Description
Potentially Misidentified Species - Mytilopsis leucophaeta (Dark False Mussel) is regarded as probably native in Chesapeake Bay, though introduced in the Hudson River, upper Mississippi and Europe (Pathy and Mackie 1992). Dreissena bugensis (Quagga Mussel) is, like D. polymorpha, a Eurasian species introduced to the Great Lakes and connected waters. Its environmental requirements may make it a less likely invader in Chesapeake Bay (Mills et al. 1996a; Pathy and Mackie 1992; Wright et al. 1996).
Planktonic larvae of Corbicula fluminea (Asiatic Clam) may be mistaken for D. polymorpha , but have a foot and siphon at all stages of development, and so are technically not veligers. Veliger larvae of D. polymorpha and D. bugensis cannot always be separated (Nichols and Black 1994).
Taxonomy
| Kingdom | Phylum | Class | Order | Family | Genus |
|---|---|---|---|---|---|
| Animalia | Mollusca | Bivalvia | Veneroida | Dreissenidae | Dreissena |
Synonyms
Invasion History
Chesapeake Bay Status
| First Record | Population | Range | Introduction | Residency | Source Region | Native Region | Vectors |
|---|---|---|---|---|---|---|---|
| 2008 | Unknown | Expanding | Introduced | Regular Resident | North America | Eurasia | Fisheries - accidental (not oysters) |
History of Spread
Dreissena polymorpha (Zebra Mussel) was described 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. Early in the 19th century, D. polymorpha invaded canals connecting with 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 to France, and in the lagoons bordering the Black Sea (Strayer and Smith 1993).
North American records are summarized below:
Great Lakes - St. Lawrence River - Dreissena polymorpha was first collected in Lake St. Clair MI (between Lakes Huron and Erie) in 1988, but was thought to have been introduced 2-3 years earlier, most likely by ship's ballast water (Ludyanskiy et al. 1993; Mackie and Schloesser 1996). By 1990, it was found in all five Great Lakes, covering a range of ~600 km north-south and 1400 km east-west, with populations being most continuous downstream of Lake St. Clair. In 1992, D. polymorpha was collected in the intertidal zone of the St. Lawrence River at Quebec City (freshwater) (Mellina and Rasmussen 1994).
Mississippi River Drainage and western North America - In 1991, D. polymorpha was collected outside the Great-Lakes-St. Lawrence system for the first time, colonizing the Ilinois 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 MS (Vicksburg MS) by 1992 (Ludyanskiy et al. 1993), New Orleans LA by 1993, and the mouth of the river by 1994. In 1997, its range in the Mississippi valley extended from PA and TN west to MN and OK. By 2013, these mussels were found in the upper Colorado and Arkansas drainages in Colorado (Grand Lake and Pueblo Reservoir), several reservoirs in Texas, a lake in Utah, and a reservoir in the Monterey Bay watershed, California (USGS Nonindigenous Aquatic Species Program 2013). 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).
East Coast
Hudson River Drainage - D. polymorpha colonized the New York State Barge (Erie) Canal, reaching Rochester NY by 1990 (Lange and Cap 1991), and the Hudson River estuary at Catskill NY, by 1991 (Mills et al. 1997). In 1992, the mussel was found throughout the tidal Hudson, and approached its limits of salinity tolerance, occurring at 2-6 ppt in Haverstraw Bay (Walton 1996).
Susquehanna River Drainage - 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. Lange (cited by Kraft 1995) suggested that a clump of mussels might have been transferred, and spawned, but without successful recruitment because of the rapid dispersal of larvae. This apparently failed invasion is anomalous because of the normal rapid pace and high success rate of D. polymorpha invasions.
In 2001, an established population of D. polymorpha was found in Eaton Brook Reservoir, on the Chenango River, in Madison County NY, in the northern reaches of the Susquehanna Watershed. Based on the size of the mussels, the invasion probably began in 1999. Some adult mussels have been found downstream of the reservoir (Blankenship 2001). By 2002, the mussels had been found in a second NY lake, Candaroga Lake, in Otsego County, and by 2004, they had reached the upper mainstem of the Susquehanna in Colliersville NY (Otsego County)(USGS Nonindigenous Aquatic Species Program 2013). In May 2007, they were recorded for the first time in the Susquehanna basin in PA, in Cowanesque Lake, Tioga County, near the NY border, well to the west of the mainstem of the Susquehanna (USGS Nonindigenous Aquatic Species Program 2013). Transport by boats or fishing gear is likely.
