Invasion History

First Non-native North American Tidal Record: 1937
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1937

General Invasion History:

Mytilopsis leucophaeata was first described by Conrad (1829), as Mytilopsis leucophaeatus, from the 'southern coast of the U.S.', and later reported as 'inhabiting the rivers of Virginia, and probably further south' (Conrad 1857). It is native from Chesapeake Bay south to Veracruz, Mexico, and is strongly associated with brackish environments, often in oligohaline water (0.5-5 PSU) (Abbott 1974; García-Cubas et al. 1992; Kennedy 2011a). Museum records from Caribbean islands (Jamaica, Cuba, Haiti, Guadeloupe, mostly as Congeria leucophaeta, Academy of Natural Sciences of Philadelphia 2012; Museum of Comparative Zoology 2012; U.S. National Museum of Natural History 2012) may represent misidentifications of Mytilopsis sallei.

Mytilopsis leucophaeata was collected in Antwerp, Belgium in 1835, and given the name Mytilus cochleatus, where it was suspected as an introduction, because of its association with barnacles and boring organisms (Nyst 1835, cited by Kennedy 2011a). In Europe, M. leucophaeata has occurred in ports and estuaries in Belgium, France, Spain, the Netherlands, Wales, Germany, Poland, Sweden, Finland, and the Black and Caspian Seas (Nikolaev 1951; Oliver et al. 1998; Escot et al. 2003; Therriault et al. 2004; Laine et al. 2007; Heiler et al. 2010; Kennedy 2011a). On the East Coast of the US, M. leucophaeata has been introduced to several estuaries north of its native range, including the Hudson River, the Housatonic River (New York), and the Charles River (Massachusetts) (Jacobson 1953). Two specimens were found in the upper Mississippi River in Illinois in the 1980s (Koch 1989, cited by Kennedy 2011a), but we know of no further records from this river system. This bivalve has recently invaded South America, being found in Recife, Brazil in 2004 (Souza et al. 2005).

Although M. leucophaeata has a preference for brackish water, it tolerates salinities of 30 PSU and above, as well as freshwater (Castagna and Chanley 1973; Siddall 1982), and so could be transported by hull fouling through seawater, or freshwater canals. Its planktonic larvae can also tolerate salinities as high as 32 PSU (Siddall 1982), facilitating transfer with ballast water (Verween et al. 2010). It also appears to tolerate a wide range of climates, having recently invaded both Finland and Brazil (Souza et al. 2005; Laine et al. 2007). In its native range, M. leucophaeata is associated with Eastern Oysters (Crassostrea virginica), so oyster transfers are a possible vector, especially for its northward invasions on the East Coast (Jacobson 1953).

North American Invasion History:

Invasion History on the East Coast:

The native range of M. leucophaeata appears to reach its northern limit in the upper Chesapeake Bay, at least as far north as the Elk River, Maryland (Bergstrom 2005; Kennedy 2011b; Florida Museum of Natural History 2012). We have no records of this bivalve occurring in the Delaware River estuary. In 1937, two specimens were collected in the tidal Hudson River, near Haverstraw, New York (NY) (Rehder 1937). In 1952, an established population of M. leucophaeata was found near Haverstraw, and as far downriver as Englewood, New Jersey (Jacobson 1953). In 1992, M. leucophaeata was found from Tarrytown, NY (49 River Km, 5-9 PSU) to Newburgh, NY (99 River Km, 0-3 PSU) (Walton 1996). It was also found to be common in the tidal Housatonic River, upstream from Long Island Sound, at Shelton, Connecticut at 0.5-2 PSU (Smith and Boss 1995). In 1995, M. leucophaeata was found in the impounded section of the Charles River in Cambridge, Massachusetts, which receives brackish water through locks, at salinities of ~1 PSU (Smith and Boss 1995). This population appears to be established, with museum specimens collected in 2008 (Museum of Comparative Zoology 2012). In the Housatonic and Charles Rivers, fouling on recreational boats appears to be the likeliest vector.

