Corbicula fluminea

Invasion History

First Non-native North American Tidal Record: 1938
First Non-native West Coast Tidal Record: 1938
First Non-native East/Gulf Coast Tidal Record: 1960

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General Invasion History:

Asian freshwater clams Corbicula spp. are native to Asia, including Indonesia and the Philippines and probably also to Africa and Australia (Counts 1986; McMahon 1983; McMahon 2000). Several hermaphroditic genetic lineages of these clams have been introduced to North America, Europe, South America, and Hawaii (Lee et al. 2005; Hedtke et al. 2008; Pigneur et al. 2011). We will refer to the most widespread form (lineage A of Siripattrawan et al. 2000; lineage R of Pigneur et al. 2011) as C. fluminea, and its origin as 'Asia', although the correct name may be C. leana, from Japan (Hedtke et al. 2008) (see Description). Corbicula fluminea is believed to have been introduced to Western North America from Asia before 1924, and then spread rapidly across the continent (McMahon 1983). The most likely vector was the transport of clams as a potential food item by Asian immigrants (Counts 1986; McMahon 1983). Its spread across North America was rapid, and indicates a wide variety of vectors. Likely modes of transport include use as food or bait; transport on barges, dredges, anchors, and digging machinery; use as an aquarium animal; and ballast water transport of pediveligers. Its spread in relatively unpopulated, undisturbed watersheds in Mexico and South America suggests that this species also has natural modes of spread, including the possibility of transport in bird and fish guts. Corbicula fluminea colonizes slow-moving rivers, lakes, and low-salinity, mostly fresh to oligohaline regions (0-5 PSU) of estuaries (McMahon 1983; McMahon 2000).

N.B. Corbicula lineage B, genetically related to C. fluminea from China, Korea, and Thailand, has been found in Utah and New Mexico (Siripattrawan et al. 2000; Lee et al. 2005), but is not known to occur in US estuaries.

North American Invasion History:

Invasion History on the West Coast:

Corbicula fluminea was first collected (only as dead shells) on the Pacific Coast in the Nanaimo River, on Vancouver Island, British Columbia, in 1924 (Kirkendale and Clare 2008). In 1938, it was found in the Columbia River at Knappton, Washington. It spread across the southern part of the state eastward into the Snake River, and was widespread in the Columbia Basin by 1969-71. It was first collected in the Sacramento River, California in 1945 and soon spread through the Delta region and fresher parts of the estuary via canals. Corbicula fluminea reached lower Colorado-Imperial Valley canals by 1953; Phoenix, Arizona by 1956; and by the 1970's had fouled irrigation systems and reservoirs in most of the lower Colorado Basin (Counts 1986), including the Colorado Delta in Mexico (Mellink and Ferreira-Bartrina 2000). By the 1970-80s, it was found in many smaller drainages of the West Coast, including the Willapa River, Washington (in 1971, Counts 1991); Siuslaw River, Oregon (in 1971, Counts 1991); Coos River, Oregon (Carlton 1989); the Smith River in northern California (Carlton 1979); the Santa Margarita River in Camp Pendleton, California (in 1983, Counts 1991); and the Sweetwater River, flowing into San Diego Bay, in southern California (2005, USGS Nonindigenous Aquatic Species Program 2007). In 2008, established populations have been found in inland lakes on Vancouver Island and isolated collections have been made in the Fraser River Basin, British Columbia (Kirkendale and Clare 2008). Collections of this freshwater clam from several marine sites (e.g. Point Loma, San Diego in 1964; Anaheim Bay in 1961; Santa Barbara Harbor in 1986; Counts 1986) probably represent shells washed into the sea by floods.

Invasion History on the East Coast:

Corbicula fluminea spread rapidly up the Atlantic Coast in the 1970s. It was first collected in Altamaha River, Georgia in 1971, and was abundant by 1974. Many collections to the north were nearly simultaneous: Pee Dee River, Savannah River, and Intracoastal Waterway, South Carolina (in 1972-76); Catawba River, North Carolina (in 1971); James River, Richmond, Virginia (in 1971, probably before 1968); Potomac River, Washington D.C. (in 1976); Susquehanna Flats, Upper Chesapeake Bay (in 1975); Delaware River, New Jersey-Pennsylvania (in 1971). Further north, this clam reached the Raritan River, New Jersey in 1982 (Counts 1986); but has not yet been reported from the Hudson River (Mills et al. 1997). However, in 1990, it colonized the tidal Connecticut River at East Haddam, Connecticut. This population is largely dependent on effluent from a nuclear power plant for winter survival (Balcom 1994; Morgan et al. 2003). The northward range of this clam continues to expand - it is known from several freshwater lakes in Rhode Island and Massachusetts, and from the Charles River in Watertown and Cambridge, Massachusetts (in 2001, 2005, USGS Nonindigenous Aquatic Species Program 2008; in 2003, Museum of Comparative Zoology 2008).

