Dreissena bugensis

Invasion History

First Non-native North American Tidal Record: 1993
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1993

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General Invasion History:

Dreissena bugensis was first described from the Bug River delta (Ukraine) of the Black Sea in 1897. It had a localized range in a small portion of the Black Sea drainage, but since the 1980s, has expanded its range to colonize many rivers in Eastern and Western Europe (Therriault et al. 2004; Orlova 2005; Son 2007; Zhulidov et al. 2009). In 1989, an unknown mussel, similar to the Zebra Mussel (D. polymorpha), was noticed in Lake Erie, dubbed the 'Quagga Mussel' and then identified as D. bugensis. Dreissena bugensis has colonized all of the Great Lakes, and the St. Lawrence River down to Quebec City, its only estuarine occurrence in North America (Mills et al. 1996). In 1995, D. bugensis was found in the Mississippi River between St. Louis, Missouri and Alton, Illinois (Mills et al. 1996). It now extends upstream from the Mississippi-Ohio junction to Wisconsin and Ohio. In January 2007, it was discovered in Lake Mead, Nevada, on the Colorado River, and subsequently from other reservoirs in Nevada, Arizona, and southern California. In the San Diego and Los Angeles areas, it has colonized at least 17 bodies of water. In 2008, it was discovered in Lake Granby, Colorado, near the headwaters of the Colorado River, and now is found in seven lakes and reservoirs in both the Colorado (Pacific) and Arkansas (Gulf) drainages (USGS Nonindigenous Aquatic Species Program 2012). In 2014, it was found in the water supply of Tijuana, Mexico, which comes from the El Carrizo Reservoir (Wakida-Kusunoki et al. 2015). While D. bugensis is a highly successful invader, its overall rate of spread has been slower than that of D. polymorpha, and its lag time from introduction to peak abundance has been about 5 times longer. This reflects many differences in environmental requirements and life history of the two species (Karatayev et al. 2011; Ram et al. 2012).

North American Invasion History:

Invasion History on the East Coast:

The earliest known collection of D. bugensis in North America was in Lake Erie, at Port Colborne, Ontario in 1989 (USGS Nonindigenous Aquatic Species Program 2008). After extensive genetic and morphological research, including comparisons with Russian and Ukrainian specimens, the new form was recognized as D. bugensis in 1991 (Spidle et al. 1994; Mills et al. 1996). By 1993, D. bugensis was found from Lake St. Claire, Michigan to Quebec City in the St. Lawrence River, in the New York State Barge Canal, and in the Finger Lakes (Cayuga and Seneca) (Mills et al. 1996). By 1997, the Quagga mussel was found in Lakes Huron and Michigan, though it was not collected in Lake Superior until 2005 (Grigorovich et al. 2008). Quagga Mussels have colonized deeper waters than Zebra Mussels, and have replaced Zebra Mussels in some of the deeper and cooler portions of the lakes. In Lake Ontario, D. bugensis and D. polymorpha coexist at 8-110 m, but only D. bugensis was found at 130 m. This is consistent with observations of the two species in Europe (Mills et al. 1996). Physiological differences between the two species, including a lower respiration rate, lower temperature threshold for spawning, and larger body size, may favor D. bugensis over D. polymorpha in cooler water (Stoeckmann 2003).

Since D. bugensis is less tolerant of high temperatures and salinities than the Zebra Mussel (D. polymorpha) (Mills et al. 1996; Wright et al. 1996), it may be a less successful invader in mid-Atlantic estuaries. Its only North American estuarine occurrence is in the tidal fresh St. Lawrence River, from the head of tide at Trois-Rivieres to Quebec City, where it was collected in 1993 (Mills et al. 1996). We do not have information on its abundance in the tidal fresh St. Lawrence River. In the nontidal river from Prescott, Ontario to Montreal, Quagga Mussels tended to prefer deeper water than Zebra Mussels. The St. Lawrence River is the only large river in North America where the two species extensively coexist (Jones and Ricciardi 2005). In the Soulange Canal, upstream of Montreal, Quagga Mussels had largely replaced Zebra Mussels in bottom sediments of the canal (Ricciardi and Whoriskey 2004).

Elsewhere in eastern North America, isolated populations have been found in a Pennsylvania quarry (Clover Creek Quarry, near Williamsburg in 2006) and a reservoir (Dutch Springs Reservoir, near Bethlehem in 2000) in the Susquehanna and Delaware watersheds, respectively where they may have been introduced from the Great Lakes, possibly with scuba diving equipment. It was discovered in the Mohawk River in Crescent, New York, near the head of tide of the Hudson River in 2005 (USGS Nonindigenous Aquatic Species Program 2008), but has not yet been reported from any US estuary.

