee, MA. The mussel is now found in the upper Housatonic around the MA-CT border, and in 2010-2011, the lower river, in Lake Zoar and Lake Housatonic, about 10-2 km from Long Island Sound (USGS Nonindigenous Aquatic Species Program 2012). Isolated populations were found in 2000 in a reservoir near Bethlehem, PA in the Delaware drainage, and in 2002, in a quarry in the Potomac watershed. The quarry population was believed to have been stocked illegally by divers to clear the water (USGS Nonindigenous Aquatic Species Program 2012). The mussels were successfully eradicated in 2006 (Virginia Department of Game and Inland Fisheries 2008 http://www.dgif.virginia.gov/zebramussels/).

Invasion History on the Gulf Coast:

In 1991, D. polymorpha was collected outside the Great Lakes-St. Lawrence system for the first time, colonizing the Illinois and Mississippi Rivers, by way of the Chicago Sanitary and Ship Canal (Tucker et al. 1993). It rapidly spread through much of the central Mississippi drainage, reaching the lower Mississippi (Vicksburg, Mississippi) by 1992 (Ludyanskiy et al. 1993), New Orleans, LA by 1993, and the mouth of the river by 1994 (USGS Nonindigenous Aquatic Species Program 2012). In 1995, Zebra Mussels were abundant in the Atchafalaya River basin, LA including the Intracoastal Canal, but subject to seasonal die-offs in summer (Mihuc et al. 1999). The mussels occur in many of the rivers and bayous of Louisiana's delta region (USGS Nonindigenous Aquatic Species Program 2012).

Invasion History Elsewhere in the World:

Dreissena polymorpha evolved in the Ponto-Caspian region, and is one of many recent and fossil Dreissena species known from the area. In preglacial times, it was widespread in Western Europe, but died off due to glaciation (Son 2007). It re-entered Western Europe very early, with the construction of canals in the 18th and early 19th centuries reaching the Rhine in Netherlands by 1826, the Elbe in Germany by 1838, and the Baltic basin by 1840 or 1850 (Karateyev et al. 1997; Bij de Vaate et al. 2002; Minchin et al. 2002; Birnbaum 2011). It was discovered in the Thames River, London, in 1826 and appeared in several other British ports in the 1830s, apparently with timber imported from mainland Europe (Aldridge et al. 2004). Zebra mussels continued to spread in the 20th century, not only through canals, but also with fishing gear, as bait, and with trailered boats, etc., to unconnected lakes and rivers. They appeared in Sweden in 1926 (Hallstan et al. 2010), Italy in 1971 (Lake Garda and Po estuary, Occhipinti Ambrogi 2002), Ireland in 1997 (Minchin 2005), and Spain in 2001 (Ebro River, Rajagopal et al. 2009), and are continuing to colonize new bodies of water.

Zebra Mussels are found in tidal fresh and oligohaline waters of many European estuaries, but are limited by intolerance of high salinity, air exposure, ice scour, and extremes of temperature. In the Baltic region, it is absent from the open sea, but is common in adjoining lagoons and estuaries, where low salinities and warmer temperatures permit reproduction (Strayer and Smith 1993). They reached Lake Ladoga, Russia, east of the Baltic by the mid-1800s. The mussels were present in the Polula River estuary, Estonia, in the Gulf of Finland, and Pärnu Bay in the Gulf of Riga. A later invasion of the eastern Gulf of Finland occurred in the 1930s, from Lake Peipsi (Estonia-Russia) via the Narva River (Birnbaum 2011). A more extensive invasion of the eastern Gulf of Finland, in the Neva River estuary, west of St. Petersburg, Russia, occurred in the 1980s. The northward expansion of the range of D. polymorpha may result from climate warming (Orlova and Panov 2004). Elsewhere in the Baltic, zebra mussels are present in the Gulf of Riga, Estonia-Latvia (in 1855, Birnbaum 2011; Leppakoski et al. 2002); the Curonian Lagoon, Lithuania (before 1850, Daunys et al. 2006); the Vistula Lagoon, Poland; and the Szczecin Lagoon, Poland (1924, Stanczykowska et al. 2010). In the Rhine Delta, in the Netherlands, D. polymorpha was limited to areas below an average of 0.6 PSU, and was absent from intertidal areas (Wolff 1969, cited by Strayer and Smith 1993).

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Description

Dreissena polymorpha belongs to the family of 'false mussels' (Dreissenidae), mostly associated with fresh and brackish (but sometimes estuarine) waters. They are distinguished from true mussels (Mytilidae) by having a small shelf-like platform, on the interior of both shells at the beak. This is the site of attachment of the adductor muscle. Like true mussels, they have an elongate, curved shell, narrowing at the umbo, and attach to hard substrates, by secreting strong threads, called byssus. Dreissena polymorpha is easily confused with D. bugensis (Quagga Mussel, native to the Ponto-Caspian region) and Mytilopsis leucophaeta (Conrad's False Mussel, native from Mexico to Chesapeake Bay) (Abbott 1974; Pathy and Mackie 1993). Mytilopsis leucophaeta is easily recognized by a prominent tooth at the dorsal corner of the platform, inside the shell's beak (Pathy and Mackie 1993).

Dreissena polymorpha has a sharply pointed umbo, and is strongly curved inward, ventroposterially, while D. bugensis curves outward. They have a ventrolateral ridge on each shell and often have a more rounded and higher dorsoanterior slope, sometimes with a winglike extension. The ventral surface of D. bugensis is arched inward, with a flattened surface where the shells join, so the shell usually stands up when placed on its ventral edge, whereas D. polymorpha has a keeled ventral edge, and falls over. The posterior end of the shell is angled ventrally, while in D. bugensis, this end is rounded. When viewed from the edge, the two valves of D. polymorpha are symmetrical, with the edges of the two shells forming a straight line, while in D. bugensis, the shells are asymmetrical and join in a curved line. The color patterns are highly variable in both D. bugensis and D. polymorpha. Dreissena polymorpha can be all brown, black, or white, or have various striped patterns on the exterior. This mussel can grow to 5 cm in length, but is typically much smaller (Pathy and Mackie 1993; USGS Nonindigenous Aquatic Species Program 2012). Larval development of D. polymorpha and D. bugensis is described by Nichols and Black (1994).

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Taxonomy

Ecology

General:

Dreissena polymorpha has separate sexes which release eggs and sperm into the water, resulting in planktonic larvae, first a trochophore, and then a shelled veliger. Fecundity varies with size, from around 500 eggs for females with shells ~10 mm long to 150-300,000 for females 20-25 mm long (Stoeckel et al. 2004). Annual fecundity is estimated at 960,000 embryos per year (Keller et al. 2007). Some females can produce more than one million eggs per spawning (Sprung 1993). Laboratory reared larvae of D. polymorpha, at 22-24?C, reached the pediveliger stage, and began to settle at about 15-23 days from fertilization (Wright et al. 1996; Stoeckel et al. 2004). Successful reproduction and larval development occurs at 12-27?C (Sprung 1993; Fong et al. 1995; Wright et al. 1996). Reproduction is most successful in freshwater, but fertilization and development take place successfully at 3.5 PSU, with acclimated animals (Fong et al. 1995). Veligers can be extremely abundant. They are the dominant form of zooplankton in the estuarine transition zone of the St. Lawrence River, but their source of nutrition, bacteria and dissolved organic carbon, were not used by native plankton. Isotopic analysis indicates that they are not heavily fed on by native fishes (Barnard et al. 2006).