Corbicula's history in the Chesapeake Bay gives an example of its spread through an estuary and watershed. It was first recorded in 1971, in a tidal river near Richmond, but the size distribution indicated some shells were at least 3 years old. By 1972, it was found from River Mile 45-80 (measured from river mouth), and the lower Appomattox River (Diaz 1974). By 1976, it comprised ~ 95% of all bivalves in the river (Diaz 1994). Further downstream, at Hog Island Point, Surry, Virginia (0-5 PSU), it was less abundant, except during periods of low salinities (Jordan and Sutton 1984). By 1984, C. fluminea had spread through almost the entire non-tidal James River system, except for a few highly polluted areas (Clarke 1986). In the Potomac River, the first record of C. fluminea was in 1977 in the tidal reaches of the river. By 1978, it was present from the center of Washington (River Mile 84.5 – 95, at the mouth of Piscataway Creek), and by 1979 it was causing problems in Potomac Electric Company plants in Alexandria (Dresler and Cory 1980). Corbicula fluminea reached a biomass peak in 1984 and declined to about one-eighth of its peak by 1992, but still comprises a substantial biomass and is the dominant mollusc in the tidal reaches of the Potomac (Phelps 1994). A 'large population' occurred at Whites Ferry, Maryland in the nontidal river (~40 km upstream of Washington D.C., 1981) (Kennedy and Huekelem 1985) and the clam now occurs throughout the entire Potomac drainage (Taylor 1985). In the upper Bay, the first record of C. fluminea was in 1977 at Susquehanna Flats, but it probably arrived by 1975, and is now present from Havre de Grace to Turkey Point (Counts 1986). Corbicula fluminea was first collected in the Susquehanna River at Conowingo Dam, in 1980, but was not found above the dam (Counts 1986; Nichols and Domermuth 1981). By 1984, it was found above the dam, and by 2001 C. fluminea had colonized the North Branch of the Susquehanna in Pennsylvania and was present along at least 135 river miles (217 km) of the Susquehanna in PA (Mangan 2002). By 2002, it had colonized the upper reaches of the Susquehanna in Chengango and Otsego counties, New York (2005, USGS Nonindigenous Aquatic Species Program 2008).

In the Great Lakes St. Lawrence Basin, Corbicula fluminea reached Lake Erie in 1978 and Lake Michigan by 1984 (Mills et al. 1993). The Asian Clam is established in Lake Erie, but in Lake Michigan and Lake Superior it is confined to power plant and warm sewage effluents, where they are vulnerable to power plant shutdowns (USGS Nonindigenous Aquatic Species Program 2008; Trebitz et al. 2012). In 2009, C. fluminea was found in the fresh tidal St. Lawrence River, in the thermal plume of a nuclear power plant, downstream of Trois Rivieres, Quebec. This is its northernmost occurrence in eastern North America. Establishment of the clam here is uncertain, but it was found at sites where the influence of the thermal plume was minimal (Simard et al. 2012).

Invasion History on the Gulf Coast:

In the Eastern U.S., C. fluminea was first found in the Ohio River at Paducah, Kentucky in 1957, and rapidly spread through the Mississippi system, and adjacent rivers. To the east, it was dominant in the Tennessee River by 1969, and moved upstream to Cincinnati (1964). To the west, the Arkansas, Black and White Rivers in Arkansas were colonized by 1970; Lake Overholser, Oklahoma was colonized by 1969; and Cherry Creek Reservoir, Arapahoe County, Colorado was colonized by the 1990s (Nelson and McNabb 1994). Corbicula fluminea rapidly moved downstream, reaching Louisiana in the Mississippi by 1962; the Calcasieu River, Louisiana in 1961; the Escambia River in Century, Florida by 1960; and Galveston Bay, Texas by 1967 (Counts 1986). By 2004, it had colonized nearly every coastal county in Texas, and was expected to be found in most tidal fresh tributaries (Karatyev et al. 2005).

Invasion History in Hawaii:

Corbicula fluminea was first collected on Kauai at the Hanalei National Wildlife Refuge, in irrigation canals in 1971 (Counts 1986; USGS Nonindigenous Aquatic Species Program 2008). It was sold as food in markets in Oahu in 1977 (Counts 1986), and found in streams on Maui in 1988 and the island of Hawaii in 1991 (USGS Nonindigenous Aquatic Species Program 2008). In a recent survey of coastal streams, it was found on Maui, Kauai, and Oahu, only in freshwater (MacKenzie and Bruland 2012). The Asian Clam may have been brought to the islands as food by immigrants or as an aquarium animal, and was probably spread further with use as bait, or with irrigated agriculture.

Invasion History Elsewhere in the World:

Corbicula fluminea is widespread in central Mexico, including drainages with little human population or disturbance (Lopez-Lopez et al. 2010). It has been introduced to Central America in Panama, including the Panama Canal system (Counts et al. 2004; Lee et al. 2005), possibly with Tilapia fish stock imported from the US for aquaculture (Counts et al. 2004). In the 1970s, it was introduced to the Rio de La Plata in Argentina (both lineages A and C, Lee et al. 2005) and soon colonized the rivers in Rio Grande do Sul, Brazil. In Brazil, it is known from 10 of 26 states (da Silva and Barros 2011). Corbicula fluminea also invaded rivers in Venezuela in the 1980s (McMahon 2000), and is now in Ecuador and Peru (Lee et al. 2005). In 1998, C. fluminea was collected in the Cayey River, Puerto Rico (Williams et al. 2001). It is now found in reservoirs in several watersheds on the island (USGS Nonindigenous Aquatic Species Program 2012).

Corbicula fluminea invaded Europe around 1980, first appearing in the Dodogne River, France or the Tajo/Tagus River in Spain and Portugal (McMahon 1983; Araujo et al. 1993). The probable source was North America, since the most widespread genotype is identical or closely related to the North American 'form A' (Pfenninger et al. 2002; Pigneur et al. 2011). In 1987, Corbicula spp. appeared in the Rhine river and quickly became very abundant in the middle and lower reaches, occurring in the tidal fresh portions of the Delta in the Netherlands by 1990 (den Hartog et al. 1992, Wolff 2005). It has invaded many rivers in Western Europe, and is expanding its range towards the Baltic and Black Sea coasts. In France, Corbicula spp. is found in many of the rivers of the Atlantic and Mediterranean basins, invading the Seine around 2000 (Marescaux et al. 2010). On the Iberian Peninsula, it now occurs in 10 rivers systems in Spain and Portugal, on the Bay of Biscay, Atlantic, and Mediterranean coasts (Araujo et al. 1993; Pérez-Quintero 2008; Oscoz et al. 2009). In 1998, it was found in brackish coastal lakes (the Norfolk Broads) in eastern England (Howlett and Baker 1999) and in 2004 was found in the tidal river Thames (Elliott and zu Ermgassen 2008). In 2010, populations were discovered in tidal freshwater portions of the rivers Barrow and Nore, on the east coast of Ireland (Caffrey et al. 2011).