Invasion History Elsewhere in the World:

Dreissena bugensis expanded from its restricted range in the Southern Bug River, Ukraine, when a series of reservoirs were constructed on the Dnieper River, which were rapidly colonized by the mussel. It spread through the adjacent Dniester River in the 1990s (Son 2007; Zhulidov et al. 2009). In the 1980s, it colonized the Don River, flowing into the Sea of Azov (Zhulidov et al. 2009). In the 1990s, it spread into much of the Volga River, from the Kuibyshev to Rybinsk reservoirs (Therriault et al. 2004; Orlova 2005; Son 2007). In 2004, D. bugensis entered the Danube River in Romania and in 2006, was collected in the Rhine River, Netherlands. Colonization probably took place through the Main-Danube Canal, by barge transport (Heiler et al. 2013). In 2006-2007, Quagga Mussels were collected in the Haringvliet, an enclosed freshwater estuary in the Rhine-Meuse delta (Schonenberg and Gittenberger 2008). River and inter-basin canal shipping has facilitated this mussel's spread through Europe (Molloy et al. 2007; bij de Vaate and Beisel 2011; Matthews et al. 2013). However, the patterns of growth and spread in the Rhine Delta indicate that ballast water release in the Delta was the likeliest source of this mussel in the Rhine (bij de Vaate 2010). By 2011, D. bugensis had spread upriver in the Rhine, and into a tributary, the Moselle, providing the first records from France (bij de Vaate and Beisel 2011). Dispersal rates in six European rivers have ranged from 23 to 383 km y-1, all in an upstream direction, and mostly aided by human vectors, such as boats (Matthews et al. 2013). In the Baltic Sea, a single record was reported in the Gulf of Finland in 2005 (Orlova et al. 2006), but its establishment here is uncertain. In 2014, established populations were found in the Szczecin Lagoon, Poland (Woznicki et al. 2016). Genetic surveys indicate that at least two invasions have occurred in Western Europe, one through south-eastern Europe via canals into the Danube, Main and Rhine Rivers, and a second by ballast water from populations introduced to the Great Lakes and released into the Meuse River system (Marescaux et al. 2015). n 2014,Dreissena bugensis was collected in the Wraysbury River, Surrey, a Thames tributary England (Aldridge et al. 2014).   Subsequently, it spread through much of England, and in 2021, was collected in the Shannon River system, Ireland (Baars et al. 2022).

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Description

Dreissena bugensis is one of a family of 'false mussels' (Dreissenidae), mostly associated with fresh and brackish (but sometimes marine) 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 bugensis is easily confused with D. polymorpha (the Zebra 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 bugensis has a pointed umbo, and is rounded ventroposterially, usually lacking the strong concave curve seen in D. polymorpha. They lack a ventrolateral ridge, and often have a more rounded and higher dorsoanterior slope, sometimes with a wing-like extension. The ventral surface of D. bugensis is convex, with a sharp keel where the shells join, so the shell topples when placed on its ventral edge, whereas D. polymorpha has a flattened ventral edge, and will usually stand up without falling over. When viewed from the edge, the two valves of D. bugensis are clearly asymmetrical, with the edges of the two shells forming a sinuous line. Color patterns are highly variable in both D. bugensis and D. polymorpha. Dreissena bugensis can be all brown, black, or white, or have various striped patterns on the exterior. A common feature is a white stripe running from the umbo along the midline of the shell. This mussel can grow to 4 cm in length, but is typically much smaller (Pathy and Mackie 1993; USGS Nonindigenous Aquatic Species Program 2012). Dreissena bugensis is limited to fresh and low-salinity brackish waters (Spidle et al. 1995; Mills 1996; Wright et al. 1996). Larval development of D. bugensis and D. polymorpha is described by Nichols and Black (1994).

Dreissenid mussels show considerable variation in color and shape, probably in response to environmental conditions. An elongated, oval, white form found in deep waters of Lakes Erie and Ontario and a Volga River reservoir was found to be a phenotype of D. bugensis. It is known as the 'profunda' morph (Spidle et al, 1994; Pavlova 2012). A recent study in the German portion of the Danube River found that morphological identifications of D. bugensis and D. polymorpha disagreed with genetic methods for about 17.5% of specimens (Beggel et al. 2014). The Quagga mussel is named after an extinct subspecies of the Plains Zebra (Equus quagga quagga) of South Africa. The name was applied in the Great Lakes, when the mussel was newly discovered, and its identity was unknown (Spidle et al. 1994).