Zebra Mussels settle on hard substrate such as rock, wood, and man-made structures, but also on vegetation (Sprung 1993; Mellina and Rasmussen 1994; Strayer et al. 1996). Soft bottom sediments including sand, silt and mud are usually regarded as unsuitable habitat. They use byssal threads to attach to hard surfaces, but can settle in clumps on soft substrates, attaching to scattered shells and other hard objects, or the sediment surface (Berkman et al. 1998). They are most often found in shallow waters, but have been reported at over 110 m depth (Mackie 1993; Martel et al. 2001; Ricciardi and Whoriskey 2004). In the St. Lawrence estuary, Quebec, some Zebra Mussels occurred and survived the winter in intertidal rock crevices (Mellina and Rasmussen 1994), but usually, they are absent in intertidal areas, due to temperature extremes and ice scour (Strayer and Smith 1993; Strayer et al. 1996).

Adult Zebra Mussels tolerate short exposures to temperatures as high as 37?C with acclimation (Spidle et al. 1995), but the upper limit for long-term survival is about 30?C (Iwanyzki and McCauley 1993; McMahon 1996). They do tolerate temperatures near zero, and survive in ice-covered lakes, but require temperatures of at least 10-12?C for optimal feeding, growth, and reproduction (McMahon 1996). North American populations appear to have a higher upper thermal limit, than European ones, ~ 30?C vs. 27-28?C (McMahon 1996), which could be due to genetic differences, acclimation or to different experimental methods.

Reported salinity tolerances vary geographically, partly with the composition of salts involved, and with experimental methods. Zebra Mussels are well-established in the northern Caspian Sea, at 6-9 g. L-1, and, formerly, in the Aral Sea at 10 g. L-1, but the salts of these inland lakes differ in composition from seawater (Strayer and Smith 1993). Zebra mussels could not tolerate pure NaCl above 5 g. L-1 (Spidle et al. 1995), but natural or artificial seawater appears to be less toxic. Adult Zebra mussels survive gradual acclimation to 10 PSU (Wright et al. 1996), but may have much lower salinity limits in areas with variable salinities (~2 PSU, Sea of Azov, Zenkevich 1965; 0.6 PSU, Rhine Delta, Wolff 1969, both cited by Strayer and Smith 1993), or additional stresses, such as air exposure.  In simulated ship trasport experiments, a few (>0.1%) survived up to 8 days in marine waters (35#% PSU) suggesting that fouling transport is possible, but improbalbe at low temperatures (Riley et al. 2022).

In fresh waters, other ions can affect the invasibility of D. polymorpha. The lower pH limit for adult mussels has been reported to be about 7.5, with dissolution of shells occurring at lower levels (Baker et al. 1994; McMahon 1996; Claudi et al. 2012). Calcium concentrations are another water-quality factor which may limit the distribution of D. polymorpha, and in the nontidal St. Lawrence River, Zebra Mussels occurred at levels as low as 7.5 mg Ca .L-1 (Jones and Ricciardi 2005). However, higher concentrations (18-21 mg L-1) greatly improve survival and growth (McMahon 1996; Baldwin et al. 2012). Zebra Mussels have high oxygen requirements, and survive severe hypoxia (3% saturation) only for 3-5 days at 25?C, which may limit their distribution in eutrophic waters (McMahon 1996; Matthews and McMahon 1999).

Dreissenid mussels are suspension feeders, pumping large quantities of water through their gills, retaining particles, and sloughing off surplus or inedible particles as pseudofeces. Zebra Mussels maintained high filtration rates for particles 10-150 µm. Large mussels were capable of capturing particles up to 900-1200 µm, which can include small zooplankton, and large, chain-forming diatoms. However, they have some degree of selectivity, and can exclude detritus, inorganic particles, or toxic items, such as toxic Microcystis colonies (Horgan and Mills 1997; Baker 1998; Vanderploeg et al. 2001). However, in the Hudson River estuary, terrestrial detritus is estimated to constitute ~40% of the Zebra Mussels' diet (Cole and Solomon 2012).

Food:

Phytoplankton, detritus, small zooplankton

Consumers:

Fishes, birds, crayfish

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Grass Bed	None
General Habitat	Coarse Woody Debris	None
General Habitat	Nontidal Freshwater	None
General Habitat	Tidal Fresh Marsh	None
General Habitat	Unstructured Bottom	None
General Habitat	Marinas & Docks	None
General Habitat	Canals	None
General Habitat	Rocky	None
Salinity Range	Limnetic	0-0.5 PSU
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	0	Baker et al. 1994; Fong et al. 1995; IIwanyzki and McCauley 1993; Ludyanskiy et al. 1993; McMahon 1996
Maximum Temperature (ºC)	34	Baker et al. 1994; Fong et al. 1995; Iwanyzki and McCauley 1993; Ludyanskiy et al. 1993; McMahon 1996
Minimum Salinity (‰)	0	None
Maximum Salinity (‰)	10	Fong et al. 1995; Kilgour et al. 1994; Strayer and Smith 1993; Wright et al. 1996. Salinity- Results presented by various authors have differed as a result of methods used, including different acclimation regimes and salt solutions used. [Spidle et al. (1995) used NaCl solutions, which may not be comparable to seawater, because of the lack of physiologically necessary ions. Observations in the species' native Caspian-Aral region are also not comparable because of differences in ionic composition.] The 10 ppt 'Survival' value was for juvenile mussels gradually acclimated to artificial seawater (1ppt/day) and then kept for 7.5 months (Wright et al. 1996). In simulated voyages, using animals from the Netherlands, D. polymorpha, had a high tolerance at 0.2-6.0 PSU (van der Gaag et al. 2016), with 164-308 days for 100% mortality.
Minimum Dissolved Oxygen (mg/l)	1.7	McMahon 1996
Minimum pH	7	Baker et al. 1994; McMahon 1996; Claudi et al. 2012
Maximum pH	9	Baker et al. 1994; McMahon 1996; Claudi et al. 2012
Minimum Reproductive Temperature	12	Sprung 1996
Maximum Reproductive Temperature	27	Sprung 1996
Minimum Reproductive Salinity	0	This is a freshwater species.
Maximum Reproductive Salinity	3.5	for fertilization of D. polymorpha eggs (Fong et al. 1995).
Minimum Duration	8	From fertilization to settlement (Sprung 1993, temperature not specified)
Maximum Duration	33	Sprung 1993 (Sprung 1993, temperature not specified)
Minimum Length (mm)	10	Size at maturity (Ludyanskiy et al. 1993)
Maximum Length (mm)	50	Ludyanskiy et al. 1993
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Nontidal Limnetic-Oligohaline

General Impacts

Dreissena polymorpha (Zebra Mussel) has been listed by the Invasive Species Specialist Group of the World Conservation Union (IUCN) as one of the '100 worst invasive species.' In North America, it has been the 'poster child' of aquatic invasive species, because of its wide-ranging economic and ecological impacts. In the 1990s, the Zebra Mussel invasion attracted much attention in the media and among the public, spurring the adoption of legislation in the US and Canada to regulate ballast water discharges and other vectors of biological introductions (Vasarhelyi and Thomas 2003). Impacts of Zebra Mussels have been widely reported from European lakes, rivers, and estuaries, and from the Hudson River, Great Lakes, and Mississippi Rivers. While many of the impacts are similar among water-bodies, some systems have shown differing responses to zebra mussel invasion, resulting from ecological and biotic differences (Ludyanskiy et al. 1993; MacIsaac 1996; Karatayev et al. 2002; Minchin et al. 2005; Strayer et al. 2011).

