Mytilopsis leucophaeata

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

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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).

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Description

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).

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Taxonomy

Ecology

General:

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).

Food:

Phytoplankton

Consumers:

Crabs, fishes

Competitors:

barnacles, Dreissena spp.

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Mangroves	None
General Habitat	Canals	None
General Habitat	Vessel Hull	None
Salinity Range	Limnetic	0-0.5 PSU
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	0	Based on geographical range
Maximum Temperature (ºC)	35	Experiment, Rajagopal et al. 2005
Minimum Salinity (‰)	0	Experimental, Castagna and Chanley (1973)
Maximum Salinity (‰)	32	Experimental, 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 Temperature	10	Experimental, 4 hr old embryos- Verween et 2007, Belgium
Maximum Reproductive Temperature	30	Field, spatfall, July, Miami FL(Siddall 1982), but 24, Experimental, 4 hr old embryos- Verween et 2007, Belgium
Minimum Reproductive Salinity	5	Experimental, 4 hr old embryos- Verween et 2007, Belgium
Maximum Reproductive Salinity	20	Experimental, 4 hr old embryros- Verween et 2007, Belgium
Minimum Duration	6	Larval duration, experimental, 26 C (Siddall 1982)
Maximum Duration	8	Larval duration, experimental, 26 C (Siddall 1982)
Minimum Length (mm)	10	Vorstman 1933, (Germany) cited by Kennedy 2011a
Maximum Length (mm)	25	Abbott 1974; Pathy and Mackie 1993; Oliver et al. 1998; Kennedy 2011a
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Oligohaline-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).

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Regional Impacts

NEA-II	None		Economic Impact	Industry
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-V	None		Economic Impact	Industry
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-III	None		Economic Impact	Fisheries
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-II	None		Ecological Impact	Herbivory
Compared to a native bivalve, Mytilopsis leucophaeta could increase its ingestion rate at high phytoplankton concentrations (Rodrigues et al. 2023)
SA-II	None		Ecological Impact	Competition

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