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Taxonomy

Ecology

General:

Dreissena bugensis is a mussel with separate sexes, which release eggs and sperm into the water column, resulting in planktonic larvae. Larvae are trochophores first and then develop into shelled veligers. Annual fecundity is estimated at 960,000 embryos per year (Keller et al. 2007). D. bugensis, but the numbers of eggs released is very large. The pattern of larval development in D. bugensis is similar to that in D. polymorpha. Laboratory reared larvae of D. bugensis, reached the pediveliger stage, and began to settle at about 30 days from fertilization, compared to 23 days for D. polymorpha at 22ºC (Nichols and Black 1994; Wright et al. 1996). In Lake Erie, newly settled D. bugensis ranged from 256- 284 µm, larger than D. polymorpha (236-249 µm) in shallow water, and were even larger in the colder, deeper, hypolimnion (mean of 314 µm, SD 25) (Martel et al. 2001).

Quagga mussels settle on hard substrates such as rock, wood, and man-made structures, but also on soft bottom sediments including sand, silt and mud. They use byssal threads to attach to hard surfaces, but can settle in clumps on soft substrates, attaching to scattered shells or other hard objects, other mussels, or the sediment surface. They range from shallow waters down to over 130 m, and are often very abundant in deep water, while Zebra Mussels, can occur down to 110 m, but are usually more abundant in shallower waters. These species differences in abundance with depth are seen even in bodies of water only 6-8 m deep (Mills et al. 1996; Martel et al. 2001; Ricciardi and Whoriskey 2004; Ram et al. 2011). Quagga Mussels are less tolerant of air exposure than zebra mussels, and so are less likely to colonize intertidal areas (Ricciardi et al. 1995). However, they are more tolerant of light, and more likely to settle on partially shaded plates, while Zebra Mussels were more ommon on completely shaded plates (DHondt et al. 2018). North American populations of D. bugensis are less tolerant of high temperatures than D. polymorpha. Quagga Mussels acclimated at 20°C had >50% mortality with abrupt transfer to 35ºC, compared to 37ºC for Zebra Mussels (Spidle et al. 1995). Spidle et al. (1995) suggested that the long-term upper limit for D. bugensis might be as low as 25ºC, compared to ~30ºC for D. polymorpha. In Netherlands estuaries, Quagga Mussels had better survival than Zebra Mussels (d'Hondt et al. 2018). Studies of thermal tolerance of Ukrainian Quagga and Zebra Mussels are somewhat contradictory, with some suggesting little difference, or greater temperature tolerance in D. bugensis compared to D. polymorpha. This could be the result either of differences in methods or genetic differences between the populations (Mills et al 1996).

The two mussel species also differ in salinity tolerance, with D. polymorpha occurring in waters reaching salinities as high as 10 PSU, and completing development at 4 PSU, while in experiments, adult D. bugensis did not survive at salinities above 6 PSU, and did not complete larval development above 2 PSU (Spidle et al. 1995; Mills et al. 1996; Wright et al. 1996). Data on estuarine distribution of D. bugensis in the Ukraine, and the pattern of recent invasions in Europe generally support the idea of a lower salinity tolerance in the Quagga Mussel, compared to the Zebra Mussel (Orlova et al. 2004; Orlova et al. 2006; Son 2007; Schonenberg and Gittenberger 2008; d'Hondt et al. 2018). Calcium concentrations are another water-quality factor which may limit distribution of D. bugensis, and affect its competition and/or co-occurrence with D. polymorpha. In the nontidal St. Lawrence River, Quagga Mussels were not found below 12 mg Ca L-1, while Zebra Mussels occurred at levels as low as 7.5 mg Ca L-1 (Jones and Ricciardi 2005).

Dreissenid mussels are suspension feeders, pumping large quantities of water through their gills, retaining particles, and sloughing off surplus or inedible particles as pseudofeces. Quagga Mussels from Lake Erie had high filtering rates, even at low temperatures (~3-4ºC), much higher than those reported for Zebra Mussels at similar temperatures. At higher temperatures, ~22ºC, weight-specific filtration rates of the two species are usually similar (Vanderploeg et al. 2010).