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 Program 2013). In November 2008, D. polymorpha was also found further upstream in Pennsylvania, in Muddy Run, a tributary near the Maryland border. In December 2008, a clump of mussels was found attached to a boat at Glen Cove Marina, just above Conowingo Dam (Thomson 2008). In May 2009, 8 mussels were found in Conowingo Pond above the dam (Halsey 2009). This mussel appears to be established in the lower Susquehanna and is likely to colonize tidal fresh regions of upper Chesapeake Bay. In October, 2011, one dead mussel was found attached to a jet-ski mooring in the Sassafras River near Betterton MD (USGS Nonindigenous Aquatic Species Program 2013). A recent report (Klauda and Ashton 2013) lists additional collections in the lower Susequhanna River and Susquehanna Flats, at the head of the Bay. 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 likely to colonize tidal fresh regions of upper Chesapeake Bay (Klauda and Ashton 2013). In 2014, more than 500 Zebra Mussels were collected from concrete blocks in the Susquehanna River, and upper Chesapeake Bay, near Havre de Grace (Ashton et al. 2014; Wheeler 2014). 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).
In 2002, a population of D. polymorpha was found in a quarry in Prince William County VA, near Broad Run, a Potomac tributary. The mussels were believed to have been stocked illegally by divers to clear the water (Center for Aquatic Resource Studies 2004). Budget difficulties have delayed attempts at eradication, planned by the Virginia state government, as of September (Harper 2004). Budget difficulties have delayed attempts at eradication, planned by the Virginia state government, as of September (Harper 2004). The mussels were successfully eradicated in 2006 (Virginia Department of Game and Inland Fisheries 2008 http://www.dgif.virginia.gov/zebramussels/)
Dreissena polymorpha are established in the Finger Lakes (Seneca, Cayuga and others), just to the north of the Susquehanna watershed, but there is no direct connection between these lakes and the Chesapeake drainage. Introductions of D. polymorpha probably took place on trailered boats, boat's live wells, or in bait buckets (Lange and Cap 1991; Blankenship 2001; Kelly et al. 2012).
Dreissena polymorpha is regarded as an inevitable invader in the Chesapeake Bay drainage. Fresh and low-salinity waters of most of the major tributaries of the Chesapeake appear to be suitable as habitat for this species, though some waters have low probability of colonization due to low calcium content or low pH (Christmas et al. 1994; Baker et al. 1994). (Baker et al. 1994; Christmas et al. 1994).
Invasion Comments
Vector(s) of Introduction - Trailered boats from the nearby Finger Lakes region were thought to be the likeliest source of the Dreissena polymorpha (Zebra Mussel) veligers reported from the Susquehanna River in 1991-1993. No direct connection exists between the Finger Lakes, where D. polymorpha is well-established, and the Susquehanna (Lange and Cap 1991).
Population Status - Adults and veligers of Dreissena polymorpha (Zebra Mussel) has been collected from several sites in the Lower Susquehanna River and uppermost Chesapeake Bay (Susquehanna Flats) from 2008 through 2013. As of December 2013, a population appears to be established, although the density remains very low (Klauda and Ashton 2013).