Economic impacts

Zebra Mussels in the Great Lakes were first noticed as a very troublesome invader in 1988-1990, fouling natural gas wells, power plants, waterworks, boats, and docks, exacting substantial costs in cleaning and removal (Kovalak et al. 1993; LePage 1993; Carlton 2008). Subsequently, drastic ecological changes occurred in the lakes, with mixed costs and benefits for fisheries and recreation, including greatly increased water clarity, changes in food webs, adversely affecting some fish species and benefiting others, increased growth of submerged vegetation, dead mussels and shells washing up on shore, blooms of toxic 'blue-green' algae, etc. (Ludyanskiy et al. 1993; MacIsaac 1996; Limburg et al. 2010). Colautti et al. (2006) estimated the costs of Zebra Mussels to power plants in Canada to be $CAN 6-7 million per year. Lovell et al. (2006) present a set of greatly diverging estimates for the economic impacts of Zebra Mussels from $83 million to $3 billion per year, depending on the time period, and what costs are included. These estimates are largely based on costs to power plants and water filtration plants. Estimating aesthetic and recreational costs and benefits is more difficult, since these depend heavily on perceptions (Lovell et al. 2006; Limburg et al. 2010).

Industry- The earliest observations of Zebra Mussels in the Great Lakes involved fouling of natural gas wells in Lake Erie and Ontario, causing damage to the wellheads by 1990 (Carlton 2008). Zebra Mussels have long been recognized as a problem for power plants and other systems requiring large-scale water use in Eastern European waters, where large reservoirs provide an especially favorable habitat (Ludyanskiy et al. 1993; Minchin et al. 2005). In the Great Lakes, D. polymorpha were observed in the 10 power plants of the Detroit Edison system in 1988. Extensive fouling first occurred in the Monroe, MI plant, at the western end of Lake Erie in the summer of 1989, requiring mechanical cleaning and the use of chlorination, costing ~$600,000 in 1989-1991. While the Monroe plant was most heavily impacted, similar problems occurred throughout the Detroit Edison system (Kovalak et al. 1992). Fouling of intakes varied greatly with the size of the intake and the water flow. Fouling was less extensive at plants that used less cooling water (Kovalak et al. 1992), while very high flow velocities can discourage settlement (MacIsaac 1996). In the tidal fresh Hudson River, fouling of power plants has required the use of various biocides, at a cost ranging from $100,000 to $1 million per year (Strayer 2006).

Health- Zebra Mussel fouling has had adverse impacts on municipal water supplies which depend on filtration of water from lakes and rivers. Water filtration plants on Lake St. Clair (Windsor, Ontario), Lake Erie (Monroe, MI), and the Hudson River had major disruptions of water supplies, and have had large expenditures on water cleaning and chlorination in order to maintain water flow, and control unpleasant tastes and odors (LePage 1993; Colautti et al. 2006; Strayer 2006). A unique health impact is an increase in cuts on the feet of people bathing and wading, due to shells in shallow water and on beaches. These lacerations may be prone to bacterial infections, especially in the vicinity of sewage discharges (Minchin et al. 2005).

Shipping and boating- Settlement of Zebra Mussels in Europe, the Great Lakes, and the Hudson River has affected recreational boats, docks, and navigational buoys (MacIsaac 1996; Minchin et al. 2005; Strayer 2006). One study found that increased cleaning, painting, maintenance, and insurance can cost recreational boat owners on Lake Eire about $600 per boat, but this was based on a small sample size (Vilaplana and Hushak 1994, cited by Lovell et al. 2006).

Aesthetic- The invasion and development of Zebra Mussel populations has had contrasting impacts to people's enjoyment of lakes and rivers. The vast biomass of the mussels, and their huge filtering capacity has increased clarity of the water, increasing its attractiveness, but at the same time has resulted in increased growth of macrophytes (often perceived as 'weeds' by fishermen and boaters), filamentous algae, and in some bodies of water, promoted the growth of large colonies of the cyanobacterium ('blue-green alga') Microcystis sp., which are too large for the mussels to filter (Vanderploeg et al. 2001; Limburg et al. 2010). On Lake Ontario, the majority of homeowners surveyed considered that water quality had improved, and that this contributed to an increase in property values, but others noted problems with algal blooms, and perceived declines in fisheries (Limburg et al. 2010). In addition, when mussels die and are dislodged by storms, the dead animals and their shells create unpleasant odors, and can make walking on beaches unpleasant (Minchin et al. 2005; Limburg et al. 2010).

Fisheries- Economic impacts of the Zebra Mussel on commercial and sport fisheries are difficult to determine, because of the complexity of food webs, and the ability of many fishes to use alternate prey. In addition, in the Great Lakes, impacts of the Zebra Mussel are difficult to separate from those of the Quagga Mussel, although the latter predominates in the colder and deeper waters of the lakes, while Zebra Mussels are most abundant in shallower and warmer waters. In the Great Lakes, the reduction of phytoplankton and zooplankton biomass could be expected to adversely affect planktivorous adult fishes, and the larvae and juveniles of many other species. Given the depth and size of the Great Lakes, grazing by the Quagga Mussel has probably had the greatest impact on open-water planktivorous fishes (Vanderploeg et al. 2001; Cuhel and Aguilar 2013). At the same time, the recovery and growth of macrophytes and algae in shallow water provides habitat for many littoral species, important to sport fisheries, such as Yellow Perch (Perca flavescens), Smallmouth Bass (Micropterus dolomieu) and Muskellunge (Esox masquinongy) (Vanderploeg et al. 2001). In the Hudson River, open-water fishes, such as American Shad (Alosa sapidissima), Alewife (Alosa pseudoharengus), and White Perch (Morone americana) were negatively affected by the Zebra Mussel invasion, while nearshore species, including Bluegill (Lepomis macrochirus), Redbreast Sunfish (L. auritus), and Smallmouth Bass (M. dolomieu), benefited from increased macrophyte growth (Strayer et al. 2004).

Ecological Impacts

Reports of the ecological impacts of Zebra Mussels from different bodies of water have many common features, including: the establishment of dense populations, increases in water clarity, decreases in the abundance and/or diversity of other macrobenthic species, and habitat changes due to the creation of new structure on the bottom (Strayer et al. 1999; Karatayev 2002; Vanderploeg et al. 2001).