Invasion History Elsewhere in the World:

As noted above, Mytilopsis leucophaeata was first collected in Antwerp, Belgium in 1835 (Nyst 1835, cited by Kennedy 2011a), and subsequently appeared in various ports and estuaries throughout Europe, often in a very spotty fashion in space and time, often in confined brackish waters, sometimes near thermal effluents, and often prone to massive settlement and die-offs (Kerckhof et al. 2007; Kennedy 2011a; Verween et al. 2010). Early invasions in northern Europe included the Amstel River, Amsterdam, Netherlands in 1895 (Wolff 2005); the Canal de Caen, Normandy in 1910 (Germain 1931, cited by Oliver et al. 1998), the Weser River and the Kiel Canal, Germany both in 1928 (Nehring 2002), and the Zuider Zee and Rhine Delta (Van Jutting 1936, 1943, cited by Wolff 2005). Beginning in the second half of the 20th century, it began to colonize wider areas of the European coast. In 1962, it was found in the isolated Russian Baltic port of Kaliningrad (surrounded by Poland), but it is likely extinct there now (Brohmer 1962, cited by Laine et al. 2007). However, in the Baltic Sea, it is established in the Warnow River, Rostock, Germany (Darr and Zettler 2000, cited by Laine et al. 2007), the Gulf of Gdansk (in 2010, Dziubinska 2011), the Gulf of Finland (in 2003, Laine et al. 2007), and the Gulf of Bothnia, Sweden (in 2011, Werner in ICES Advisory Committee on the Marine Environment 2012). The Finnish and Swedish populations are in areas affected by thermal effluents. However, in 2011-2015, established populations were found in Finnish areas unaffected by thermal plumes (Forsstrom et al. 2016). In the British Isles, M. leucophaeata was first found on an enclosed dock in Cardiff, Wales in 1996 (Oliver et al. 1998), and later found in the Thames estuary in 1999 (Bamber and Taylor 2002, cited by Heiler et al. 2010).

The recent spread of M. leucophaeta to scattered localities in southern Europe has been remarkable, although it is possible that this small bivalve has been overlooked in the past, and is still undiscovered at many intervening locations. In 1993, it was found near Seville, on the Guadalquivir River estuary, on the southern Atlantic coast of Spain (Escot et al. 2003), and in 2002, in a canal at d’Aigues-Mortes on the Mediterranean coast of France (Girardi 2003, cited by Heiler et al. 2010). In 2002, it was identified by molecular means in the Dniester lagoon on the Black Sea in Ukraine (Therriault et al. 2004). In 2009, it was found to be abundant in the southern Caspian Sea, near Bandar Anzali, Iran (Heiler et al. 2010). Possible invasion routes are from the Black Sea, by the Volga-Don Canal, or from the Baltic Sea by the Neva River, Lake Ladoga and Lake Onega, and the Baltic-Volga canal system, to Volga and the Caspian Sea (Heiler et al. 2010).


Mytilopsis leucophaeata is a small mussel-like bivalve which belongs to a family (Dreissenidae), found in fresh, brackish, and marine waters. Dreissenids are distinguished from true mussels (Mytilidae) by having a small shelf-like platform (septum) 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. Mytilopsis leucophaeata is easily confused with Dreissena polymorpha (Zebra Mussel) and D. bugensis (Quagga Mussel), both native to the Ponto-Caspian region (Abbott 1974; Pathy and Mackie 1993). Mytilopsis leucophaeata is recognized by a prominent, downward projecting tooth at the dorsal corner of the platform, inside the left valve's beak. The shell of M. leucophaeata is roughly elliptical in the posterior region, but has a curved beak, ending in a rounded knob, in contrast to the more pointed umbo and more wedge like appearance of D. polymorpha and D. bugensis. The right valve is larger than the left. It lacks the strongly arched appearance and ventrolateral shoulder seen in D. polymorpha. The ventral pallial line is slightly curved inward, forming a very shallow pallial sinus, which is absent in the Zebra and Quagga mussels. The hinge plate is low and wide, with a ligament and a nymph (a bar, or platform) extending to a point about 30-40% of the distance from the beak to the posterior edge of the shell. Maximum reported sizes range from 18 to 25 mm. The shell is marked with concentric lines. The color of the shell is cream-colored to brown, sometimes bluish-brown. The shell is sometimes marked with zigzag stripes, resembling those of D. polymorpha (Zebra Mussel), especially in juveniles. The shell interior is white to grayish, and somewhat pearly. Larval development of M. leucophaeata is described by Siddall (1982) and by Verween et al. (2010). (Description from: Abbott 1974; Siddall 1982; Marelli and Gray 1985; Pathy and Mackie 1993; Oliver et al. 1998; Kennedy 2011a)

Nomenclature of Mytilopsis spp. has been confused, because of the diversity of tropical species, and of fossil species known from Europe, and frequent invasions, including a very early one in Europe. Mytilopsis leucophaeata was described from Belgium in 1835, under the name Mytilus cochleatus, later known as M. cochleata, and frequently treated as a separate species, either native, or introduced from Africa (Oliver et al. 1998; Marelli and Gray 1985a; Marelli and Gray 1985b; Kennedy 2011a). Marelli and Gray (1985a; 1985b) concluded that M. cochleata was a junior synonym of M. leucophaeata.  Recent genetic and morphological studdies sugges that M. leucophaeta may consist of two species, s northern population (e.g. Chesapeake Bay, introduced to New Yotk and Europe) and southern populaitons (e.g. Flrida, introduced to Venezueala and Brazil) (Fernandes et al. 2022).