Ecology
Environmental Tolerances
| For Survival | For Reproduction | |||
|---|---|---|---|---|
| Minimum | Maximum | Minimum | Maximum | |
| Temperature (ºC) | 0.0 | 34.0 | 12.0 | 27.0 |
| Salinity (‰) | 0.0 | 10.0 | 0.0 | 3.5 |
| Oxygen | well-oxygenated | |||
| pH | 7.0000000000 | 9.0000000000 | ||
| Salinity Range | fresh-oligo |
Age and Growth
| Male | Female | |
|---|---|---|
| Minimum Adult Size (mm) | 3.0 | 3.0 |
| Typical Adult Size (mm) | 30.0 | 30.0 |
| Maximum Adult Size (mm) | 50.0 | 50.0 |
| Maximum Longevity (yrs) | 9.0 | 9.0 |
| Typical Longevity (yrs | 2.0 | 2.0 |
Reproduction
| Start | Peak | End | |
|---|---|---|---|
| Reproductive Season | |||
| Typical Number of Young Per Reproductive Event |
|||
| Sexuality Mode(s) | |||
| Mode(s) of Asexual Reproduction |
|||
| Fertilization Type(s) | |||
| More than One Reproduction Event per Year |
|||
| Reproductive Startegy | |||
| Egg/Seed Form |
Impacts
Economic Impacts in Chesapeake Bay
The arrival of (Dreissena polymorpha (Zebra Mussel) has been long anticipated in Chesapeake Bay (Baker 1994; Christmas et al. 1994). Dreissena polymorpha (Zebra Mussel) is now established in the Susquehanna River from its headwaters in New York states to uppermost Chesapeake Bay, although densities in the lower river and Bay are too low to cause detectable impacts (Klauda and Ashton 2013). Some research has been done in the Chesapeake Bay region, in an attempt to predict the future distribution and abundance of D. polymorpha (Christmas et al. 1994; Wright et al. 1996). In 1992, boat angling on the city of Baltimore’s reservoirs was temporarily banned, while D. polymorpha control devices were selected and installed, costing local economies hundreds of thousands of dollars (Chesapeake Bay Commission 1995). Monitoring for possible occurrence of these bivalves is continuing at power plants (Krueger 1997, pers. comm.) and reservoir systems. Restrictions on the use of trailered boats on some Virginia reservoirs have prompted public protests (Cairns 1996). In 2002, an infestation of mussels was found in Milbrook Quarry, Prince William County VA, near Broad Run, a Chesapeake Bay tributary. The mussels were introduced by scuba divers, in order to clear the water. Funding problems delayed control measures, but in 2006, the mussels were eradicated using 174,000 gallons of potassium chloride solution. The eradication, plus monitoring cost ~400,000 (Virginia Department of Game and Inland Fisheries 2006). Control measures of this kind are possible only in relatively small, confined, bodies of water.
In the Chesapeake Bay region, although the D. polymorpha has not yet arrived, the possibility of its invasion has already resulted in significant economic impacts. Based on experiences in Europe, the Great Lakes, and the Hudson Rivers, substantial impacts can be expected in tidal and nontidal fresh waters, and in lower-salinity parts of the Bay (Baker et al. 1994;Christmas 1994). These impacts are reviewed in 'Economic Impacts beyond Chesapeake Bay' below.
Baker et al. 1994; Cairns 1996; Chesapeake Bay Commission 1995; Christmas et al. 1994; Kraft 1995; Kraft 1995; Krueger 1997, pers. comm; Wright et al. 1996
Economic Impacts Outside of Chesapeake Bay
Invasions of Dreissena polymorpha (Zebra mussel) have had dramatic economic impacts in Europe and North America.
Industry - Dreissena polymorpha has caused widespread fouling of powerplants in Europe and the Great Lakes region (Ludyanskiy et al. 1993; MacIsaac 1996). In the Great Lakes, D. polymorpha were first observed in the 10 powerplants of the Detroit Edison system in 1988. Extensive fouling first occurred in the Monroe MI plant in the summer of 1989. Mussels blocked slats in the trash bars, restricting flow, and were frequently impinged on the travelling screens. Masses of dislodged mussels and large individual mussels blocked up to 35% of condenser tubes, reducing efficiency of the plant. Mechanical cleaning of the trash bars took 8-10 days and cost $35,000. Cleaning of the screenhouses, to reduce blocking of the condensers, cost ~$600,000 in 1989-1991. [Four screenhouses required 6 cleanings, at $25,000-35,000 each.] Chlorination was used to control buildup of mussels in the fire protection system. While the Monroe plant was most heavily impacted, similar problems occurred throughout the Detroit Edison system (Kovalak et al. 1992). Fouling of intakes varies 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 may discourage settlement (MacIsaac 1996).