To a greater extent than in the Americas, an understanding of the invasion in Europe is complicated by the involvement of multiple genetic lineages. While the lineage R (related to or identical with the American A, and Japanese C. leana) is most widespread, it occurs sympatrically with lineage S (morphologically similar to Middle Eastern C. fluminalis, and genetically related to South American lineage C) in the Rhine, Meuse, and Seine rivers. A third lineage, Rlc, from the Rhone River, France, is related to the American lineage B and Chinese and Korean C. fluminea (Pigneur et al. 2011). These occurrences imply multiple introductions. The rapid spread of these clams in Europe has likely been aided by Europe's extensive canal system, as well as many other human vectors.

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Description

The common North American bivalve, currently known as Corbicula fluminea is a freshwater clam with a relatively thick, massive shell, compared to most other freshwater bivalves. The shell is triangular to ovate, with a distinct umbo, raised above the dorsal shell margin. The shell hinge has three distinct cardinal teeth, and two lateral teeth. The shell has many concentric ridges, about ~1.5 per mm of shell height. The ratio of shell length to shell height is ~1.06, and shell length to shell width is ~ 1.47. The shell interior is glossy white to pale gray with light blue, rose, or purple highlights. The periostracum in healthy, growing shells is yellow-green, but in old, eroded shells is dark brown and white. While North American populations are hermaphroditic and show little genetic variability, phenotypic variation in shell color and shape is considerable (McMahon 1991, in Thorp and Covich 1991; Lippson and Lippson 1997; Coan et al. 2000). The clams mature at sizes as small as 6.6 mm, but occasionally reach 60 mm in length (McMahon 1983).

The systematics of the genus Corbicula is uncertain. Morton (1986) lumped ~200 synonyms, and then divided Asian populations into two species C. fluminea a freshwater, hermaphroditic clam, and C. fluminalis a dioecious (2 sexes) brackish water clam. He classified North American populations, which are all hermaphroditic, as C. fluminea, the name provisionally used here. Subsequent genetic analysis indicates that most North and South America populations belong to a single genetic lineage, lineage A, with a whitish shell interior (Lee et al. 2005). However, three other morphologically and genetically different forms were also found, lineage B, from the Southwestern US (purple shell interior), lineage C (purple interior, finer shell sculpture), from La Plata, Argentina, and a 4th lineage from Igazu Falls, Brazil (Lee et al. 2005). Forms A and B were found co-occurring in the Illinois River, together with a new form D, possibly an androgenetic hybrid of forms A and B, formed by male sperm fertilizing hermaphroditic clams, with only male chromosomes being retained (Tiemann et al. 2017). Worldwide, some of the names synonymized by Morton (1986) have been revived. Corbicula lineage A from North America appears to be genetically most similar to C. leana from Japan, while lineage B resembles populations of C. fluminea from China and Korea (Siripattrawan et al. 2000; Hedtke et al. 2008). Pigneur et al. (2011) found three morphotypes of Corbicula in Europe: lineage R, sharing haplotypes with the American lineage A and the Japanese C. leana; lineage S, genetically resembling the South American C, but differing in morphology; and a form Rlc, resembling the American lineage B. Bespalya et al. studied Corbicula spp. in Pacific \Russia and Korea, generally supporting this classification of the genus and the native and exported genotypes.

The species status of these varying invasive lineages is unclear. All are hermaphroditic, and share androgenesis, in which the offspring retain only male chromosomes, resulting in clonal populations (Hedtke et al. 2008; Tiemann et al. 2017). It is possible that the names will be revised in the future, in which case the most widespread form (American lineage A; European lineage R) will probably be known as C. leana, as suggested by Hedtke et al. (2008). However, for purposes of continuity, we will use the name C. fluminea for lineage A until the change is formally made.

The brooded and pediveliger larvae of C. fluminea are described and illustrated by Nichols and Black (1994) and compared with larvae of the Zebra and Quagga Mussels (Dreissena polymorpha and D. bugensis).

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Taxonomy

Ecology

General:

The Asian clams known as Corbicula fluminea in North America and Europe are simultaneous hermaphrodites and frequently have male and female gametes together in their gonadal follicles. Self-fertilization is common (McMahon 1983; Kennedy and Huekelem 1985), but mucosal strands containing sperm have been seen connecting individuals. In invasive Corbicula lineages, sperm contains a full set of nuclear chromosomes, and the oocyte ejects the maternal chromosomes after fertilization, so only the male genome is transmitted, a phenomenon known as androgenesis (Hedtke et al. 2008). Embryos are brooded in the adult's gills, and are released at varying stages of development, sometimes as D-shaped non-swimming veligers, and sometimes as crawling postlarvae (McMahon 1983; Nichols and Black 1994). Many temperate populations have two breeding peaks a year (Kennedy and Heukelem 1985; Doherty et al. 1986; Phelps 1994). The number of larvae released are highly variable. Examples include: 1050-1900 larvae/clam in New River, Virginia (VA) (Doherty et al. 1986); and 480-1919 larvae/clam in Mechum's River, VA (Hornbach 1992). Lifetime fecundity was estimated at 68,678 larvae per year (Keller et al. 2007). Two types of larvae are known; one lacks a velum, has a well-developed foot and shell, and is best described as a 'benthic juvenile' stage (non-swimming, but can be carried by strong currents). However, D-shaped veligers, capable of swimming, have also been reported (McMahon 1983). In the Columbia River, planktonic veligers of C. fluminea comprised 13% of zooplankton individuals from 2005 to 213 (Dexter et al. 2015). Dispersal by birds in mud on feet or feathers, or in the gut, is unlikely over long distances (Counts 1986). Juveniles mature rapidly, in three months to a year after birth (McMahon 2000).

