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    Taxonomic name: Dreissena bugensis Andrusov, 1897
    Common names: quagga mussel
    Organism type: mollusc
    Dreissena bugensis is native to parts of Ukraine. This small freshwater mussel is an active filter feeder, which competes for food resources with filter-feeding zooplankton by accelerating sedimentation of suspended matter, including organic substances. It is also a nuisance and economic problem when it grows on recreational or commercial ships/boats, potable water treatment plants and electric power stations.
    Dreissena bugensis commonly has alternating light and dark brown stripes, but can also be solid light brown or dark brown. It has two smooth shells that are shaped like the letter “D”. These mussels are usually less than 2 inches in length. In new populations, most mussels are young and therefore very small (under ¼ -inch long) (California Department of Fish and Game 2008).

    There are two phenotypes of D. bugensis that have been reported in the Great Lakes: the "epilimnetic" form, which has a high flat shell, and the "profunda" form, which has an elongate modioliform shell and has invaded soft sediments in the hypolimnion. The epilimnetic form uses its byssal threads to attach to objects and particles and form druses or colonies. The profunda morph can form colonies and attach to objects with its byssal threads or it can partially bury itself in soft sediments and extend its very long incurrent siphon above itself to bring in suspended food particles (Vanderploeg et al. 2002).

    Similar Species
    Corbicula fluminea, Dreissena polymorpha, Dreissena rostriformis

    Occurs in:
    estuarine habitats, lakes, water courses
    Habitat description
    Adult D. bugensis attach to natural hard substrata including rocks, wood, and macrophytic plants and to man-made materials including concrete, metal piping, steel, nylon, fiberglass and wood. Mussels attach to substrates via proteinaceous byssal threads produced from a gland posterior to the foot. D. bugensis typically occur in fresh water but thrive in salinities up to 1‰ and can reproduce in salinities below 3‰. Salinities exceeding 6‰ cause mortality (Ussery & McMahon 1995; Wright et al. 1996).
    General impacts
    Nutrient loading and species introductions are thought to be two of the major environmental problems currently facing freshwater ecosystems (Richter et al. 1997, Hall et al. 2003 in Haynes et al. 2005), and both of these anthropogenic factors are of concern in the Great Lakes, USA (Haynes et al. 2005).

    Reduction in Native Biodiversity: D. bugensis causes changes in the structural characteristics of zooplankton including total abundance, biomass and species composition. Specifically, there is an inverse relationship between zooplankton abundance/biomass and density of Dreissena mussels (Grigorovich & Shevtsova, 1995). Dreissena infestations have caused upwards of 95% reduction in unionid numbers and extirpated eight species of unionids in some areas of the Great Lakes (Schloesser et al. 1998; Schloesser & Masteller 1999). Individuals attach themselves to the shells of other mussels, forming encrusting mats many shells thick (10-30mm).

    Modification of Natural Benthic Communities: Dreissena negatively affects benthic invertebrate communities, especially filter-feeding or deep-dwelling invertebrates that rely on detrital rain (Dermott and Munawar 1993, Strayer et al. 1998, Johannsson et al. 2000, in Haynes et al. 2005). Predicting benthic invertebrate community response to a change in nutrient levels is very difficult, and the potential synergistic effects of nutrient alterations and exotics such as Dreissena are complex (Haynes et al. 2005).

    Economic: Thick encrustations of mussels form on man-made structures or within raw water systems, impacting on operation and efficiency. D. bugensis can have major detrimental impacts on recreational and commercial shipping/boating as well as on water-using industries, potable water treatment plants and electric power stations (Ussery & McMahon, 1995).

    Because they are long-lived and sessile, quagga mussels can be used as bioindicators of hazardous substances such as radionuclides (Lubianov 1972, in Orlova 2009).
    In both North America and its original range in Europe, D. bugensis is replacing zebra mussel (D. polymorpha) populations (Domske & Oneill 2003; Diggins et al. 2004). Some industries build intake structures at depths too low for D. polymorpha to grow in; however, D. bugensis is able to colonise surfaces at greater depths, rendering these new structures vulnerable to mussel colonisation (Mills et al., 1999; and Richerson and Maynard, 2004).
    Geographical range
    Native range: Dreissena bugensis originally had a very restricted distribution: in just the Dnieper Bug estuary and lower Inguletz river in Ukraine where it was discovered in 1890 (Andrusov 1890 in Orlova 2009).

    Known Introduced range: From the 1940s the quagga mussel began to expand its range throughout Eastern Europe into the Black Sea, the Dnieper River, the lower reaches of the Pripiat (Pripyat) River, the Dniester, the Main and Rhine Rivers, and to the Don and Manych Rivers. It invaded the Volgo-Caspian basin 40 years after the opening of the Volga-Don canal in 1952, which links the two rivers. It was first recorded in the Volga River in 1992 and in the Caspian Sea delta and avan-delta of the Volga River in 1994. In the Volgo-Caspian area it has also expanded its range northward to the Sheksna reservoir. D. bugensis is now commonly found in the large inland lakes and rivers of European Russsia, Ukraine, Moldova and Belarus (Orlova 2009 and references therein).

    The quagga mussel has also been introduced to Great Lakes region of North America where it is considered an invasive species. It is also present in the Mississippi and Ohio Rivers and is expanding west of the 100th Meridian with established populations in Nevada, Arizona and Utah.