Health - Fouling of water intake pipes has interfered with municipal water supplies in the Great Lakes region. In Monroe MI, on Lake Erie, Dreissena polymorpha were first discovered in the waterworks in January 1989. By July, 1989, the mussels were seriously interfering with pumping operations. From 1989 to 1991, there were 3 water outages, ranging from 9 to 56 hours, and causing varying degrees of disruption of activities in the city. Control measures included release of chlorine, cleaning by divers, and redesign and replacement of the intake pipes. Overall costs due to D.polymorpha were ~$310,000 in 1989-1991, with $100,000 to 3 million dollars in projected costs (Lepage 1992).
Fisheries - Dreissena polymorpha directly affects commercial and sport fisheries operations by fouling fishing boats, fixed gear (fish traps), fishing nets, etc. (MacIsaac 1996a). The D. polymorpha invasion has been expected to have widespread effects on Great Lakes fisheries because of its drastic alteration of foodwebs, shifting much of the phytoplankton production to the benthos, and potentially decreasing abundance of zooplankton, the primary food of larval fishes. Reduced recruitment of Perca flavescens (Yellow Perch) in the Great Lakes has been attributed to reduced zooplankton abundance, although competition with introduced Morone americana (White Perch) may be an important factor (MacIsaac 1996a). Studies of spawning reefs used by Stizostedion vitreum (Walleye) and Coregonus spp. (Whitefishes) suggest that the abundance of D. polymorpha did not affect use of the reefs, water quality, or reproductive success of S. vitreum (Fitzsimmons et al. 1994; Leach 1999). D. polymorpha may have positive impacts on some benthic feeding fishes, either by serving directly as a food, or by enriching sediments and increasing the overall diversity of benthic invertebrates (Karatayev et al. 1997). Dreissena polymorpha have become an important food resource for wintering diving ducks in Europe and the Great Lakes, resulting in alteration of migration patterns (Hamilton et al. 1994), and potential benefits to local duck hunters.
Habitat Change - The invasion of bodies of water by Dreissena polymorpha has had a wide range of effects on habitats. Large filtering biomasses of D. polymorpha can result in drastic increases in water clarity (MacIsaac 1996a), although these changes may be small in rivers and estuaries, such as the Hudson or the Detroit River (or Chesapeake Bay), with high densities of suspended inorganic particles (MacIsaac 199a6; Roditi et al. 1995). Increased water clarity in the Great Lakes has favored increased growth of submerged macrophytes, enhancing habitat quality for some fishes [Esox lucius (Northern Pike); E. masquinongy (Muskellunge); Micropterus spp. ('Black Basses')]. On the other hand, increased light levels may have driven Stizistedion vitreum (Walleye) into more turbid waters (MacIsaac 1996a).
Aesthetic - Dreissenia sp. invasions typically result in increased water clarity, due to filtration by large biomasses of mussels (Karatayev et al. 1996; MacIsaac 1996). People generally perceive this as an improvement in water quality. However, increased light penetration can result in development of large biomasses of submerged vegetation, resulting in seasonal die-offs and strandings of unpleasant quantities of dying plants (MacIsaac 1996a). Extensive strandings of washed-up mussels (MacIsaac 1996b) also result in unpleasant odors, and the sharp-edged shells can interere with beach recreation (Ludyanskiy et al. 1993).
Boating - Dreissenia polymorpha has caused extensive fouling of boats, ships, barges, piers, navigational structures, and buoys in the Great Lakes (Ludyanskiy et al. 1993).
References - Fitzsimmons et al. 1994; Hamilton et al. 1994; Karatayev et al. 1997; Leach 1999; Lepage 1992; Ludyanskiy et al. 1993; MacIsaac 1996a; MacIsaac 1996b; Roditi et al. 1995
Ecological Impacts on Chesapeake Native Species
Dreissena polymorpha (Zebra Mussel) has no established populations in Chesapeake Bay tidal waters, but populations are now established in the upper watershed (Susquehanna drainage) in New York state, and downstream spread is likely. Chesapeake populations would be concentrated in tidal fresh-oligohaline regions of rivers and the uppermost Bay. Based on experiences in the Great Lakes and elsewhere, impacts of this species in Chesapeake Bay may include the following (a selective summary):
Competition - Dreissena polymorpha (Zebra Mussel) has competed with native North American fauna in a variety of ways, including competition with other fouling taxa for settling sites, by fouling native molluscs, and competition for food with other filter-feeding taxa. In the Hudson River, ranges of the dreissenid Mytilopsis leucophaeta (probably native in the Chesapeake, introduced in the Hudson) overlap, but sympatric settlement was not seen (Walton 1996). Competition for space between these related species has been suggested to occur in the Netherlands (Jenner and Jansen-Mommen 1993), but has not been experimentally investigated. Competition with native species of freshwater bryozoans and other freshwater fouling organisms is possible, but we are not aware of such studies.