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Mollusca
Class:   Bivalvia
Subclass:   Heterodonta
Order:   Veneroida
Superfamily:   Dreissenoidea
Family:   Dreissenidae
Genus:   Mytilopsis
Species:   leucophaeata


Congeria cochleata (Dall, 1898)
Congeria leucophaeta (Dall, 1898)
Dreissena cochleata (Nyst, 1843)
Dreissena cumingiana (Dunker, 1853)
Mytilina cochleata (Cantraine, 1837)
Mytilus americanus (Reeve, 1858)
Mytilus cochleatus (Kickx, in Nyst, 1835)
Mytilus leucophaetua (Conrad, 1829)
Mytilus tenebrosus (Reeve, 1858)
Tichogonia americana (Kuster, 1899)
Tichogonia cochleata (Dunker, 1853)

Potentially Misidentified Species

Dreissena bugensis
The Quagga Mussel, native to Black Sea drainages has invaded the Great Lakes-St. Lawrence basin, and European waterways.

Dreissena polymorpha
The Zebra Mussel, with a broad native range in the Ponto-Caspian basin, has invaded many North American River systems, and some estuaries, including the St. Lawrence River, the Hudson River, and Chesapeake Bay. In the Hudson and Chesapeake, it co-occurs with M. leucophaeata.

Mytilopsis adamsi
This tropical East Pacific species has invaded Pacific estuaries in Mexico, Australia, and East Asia.

Mytilopsis sallei
This Caribbean species overlaps with M. leucophaeata in Florida and the West Indies (Marelli and Gray 1985).



Mytilopsis leucophaeata is a small mussel which has separate sexes that release eggs and sperm into the water column, resulting in planktonic larvae. The first stage of larval development is a trochophore, followed by a shelled veliger. Spawning in Belgium began at 16-19ºC, and was prolonged, with individuals spawning several times from spring to fall, peaking in summer (Verween et al. 2010). The trochophore stage was reached within 8-24 h, and by 21- 48 h larvae had a D-shaped shell (Verween et al. 2010; Kennedy 2011b). Laboratory-reared larvae at 21-26°C began to settle at about 6 to 15 days from fertilization, and 194-210 μm in length (Siddall et al. 1982; Kennedy 2011b). Successful reproduction has been reported at 0.8 to 32 PSU (Siddall et al. 1982; Kennedy 2011b; Verveen et al. 2010). Field and laboratory observations suggest that spawning and settlement can be stimulated by sudden decreases in salinity, such as occurs following heavy rains (Kennedy 2011a; Kennedy 2011b). Mussels in the Netherlands grew at temperatures above 9.1 C, and had a maximum lifetime at 4.5 years.

Postlarvae of M. leucophaeata often settle on a filamentous surface, such as vegetation, after which larvae move to hard surfaces such as logs, stones, shells, and artificial structures (Verveen et al. 2010; Kennedy 2011b). Growth occurs mostly during the warmer months. Data from Belgium show the most growth in July-August. The age and size at reproductive maturity is not known, but Belgian populations grew from 8 to 10 mm during their second year, a possible time of first spawning. Growth patterns suggested longevity of about 5 years, reaching a mean size of about 16 mm (Verween et al. 2006; Verween et al. 2010).

Mytilopsis leucophaeata is a suspension feeder, capable of ingesting particles as small as 4 μm. In the laboratory, they have been maintained on the flagellate Isochrysis galbana (Verween et al. 2010). They are vulnerable to predation by crabs and fishes, which may account for their frequent scarcity and the 'boom and bust' nature of their occurrences in Chesapeake Bay (Kennedy 2011b).




Crabs, fishes


barnacles, Dreissena spp.