Food:

Phytoplankton; Zooplankton

Consumers:

Fishes; Ducks

Competitors:

Dreissena polymorpha

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Nontidal Freshwater	None
General Habitat	Coarse Woody Debris	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Canals	None
Salinity Range	Limnetic	0-0.5 PSU
Tidal Range	Subtidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	0	Based on geographical distribution
Maximum Temperature (ºC)	25	D. bugensis shows rapid mortality above 30 C and appears to be more temperature sensitive than D. polymorpha (at least for North American populations) (Spidle et al. 1995; Mills et al. 1996). The upper temperature limit of this species may be as low as 25 C (Spidle et al. 1995).
Minimum Salinity (‰)	0	D. bugensis is primarily a freshwater species.
Maximum Salinity (‰)	6	Spidle et al. 1995; Mills et al. 1996
Minimum Dissolved Oxygen (mg/l)	3.2	Netherlands, Matthews et al. 2013
Minimum pH	7.1	Experimental data, Claudi et al. 2012, mixed population but predominantly quagga mussels.
Maximum pH	10.4	Netherlands, Matthews et al. 2013
Minimum Reproductive Temperature	9	Claxton and Mackie 1998, cited byUSGS Nonindigenous Aquatic Species Program 2008 http://fl.biology.usgs.gov/Nonindigenous_Species/Zebra_mussel_FAQs/Dreissena_FAQs/dreissena_faqs.html#Q15)
Maximum Reproductive Temperature	22	Laboratory, highest tested? (Wright et al. 1996)
Minimum Reproductive Salinity	0	This is a freshwater species.
Maximum Reproductive Salinity	2	Wright et al. 1996
Minimum Duration	30	Larval development, at 22 C, laboratory, (Nichols and Black 1994; Wright et al. 1996).
Minimum Length (mm)	10	Size at maturity, Ram et al. 2011
Maximum Length (mm)	38	Pathy and Mackie 1993; Mills et al. 1996; USGS Nonindigenous Aquatic Species Program 2012
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Nontidal Limnetic-Oligohaline

General Impacts

Dreissena bugensis has many similarities to Zebra Mussels (D. polymorpha) in their ecological and economic impacts in North American waters, but differences in their tolerances and life history affect the magnitude and location of their impacts. Major ecological impacts of D. bugensis include the replacement of D. polymorpha especially in cool, deep-water habitats, and the colonization of deep-water and soft-bottom habitat where Zebra Mussels are scarce (Mills et al. 1996; Ram et al. 2011). We have no information on the abundance of Quagga Mussels in the St. Lawrence River estuary (the only North American estuary colonized so far), but their lower maximum-temperature and salinity tolerances are likely to make them less competitive, at least in the shallow, less saline parts of estuaries (Mills et al. 1996; Ram et al. 2011). In addition, their low tolerance of air exposure is likely to limit their presence in the intertidal zone (Ricciardi et al. 1995).

Economic impacts

Quagga Mussels are likely to have many of the same economic impacts as Zebra Mussels, but the magnitude of their impact as fouling organisms on boats, ships, and power plants, in estuaries and shallow waters is likely to be lower because of their preference for deeper water, and their lower tolerance to high temperatures. However, as filter feeders with large biomasses in deeper waters of Lakes Michigan, Erie, and Ontario, they have dramatically affected phytoplankton biomass, nutrient cycles, and ultimately, fisheries and perceived water quality (Limburg et al. 2010; Vanderploeg et al. 2010; Evans et al. 2011; Zhang et al. 2011; Cuhel and Aguilar 2013).

Ecological Impacts

Competition- In the Great Lakes and St. Lawrence River, the most obvious impact of D. bugensis is the partial or complete replacement of D. polymorpha in deeper waters of the Great Lakes and St. Lawrence River, especially in waters more than 30 m deep, but also in rivers and canals only 2-6 m deep (Mills et al. 1996; Ricciardi and Whoriskey 2004; Jones and Ricciardi 2005; Vanderploeg et al. 2010). Quagga mussels have a lower threshold for spawning (below 12?C), spend more time in the plankton as larvae, and settle at a larger size. They are superior competitors in deep water, where temperatures are low and phytoplankton concentrations are low (Martel et al. 2001; Ram et al. 2011). When reared in Lake Erie water at surface and bottom temperatures, Quagga Mussels grew faster than Zebra Mussels in both treatments, and had higher winter survivorship (Karatayev et al. 2011). Dreissena bugensis feeding and filtration is less affected by wave action, including that due to ship travel, than D. polymorpha and several native European unionid mussels (Lorenz and Pusch 2013). In Europe, in the Main and Rhine Rivers, and the Rhine-Danube Canal, the ratio of Quagga to Zebra mussels is increasing, but the degree to which Zebra Mussels are being replaced is unclear (Matthews et al. 2013). Filter-feeding by D. bugensis may reduce the amount of phytoplankton settling in deeper parts of the lakes, contributing to the decline of the formerly dominant benthic deposit feeding amphipod Diporeia (Vanderploeg 2002; Vanderploeg et al. 2010).