Juvenile C. fluminea are restricted to shallow nearshore waters and well oxygenated sediments. However, it is now clear that human activities in North American waterways such as stream canalization and dredging are not only detrimental to habitats of native species, but also optimize the environment for C. fluminea by increasing current flow and eliminating mud and silt (McMahon 1983). In Meyers Branch, South Carolina, a Savannah River Coastal Plain tributary, C. fluminea was limited to gravel beds and was not found in sand (Leff et al. 1990).

Corbicula fluminea tolerates a wide range of environmental conditions, permitting it to colonize many of the watersheds of North America, South America and Europe. In many colder areas, initial populations were associated with thermal effluents, but populations have spread into colder waters (Kreiser and Mitton 1995; Müller and Bauer 2011; Simard et al. 2012). Winter die-offs are common, and the lower temperature limit for most populations was considered to be around 2C (McMahon 1983), but some individuals, especially larger ones (>15 mm length), survived 9 weeks at 0C (Müller and Bauer 2011). The upper temperature limit is around 34-35°C for prolonged exposure, while reproduction is greatly reduced above 30C (McMahon 1983).

There has been some uncertainty concerning salinity tolerance in Corbicula sp., in part due to the unresolved taxonomy of the genus and the presence of cryptic species. Corbicula fluminea from Hong Kong has been reported to tolerate salinities up to 13 PSU, if the salinity is increased gradually (McMahon 1983; Morton and Tong 1985). They cite experiments by Kado and Murata (1974) on Japanese C. leana, which may be the 'lineage A' widely introduced to the US and Europe (Lee et al. 2005; Hedtke et al. 2008), and C. japonica a brackish-water form. The upper limits for these two species were reportedly 5 and 24 PSU, respectively. In North America, Corbicula sp. is rare above 2-5 PSU (Carlton 1979; McMahon 1983; Montagna et al. 2008). To our knowledge, the salinity tolerances of the widespread North American lineage A (C. leana) and lineage B (the real C. fluminea, confined to the southwest of the US) have not been compared. The lower limit for pH is about 5.6 (Karatayev et al. 2005), but this clam is common in the somewhat acidic tidal fresh Pocomoke River (Fofonoff, personal observation).

Corbicula fluminea is a suspension feeder, feeding on phytoplankton and suspended detritus, with a high filtration rate (McMahon 1983; Cohen et al. 1984; Way et al. 1990; Bolam et al. 2019). In river and reservoir habits, show varying prefereces for diatoms and flagellates, but tend to avoid cyanobacteria, which could promote blue-green blooms (Bolam et al. 2019). It is also capable of deposit-feeding on organic matter and bacteria in sediments (Hakenkamp et al. 2001). In experiments, filtration rate was highest on the smallest particles tested, about 3 µm in diameter. Pseudofeces, masses of rejected, undigested food, wrapped in mucus are discharged from the gills into the sediment (Way et al. 1990).

Food:

Phytoplankton, detritus

Consumers:

Fishes, Birds, Benthic invertebrates

Competitors:

Freshwater bivalves

Trophic Status:

Deposit Suspension Feeder

DepSusFed

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Habitats

General Habitat	Fresh (nontidal) Marsh	None
General Habitat	Grass Bed	None
General Habitat	Swamp	None
General Habitat	Nontidal Freshwater	None
General Habitat	Tidal Fresh Marsh	None
General Habitat	Unstructured Bottom	None
General Habitat	Canals	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	Endobenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	2	The lower limit for most populations is 2 C, but some populations (NE, CO) seem to have survived in colder locations, suggesting acclimation. However, many northern populations are prone to winter die-offs. 'Though observations suggest that this invading species has become established in numerous northern environments, these locations are protected from winter temperatures by industrial thermal effluents, usually from power plants '...suggesting that these thermally protected populations may serve as stepping stones in further northern expansion' (Kreiser and Mitton 1995).
Maximum Temperature (ºC)	34	Short term limits for survival of Corbicula fluminea are ~40 C, but for long-term survival, with acclimation at 5-30 C, upper LD 50's (50% Lethal Doses) are 24-34 C (McMahon 1983)
Minimum Salinity (‰)	0	This is a freshwater organsim.
Maximum Salinity (‰)	13	Salinity- Corbicula fluminea can tolerate gradual increase to 24 PSU and sudden increases to 14 ppt, but the usual limit for reproducing populations is 2-5 PSU (McMahon 1983). Specimens from Hong Kong tolerated salinities up to 13 PSU with little mortality (Morton and Tong 1985). In estuaries of southwest Florida (Charlotte Harbor, Tampa Bay, and smaller coastal rivers, C. fluminea was most abundant at salinities below 2 PSU, and was absent above 7 PSU (Montagna et al. 2008).
Minimum Dissolved Oxygen (mg/l)	6	~70% saturation (McMahon 2000), at temperatures of 20-25 C
Minimum pH	5.6	Florida, Kat 1983, cited by Karatayev et al. 2007
Maximum pH	8.2	Mino River, Portugal (Araujo et al. 1993)
Minimum Reproductive Temperature	18	Field, McMahon 1983
Maximum Reproductive Temperature	30	Field, McMahon 1983
Minimum Length (mm)	6.5	Size at maturity, McMahon 1983
Maximum Length (mm)	60	McMahon 1983
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Fresh-Mesohaline

General Impacts

Corbicula fluminea's economic and ecological impacts on freshwater and estuarine systems have been diverse and complex, owing to its ability for rapid population growth creating large biomass and quick geographic spread. Many of its ecological impacts, including impacts on water clarity, benthic/pelagic partitioning of biomass, and providing new food resources also have economic implications.