Trophic Status:

Suspension Feeder



General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatMangrovesNone
General HabitatCanalsNone
General HabitatVessel HullNone
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone

Life History

Tolerances and Life History Parameters

Minimum Temperature (ºC)0Based on geographical range
Maximum Temperature (ºC)35Experiment, Rajagopal et al. 2005
Minimum Salinity (‰)0Experimental, Castagna and Chanley (1973)
Maximum Salinity (‰)32Experimental, Castagna and Chanley (1973); (Siddall 1982), but a more normal range is 0-18, with greatest abundances at 1-10 PSU (Kennedy 2011a). In simulated voyages, using animals from the Netherlands, M. leucophaeta, had a high tolerance at 0.2-17.5 PSU (van der Gaag et al. 2016), with 655-781 days for 100% mortality.
Minimum Reproductive Temperature10Experimental, 4 hr old embryos- Verween et 2007, Belgium
Maximum Reproductive Temperature30Field, spatfall, July, Miami FL(Siddall 1982), but 24, Experimental, 4 hr old embryos- Verween et 2007, Belgium
Minimum Reproductive Salinity5Experimental, 4 hr old embryos- Verween et 2007, Belgium
Maximum Reproductive Salinity20Experimental, 4 hr old embryros- Verween et 2007, Belgium
Minimum Duration6Larval duration, experimental, 26 C (Siddall 1982)
Maximum Duration8Larval duration, experimental, 26 C (Siddall 1982)
Minimum Length (mm)10Vorstman 1933, (Germany) cited by Kennedy 2011a
Maximum Length (mm)25Abbott 1974; Pathy and Mackie 1993; Oliver et al. 1998; Kennedy 2011a
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNoneOligohaline-Euhaline

General Impacts

Mytilopsis leucophaeata is often sporadic in its occurrence in the Chesapeake Bay, and has not been well-studied elsewhere in its native range (Kennedy et al. 2010). During its short periods of abundance, it has been associated with improved water quality in low-salinity tributaries of the Bay (Bergstrom 2005). However, significant ecological impacts have not been observed in invaded waters, either in the Northeast US (Hudson, Housatonic, and Charles Rivers), or in invaded European waters. However, it has been a significant industrial fouling organism in Belgium, the Netherlands, and Spain (Escot et al. 2003; Rajagopal et al. 2005; Verween et al. 2010).

Economic Impacts

Mytilopsis leucophaeata has caused significant problems in industrial water systems in Belgium, the Netherlands, and Spain (Escot et al. 2003; Rajagopal et al. 2005; Verween et al. 2010). These bivalves have higher temperature tolerances than Blue Mussels (Mytilus edulis) or Zebra Mussels (Dreissena polymorpha), probably due to their tropical origin, and are more likely to survive in cooling systems (Rajagopal et al. 2005). They are also more tolerant to chlorination than other fouling organisms. Problems due to Mytilopsis fouling were noticed when environmental regulations limited the amount of chlorine that could be used in biofouling control (Rajagopal et al. 2005; Verween et al. 2010). The fouling observed in the water-pumping system in the Guadalquivir River, Seville, Spain, occurred in water used for industry and agriculture, where biocides cannot be used (Escot et al. 2003).

Regional Impacts

NEA-IINoneEconomic ImpactIndustry
Mytilopsis leucophaeata is known as a major power plant fouling organism in the Netherlands and Belgium. It is more tolerant of high temperatures, chlorination, and salinity than the Zebra Mussel (Dreissena polymoirpha), which makes it more difficult to eradicate (Rajagopal et al. 2005; Verween et al. 2010).
NEA-VNoneEconomic ImpactIndustry
Fouling by M. leucophaeata, together with Corbicula fluminea (Asian Freshwater Clam) and Cordylophora caspia (Freshwater Hydroid) interfered with a system for pumping industrial and agricultural water from the Guadalquivir River in the city of Seville (Escot et al. 2005).
CAR-IIINoneEconomic ImpactFisheries
High abundances of Mytilopsis leucophaeta were associated with up to 30% reduction in shrimp production in shrimp ponds next to Lake Maracaibo, Venezuela (Lodeiros et al. 2019). Mechanisms of interfierence with shrimp production were not discussed.
SA-IINoneEcological ImpactHerbivory

Compared to a native bivalve, Mytilopsis leucophaeta could increase its ingestion rate at high phytoplankton concentrations (Rodrigues et al. 2023)