Physical fouling of animals can be considered an extreme form of competition for space. Fouling of unionid clams, with adverse affects on their growth and survival, has been observed in North America and Europe (Karatayev et al. 1997). Increasing D. polymorpha populations in the Great Lakes, Mississippi River, and Hudson River have been associated with dramatic population declines of unionids. D. polymorpha has been found on 37 species of North American unionids, with as many as 10,500 mussels on a unionid in Lake St. Clair (Schloesser et al. 1996). Causes of mortality may include obstruction of valves, reduction of ambient food concentrations, and interference with movement, including burrowing (Mackie 1993; Schloesser et al. 1996). In the Hudson River estuary, the arrival of D. polymorpha was associated with declines of 90-36% (mean of 56%) for 4 species of unionids, with ~30% of clams infested with mussels, and a mean of ~4 D. polymorpha per clam. However, in one portion of the estuary, unionids had ~30 mussels per clam. In general, Hudson infestations were lighter than those seen in other locations, so that competition for phytoplankton may have played more of a role (Strayer and Smith 1996). In the Mississippi River, unionids and two species of snails, heavily colonized by D. polymorpha, were found to be prone to being washed ashore and stranded by waves. However, the significance of this mortality to the populations is not known (Tucker 1994a; Tucker 1994b).
As a filter-feeder which has developed extraordinary biomasses in short periods of time, D. polymorpha is a severe competitor with other planktivores, including unionid clams, zooplankton, suspension-feeding fouling organisms such as bryozoans and hydroids, and planktivorous fishes. Invasions of D. polymorpha are frequently accompanied by sharp decreases in phytoplankton biomass (see below). Such reductions in phytoplankton can be expected to affect zooplankton through reductions in food supply. In the Hudson River estuary, however egg production of the cladoceran Bosmina freyi does not appear to have been significantly affected by the D. polymorpha invasion and reductions in zooplankton abundance and biomass appear to have resulted from direct predation (Pace et al. 1998). Similarly, sharp declines in rotifers and copepod nauplii abundance in western Lake Erie probably reflect both competition for food and direct predation on these small organisms. By directly suppressing zooplankton, D. polymorpha can also be regarded as a competitor of larval fishes. Diminished recruitment of Perca flavescens and Stizostedion vitreum (native in the Great Lakes, introduced in Chesapeake Bay) has been attributed to decreased larval food supplies. However, other mussel-related habitat changes, as well as unrelated changes in other fish populations may have influenced the decline of these fish species (MacIsaac1996a).
Herbivory - As a suspension feeding bivalve, the primary food of Dreissena polymorpha (Zebra Mussel) is phytoplankton. In areas which have been extensively colonized, D. polymorpha has reduced chlorophyll concentrations by 23% (Lake Erie, west-central basin, Leach 1993) to 81-85% (Hudson River, Roditi et al. 1996; Ukrainian lakes, Karatayev et al. 1997). Biomasses of D. polymorpha in European lakes vary greatly in their filtering ability, but some populations can filter the entire water column of a lake in several days (Karatayev et al. 1997). The biomass in the Hudson River estuary is estimated to filter the water column in ~2 days (Roditi et al. 1996). Reports vary on the size and taxonomic selectivity of D. polymorpha's phytoplankton feeding, in some cases, blooms of cyanobacteria have been reported after the establishment of the mussels, suggesting that some phytoplankton may be avoided due to size, mechanical difficulties (e.g. spines, colony shape, etc.), or toxicity (MacIsaac 1996a).
Predation - In addition to phytoplankton, Dreissena polymorpha (Zebra Mussel) directly ingests smaller zooplankton (rotifers, nauplii, tintinnids) which lack the ability to escape the intake current. These may be digested or ejected, entangled, in the pseudofeces (MacIsaac 1996a). Decreased abundance of rotifers and nauplii in the Hudson River estuary has been attributed to predation by D. polymorpha, but older copepods and the cladoceran Bosmina freyi were not affected by D. polymorpha predation (Pace et al. 1998).