Economic impacts

Industry- In the United States, Corbicula fluminea caused fouling problems in electric generating plants, and in water treatment and water-filtration plants, as well as many other industrial operations using river water (McMahon 1983; Potter and Liden 1986; McMahon 2000). It caused shutdowns of a nuclear generating plant in Arkansas in 1980. Overall costs of Corbicula to the electric power industry probably exceed $1 billion per year (Isom 1986). 'For facilities already in use, biofouling by C. fluminea continues to be an expensive and exasperating problem for which there are now no universally accepted remedies' (McMahon 1983). Fouling problems in power plants have also been noted in Brazil (Darrigan 2002) and Europe. Corbicula fluminea caused fouling in irrigation canals including deposition of dead clams and shells, and increased sedimentation rates. This clam also clogged irrigation pipes (Isom 1986), interfered with riverbed gravel-mining operations (Diaz 1974), and fouled gravel aggregate which is used in making cement. When cement is poured and begins to set, the clams burrow to the surface, causing the cement to become porous and structurally weakened (McMahon 1983). In California, C. fluminea formed extensive bars, trapping sediment in the Delta-Mendota Canal, requiring dewatering and removal of 50,000 cubic yards of sediment. It has caused extensive problems in California irrigation systems (Cohen and Carlton 1995). In Portugal, cement plants were not affected, but 2 of the 6 power plants, 4 of 16 irrigation systems, and 6 of 420 drinking water facilities surveyed, reported problems due to fouling by C. fluminea. Overall, economic impacts due to this clam in Portugal were considered moderate, estimated at ~ 200,000 euros. However, it is possible that the invasion there is still at its early stages. (Rosa et al. 2011).

Likely beneficial uses of C. fluminea include: as a bioassay or bioindicator organism; as a protein and calcium supplement in domestic livestock feed; as a source of lime for poultry feeds and fertilizers; as a source of live and preserved bivalve material for commercial biological suppliers; and as a clarifier for tertiary sewage treatment systems by the removal of particulate organics (McMahon 1983; McMahon 2000).

Aesthetic - Die-offs and strandings due to floods and other causes produce bad odors (McMahon 1983), but C. fluminea grazing can increase water clarity (Cohen et al. 1984; Phelps 1994).

Fisheries and Hunting- Increased light penetration and vegetation growth, believed to be caused by C. fluminea's filtering (Phelps 1994), may have been responsible for increased fish populations (Killgore et al. 1989). This increased catches of Micropterus salmoides (Largemouth Bass) by sportsmen, and local aggregations of dabbling and diving ducks feeding on the clams, benefiting hunters (Perry 1981; Phelps 1994). In 1970, 2.2 million pounds of C. fluminea were sold as bait in California alone, at a value of $234,000 (Isom 1986).

Ecological Impacts

Herbivory- Corbicula fluminea can produce vast filter-feeding biomasses, with the potential to reduce and alter phytoplankton communities and suspended organic matter (seston) concentrations. Significant filtering populations occur in the tidal fresh reaches of the Potomac River (Cohen et al. 1984; Gerritsen et al. 1994; Phelps 1994; Cerco and Noel 2010), tidal fresh regions of the Sacramento-San Joaquin Delta (Lucas et al. 2002), in the nontidal Savannah River (Leff et al. 1990), and probably in many other bodies of water.

Competition- Corbicula fluminea's ability to rapidly colonize fresh waters, and vastly outnumber and outweigh native bivalves, creates concern that it will outcompete and replace native species. Corbicula fluminea reaches densities, biomasses, reproductive rates, and population filtration rates rarely reached by native molluscs (McMahon 1983). Effects of C. fluminea invasions on native bivalve populations appear to vary. Factors limiting C. fluminea's dominance include its lesser tolerance of extreme temperatures, low oxygen concentrations, and dessication compared to many native species (McMahon 1983). Competition for space with the sphaeriid Musculium partumeium (Swamp Fingernail clam) has been noted, and in some cases reductions in native unionid and sphaeriid populations have been noted (McMahon 1983). In addition to filter-feeding, C. fluminea can also deposit-feed on buried organic matter, decreasing the abundance of bacteria and flagellates in sediment (Hakenkamp 2001), and raising the potential for competition with sphaeriids (pea clams). A decline of the native sphaeriid Pisidium amnicum in the Minho estuary, Portugal, has been attributed, in part to competition with C. fluminea, and to stress caused by mass die-offs of the invading clam during heat waves (Sousa et al. 2011). The importance of C. fluminea in the decline of native mussels is unresolved (Strayer 1999; Vaughn and Spooner 2006).