Herbivory- Many of the wide-ranging ecological impacts of Zebra Mussels can be tied to their combined capacity for high rates of filtration, with rapid reproduction and dispersal, creating large biomasses capable of removing large portions of phytoplankton per day. At the peak of its invasion, the Zebra Mussel population filtered the whole volume of the tidal Hudson River in 2 days (Roditi et al. 1996), while in Saginaw Bay, Lake Huron, the estimated filtering time was 1-5 days (Budd 2001). At the same time, Zebra Mussels do have some ability to select their food, by varying filtration rates, and by rejecting particles such as detritus, inorganic sediment, or less desirable phytoplankton, as pseudofeces (Ludyanskiy et al. 1993; Horgan and Mills 1997; Baker 1998). In many bodies of water, Zebra Mussel invasions often result in dramatic reductions in phytoplankton biomass (measured as chlorophyll a) (Leach 1993; Caraco et al. 1997; Budd 2001).

A notable difference among systems is that some invaded areas, such as Lake Erie and Lake Ontario, develop blooms of large colonies of cyanobacteria (often dominated by Microcystis spp.), too big for mussels to filter, using the nutrients which formerly fueled blooms of edible phytoplankton (Karatyev et al. 2002; Vanderploeg et al. 2001). These blooms are not seen in the Hudson River estuary, where Microcystis is present, but the colonies apparently do not reach an inedible size, and are controlled by Dreissena's grazing (Fernald et al. 2007; Strayer et al. 2008). For reasons which are unclear, Dreissena spp. favor Microcystis blooms in low-nutrient, but not high-nutrient lakes (Raikow et al. 2004).

Predation- Zebra Mussels' strong filtration currents capture small zooplankton, such as rotifers, tintinnids, copepod nauplii, and their own veligers. Reductions in microzooplankton in the presence of mussels were seen in the tidal Hudson River (Pace et al. 1998) and the nontidal St. Lawrence River (Thorp and Casper 2002).

Competition- One of the widespread impacts on native species has been the effect of Zebra Mussel settlement on the larger native freshwater mussels of the family Unionidae. This has been seen in European lakes (Karatyev et al. 2002) and Baltic lagoons (Orlova et al. 2006; Zaiko et al. 2009) in the Hudson River (Strayer and Smith 1996; Strayer et al. 1999), Great Lakes-St. Lawrence River (Schloesser et al. 1996; Ricciardi et al. 1998), and in the Mississippi Basin (Tucker et al. 1993). Fouling can interfere with burrowing, and reduce feeding and growth. This is a particular concern in North America where the diversity of unionids is especially great, and where many species are already threatened by pollution and disturbance of streams and lakes (Schloesser et al. 1996; Ricciardi et al. 1998). At least one species of native mussel (Amblema plicata) in Lake Erie was able to eliminate fouling by Zebra Mussels by burrowing in mud (Nichols and Wilcox 1997). In the Hudson River, there was a sharp decline in native unionid mussels and in pea-clams (Sphaeriidae) with the onset of the Zebra Mussel invasion. The decline in the pea-clams, which are too small to be fouled, suggests that competition for food, and reduction of the phytoplankton biomass was the major mechanism of decline (Strayer and Smith 1996; Strayer et al. 1999). In later years, unionid mussels and sphaeriid clams showed some recovery, but the mechanism for this was not clear (Strayer and Malcolm 2006). During the invasion, a decline was also seen in two groups of filter-feeding midge larvae, the attached case-building tanytarsines, and the planktonic Chaoborus spp. (Strayer and Smith 2001).

Habitat Change- Zebra Mussel invasions have dramatically altered habitats in two major ways. First, by attaching themselves to substrates they alter the structure and complexity of the benthos and second, through intense filtration they change the properties of whole water bodies by removing phytoplankton and other particles from the water column, and depositing them in the sediment. As a result, light penetration often becomes greatly increased, and is accompanied by the growth of vascular macrophytes and filamentous algae, increasing the amount of shelter for small invertebrates and fishes. Increased light penetration has resulted in an increase in shallow-water macrophytes (Griffiths 1993; Leach 1993; Caraco et al. 1997; Budd 2001; Strayer et al. 1999; Strayer and Smith 2001; Strayer et al. 2011). The increase in macrophytes in the littoral zones of the Great Lakes and the Hudson River has favored shallow-water invertebrates and fishes (Strayer et al. 1998; Strayer and Smith 2001; Strayer et al. 2004).

Zebra Mussels alter hard substrates by increasing their complexity and by depositing pseudofeces, increasing sedimentation (Daunys et al. 2006; Zaiko et al. 2009). On soft substrates, the transformation can be dramatic, adding a three-dimensional aspect to a flat habitat, providing shelter for prey, and stabilizing the sediment (Berkman et al. 1998; Beekey et al. 2004a). The cryptogenic amphipod Gammarus fasciatus and the introduced Echinogammarus ischnus both used Zebra Mussel beds as shelter, with about equal frequency (Palmer and Ricciardi 2005; Kang et al. 2007). Again, deposition of pseudofeces increases the organic content of the sediment, and may increase the diversity of macroinvertebrates (Beekey et al. 2004a). However, in the Hudson River, a negative effect was seen on benthic macroinvertebrates in deeper water, attributed to the overall decrease in phytoplankton sinking to the bottom (Strayer et al. 1998; Strayer and Smith 2001).

Food/Prey- The huge biomasses developed by Zebra Mussel populations present a large food resource for predators capable of cracking the mussel shells. In the Hudson River estuary, Blue Crabs (Callinectes sapidus) are significant predators of Zebra Mussels (Boles and Lipcius 1997; Carlsson et al. 2011). Increased mortality due to predation seems to have played a major part in the decline of Zebra Mussels in the Hudson River, but effects on the abundance of Blue Crabs or other predators are unknown (Strayer et al. 2011). The Rusty Crayfish (Orconectes rusticus) sharply reduced Zebra Mussel abundance in a Minnesota stream (Christiana Creek) (Perry et al. 2000), but had negligible impact on Zebra Mussel populations in nearshore rocky habitat in Lake Erie (Stewart et al. 1998). Round Gobies (Neogobius melanostomus) also of Ponto-Caspian origin, feed extensively on young Zebra Mussels (Ray and Corkum 1997). In the Great Lakes and Baltic Sea, they are the only small forage fish which preys heavily on Zebra and Quagga Mussels, so the invasions of these species are linked (Kornis et al. 2012). In the Great Lakes, Zebra and Quagga Mussels have become major prey for Lake Whitefish (Coregonus clupeaformis), an important fisheries species, as well as forage for large game fishes (Cuhel and Agular 2013). Other fishes, such as Common Carp (Cyprinus carpio) and Redear Sunfish (Lepomis microlophus) do feed on Zebra Mussels, but consume a wide variety of other prey, so the effect on their populations is probably small (French and Morgan 1995; Tucker et al. 1996). Large flocks of diving ducks (mostly Greater and Lesser Scaup, Athya marila and A. affinis) have been found to be feeding on Zebra Mussels in the Great Lakes, but effects on these migratory bird populations are unknown (Hamilton et al. 1994).