Food/Prey - In areas where it has invaded, Dreissena polymorpha (Zebra Mussel), predation has not prevented the buildup of massive populations. However, a variety of animals are known to feed on this mussel, and it has become a significant food source for some. Enclosure experiments in the Hudson River estuary showed that Callinectes sapidus (Blue Crabs) at high densities (over 0.1 m-2) were capable of reducing D. polymorpha populations. In the Hudson, C. sapidus was rarely abundant enough in the field to regulate mussel populations, but such densities are common in low-salinity tributaries of Chesapeake Bay (Boles and Lipscius 1997). Fishes native to the Chesapeake region which are known or expected to feed on D. polymorpha include: Lepomis auritus (Redbreast Sunfish), Lepomis gibbosus (Pumpkinseed, MacIsaac 1996a) and Nocomis raneyi (Bull Chub), Cloe et al. 1995), and possibly Moxostoma macrolepidotum (Shorthead Redhorse), since congeners feed on D. polymorpha in the Great Lakes (MacIsaac 1996a). In the Great Lakes, D. polymorpha populations have been heavily exploited by diving ducks, including Aythya marila (Greater Scaup), A. affinis (Lesser Scaup), Bucepha clangula (Common Goldeneye) and B. albeola (Bufflehead), which consumed up to 57% of standing biomass in winter. Impacts in Lake Erie were limited by ice-cover (Hamilton et al. 1994), so diving duck predation could be greater in Chesapeake tributaries. A major consequence of D. polymorpha invasions seems to be the shifting of foodwebs in favor of higher trophic levels and larger predators, including benthivorous fishes and birds (Karatayev et al. 1997; Ludyanskiy et al. 1993; MacIsaac 1996a).
Habitat Change - Dreissena polymorpha's huge populations in many invaded areas have had dramatic effects on water clarity, sediment composition, and biogeochemical cycling, as well as providing a new habitat structure for many small benthic organisms (Karatayev et al. 1997; Ludyanskiy et al. 1993; MacIsaac 1996a). In lakes, dramatic increases in water transparency have been observed, e.g., from 0.5-1.5 m Secchi Disk depth before 1988 to 1.8-2.8 m in 1991 in Lake St. Clair MI-Ontario. Such increases in light penetration have led to the resurgence of macrophytes in parts of the Great Lakes (MacIsaac 1996a) and in European lakes (Karatayev et al. 1997). In the Hudson River estuary, while chlorophyll concentrations have decreased sharply (81-85%), effects on water transparency and total seston (suspended particles) have been small (12% and 15% respectively), owing to the heavy load of suspended non-phytoplankton particles (Roditi et al. 1996). D. polymorpha populations rapidly move particles from the water column to the sediments, as pseudofeces, increasing the organic matter content of sediments, and so tending to increase the biomass and diversity of the detritivore community (Griffiths 1993; Karatayev et al. 1997).
Potential habitat changes can be expected to be even more widespread, and many such as changes in thermal structure due to increased water clarity, effect of increased light penetration on predator-prey relations, changes in mineral and nutrient cycling, etc., remain to be investigated. Qualitative observations suggest that increased macrophyte growth may have favored some species of predatory fishes in the Great Lakes [Esox lucius (Northern Pike), E. masquinongy (Muskellunge), Micropterus dolomieu (Smallmouth Bass) , M. salmoides (Largemouth Bass)], while increased light levels may have adversely affected Stizostedion vitreum (Walleye) (MacIsaac 1996a). All these species, native to the Great Lakes, are introduced in Chesapeake Bay, but similar divergent responses to habitat change can be expected in native Chesapeake Bay species.