Habitat Change - Phelps (1994) suggested that the invasion of the tidal Potomac River by C. fluminea caused wide-ranging changes in water quality, contributing to a resurgence of submerged vegetation and alterations in sediment, which improved habitat for many species of fish and waterfowl (see below). Additional factors, such as reduction of nutrient inputs are likely involved in these ecosystem-level changes, and the relative importance of the C. fluminea invasion remains to be determined. Corbicula fluminea populations are prone to mass die-offs, releasing ammonia and creating hypoxia, which can be stressful to other aquatic organisms (Strayer 1999; Sousa et al. 2011). The empty shells can be an important source of structure in freshwater benthic communities, increasing the numbers of mayfly (Caenis spp.) larvae in experiments in Lake Constance, Switzerland, while live clams had no effect on mayfly larva abundance (Werner and Rothhaupt 2007). In fresh and brackish waters of the Minho estuary, Portugal, the abundance of benthic invertebrates, including oligochaetes, amphipods, isopods, and gastropods was positively correlated with the abundance of C. fluminea (Ilarri et al. 2012)

Food/Prey - A wide variety of fishes are known to eat C. fluminea, including many widely introduced species, such as Lepomis macrochirus (Bluegill), L. microlophus (Red ear Sunfish), L. megalotis, Cyprinus carpio (Common Carp), Ictalurus punctatus (Channel Catfish), and I. furcatus (Blue Catfish) (McCrady 1990). Corbicula fluminea is an important food source for the endangered Shortnose Sturgeon, Acipenser brevirostris, and rare Atlantic Sturgeon, A. oxyrynchus, on the East Coast (Horwitz 1986). In addition, Corbicula provides food for many species of dabbling and diving ducks (Perry 1981; Phelps 1994: Perry and Deller 1996).

Trophic Cascade- The changes caused by C. fluminea's high filtering rates in the Potomac altered the habitat but also changed patterns of energy and nutrient flow – shifting production from the plankton to the benthos through increased light penetration, resulting in growth of submerged aquatic vegetation, decreased down-bay transport of phosphates, disappearance of blooms of the blue-green alga Microcystis, and increased organic content of sediments due to deposition of pseudofeces (Phelps 1994). Phelps (1994) argued that increased water clarity resulting from C. fluminea's filtering facilitated the invasion of the Potomac River by Hydrilla verticillata in the 1980's. Other introduced submerged aquatic vegetation, such as Myriophyllum spicatum (Eurasian Watermilfoil), Potamogeton crispus (Curly Pondweed), and Najas minor (Eurasian Water-Nymph), also would have benefited from these effects. Regrowth of native and introduced submerged aquatic vegetation in turn has positively affected waterfowl and fish populations (Killgore et al. 1989; Perry and Deller 1996; Phelps 1994). Additional factors, such as reduction of nutrient inputs are likely involved in these ecosystem-level changes, and the relative importance of the C. fluminea invasion remains to be determined.

Corbicula fluminea's capacity to deposit-feed, and to filter suspended particulate organic matter (POM) from terrestrial sources has the potential to affect the productivity of estuaries. In the Minho River, Portugal, stable isotope ratios in C. fluminea showed a shift from feeding on refractory terrestrial-derived (POM) to feeding on benthic microalgae and phytoplankton along a seaward gradient in the estuary. This clam's ability to use refractory organic matter in the water and sediments, favors its invasion in rivers and estuaries with low phytoplankton production. It also makes this terrestrial carbon available to predators and benthic fauna feeding on pseudofeces, which has the potential to alter foodwebs and increase the secondary production of estuaries (Dias et al. 2014).

Toxicity - While not inherently toxic, as very efficient filterers C. fluminea accumulate toxicants from agriculture and industry in their tissue (Baudrimont et al. 1997; Leland and Scudder 1990). Corbicula fluminea could provide a new nutritious food source for waterfowl, but cause adverse effects on the birds by increasing body levels of toxicants (Perry 1981).

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Regional Impacts

P090	San Francisco Bay		Economic Impact	Industry
In California, C. fluminea formed extensive bars, trapping sediment in the Delta-Mendota Canal, requiring dewatering and removal of 50,000 cubic yards of sediment. It has caused extensive problems in California irrigation systems (Cohen and Carlton 1995).
P090	San Francisco Bay		Ecological Impact	Toxic
While not inherently toxic, as very efficient filterers, C. fluminea accumulate toxicants in their tissue, such as selenium, arsenic and mercury. Concentrations were highest upstream near agricultural areas, and tended to decrease downstream (Leland and Scudder 1990)
P090	San Francisco Bay		Ecological Impact	Herbivory
Corbicula biomasses and filtering rates in some regions of the Sacramento-San Joaquin Delta were sufficient to sharply decrease phytoplankton biomass (Lucas et al. 2002; Lopez et al. 2006).
CA	California		Ecological Impact	Herbivory
Corbicula biomasses and filtering rates in some regions of the Sacramento-San Joaquin Delta were sufficient to sharply decrease phytoplankton biomass (Lucas et al. 2002; Lopez et al. 2006).
CA	California		Ecological Impact	Toxic
While not inherently toxic, as very efficient filterers, C. fluminea accumulate toxicants in their tissue, such as selenium, arsenic and mercury. Concentrations were highest upstream near agricultural areas, and tended to decrease downstream (Leland and Scudder 1990)
CA	California		Economic Impact	Industry
In California, C. fluminea formed extensive bars, trapping sediment in the Delta-Mendota Canal, requiring dewatering and removal of 50,000 cubic yards of sediment. It has caused extensive problems in California irrigation systems (Cohen and Carlton 1995).