Habitat Change- Quagga Mussel's ability to settle on bare, soft sediment provides new habitat for a wide variety of organisms requiring solid structures for attachment or shelter. In Lake Erie, D. bugensis comprised most of the mussel biomass on soft sediments. Invertebrate diversity and abundance was strongly correlated with mussel abundance and biomass. Among species congregating on and near mussel colonies were the amphipods Gammarus fasciatus (cryptogenic) and Echinogammarus ischnus (introduced), copepods, ostracods, flatworms, and Hydra sp. (Bially and MacIsaac 2000). The burrowing mayfly (Hexageneia), in experiments, strongly preferred to settle near mussel colonies, versus bare sediment (Devanna et al. 2011). In Lake Michigan, D. bugensis greatly altered rocky reefs surrounding soft sediments and cobble habitats. On rocky reefs, the added structure of mussels increased sedimentation, while on soft sediment it stabilized sediment and greatly decreased scouring by currents. In cobble habitats, used for spawning by Lake Trout (Salvelinus namaycush), increased light penetration led to algal growth, and made the habitat less attractive (Cuhel and Aguilar 2013).

Herbivory- In the deeper waters of the Great Lakes, the large biomass of Quagga Mussels has a potentially large impact on phytoplankton biomass. In Lake Michigan, the filtering capacity of D. bugensis at intermediate depths was sufficient to eliminate the spring phytoplankton bloom, and reduce the deposited phytoplankton available for the offshore benthic community (Vanderploeg et al. 2010). In Lake Erie, the grazing impact of deepwater dreissenids (mostly D. bugensis) was estimated to range from about equal (in the shallow western basin) to much less than that of the zooplankton population (in the deeper central basin). The impact of nutrients released into the water column by the mussels was considered to exceed that of mussel grazing (Zhang et al. 2011).

Food/Prey- Quagga and Zebra Mussels present a potential major food resource for predators in the Great Lakes. The Round Goby (Neogobius melanostomus), also of Ponto-Caspian origin, feeds heavily on dreissenid mussels. In western Lake Erie, D. bugensis was the primary food of adult Round Gobies (Campbell et al. 2009, cited by Kornis et al. 2012). The Lake Whitefish (Coregonus clupeaformis) appears to be in the process of incorporating dreissenid mussels into its diet. Whitefish in Lake Huron consume mussels at a rate 8X higher than those in Lake Michigan, which may be contributing to a higher whitefish biomass (2X) in Lake Huron (Madenjian et al. 2010).

Trophic Cascades- The large biomass of D. bugensis in portions of the Great Lakes system has the potential to remove much of the phytoplankton biomass from the water, altering the food web. In a slackwater area of the nontidal St. Lawrence River, in enclosure experiments, D. bugensis greatly reduced the abundance of a rotifer (Polyarthra sp.), probably through predation, apparently resulting in an increased abundance of the introduced copepod Eurytemora carolleeae (Thorp and Casper 2002). In Lake Michigan, the large biomass of Quagga mussels in the mid-depth regions (35-50 m) may be involved in a reduction of the formerly dominant deposit-feeding amphipod Diporeia sp., by reducing the amount of phytoplankton reaching sediments of the deeper regions of the bay. This amphipod is an important food item for many species of Great Lakes fishes (Vanderploeg 2002; Vanderploeg et al. 2010). The decrease of the amphipod led to decline in condition of the Lake Whitefish (Coregonus clupeiformis) (Cuhel and Aguilar 2013). Grazing of phytoplankton by Quagga Mussels has also led to bottom-up effects, eliminating the spring diatom bloom, and altering nutrient cycles, particularly a spring drawdown in silica. Nutrient and phytoplankton cycles in Lakes Michigan and Huron are now similar to that of oligotrophic Lake Superior (Evans et al. 2011).