References - Boles and Lipscius 1997; Cloe et al. 1995; Griffiths 1993; Hamilton et al. 1994;; Karatayev et al. 1997; Leach 1993; Ludyanskiy et al. 1993; MacIsaac 1996a; Mackie 1992; Pace et al. 1998; Roditi et al. 1996; Schloesser et al. 1996; Strayer and Smith 1996; Tucker 1994a; Tucker 1994b; Walton 1996
Ecological Impacts on Other Chesapeake Non-Native Species
Potential effects of a Dreissena polymorpha (Zebra Mussel) invasion on introduced biota in the Chesapeake Bay region may include the following:
Competition - Dreissena polymorpha (Zebra Mussel) may compete with the hydroid Cordylophora caspia for settling sites, since both species co-occur closely in the Hudson River estuary (Walton 1996). Competition between these introduced fouling forms has been suggested to occur in the Netherlands (Walton 1996), but has not been experimentally investigated. Competition, particularly for phytoplankton, seems likely to occur between D. polymorpha and Corbicula fluminea (Asiatic Clam). The latter is now a dominant bivalve in many tidal fresh Chesapeake tributaries. Interactions between these species have not yet been investigated, to our knowledge, at least in North America. Some displacement of D. polymorpha by its congener D. bugensis (Quagga Mussel), has occurred in the Great Lakes and the Ukraine, suggesting that these two species may be competitors. D. bugensis has replaced D. polymorpha in deeper, colder parts of the lower Great Lakes, but may be increasingly moving into shallow water (Mills et al. 1996a).
Food/Prey - A number of species introduced to the Chesapeake Bay region are known predators of D. polymorpha. Orconectes virilis (Virile Crayfish) feeds on D. polymorpha in aquaria, but preferred rainbow trout eggs, which require less handling time, when the later were available (Love and Savino 1993). Cyprinus carpio (Common Carp) feeds on D. polymorpha at high frequencies [a mean of 112 beaks (range 1-107) per fish stomach] in the Mississippi River, but its impact on mussel abundances is not clear (Tucker et al. 1996). The introduced Lepomis microphus (Redear Sunfish; Shellcracker) is well-known as a molluscivore, but preferred a snail (Helisoma anceps, Rams-Horn Snail), in choice experiments in aquaria (French and Morgan 1995).
Habitat Change - Increases in light penetration due to D. polymorpha's clearing of the water column have led to the resurgence of macrophytes in parts of the Great Lakes (MacIsaac 1996a) and in European lakes (Karatayev et al. 1997). This could benefit introduced macrophytes, such as Hydrilla verticillata (Hydrilla) and Myriophyllum spicatum (Eurasian Watermilfoil), as well as native plants. Phelps (1994) suggested that the filter-feeding of large biomasses of Corbicula fluminea (Asiatic Clam) favored the invasion of the Potomac River by H. verticllata. Invasions of D. polymorpha could also benefit introduced macrophytes, although high concentrations of suspended particles may limit changes in light regime (Carter et al. 1994a; Roditi et al. 1995). Qualitative observations suggest that increased macrophyte growth may have favored some species of predatory fishes (native in the Great Lakes, introduced in Chesapeake Bay) in the Great Lakes [Esox lucius (Northern Pike), E. masquinongy (Muskellunge), Micropterus dolomieu (Smallmouth Bass) , M. salmoides (Largemouth Bass)], while increased light levels may have adversely affected Stizostedion vitreum (Walleye) (MacIsaac 1996a).
References Carter et al. 1994a; French and Morgan 1995; ; Karatayev et al. 1997; Love and Savino 1993; MacIsaac 1996a; Mills et al. 1996a; Phelps 1994; Tucker et al. 1996; Walton 1996
References
Baker, Patrick; Baker, Shirley; Mann, Roger (1994) Potential range of the Zebra Mussel, Dreissena polymorpha, in and near Virginia., In: (Eds.) Zebra Mussels and the Mid-Atlantic: Reports from the Sea Grant Programs of New Jersey, Delaware,Maryland, Virginia, and North Carolina.. , College Park. Pp. 5-18Blankenship, Karl (2001) Zebra mussels gain toehold in northern fringe of watershed, Bay Journal 11:
Boles, Larry C.; Lipcius, Romuald N. (1997) Potential for population regulation of the zebra mussel by finfish and the blue crab in North American estuaries, Journal of Shellfish Research 16: 179-186
Cairns, John, Jr.; Bidwell, Joseph R. (1996) Discontinuities in technological and natural systems caused by exotic species, Biodiversity and Conservation 5: 1085-1094
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