References

Shalovenko, N. N. (2020) Tendencies of Invasion of Alien Zoobenthic Species into the Black Sea, Russian Journal of Biological Invasions 11(2): . 164–171

Agudo-Padrón, A. Ignacio (2011) Exotic molluscs (Mollusca, Gastropoda et Bivalvia) in Santa Catarina State, Southern Brazil region: check list and regional spatial distribution, Biodiversity Journal 2: 53-58

Alexandre, Ana; Collado-Vides, Ligia; Santos, Rui (2021) The takeover of Thalassia testudinum by Anadyomene sp. at Biscayne Bay, USA, cannot be simply explained by competition for nitrogen and phosphorous, Marine Pollution Bulletin 167(112326): Published online
https://doi.org/10.1016/j.marpolbul.2021.112326

Araujo, R.; Moreno, D.; Ramos, M. A. (1993) The Asiatic clam Corbicula fluminea (Müller, 1774) (Bivalvia: Corbiculidae) in Europe, American Malacological Bulletin 10(1): 39-49

Balcom, N.C. 1994 Aquatic Immigrants of the Northeast, No. 4: Asian Clam, <i>Corbicula fluminea</i>. <missing URL>



Barnett, Rachel; Bell, Sabrina; Floerke, Wyatt; Templin, Bill (2011) <missing title>, California Interagency Ecological Program, Sacramento CA. Pp. 13

Baudrimont, Magalie, Metivaud, Jacqueline, Maury-Brachet, Regine, Ribeyre, Francis, Boudou, Alain (1997) Bioaccumulation and metallothionein response in the Asiatic clam (Corbicula fluminea) after experimental exposure to cadmium and inorganic mercury, Environmental Toxicology and Chemistry 16(10): 2096-2105

Baur, Bruno; Schmidlin, Stephanie (2007) Biological Invasions, 193 Springer, Berlin. Pp. 257-271

Belz, Carlos Eduardo; Darrigran, Gustavo; Netto,Otto Samuel Mader; Boeger, Walter A.; Ribeiro, Paulo Justiniano Junior (2012) Analysis of four dispersion vectors in inland waters: the case of the invading bivalves in South America, Journal of Shellfish Research 31(3): 777-784

Bespalaya, Yulia (2022) A taxonomic reassessment of native and invasive species of Corbicula clams (Bivalvia: Cyrenidae) from the Russian Far East and Korea, Zoological Journal of the Linnean Society 20: 1-23

bij de Vaate, Abraham ; Hulea, Orieta (2000) Range extension of the Asiatic clam Corbicula fluminea ( Müller 1774) in the River Danube: first record from Romania, Lauterbornia 38: 23-26

Bodis, E.; Toth, B.; Sousa, R. (2013) Impact of Dreissena fouling on the physiological condition of native and invasive bivalves: interspecific and temporal variations, Biological Invasions published online: <missing location>

Boward, Daniel M., Dail, Helen M., Kazyak, Paul F. (1997) Chester River Basin: Environmental Assessment Stream Conditions, Maryland Department of Natural Resources, Annapolis. Pp. <missing location>

Boward, Daniel; Dail, Helen M.; Kazyak, Paul F. (1997) <missing title>, Maryland Department of Natural Resources, Annapolis. Pp. <missing location>

Caffrey, Joseph M.; Evers, Stephanie; Millane, Michael; Moran, Helen (2011) Current status of Ireland’s newest invasive species: the Asian clam Corbicula fluminea (Müller, 1774), Aquatic Invasions 6: corrected proof

Cairns, John, Jr.; Bidwell, Joseph R. (1996) Discontinuities in technological and natural systems caused by exotic species, Biodiversity and Conservation 5: 1085-1094

Carlton, James T. (1979) History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific Coast of North America., Ph.D. dissertation, University of California, Davis. Pp. 1-904

Carlton, James T. (1989) <missing title>, <missing publisher>, <missing place>. Pp. <missing location>

Cazzaniga, Nastor J. (1997) Asiatic clam, Corbicula fluminea, William T.; Flynn, Kevin C., reaching Patagonia, Journal of Freshwater Ecology 12(4): 629-630

Cerco, Carl F.; Noel, Mark R. (2010) Monitoring, modeling, and management impacts of bivalve filter feeders in the oligohaline and tidal fresh regions of the Chesapeake Bay system, Ecological Modelling 221: 1054-1064

Chainho, Paula and 20 additional authors (2015) Non-indigenous species in Portuguese coastal areas, lagoons, estuaries, and islands, Estuarine, Coastal and Shelf Science <missing volume>: <missing location>

Clarke, Arthur H. (1986) Competitive exclusion of Canthyria (Unionidae) by Corbicula fluminea (Müller), Malacology Data Net 1: 3-10

Coan, Eugene V.; Valentich-Scott, Paul; Bernard, Frank R. (2000) Bivalve Seashells of Western North Ameira, Santa Barbara Museum of Natural history, Santa Barbara CA. Pp. <missing location>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Cohen, R. R. H.; Dresler, P. V.; Phillips, E. J. P.; Cory, R. L. (1984) The effect of the Asiatic clam, Corbicula fluminea, on phytoplankton of the Potomac River, Maryland, Limnology and Oceanography 29(1): 170-180

Counts, Clement L. III (1986) The zoogeography and history of the invasion of the United States by Corbicula fluminea (Bivalvia: Corbiculidae), American Malacological Bulletin, Special Edition 2: 7-39

Counts, Clement L. III (1991) Corbicula (Bivalvia: Corbiculidae): Part II. Compendium of zoogeographical records of North America and Hawaii, 1924-1984, Tryonia 21(69): 132

Counts, Clement L., III (1991) Corbicula (Bivalvia: Corbiculidae): Part I. Catalog of fossil and recent nominal species, Tryonia 21(ii): 1-134

Counts, Clement L., III, Handwerker, Thomas S., Jesien, Roman V. (1991) The Naiades (Bivalvia: Unionoidea) of the Delmarva Peninsula, American Malacological Bulletin 9(1): 27-37