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Regional Impacts

L115	_CDA_L115 (Salmon-Sandy)		Ecological Impact	Competition
Replacement of Zebra Mussels (Dreissena polymorpha) by Quagga Mussels (D. bugensis) in deeper water (>25 m) of Lake Ontario, off Oswego, NY was seen as early as 1994 (Mills et al. 1996).
L105	_CDA_L105 (Buffalo-Eighteenmile)		Ecological Impact	Competition
Replacement of Zebra Mussels (Dreissena polymorpha) by Quagga Mussels (D. bugensis) in deeper water of Lake Erie, off Dunkirk, NY was seen as early as 1994 (Mills et al. 1996).
L123	_CDA_L123 (St. Lawrence River)		Ecological Impact	Competition
In the Soulanges Canal, Quebec, connecting the Ottawa and St. Lawrence Rivers, upstream of Montreal, Quagga Mussels were observed on the canal walls in 1992. By 2002, they partially replaced Zebra Mussels, comprising 79% of the mussels on the canal walls, and almost completely replaced Zebra Mussels on the canal bottom by 2003. Zebra Mussels remained abundant only at shallow depths (1-2 m), probably because of their greater tolerance of high temperatures (Ricciardi and Whoriskey 2004). At 2 of 20 sites along the St. Lawrence River, in the Montreal area, Quagga Mussels greatly exceeded Zebra Mussels in abundance. Quaggas were positively correlated with deeper water and higher calcium concentrations (Jones abd Ricciardi 2005).
L054	_CDA_L054 (Muskegon)		Ecological Impact	Competition
By 2008, D. bugensis was the overwhelming dominant at a sampling site at 45 m depth, off Muskegon MI (Vanderploeg et al. 2010), where D. polymorpha was the sole dreissenid in 1999 (Nalepa et al. 2001).
L054	_CDA_L054 (Muskegon)		Ecological Impact	Herbivory
Grazing experiments with the 'profunda' morph of D. bugensis (from the 45m site off Musekegon MI) indicated that the filtering capacity of the mussel biomass in the 30-50 m zone exceeded the growth rate of the phytoplankton by several times, resulting in a disappearance of the spring bloom, and a nutrient sink, which decreased the resources available in deeper offshore waters (Vanderploeg et al. 2010).
L098	_CDA_L098 (Black-Rocky)		Ecological Impact	Herbivory
In the western basin of Lake Erie, dreissenid mussels (predominantly D. bugensis) were estimated to graze 4-10% of edible non-diatom phytoplankton and 7-8% of diatoms per day, about comparable to that of zooplankton (6-11% and 7-8%). However, mussel grazing was largely confined to the benthic boundary layer, and was offset by their nitrogen and phosphorus excretion (Zhang 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 bugensis, 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, off the St. Lawrence River, near Massena, NY, in mesh enclosures containing Dreissena bugensis, abundances of the copepods Eurytemora carolleeae (reported as E. affinis) increased dramatically, presumably due to the reduction of competition from rotifers (Thorp and Casper 2002).
L054	_CDA_L054 (Muskegon)		Ecological Impact	Trophic Cascade
Reduction of the phytoplankton sinking to the deeper parts of the lake bottom, as a result of Dreissena grazing, is a possible factor in the decline of the benthic amphipod Diporeia, a dominant deposit-feeder, and an important food for many great lakes fishes, including the commercially important Lake Whitefish (Coregonus clupeiformis) and Bloater (C. hoyi) (Vanderploeg et al. 2002).
L094	_CDA_L094 (Maumee River)		Ecological Impact	Habitat Change
Larvae of the mayfly Hexageneia spp., which burrow in soft sediments of the Great Lakes, in mesocosm experiments, showed a strong preference to settle near clusters of Dreissena (primarily D. bugensis) (Devanna et al. 2011).
L095	_CDA_L095 (Cedar-Portage)		Ecological Impact	Habitat Change
In transects off North Bass Island, abundance of benthic invertebrates was much greater in and around dreisssenid mussel colonies (dominated in biomass by D. bugensis, numerically by smaller D. polymorpha) compared with bare sediment. Among organisms strongly associated with mussels were amphipods (Gammarus fasciatus; Echinogammarus ischnus), copepods, ostracods, and Hydra sp. Dreissenid mussels provide shelter, structure for attachment, and also enrich the sediment with pseudofeces (Bially and MacIsaac 2000).
L045	_CDA_L045 (Milwaukee)		Ecological Impact	Competition