    Introduction pathways to new locations
    Ship ballast water: Its release into Great Lakes waters is linked to discharge of ship ballast water (Mills et al., 1999).
    Translocation of machinery/equipment: A study conducted by Ricciardi and colleagues (1995) revealed that under temperate summer conditions adult D. bugensis may survive on overland transport (e.g. small trailer-boats) for up to 5 days. Veligers can be transported in fish and bait wells as well as in cooling ports of inboard and outboard motors. Most or all the introductions of quagga mussels beyond the 100th Meridian in North America are purported to be via trailered boats (Mackie & Claudi 2009).
    Management information
    Compared to the zebra mussel (Dreissena polymorpha) there has been little research carried out on the biology, ecological requirements and tolerances of quagga mussels (Dreissena bugensis (Mackie & Claudi, 2009). Indeed most research on the control of mussels has focused on D. polymorpha (McEnnulty et al., 2001). However it is thought that most of the control methods would also apply to quagga mussels (G.L. Mackie, pers. comm.; Virginia Department of Game and Inland Fisheries, 2005).

    Prevention: Studies suggest that humans are responsible for most introductions of zebra and quagga mussels into new areas. The best way to prevent and manage dreissenid invasions in open waters is thought to be prevention through public outreach and education. Examples of this include public signage and wash stations at boat launches and other potential introduction points (Frischer et al. 2005).

    Detection: One of the most important criterions for successful eradication of a species is early detection allowing control measures to take place while the incursion is still relatively small. Detection relies on monitoring and education. In Lake George, NY zebra mussels were detected in 1999 while the population was relatively small. Control efforts between 1999 and 2007, mainly using physical means and SCUBA, were successful in eradicating zebra mussels from the lake (Wimbush et al. 2009).

    Chemical Control: Chemical control is one of the most common methods for control or eradication. Chlorination is often used; D. bugensis is more sensitive to chlorination than D. polymorpha. Thus chlorination programs currently in use to combat D. polymorpha are more than sufficient to simultaneously control D. bugensis. Another alternative has been potassium permanganate, especially for drinking water sources, even though chemical controls are not environmentally sound solutions. D. polymorpha was recently eradicated from Millbrook Quarry, Virginia using 174,000 gallons of potassium chloride solution over a 3 week period in 2006 (Virginia Department of Game and Inland Fisheries, 2005). Other chemical control options include chlorine dioxide, sodium hypochloride, ozone, mollusicides and polymers (D’Itri, 1996).

    Physical: Decreasing water levels of water bodies to cause desiccation of D. bugensis is an effective, readily applied and environmentally neutral technique. It would be most effective in raw water systems such as navigation locks and water intake structures, which are designed to be periodically dewatered for maintenance. This is a particularly attractive method of control because it could be utilized to mitigate fouling not just by D. bugensis but also mixed populations of this species and D. polymorpha (Brady et al., 1996; Ussery & McMahon, 1995). Other physical methods include manual scraping, high-pressure jetting, antifouling coatings and mechanical filtration.

    Biological Control: Research is currently underway to test the effectiveness of the CL145A strain of the bacteria Pseudomonas fluorescens which produces a toxin that destroys the digestive system of Dreissena spp. (Molloy & Mayer 2007).

    Other: A variety of other control methods in use or being developed are oxygen deprivation, thermal treatment, radiation, molluscicides, ozone, antifouling coatings, electric currents, and sonic vibration (D’Itri, 1996; Mackie & Claudi, 2009). Fears and Mackie (1995) investigated the use of low-voltage currents for preventing settlement and attachment by D. bugensis by using steel rods and plates with the current running through them placed near the intake of a pulp and paper plant. Complete prevention of settlement was achieved at 8 volts/in with steel rods on both wood and concrete surfaces (Fears & Mackie, 1995).

    D. bugensis are filter feeders which use cilia to pull water into their shell cavity from where it passes through an incurrent siphon. Desirable particulate matter is removed in the siphon. Each adult mussel is capable of filtering one or more liters of water each day, removing phytoplankton, zooplankton, algae and even their own veligers (larvae) (Snyder et al. 1997). Any undesirable particulate matter is bound with mucus, known as pseudofeces, and ejected out the incurrent siphon. The particle-free water is then discharged out the excurrent siphon (Richerson 2002, D’Itri 1996, Nalepa & Schloesser 1993).
    D. bugensis is a prolific breeder. It is dioecious and exhibits external fertilisation. A fully mature female mussel is capable of producing up to one million eggs per season (Richerson 2002; D’Itri 1996).
    Lifecycle stages
    After fertilisation veligers (pelagic microscopic larvae) develop within a few days and soon acquire minute bivalve shells. Free-swimming veligers drift with the currents for three to four weeks, feeding using their hair-like cilia while trying to locate suitable substrata to settle and secure byssal threads. Mortality in this transitional stage from planktonic veliger to settled juveniles may exceed 99% (Stanczykowska 1977, in Bially & MacIsaac 2000). Macrophytes, mussel colonies and pebbles were found to be more suitable substrates for settling than gravel, sand or mud (Lewandowski 1982, in Bially & MacIsaac 2000
    Reviewed by: Gerald L. Mackie, Professor, Department of Zoology, University of Guelph, Guelph, Ontario Canada
    Compiled by: Profile revision: National Biological Information Infrastructure (NBII) & IUCN/SSC Invasive Species Specialist Group (ISSG)
    Last Modified: Wednesday, 27 October 2010

ISSG Landcare Research NBII IUCN University of Auckland