SA-IINoneEcological ImpactCompetition

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 0 Native Estab
CAR-VII Cape Hatteras to Mid-East Florida 0 Native Estab
NA-ET3 Cape Cod to Cape Hatteras 1937 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 1995 Def Estab
NEA-II None 1835 Def Estab
B-III None 1928 Def Estab
B-IV None 2000 Def Estab
MED-IX None 2003 Def Estab
NEA-V None 1993 Def Estab
B-IX None 2003 Def Estab
M040 Long Island Sound 1992 Def Estab
M060 Hudson River/Raritan Bay 1937 Def Estab
M130 Chesapeake Bay 1831 Native Estab
G070 Tampa Bay 0 Native Estab
G130 Pensacola Bay 0 Native Estab
G170 West Mississippi Sound 0 Native Estab
G210 Terrebonne/Timbalier Bays 0 Native Estab
G250 Sabine Lake 0 Native Estab
G260 Galveston Bay 0 Native Estab
N170 Massachusetts Bay 1995 Def Estab
S050 Cape Fear River 0 Native Estab
S120 Savannah River 0 Native Estab
S180 St. Johns River 0 Native Estab
S190 Indian River 0 Native Estab
S200 Biscayne Bay 0 Native Estab
B-VII None 1962 Def Estab
B-XI None 2011 Def Unk
MED-II None 2002 Def Estab
CASP Caspian Sea 1996 Def Estab
S010 Albemarle Sound 0 Native Estab
S030 Bogue Sound 0 Native Estab
SA-II None 2014 Def Estab
B-X None 2011 Def Estab
MED-X None 2004 Def Unk
NEA-III None 1996 Def Estab
CAR-III None 2017 Def Estab
CASP Caspian Sea 2017 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Abbott, R. Tucker (1974) American Seashells, Van Nostrand Reinhold, New York. Pp. <missing location>

Academy of Natural Sciences of Philadelphia 2002-2024a Malacology Collection Search. <missing URL>

Associated Press (12/2021) Lummi Nation declares disaster after invasive crab arrives, Seattle Times <missing volume>: <missing location>

Baldwin, Andy; Leason, Diane (2016) Potential Ecological impacts of Emerald Ash Borer on Maryland's Eastern Shore, In: None(Eds.) None. , <missing place>. Pp. <missing location>

Bergstrom, Peter 2005 An introduction... to Dark False Mussels. <missing URL>

Bilger, Michael D.; Riva-Murray, Karen;Wall, Gretchen L. (2005) <missing title>, U. S. Geological Survey, Reston, VA. Pp. <missing location>

Buschbaum, Christian; Lackschewitz, Dagmar; Reise, Karsten (2012) Nonnative macrobenthos in the Wadden Sea ecosystem, Journal of Ocean Management 68: 89-101

Castagna, M.; Chanley, P. (1973) Salinity tolerance of some marine bivalves from inshore and estuarine environments in Virginia waters on the western mid-Atlantic coast., Malacologia 12(1): 47-96

Conrad, T. A. (1857) Description of a new genus of the family Dreissenidae, Proceedings of the Academy of Natural Sciences of Philadelphia 9: 167

Conrad, Timothy A. (1829) Description of fifteen new species of recent, and three of fossil shells, chiefly from the coast of the United States, Journal of the Academy of Natural Sciences of Philadelphia 6: 256-268

Crandall, Marc. E. (1977) Epibenthic Invertebrates of Croton Bay in the Hudson River, New York Fish and Game Journal 24(2): 178-196

Crumb, Stephen E. (1977) Macrobenthos of the tidal Delaware River between Trenton and Burlington, New Jersey, Chesapeake Science 18(3): 253-265

da Rocha, Clarisse Duarte; Fernandes, Maurício Romulo; Miyahira, Igor Christo (2023) One more place to call home: the invasive bivalve Mytilopsis leucophaeata reaches the Maricá-Guarapina lagoon system (Rio de Janeiro, Brazil), Biota Neotropica 23(3): e20231503

da Rocha, Rosana Moreira; Antunes, Antunes Gamba, Gustavo; Zanaya. Thais Bastos (2012) Aplidium (Ascidiacea, Polyclinidae) in central coast, Brazil, Zootaxa 3565: 18-30

da Silva, Eder Carvalho; Barros, Francisco (2011) [Benthic macrofauna introduced in Brazil: List of marine and freshwater species and actual distribution], Oecologia Australis 15(2): 326-344

Dean, Thomas A.; Bellis, Vincent J. (1975) Seasonal and spatial distribution of epifauna in the Pamlico River Estuary, North Carolina, Journal of the Elisha Mitchell Scientific Society 91(1): 1-12

Dziubinska, Anna (2011) Mytilopsis leucophaeata, an alien dreissenid bivalve discovered in the Gulf of Gdansk (southern Baltic Sea), Oceanologia 53(2): 651-655

Eno, N. Clare; Clark, Robin A.; Sanderson, William G. (1997) <missing title>, Joint Nature Conservation Committee, Peterborough. Pp. <missing location>

Eno, N.C., Clark, R.A., Sanderson, W.G. 1997-2012 Directory of Non-Native Marine Species in British waters. <missing URL>