Quagga Mussels developed populations in deeper waters around offshore reefs in Lake Michigan, and kept filtering through the winter at low temperatures. This filtering lowered phytoplankton concentration in spring, when Zebra Mussels, which are dormant in winter, began to start feeding again. The difference in the seasonality of feeding led to the complete replacement of Zebra Mussels by Quagga Mussels on offshore reefs (Cuhel and Aguilar 2013).
L045	_CDA_L045 (Milwaukee)		Ecological Impact	Habitat Change
Heavy settlement of Quagga Mussels around offshore reefs in Lake Michigan, led to increased structure and sedimentation on rocky surfaces, and limited scouring of sediments. On level, soft silt/clay sediments, dense colonies of mussels created new, complex structure. The large filtering biomass led to increased water clarity, and increased depth of algal growth, dimnishing spawning habitat for Lake Trout (Salvelinus namaycus), which prefer to spawn in darkness, on bare cobble bottoms (Cuhel and Aguilar 2013).
GL-I	Lakes Huron, Superior and Michigan		Ecological Impact	Competition
By 2008, D. bugensis was the overwhelming dominant at a sampling site at 45 m depth, off Muskegon, MI (Vanderploeg et al. 2010), where D. polymorpha was the sole dreissenid in 1999 (Nalepa et al. 2001). Quagga Mussels developed populations in deeper waters around reefs offshore of Milwaukee, WI, in Lake Michigan, and kept filtering through the winter at low temperatures. This filtering lowered phytoplankton concentration in spring, when Zebra Mussels, which are dormant in winter, begin to start feeding again. The difference in the seasonality of feeding led to the complete replacement of Zebra Mussels by Quagga Mussels on offshore reefs (Cuhel and Aguilar 2013).
GL-I	Lakes Huron, Superior and Michigan		Ecological Impact	Herbivory
Grazing experiments with the 'profunda' morph of D. bugensis (from the 45m site off Musekegon, MI) indicated that the filtering capacity of the mussel biomass in the 30-50 m zone exceeded the growth rate of the phytoplankton by several times, resulting in a disappearance of the spring bloom, and a nutrient sink, which decreased the resources available in deeper offshore waters (Vanderploeg et al. 2010; Evans et al. 2011).
GL-I	Lakes Huron, Superior and Michigan		Ecological Impact	Trophic Cascade
Reduction of the phytoplankton sinking to the deeper parts of the lake bottom, as a result of Dreissena grazing, is a possible factor in the decline of the benthic amphipod Diporeia, a dominant deposit-feeder, and an important food for many great lakes fishes, including the commercially important Lake Whitefish (Coregonus clupeiformis) and Bloater (C. hoyi) (Vanderploeg et al. 2002; Cuhel and Aguilar 2013). Because of the year-round decrease in phytoplankton biomass, the biomass of zooplankton is expected to decrease, in turn decreasing the abundance of forage fishes (Cuhel and Aguilar 2013). Grazing by D. bugensis through the winter and early spring has eliminated the spring diatom bloom, and altered nutrient cycles, so that a spring decrease in silica (a nutrient required by diatoms) no longer occurs. The planktonic foodweb in Lakes Michigan and Huron now resemble that of the more oligotrophic Lake Superior (Evans et al. 2011). Quagga Mussels (Dreissena bugensis ) now domijnate phoisphoirus cycling in the lower Great Lakes.. 'The tissues and shells of quagga mussels now contain nearly as much phosphorus as the entire water columns of the impacted Great Lakes' (Li et al. 2021)
GL-I	Lakes Huron, Superior and Michigan		Ecological Impact	Habitat Change
Heavy settlement of Quagga Mussels around offshore reefs in Lake Michigan, led to increased structure and sedimentation on rocky surfaces, and limited scouring of sediments. On level, soft silt/clay sediments, dense colonies of mussels created new, complex structure. The large filtering biomass led to increased water clarity, and increased depth of algal growth, dimnishing spawning habitat for Lake Trout (Salvelinus namaycus), which prefer to spawn in darkness, on bare cobble bottoms. Condition index of Lake Whitefish decreasesd, but began to recover, as fish started to feed on mussels (Cuhel and Aguilar 2013).
L045	_CDA_L045 (Milwaukee)		Ecological Impact	Trophic Cascade
Heavy grazing of phytoplankton, and reduction of settlement, diminshed food for the previously abundant deposit-feeding amphipod Diporeia hoyi, a major food item of Lake Whitefish (Coregonus clupeiformis) and Cisco (Coregonus spp). Because of the year-round decrease in phytoplankton biomass, the biomass of zooplankton is expected to decrease, in turn decreasing the abundance of forage fishes (Cuhel and Aguilar 2013).