Trophic Cascade- The invasion of Zebra Mussels has caused dramatic changes throughout the food webs of the Great Lakes, the fresh tidal Hudson River, and to a lesser extent, Baltic lagoons. These effects are not just on their food (phytoplankton, microzooplankton) or their direct predators, but also on indirectly connected components, such as macrophytes and filamentous algae, top-predator fishes, inorganic nutrients, and light penetration (Evans et al. 2011; Strayer et al. 2011; Cuhel and Aguilar 2013). In these systems, dreissenid mussels remove large quantities of phytoplankton from the water, and excrete their accumulated nitrogen and phosphorus into the water, where it can be utilized by filamentous algae, macrophytes, and inedible phytoplankton, such as Microcystis sp. (Raikow et al. 2004; Conroy and Culver 2005; Strayer et al. 2011). The intense filtration pressure of Zebra mussels changes the distribution of light energy, from absorbance in the water column by phytoplankton, to penetration on the bottom, promoting the colonization (or re-colonization, in many eutrophic bodies of water) of macrophytes and filamentous algae (Budd 2001; Strayer et al. 2011). The macrophyte and algal communities in turn provide food and shelter for invertebrates, and support smaller forage fishes (e.g. killifishes, sunfishes) and larger predators, such as Smallmouth Bass (Micropterus dolomieu) (Vanderploeg et al. 2001; Strayer et al. 2004). These changes in the food web are complex- in some respects they have reduced the impact of human-caused eutrophication, and led to a partial oligotrophication of these systems (Evans et al. 2011; Cuhel and Aguilar 2013). At the same time, the dreissenid invasions may have reduced the resiliency of ecosystems, by creating a large biomass component, subject to limited predation, which slows the transfer of energy between trophic levels (Conroy and Culver 2005). These changes in food web dynamics may have to be incorporated into regional policies of nutrient management (Evans et al. 2011).