Escot, C.; Basanta, A.; Cobo, F.; González, M. A. (2003) Sobre la presencia de Mytilopsis leucophaeta (Conrad, 1831) (Bivalvia, Dreissenacea, Dreissenidae) en el río Guadalquivir (sur de la Península Ibérica)., Graelsia 59(1): 91-94

Farrapeira, Cristiane Maria Rocha; de Melo,Arthur Vinícius de Oliveira Marrocos; Barbosa, Débora Ferreira; da Silva, Karla Maria Euzebio (2007) Ship hull fouling in the port of Recife, Pernambuco, Brazilian Journal of Oceanography 55(3): 207-221

Farrapeira, Cristiane Maria Rocha; Tenório, Deusinete de Oliveira ; do Amaral, Fernanda Duar (2011) Vessel biofouling as an inadvertent vector of benthic invertebrates occurring in Brazil, Marine Pollution Bulletin 62: 832-839

Feldheim, Kevin A.; Brown, Joshua E.; Murphy, Douglas J.; Stepien, Carol A. (2011) Microsatellite loci for dreissenid mussels (Mollusca: Bivalvia: Dreissenidae) and relatives: markers for assessing exotic and native populations, Molecular Ecology Resources 11: 725-732

Florida Museum of Natural History 2009-2013 Invertebrate Zoology Master Database. <missing URL>

Florin, Ann-Britt and 5 authors (2013) First records of Conrad’s false mussel, Mytilopsis leucophaeata in the southern Bothnian Sea, Sweden, near a nuclear power plant, Bioinvasion Records 2: in press

Forsström, Tiia; Fowler, Amy E.; Lindqvist, Meri; Vesakoski, Outi (2016) The introduced dark false mussel, Mytilopsis leucophaeata (Conrad, 1831) has spread in the northern Baltic Sea, BioInvasions Records 5: In press

Forster, S.; Zettler, M. L. (2004) The capacity of the filter-feeding bivalve Mya arenaria L. to affect water transport in sandy beds, Marine Biology 144: 1183-1189

García-Cubas, Antonio; Reguero, Martha; Elizarrarás, Rafael (1992) [Mollusks from Chica-Grande lagoon system, Veracruz, Mexico: systematics and ecology], Anales Del Instituto de Ciencias Del Mar y Limnologia 19(1): 71-101

Harvard Museum of Comparative Zoology 2008-2021 Museum of Comparative Zoology Collections database- Malacology Collection. <missing URL>

Heiler, Katharina C. M.; Nahavandi, Nahid; Albrecht, Christian (2010) A new invasion into an ancient lake - the invasion history of the dreissenid mussel Mytilopsis leucophaeata (Conrad, 1831) and its first record in the Caspian Sea, Malacologia 53: 185-192

Holmes, Samuel; Callaway, Ruth (2021) Fouling communities and non-native species within five ports along the Bristol Channel, South Wales, UK, Estuarine Coastal and Shelf Science 252(1107295): Publiished online

Holopainen, Reetta; Lehtiniemi, Maiju; Meier, H. E. Markus; Albertsson, Jan; Gorokhova, Elena; Kotta, Jonne; Viitasalo, Markku (2016) Impacts of changing climate on the non-indigenous invertebrates in the northern Baltic Sea by end of the twenty-first century, Biological Invasions Published online: <missing location>

ICES Advisory Committee on the Marine Environment (2012) <missing title>, International Council for the Exploration of the Seas, Copenhagen. Pp. <missing location>

Invasive Species Specialist Group 2001-2016 100 Of The World's Worst Invasive Species. <missing URL>

Jacobson, Morris K. (1953) Congeria leucophaeata (Conrad) in the Hudson River, Nautilus 66(2): 125-127

Japoshvili, George; Higashiura, Yoshimitsu; Kamitani, Satoshi (2016) A review of Japanese Encyrtidae (Hymenoptera), with descriptions of new species, new records and comments on the types described by Japanese authors, Axta Entomological Musei Nationalis Pragae 36: 345-401

Kennedy, Victor S. (2011a) The invasive dark falsemussel Mytilopsis leucophaeata (Bivalvia: Dreissenidae): a literature review, Aquatic Ecology 45: 163-183

Kennedy, Victor S. (2011b) Biology of the uncommon dreissenid bivalve Mytilopsis leucophaeata (Conrad, 1831) in central Chesapeake Bay, Journal of Molluscan Studies 77: 154-164

Kerckhof, Francis; Haelters, Jan; Gollasch, Stephan G. (2007) Alien species in the marine and brackish ecosystem: the situation in Belgian waters., Aquatic Invasions 2(3): 243-257