GL-II	Lake Erie		Ecological Impact	Habitat Change
In transects off North Bass Island, the abundance of benthic invertebrates was much greater in and around dreissenid mussel colonies (dominated in biomass by D. bugensis, numerically by smaller D. polymorpha) compared with bare sediment. Among organisms strongly associated with mussels were amphipods (Gammarus fasciatus; Echinogammarus ischnus), copepods, ostracods, and Hydra sp. Dreissenid mussels provide shelter, structure for attachment, and also enrich the sediment with pseudofeces (Bially and MacIsaac 2000). Larvae of the mayfly Hexageneia spp., which burrow in soft sediments of the Great Lakes, in mesocosm experiments, showed a strong preference to settle near clusters of Dreissena (primarily D. bugensis) (Devanna et al. 2011).
GL-II	Lake Erie		Ecological Impact	Herbivory
In the western basin of Lake Erie, dreissenid mussels (predominantly D. bugensis) were estimated to graze 4-10% of edible non-diatom phytoplankton and 7-8% of diatoms per day, about comparable to that of zooplankton (6-11% and 7-8%). However, mussel grazing was largely confined to the benthic boundary layer, and was offset by their nitrogen and phosphorus excretion (Zhang et al. 2011). 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	Competition
Replacement of Zebra Mussels (Dreissena polymorpha) by Quagga Mussles (D. bugensis) in deeper water of Lake Erie, off Dunkirk, NY was seen as early as 1994 (Mills et al. 1996). Replacement of Zebra Mussels by Quagga Mussels resulted in a decrease (up to 90%) of the number of dreissenids attached to native unionid mussels in Lakes Ontario, Erie, and St. Clair (Burlakova et al. 2014).
GL-III	Lake Ontario		Ecological Impact	Competition
Replacement of Zebra Mussels (Dreissena polymorpha) by Quagga Mussels (D. bugensis) in deeper water (>25 m) of Lake Ontario, off Oswego, NY was seen as early as 1994 (Mills et al. 1996). Replacement of Zebra Mussels by Quagga Mussels resulted in a decrease (up to 90%) of the number of dreissenids attached to native unionid mussels in Lakes Ontario, Erie, and St. Clair (Burlakova et al. 2014).
GL-I	Lakes Huron, Superior and Michigan		Ecological Impact	Food/Prey
The Lake Whitefish (Coregonus clupeaformis) appears to be in the process of incorporating dreissenid mussels into its diet. Whitefish in Lake Huron consume mussels at a rate 8X higher than those in Lake Michigan, which may be contributing to a higher whitefish biomass (2X) in Lake Huron (Madenjian et al. 2010).
GL-II	Lake Erie		Ecological Impact	Food/Prey
In western Lake Erie, D. bugensis was the primary food of introduced adult Round Gobies (Neogobius melanostomus) (Campbell et al. 2009, cited by Kornis et al. 2012).
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).
L103	_CDA_L103 (Chautauqua-Connaut)		Ecological Impact	Parasite/Predator Vector
Dreissena bugensis was found to be an important host for trematode parasites, inclduing the cosmopolitan Echinoparyphium recurvatum which can cause fatal infections in waterfowl (Karatayev et al. 2012).
L098	_CDA_L098 (Black-Rocky)		Ecological Impact	Parasitism
Dreissena bugensis 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).
GL-II	Lake Erie		Ecological Impact	Parasite/Predator Vector
Dreissena bugensis was found to be an important host for trematode parasites, inclduing the cosmopolitan Echinoparyphium recurvatum which can cause fatal infections in waterfowl (Karatayev et al. 2012).
GL-I	Lakes Huron, Superior and Michigan		Economic Impact	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 (Delta 199 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 (delta 15N) and increasing in heavy carbon isotope (lipid-corrected delta 13C). This was associated a decrease in Delta 199Hg 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.
GL-III	Lake Ontario		Ecological Impact	Trophic Cascade
Quagga Mussels (Dreissena bugensis ) now domijnate phoisphoirus cycling in the lower Great Lakes.. 'The tissues and shells of quagga mussels now contain nearly as much phosphorus as the entire water columns of the impacted Great Lakes' (Li et al. 2021)
GL-II	Lake Erie		Ecological Impact	Competition
Quagga Mussels (Dreissena bugensis ) now domijnate phoisphoirus cycling in the lower Great Lakes.. 'The tissues and shells of quagga mussels now contain nearly as much phosphorus as the entire water columns of the impacted Great Lakes' (Li et al. 2021).

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