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Regional Impacts

B-IX	None		Ecological Impact	Competition
Fouling of freshwater mussels (Unionidae) by Zebra Mussels occurred in the inner Gulf of Finland, but at lower frequency than seen in North America (Orlova et al. 2006). Impacts on benthic communities were characterized as 'moderate' (Zaiko et al. 2011).
B-IX	None		Ecological Impact	Habitat Change
Zebra Mussels were reported to be increasing water clarity and promoting the growth of benthic algae (Orlova et al. 2006). The scale of these habitat impacts was characterized as moderate (Zaiko et al. 2011).
B-IX	None		Ecological Impact	Trophic Cascade
In the inner Gulf of Finland, Zebra Mussels deposited large quantities of excreted nutrients in sediments, providing nutrition for an increased abundance of deposit-feeding benthos, and promoting the growth of macroalgae (Cladophora sp.) (Orlova et al. 2006). The scale of these impacts on ecosystem function was reported as 'moderate' (Zaiko et al. 2011).
B-V	None		Ecological Impact	Competition
Zebra Mussels were considered to have moderate community impacts in the Szczecin Lagoon and Oder/Odra estuary (Zaiko et al. 2011)
B-V	None		Ecological Impact	Herbivory
Zebra Mussels were considered to have moderate ecosystem impacts, in the Szczecin Lagoon and Oder/Odra estuary, assumed to include suspension-feeding (Zaiko et al. 2011).
B-VIII	None		Ecological Impact	Habitat Change
Moderate habitat impacts (Zaiko et al. 2011).
B-VIII	None		Ecological Impact	Trophic Cascade
Moderate ecosystem impacts (Zaiko et al. 2011).
B-VII	None		Ecological Impact	Competition
In the Curonian Lagoon, Zebra Mussels contributed up to 95% of total benthic community biomass and fouled native unionid mussels (Zaiko et al. 2009). Dreissena polymorpha was also a biomass dominant in fresher portions of the Vistula Lagoon, Poland (Ezhova et al. 2005). It was considered to have strong community impacts in the Curonian and Vistula Lagoons (Zaiko et al. 2011).
B-VII	None		Ecological Impact	Habitat Change
In the Curonian Lagoon, Zebra Mussel shell deposits and living beds had higher benthic invertebrate biomass and species richness than bare sediment. The effect of living mussels was greater than that of dead shells (Zaiko et al. 2009). Zebra mussels were considered to have strong habitat impacts (Zaiko et al. 2011).
B-VII	None		Ecological Impact	Trophic Cascade
Zebra Mussels were considered to have moderate ecosystem impacts in the Curonian Lagoon. Data were not available for an assessment in the Vistula Lagoon (Zaiko et al. 2011).
L123	_CDA_L123 (St. Lawrence River)		Ecological Impact	Predation
In Robinson Bay, off the St. Lawrence River, near Massena NY, in mesh enclosures containing Dreissena polymorpha, the abundance of the rotifer Polyarthra sp. declined drastically, indicating predation on this and other microzooplankton. Enclosures with the native mussel Elliptio complanata showed no change in rotifer abundance. Chlorophyll levels in the treatments did not differ, indicating that the effect was due to predation (Thorp and Casper 2002).
L123	_CDA_L123 (St. Lawrence River)		Ecological Impact	Trophic Cascade
In Robinson Bay, of the St. Lawrence River, near Massena NY, in mesh enclosures containing Dreissena polymorpha, abundances of the copepods Eurytemora carolleeae (reported as E. affinis) increased dramatically, presumably due to reduction of competition from rotifers (Thorp and Casper 2002).
GL-II	Lake Erie		Economic Impact	Industry
Fouling of natural gas wellheads by zebra mussels, off Ontario, in Lake Erie, caused maintenance problems by 1990 (Carlton 2008). Zebra Mussels caused extensive fouling of the Detroit Edison's Monroe, MI coal-fired power plant at the western end of Lake Erie. Mussels covered the intake surfaces, blocked the trash bars, and fouled the condenser tubes. The fouled parts of the plants were cleaned with high-pressure water at a cost of $25,000-35,000 for each cleaning. Service water lines for fire-protection systems were also fouled, and cleared with chlorination, but regular use is limited by environmental concerns (Kovalak et al. 1993).
L098	_CDA_L098 (Black-Rocky)		Economic Impact	Industry
Fouling of natural gas wellheads by zebra mussels, off Ontario, caused maintenance problems by 1990 (Carlton 2008).
L099	_CDA_L099 (Cuyahoga)		Economic Impact	Industry
Fouling of natural gas wellheads by zebra mussels, off Ontario, caused maintenance problems by 1990 (Carlton 2008).
L106	_CDA_L106 (Niagara)		Economic Impact	Industry
Fouling of natural gas wellheads by zebra mussels, off Ontario, caused maintenance problems by 1990 (Carlton 2008).
L105	_CDA_L105 (Buffalo-Eighteenmile)		Economic Impact	Industry
Fouling of natural gas wellheads by zebra mussels, off Ontario, caused maintenance problems by 1990 (Carlton 2008).
L103	_CDA_L103 (Chautauqua-Connaut)		Economic Impact	Industry
Fouling of natural gas wellheads by zebra mussels, off Ontario, caused maintenance problems by 1990 (Carlton 2008).
M060	Hudson River/Raritan Bay		Ecological Impact	Herbivory
The Zebra Mussel invasion in the tidal fresh Hudson River resulted in an 85% decline in average phytoplankton biomass from 1987-1991 to 1993-1994. Light availability increased, as did phosphorus concentrations, while some planktonic grazers decreased. Flow characteristics of the river had not changed, supporting the hypothesis that grazing by the mussels was responsible (Caraco et al. 1997; Strayer et al. 1999). Laboratory grazing studies indicated that the biomass of zebra mussels could filter the tidal freshwater Hudson River in about two days (Roditi et al. 1996).The filtration rate has declined about 82% from its highest peak, in 1996, apparently due to increased mortality of mussels, and decreasing body size and biomass (Strayer et al. 2011). The toxic bloom-forming cyanobacterium Microcystis sp. was positively correlated with Zebra Mussel filtration rate, which is at odds with its behavior in other eutrophic systems, where high rates of Zebra Mussel grazing and nutrient release have promoted blooms of large, inedible colonies. Reasons for the absence of these cyanobacterial blooms in the Hudson River are not clear (Fernald et al. 2007).
M060	Hudson River/Raritan Bay		Ecological Impact	Habitat Change
Grazing by the Zebra Mussels resulted in a 39% decrease in the light extinction coefficient, which indicates a sharp increase in light penetration, due to the removal of phytoplankton (Caraco et al. 1997; Strayer et al. 1999). Increased light penetration has resulted in an increase in shallow-water macrophytes (Strayer and Smith 2001; Strayer et al. 2011).
M060	Hudson River/Raritan Bay		Ecological Impact	Predation
Filter-feeding by Zebra Mussels in the Hudson River resulted in a sharp decrease in the abundance of ciliates, rotifers, and copepod nauplii, apparently due to direct predation. The total biomass of zooplankton declined by about 70% after the invasion, due in part to predation (Pace et al. 1998).
M060	Hudson River/Raritan Bay		Ecological Impact	Competition
Zebra mussels adversely affected native mussels of the family Unionidae (especially Anodonta implicata and Leptodea ochracea) by settling on them and fouling them. Densities of the native mussels, during the invasion, fell by 56% and numbers of recruits fell by 90% during 1992-1995. A decrease in condition of unionid mussels, and a decline in small sphaeriid (Pisidium spp., Sphaerium spp.) clams, not subject to fouling, suggests that competition for phytoplankton food was also affecting native bivalves (Strayer and Smith 1996; Strayer et al. 1999). Later analyses suggested that declines in recruitment and condition during the early years of the invasion were more closely related to zebra mussel filtration, and thus food competition, rather than fouling. From 2000 to 2005, the decline of native bivalves stopped, and abundances stabilized, even showing some recovery, but the mechanism for this is not clear (Strayer and Malcom 2006).Two groups of filter-feeding midge larvae, tanytarsini midges, and Chaoborus spp. declined during the zebra mussel invasion (Strayer and Smith 2001).
M060	Hudson River/Raritan Bay		Ecological Impact	Trophic Cascade
The Zebra Mussel invasion in the Hudson River had wide-ranging effects on the estuary's food web. Effects on macrobenthos were complex. In deep-water samples, the abundance of benthic animals, mostly deposit-feeders, declined, because of the reduction of edible particles reaching the bottom. However, in shallow water, many groups of benthic invertebrates increased in abundance, probably because of increased growth of algae and macrophytes (Strayer et al. 1998; Strayer and Smith 2001). By 2000, populations of most deepwater macrobenthic species had recovered, apparently due to reduced biomass and decreased filtration rates of the Zebra Mussel population. However, shallow-water invertebrates remained at post-invasion levels (Strayer et al. 2011). The abundances of some fishes appear to have been affected by the mussel invasion. Some open-water species, particularly juveniles of Alewife (Alosa pseudoharengus) and White Perch (Morone americana) decreased during the invasion, while several littoral species increased, including Banded Killifish (Fundulus diaphanus), Bluegill (Lepomis macrochirus), Redbreast Sunfish (L. auritus), Smallmouth Bass (Micropterus dolomieu), and Tessellated Darter (Etheostoma olmstedi). Open-water fishes tended to shift their distribution downriver, while littoral fishes shifted upriver. Reduction in phytoplankton biomass and the planktonic part of the food web is believed to be the major factor in the shift in distribution and abundance of the open-water fishes, while the increase of shallow-water macrophytes and algae, as shelter for fishes and their prey, due to increased light penetration has benefited the littoral fishes (Strayer et al. 2004). Regions of the upper Hudson estuary witn more intense Zebra Mussel grazing had poor condtion and lower gut volume of Striped Bass (Morone saxatilis) larvae (Smircich et al. 2017). The general impact of Zebra mussel grazing has been to strengthen littoral food webs and increase biomasses, while weakening the planktonic and deepwater benthic food webs, and decreasing biomasses there (Strayer et al. 2008; Strayer et al. 2011).