Khamassi, F.,; Rjiba Bahri, W.; Mnari Bhouri, A., Chaffai, A.; Soufi Kechaou, E.; Ghanem, R.,; Ben Souissi, J. (2022) Biochemical composition, nutritional value and socio-economic impacts of the invasive crab Callinectes sapidus Rathbun, 1896 in central Mediterranean Sea. , Mediterranean Marine Science 23(3): 650–663

Laine, Ari O.; Mattila, Jukka; Lehikoinen, Annukka (2007) First record of the brackish water dreissenid bivalve Mytilopsis leucophaeata in the northern Baltic Sea., Aquatic Invasions 1: 38-41

Larsen, Peter F. (1985) The benthic fauna associated with the oyster reefs of the James River estuary, Virginia, U. S. A., Internationale Revue der Gesamten Hydrobiologie 70(9): 707-814

Leidenberger, Sonja; Obst, Matthias; Kulawik, Robert; Stelzer, Kerstin; Heyer, Karin; Hardisty, Alex; Bourlat, Sarah J. (2015) Evaluating the potential of ecological niche modelling as a component in marine non-indigenous species risk assessments, Marine Pollution Bulletin 97: 470-487

Lopes, Rubens M. (Ed.) (2009) <missing title>, Ministry of the Environment, Brasilia, Brazil. Pp. 1-440

Marelli, Dan C.; Gray, Susan (1985a) Conchological redescriptions of Mytilopsis sallei and Mytilopsis leucophaeta of the brackish Western Atlantic., Veliger 25(3): 185-193

Marelli, Dan C.; Gray, Susan (1985b) Comments on the status of recent members of the genus Mytilopsis (Bivalvia: Dreissenidae), Malacological Review 18: 117-122

Mathieson, Arthur C.; Dawes, Clinton J. (2017) Seaweeds of the Northwest Atlantic, University of Massachusetts Press, Amherst MA. Pp. <missing location>

Mohr, J. L.; J. A. Leveque (1948) Folliculinids Associated with Limnoria in California and Washingotn, Journal of Parasitology 34(6): 26

Mosyakin, Sergei L.; Freitag, Helmut (2023) Proposal to conserve the name Soda against Sevada (Chenopodiaceae s.str. / Amaranthaceae s.l.), Taxon 73(3): 1371-1373

Nehring, S.; Leuchs, Heiko (1999) Introduced macrozoobenthic species at the German North Sea Coast--a review, Wadden Sea Newsletter 1: 919-924

Nehring, Stefan (2002) Invasive aquatic species of Europe: Distribution, impacts and management, Kluwer Academic Publishers, Dordrecht, Boston, London.. Pp. 373-383

Nikolaev, I. N. (1951) [On new additions to the fauna and flora of the North sea and Baltic from distant regions], Zoologicheskii Zhurnal 30(8): 556-561

Nydam, Marie L.; Nichols, Claire L.; Lambert, Gretchen (2011) First record of the ascidian Ascidiella aspersa (Müller, 1776) in southern California , BioInvasions Records 11: 416-421

Olenin, S., Leppakoski, E. 2000-2016 Inventory of Baltic Sea alien species. <missing URL>

Oliver, P. G.; Holmes, A. M.; Mettam, C. (1998) Mytilopsis leucophaeta, (Conrad, 1831) [Bivalvia: Dresseinoidea]. A species new to the British fauna., Journal of Conchology 36(2): 13-18

Pathy, Diane A.; Mackie, Gerald L. (1993) Comparative shell morphology of Dreissena polymorpha, Mytilopsis leucophaeta, the 'quagga' mussel (Bivalvia: Dreissenidae) in North America, Canadian Journal of Zoology 71: 1012-1023

Pearse, A. S. (1936) Estuarine animals at Beaufort, North Carolina, Journal of the Elisha Mitchell Scientific Society 52(2): 174-224

Power, Alan; Mitchell, Marcy; Walker, Randal; Posey, Martin; Alphin, Troy; Belcher, Carolyn (2006) <missing title>, University of Georgia Marine Extension Service, Athens. Pp. <missing location>

Rajagopal, S.; van der Gaag, M.; van der Velde, G.; Jenner, H. A. (2002) Control of brackish water fouling mussel, Mytilopsis leucophaeata (Conrad), with sodium hypochlorite, Archives of Environmental Contamination and Toxicology 43: 296-300

Rajagopal, S.; Van der Gaag. M.; Van der Velde, G.; Jenner, H. A. (2005) Upper temperature tolerances of exotic brackish-water mussel, Mytilopsis leucophaeata (Conrad): An experimental study, Marine Environmental Research 60: 512-530

Rehder, H. A. (1937) Congeria leucophaeata (Con.) in the Hudson River, Nautilus 50: 143