M060	Hudson River/Raritan Bay		Ecological Impact	Food/Prey
Field and experimental studies indicate that predation by Blue Crabs (Callinectes sapidus) causes extensive mortality to Zebra Mussels in the Hudson River estuary (Boles and Lipcius 1997; Carlsson et al. 2011). Increased mortality has apparently stabilized the mussel population. However, it is not known if the Zebra Mussel has affected the abundance or distribution of Blue Crabs, or other predators in the Hudson River (Strayer et al. 2011).
L123	_CDA_L123 (St. Lawrence River)		Ecological Impact	Habitat Change
In Lake Champlain VT-NY, Zebra Mussels have extensively colonized soft sediment. Colonized sediment supported communities with a greater abundance and diversity of benthic invertebrates than adjacent sediments, lacking mussels. Experiments in which mussels were added to uncolonized sediment, or removed from colonized sediment also showed that mussels promoted increased abundance and diversity of macrobenthos (Beekey et al. 2004a). Zebra Mussels also adversely affected the foraging success of 3 benthic fishes and a crayfish, by providing shelter to prey organisms. However, the shelter effect may be offset by the increase in the density of prey (Beekey et al. 2004b). In the St. Lawrence River, near Montreal, both the introduced amphipod Echinogammarus ischnus and the native Gammarus fasciatus used Zebra Mussel colonies as shelter, about equally (Palmer and Ricciardi 2005).
GL-III	Lake Ontario		Economic Impact	Industry
Fouling of natural gas wellheads by zebra mussels, in Lake Ontario, caused maintenance problems by 1990 (Carlton 2008).
L071	_CDA_L071 (Saginaw River)		Ecological Impact	Herbivory
By 1992-1993, the biomass of Zebra Mussels in inner Saginaw Bay, Lake Huron, had a filtering capacity of 0.2-1.2 X the volume of the inner Bay per day. Chlorophyll and suspended solids were greatly reduced, and the decreased reflectivity (increased transparency) of the water was detectable by satellite imagery (Budd et al. 2001). Filtration by Zebra Mussels was selective - mussels ingested small, desirable flagellates, while rejecting large colonies of toxic Microcystis cyanobacteria in Lake Saginaw water (Vanderploeg et al. 2001).
L071	_CDA_L071 (Saginaw River)		Ecological Impact	Habitat Change
Filtration of the water by Zebra Mussels in inner Saginaw Bay, Lake Huron, resulted in greatly increased transparency and light penetration of the water, within 2-3 years after the initial invasion (Budd et al. 2001).
GL-I	Lakes Huron, Superior and Michigan		Ecological Impact	Herbivory
By 1992-1993, the biomass of Zebra Mussels in inner Saginaw Bay, Lake Huron, had a filtering capacity of 0.2-1.2 X the volume of the inner Bay per day. Chlorophyll and suspended solids were greatly reduced, and the decreased reflectivity (increased transparency) of the water was detectable by satellite imagery (Budd et al. 2001). Filtration by Zebra Mussels was selective- mussels ingested small, desirable flagellates, while rejecting large colonies of toxic Microcystis cyanobacteria in Lake Saginaw water (Vanderploeg et al. 2001).
GL-I	Lakes Huron, Superior and Michigan		Ecological Impact	Habitat Change
Filtration of the water by Zebra Mussels in inner Saginaw Bay, Lake Huron, resulted in greatly increased transparency and light penetration of the water, within 2-3 years after the initial invasion (Budd et al. 2001). The introduced amphipod Echinogammarus ischnus was strongly associated with dreissenid mussels, mostly D. polymorpha (Kang et al. 2007).
GL-II	Lake Erie		Economic Impact	Health
The city of Windsor, Ontario, spent between $CAN 400,000-450,000 on charcoal filtration of water from Lake St. Clair, to control taste and odor problems after the Zebra Mussel invasion (Colautti et al. 2006). A similar case of fouling in the intakes of the Monroe, MI public water-filtration plant reduced the supply of raw water by 20% by the summer of 1989. Several outages and water emergencies in the city of Monroe occurred. Mechanical cleaning and chlorination was required to clear the pipes and maintain water flow. Estimated costs for this episode of fouling were $US 300,000 (LePage 1993).
GL-II	Lake Erie		Ecological Impact	Trophic Cascade
Zebra Mussels have profoundly affected the food web and nutrient budget of Lake Erie. Because this lake is shallow, and is surrounded by cities and agricultural land, with high nutrient inputs, the addition of a large biomass of benthic suspension-feeders has had dramatic impacts. Dreissenid mussels remove an estimated 25% of the phytoplankton biomass per day, and excrete large quantities of nitrogen and phosphorus into the water column. The low nitrogen-to-phosporus ratio of the excreted nutrients favors the growth of nitrogen-fixing cyanobacteria, such as blooms formed by Microcystis spp. Conroy and Culver (2005) argue that the mussels slow the transfer of nutrients between trophic levels, decreasing the resilience of the system to disturbances.
GL-II	Lake Erie		Ecological Impact	Herbivory
Dreissenid mussels remove an estimated 25% of the phytoplankton biomass per day (Edwards et al., 2004, cited by Conroy and Culver 2005). In the western basin of Lake Erie, average chlorophyll a concentrations declined by 43% from 1988 to 1989, with the onset of the Zebra Mussel invasion (Leach 1993). Filtration by Zebra Mussels was selective- mussels ingested small, desirable flagellates, while rejecting large colonies of toxic Microcystis cyanobacteria in western Lake Erie water (Vanderploeg et al. 2001). Reduction in chlorophyl a and increased light penetration, since the onset of the dreissenid invasions, was also seen in the eastern basin of Lake Erie (North et al. 2012).
B-VII	None		Ecological Impact	Herbivory
In the Curonian Lagoon, Lithuania, Dreissena polymorpha is estimated to filter 10-30% of the total suspended particulate material per day, but the overall impact is considered small, because of the short residence time of the lagoon. However, within the mussel bed, the deposition of organic matter is significant, resulting in local enrichment of the benthic community (Daunys et al. 2006).
L084	_CDA_L084 (Lake St. Clair)		Ecological Impact	Habitat Change
After the invasion of Lake St. Clair, the abundance and diversity of macrobenthos increased. Water clarity increased, and macrophytes (Potamogeton sp., Vallisneria americana, and Elodea canadensis), and filamentous algae became abundant (Griffiths 1992).
L084	_CDA_L084 (Lake St. Clair)		Economic Impact	Health
The city of Windsor, Ontario, spent between $CAN 400,000–450,000 on charcoal filtration of water from Lake St. Clair, to control taste and odor problems after the Zebra Mussel invasion (Colautti et al. 2006).
GL-II	Lake Erie		Ecological Impact	Habitat Change
After the invasion of Lake St. Clair, the abundance and diversity of macrobenthos increased. Water clarity increased, and macrophytes (Potamogeton sp., Vallisneria americana, and Elodea canadensis) and filamentous algae became abundant (Griffiths 1992). In the western basin of Lake Erie, Secchi disk depth (an estimate of transparency) increased by 85% from 1988 to 1989 (Leach 1993). Although the light conditions and substrate of the lakes rocky reefs had been greatly altered, no change was seen in the spawning of Walleye (Sander vitreum), an important commercial and sport fish (Leach 1993). The introduced amphipod Echinogammarus ischnus was strongly associated with dreissenid mussels, mostly D. polymorpha (Kang et al. 2007)
GL-II	Lake Erie		Ecological Impact	Food/Prey
Diving ducks of several species (mostly Greater and Lesser Scaup, Athya marila, A. affinis) appeared in large flocks in late fall and early spring at Point Pelee, Ontario in 1991-1992. Caging experiments indicated that they sharply reduced Zebra Mussel abundance, but these effects disappeared in a few months. Ice cover prevented predation in winter (Hamilton et al. 1994). Round Gobies (Neogobius melanostomus) in the Detroit River fed largely on Zebra Mussels. The size and numbers of mussels eaten were proportional to the length of the fish (Ray and Corkum 1997).
L098	_CDA_L098 (Black-Rocky)		Ecological Impact	Food/Prey