Reise, K.; Gollasch, S.; Wolff, W.J. (1999) Introduced marine species of the North Sea coasts., Helgoländer Meeresuntersuchungen 52: 219-234

Ristich, S. S., Crandall, M., Fortier, J. (1977) Benthic and epibenthic macroinvertebrates of the Hudson River I. Distribution, natural history, and community structure, Estuarine and Coastal Marine Science 5: 255-266

Rizzo, Alexandra E.; Miyahira, Igor Christo; Moser, Gleyci; dos Santos, Sonia Barbosa (2014) A new record of Mytilopsis leucophaeata (Bivalvia: Dreissenidae) in Rio de Janeiro (Brazil), Marine Biodiversity Records 7: e12

Rodrigues, Nathalia; Ribeiro, Danielle; Miyahira, Igor C. Samira; Portugal, Luciano N.; Santos, G. M.; Neves, Raquel A. F. (2023) Do feeding responses of a non-native bivalve outperform the native one in a coastal lagoon? A possible explanation for the invasion success of the dark false mussel Mytilopsis leucophaeata, PeerJ 13(15848): Published online
DOI 10.7717/peerj.15848

Siddall, Scott E. (1982) Early development of Mytilopsis leucophaeata, Veliger 22(4): 378-379

Smith, Douglas G., Boss, K. J. (1995) The occurrence of Mytilopsis leucophaeta (Conrad, 1831) (Veneroidea: Dreissenidae) in southern New England, The Veliger 39: 359-360

Soors, Jan; Faasse, Marco; Stevens, Maarten; Verbessem, Ingrid; De Regge, Nico;Van den Bergh, Ericia (2010) New crustacean invaders in the Schelde estuary (Belgium), Belgian Journal of Zoology 140: 3-10

Souza, Jose R. B.; da Rocha, Cleia M. C.; Lima, Maria dos P. R. (2005) [Occurrence of exotic bivalve Mytilopsis leucophaeta (Conrad) (Mollusca, Bivalvia) in Brazil], Revista Brasileira de Zoologia 22(4): 1204-1206

Therriault, Thomas W.; Docker, Margaret F.; Orlova, Marina I. (2004) Molecular resolution of the family Dreissenidae (Mollusca: Bivalvia) with emphasis on Ponto-Caspian species, including first report of Mytilopsis leucophaeta in the Black Sea basin., Molecular Biology and Evolution 30: 479-489

U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database.

Verween, A.; Vincx, M.; Degraer, S. (2007) The effect of temperature and salinity on the survival of Mytilopsis leucophaeata larvae (Mollusca, Bivalvia): The search for environmental limits., Journal of Experimental Marine Biology and Ecology 348: 111-120

Verween, A.; Vincx, M.; Degraer, S. (2006) Growth patterns of Mytilopsis leucophaeata, an invasive biofouling bivalve in Europe, Biofouling 22(4): 221-231

Verween, Annick; Vincx, Magda; Degraer, Steven (2009) Seasonal variation in gametogenesis and spawning of Mytilopsis leucophaeata, an invasive bivalve in Europe, Journal of Molluscan Studies 75: 307-310

Verween, Annick; Vincx, Magda; Degraer, Steven (2010) The Zebra Mussel in Europe, Backhuys Publishers, Leiden, The Netherlands. Pp. 29-44

Verween, Annick; Vincx, Magda; Mees, Jan; Degraer, Steven (2005) Seasonal variability of Mytilopsis leucophaeata larvae in the harbour of Antwerp : implications for ecologically and economically sound biofouling control, Belgian Journal of Zoology 135: 91-93

Walton, William C. (1996) Occurrence of zebra mussel (Dreissena polymorpha) in the oligohaline Hudson River, New York., Estuaries 19(3): 612-618

Williams, Bolton S.; Hogan, Terry; Zo, Zinntae (1973) Hudson River Ecology: Proceedings of a Symposium, Hudson River Environmental Society, <missing place>. Pp. unpaged

Willing, Martin J. (2015) Two invasive bivalves, Rangia cuneata (G. B. Sowerby I 1831) and Mytilopsis leucophaeta (Conrad, 1831), living in freshwater in Lincolnshire, eastern England, Journal of Conchology 42(2): 189-192

Wolff, W. J. (2005) Non-indigenous marine and estuarine species in the Netherlands., Zoologische Verhandelingen 79(1): 1-116

Wolff, Wim J. (1999) Exotic invaders of the meso-oligohaline zone of estuaries in the Netherlands: why are there so many?, Helgoländer Meeresuntersuchungen 52: 393-400