Diving ducks of several species (mostly Greater and Lesser Scaup, Athya marila, A. affinis) appeared in large flocks in late fall and early spring at Point Pelee, Ontario in 1991-1992. Caging experiments indicated that they sharply reduced Zebra Mussel abundance, but these effects disappeared in a few months. Ice cover prevented predation in winter (Hamilton et al. 1994).
GL-III	Lake Ontario		Ecological Impact	Habitat Change
The Zebra Mussel invasion was accompanied by greatly increased transparency in Lake Ontario, along with a great increase in submerged macrophytes and filamentous algae (Limburg et al. 2010). The introduced amphipod Echinogammarus ischnus was strongly associated with dreissenid mussels, mostly D. polymorpha (Kang et al. 2007).
L085	_CDA_L085 (Detroit)		Economic Impact	Industry
Zebra Mussels caused extensive fouling of the Detroit Edison's Monroe, MI coal-fired power plant at the western end of Lake Erie. Mussels covered the intake surfaces, blocked the trash bars, and fouled the condenser tubes. The fouled parts of the plant were cleaned with high-pressure water at a cost of $25,000-35,000 for each cleaning. Sevice water lines for fire protection systems were also fouled, and cleared with chlorination, but regular use is limited by environmental concerns (Kovalak et al. 1993).
B-VIII	None		Ecological Impact	Herbivory
Rates of feeding and deposition of feces and pseudofeces in the brackish Gulf of Riga were about 1/10 of those of Zebra Mussels in freshwater lakes, so impacts are expected to be smaller (Lauringson et al. 2007). Feeding rates are affected by salinity, windspeed, and chlorophyll concentrations (Oganjan and Lauringson 2014).
L085	_CDA_L085 (Detroit)		Economic Impact	Health
Fouling by Zebra Mussels in the intakes of the Monroe MI public water-filtration plant reduced the supply of raw water by 20% by the summer of 1989. Several outages and water emergencies in the city of Monroe occurred. Mechanical cleaning and chlorination was required to clear the pipes and maintain water flow. Estimated costs for this episode of fouling were $300,000 (LePage 1993).
GL-III	Lake Ontario		Economic Impact	Aesthetic
Limburg et al. (2010) surveyed home and business owners about perceptions of water quality changes in Lake Ontario, caused by zebra mussels. There was a general positive assessment of increased water clarity, but negative perceptions of an increase in filamentous algae (Cladophora). These two changes were perceived to have opposite effects on property values, and businesses associated with recreation (Limburg et al. 2010).
L113	_CDA_L113 (Irondequoit-Ninemile)		Economic Impact	Aesthetic
Limburg et al. (2010) surveyed home and business owners about perceptions of water quality changes in Lake Ontario, caused by zebra mussels. There was a general positive assessment of increased water clarity, but negative perceptions of an increase in filamentous algae (Cladophora). These two changes were perceived to have opposite effects on property values, and businesses associated with recreation (Limburg et al. 2010)
L095	_CDA_L095 (Cedar-Portage)		Ecological Impact	Herbivory
In the western basin of Lake Erie, average chlorophyll a concentrations declined by 43% from 1988 to 1989, with the onset of the Zebra Mussel invasion (Leach 1993). Filtration by Zebra Mussels was selective- mussels ingested small, desirable flagellates, while rejecting large colonies of toxic Microcystis cyanobacteria in western Lake Erie water (Vanderploeg et al. 2001).
L095	_CDA_L095 (Cedar-Portage)		Ecological Impact	Habitat Change
Although the light conditions and substrate of the lake's rocky reefs had been greatly altered, no change was seen in the spawning of Walleye (Sander vitreum), an important commercial and sport fish (Leach 1993).
B-IX	None		Ecological Impact	Herbivory
In the inner Gulf of Finland, Zebra Mussels were reported to have a high water clearance capacity, although the effect on phytoplankton biomass was not reported (Orlova et al. 2006).
L084	_CDA_L084 (Lake St. Clair)		Ecological Impact	Food/Prey
Round Gobies (Neogobius melanostomus) in the Detroit River fed largely on Zebra Mussels. The size and numbers of mussels eaten were proportional to the length of the fish (Ray and Corkum 1997).
GL-II	Lake Erie		Ecological Impact	Competition
In the western basin of Lake Erie, Presque Isle Bay, and Lake St. Clair, fouling by Zebra Mussels was reported to cause declines of 89-100% in native Unionid mussels (Schloesser et al. 1996; Ricciardi et al. 1998).
L095	_CDA_L095 (Cedar-Portage)		Ecological Impact	Competition
In the western basin of Lake Erie, fouling by Zebra Mussels was reported to cause a complete disappearance of native unionid mussels (Schloesser 1996; Schloesser and Nalepa 1994, cited by Ricciardi et al.1998).
L103	_CDA_L103 (Chautauqua-Connaut)		Ecological Impact	Competition
In Presque Isle Bay (PA), fouling of native Unionid mussels by Zebra Mussels is reported to have caused an 89% reduction in their population (Maleski & Masteller, cited by Ricciardi et al.1998).
L084	_CDA_L084 (Lake St. Clair)		Ecological Impact	Competition
In Lake St. Clair, fouling of native unionid mussels by Zebra Mussels has caused an estimated 97% decline in abundance (Schloesser et al. 1996; Ricciardi et al.1998).
L123	_CDA_L123 (St. Lawrence River)		Ecological Impact	Competition
Fouling by Zebra Mussels is reported to have caused a >90% decline in native unionid mussels in the St. Lawrence River near Montreal (Ricciardi et al. 1998).
B-V	None		Ecological Impact	Trophic Cascade
Zebra Mussels were considered to have moderate ecosystem impacts, in the Szczecin Lagoon and Oder/Odra estuary, assumed to include impacts on other trophic levels (Zaiko et al. 2011).
M060	Hudson River/Raritan Bay		Economic Impact	Shipping/Boating
Zebra Mussels have caused significant fouling to boats and docks in the Hudson (Strayer 2006).
M060	Hudson River/Raritan Bay		Economic Impact	Industry
Zebra Mussels have caused significant fouling to power plants and water treatment plants, in the Hudson River estuary. Fouling problems have required increased inspection and cleaning, and the use of biocides, such as chlorine, potassium permanganate, or polyquaternary ammonium compounds. The cost of these treatments probably varies from $100,000 to $1 million per year (Strayer 2006).
L105	_CDA_L105 (Buffalo-Eighteenmile)		Ecological Impact	Herbivory
Reduction in chlorophyl a and increased light penetration, since the onset of the dreissenid invasions, was also seen in the eastern basin of Lake Erie (North et al. 2012).
L103	_CDA_L103 (Chautauqua-Connaut)		Ecological Impact	Herbivory
Reduction in chlorophyl a and increased light penetration, since the onset of the dreissenid invasions, was also seen in the eastern basin of Lake Erie (North et al. 2012).
GL-II	Lake Erie		Ecological Impact	Parasite/Predator Vector
Dreissena polymorpha was found to be an important host for trematode parasites, including the cosmopolitan Echinoparyphium recurvatum which can cause fatal infections in waterfowl (Karatayev et al. 2012).
L103	_CDA_L103 (Chautauqua-Connaut)		Ecological Impact	Parasite/Predator Vector
Dreissena polymorpha was found to be an important host for trematode parasites, including the cosmopolitan Echinoparyphium recurvatum which can cause fatal infections in waterfowl (Karatayev et al. 2012).
L098	_CDA_L098 (Black-Rocky)		Ecological Impact	Parasitism
Dreissena polymorpha was found to be an important host for trematode parasites, including the cosmopolitan Echinoparyphium recurvatum which can cause fatal infections in waterfowl (Karatayev et al. 2013).
B-V	None		Ecological Impact	Food/Prey
Zebra Mussels have become a major food source for most of the European wintering populaiton of a duck, Greater Scaup (Athya marila, a bird of conservation concern, in the Szczecin Lagoon (Marchowski et al. 2015)
GL-I	Lakes Huron, Superior and Michigan		Economic Impacts	Toxic
The invasion of dreissenid mussels into the Great Lakes caused major changes in the foodwebs of the lakes, which also affected the passage of toxic metals and chemical through the foodweb. Mercury inputs to Lake Michigan declined, due to pollution laws enacted in the 1970s. This was reflected in dropping mercury concentrations in the flesh of Lake Trout (Salvelinus namaycush) from 1978 to the early 1990s. The Zebra-Quagga Mussel invasion led to a drop in Secchi disk depth (increased water clarity) and a decrease in the availability of high-quality pelagic prey, and an increased reliance on benthic prey. Increased light penetration and photodegradation of methylmercury leads to mass-independent fractioning of mercury isotopes, resulting in increased ratios of lighter isotopes (Delta199 Hg) in pelagic prey. As the fish relied more on dreissenid mussels and associated benthic prey (e.g. Round Goby, Neogobius melanostomus, they consumed less pelagic prey, resulting in decreases in a nitrogen isotope (delta15N) and increasing in heavy carbon isotope (lipid-corrected delta13C). This was associated a decrease in Delta199Hg ratios, and increasing ratios of heavier mercury isotopes (Delta202Hg), even as outside inputs decreased. These results suggest that the mussel invasions offset the decrease in mercury inputs by using organic mercury stored in the sediments (Lepak et al. 2019). Increased mercury in Lake Trout results in health risks to people eating the fish.

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