Counts, Clements L.; Janzel, R. Villalaz; Gomez H. (2004) Occurrence of Corbicula flumimea (Bivalvia: Corbiculidae) in Panama., Journal of Freshwater Ecology 18(3): 497-498

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

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

DAISIE (Delivering Alien Invasive Species Inventories to Europe) (2009) Handbook of alien species in Europe, Springer, Dordrecht, Netherlands. Pp. 269-374

Darrigran, Gustavo. (2002) Potential impact of filter-feeding invaders on temperate inland freshwater environments., Biological Invasions 4: 145-156

den Hartog, C.; van den Brink, F. W. B.; van der Velde, G. (1992) Why was the invasion of the river Rhine by Corophium curvispinum and Corbicula species so successful?, Journal of Natural History 26: 1121-1129

Dexter, Eric; Bollens, Stephen M.; Rollwagen-Bollens, Gretchen; Emerson, Josh; Zimmerman, Julie (2015) Persistent vs. ephemeral invasions: 8.5 years of zooplankton community dynamics in the Columbia River, Limnology and Oceanography 60: 527-539

Dias, Ester; Morais, Pedro; Antunes, Carlos; Hoffman, Joel C. (2014) Linking terrestrial and benthic estuarine ecosystems: Organic matter sources supporting the high secondary production of a non-indigenous bivalve, Biological Invasions 16(10): 2163-2179

Diaz, R. J. (1974) Asiatic clam, Corbicula manilensis (Philippi), in the tidal James River, Virginia, Chesapeake Science 15: 118-120

Diaz, Robert J. (1977) <missing title>, Ph. D. Dissertation, University of Virginia, Charlottesville. Pp. <missing location>

Diaz, Robert J. (1989) Pollution and tidal benthic communities of the James River estuary, Hydrobiologia 180: 195-211

Diaz, Robert J. (1994) Response of tidal freshwater macrobenthos to sediment disturbance, Hydrobiologia 278(1-3): 201-212

Doherty, F.G.; Cherry, D. S.; Cairns, Jr., J. (1986) Spawning periodicity of the Asiatic clam, Corbicula fluminea, in the New River, Virginia, American Malacological Bulletin 4: 116

Dresler, Paul V.; Cory, Robert L. (1980) The Asiatic clam Corbicula fluminea (Müller) in the tidal Potomac River, Maryland, Estuaries 3(1): 150-151

Dundee, Dee S. (1974) Catalog of introduced molluscs of eastern North America (North of Mexico), Sterkiana 55: 1-37

Elliott, Paul; zu Ermgassen, Philine S.E (2008) The Asian clam (Corbicula fluminea) in the River Thames, London, England., Aquatic Invasions 3(1): 54-60

Ferreira-Rodriguez, Noe; Pavel, Ana Bianca; Cogalniceanu, Dan (2021) Integrating expert opinion and traditional ecological knowledge in invasive alien species management: Corbicula in Eastern Europe as a model, Biological Invasions 23: 1087-1099

Foss, Stephen (2009) <missing title>, California Department of Fish and Game, Sacramento CA. Pp. <missing location>

Franco, J. N. and 9 authors (2012) Population dynamics of Corbicula fluminea (Müller, 1774) in mesohaline and oligohaline habitats: Invasion success in a Southern Europe estuary, Estuarine, Coastal and Shelf Science 112: 31-39

Fuller, Samuel L. H.; Powell, Charles E., Jr. (1973) Range extensions of Corbicula manilensis (Philippi) in the Atlantic drainage of the United States, The Nautilus 87(2): 59

Gatlin, Michael R.;Shoup, Daniel E.; Long, James M. (2012) Invasive zebra mussels (Dreissena polymorpha) and Asian clams (Corbicula fluminea) survive gut passage of migratory fish species: implications for dispersal, Biological Invasions published online: <missing location>

Gerritsen, Jeroen; Holland, A Frederick; Irvine, David E. (1994) Suspension-feeding bivalves and the fate of primary production: an estuarine model applied to Chesapeake Bay., Estuaries 17(2): 403-416

Glisson, Wesley J.; Larkin, Daniel J. (2021) Hybrid watermilfoil (Myriophyllum spicatum X Myriophyllum sibiricum) exhibits traits associated with greater invasiveness than its introduced and native parental taxa, Biological Invasions Published online: <missing location>

Grajales, Alejamdro ; Rodriguez, Estefania (2014) Morphological revision of the genus Aiptasia and the family Aiptasiidae (Cnidaria, Actiniaria, Metridioidea), Zootaxa 3826(1): 55-100
http://dx.doi.org/10.11646/zootaxa.3826.1.2

Hakenkamp, Christine C. and 5 other authors (2001) The impact of introduced bivalve (Corbicula fluminea) on the benthos of a sandy stream, Freshwater Biology 46: 491-501

Hakenkamp, Christine C.; Palmer, Margaret A. (1999) Introduced bivalves in freshwater ecosystems: the impact of Corbicula on organic matter dynamics in a sandy stream., Oecologia 119: 445-451

Hanna, G. Dallas (1966) Introduced mollusks of Western North America, Occasional Papers of the California Academy of Sciences 48: <missing location>

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



Hedtke, Shannon M.; Stanger-Hall, Kathrin; Baker, Robert J.; David M. Hillis (2008) All-male asexuality: Origin and maintenance of androgenesis in the Asian clam Corbicula, Evolution 62(5): 1119-1136

Henricksen, Summer; Bollens, Stephen M. (2021) Abundance and growth of the invasive Asian clam, Corbicula fluminea, in the lower Columbia River, USA, Aquatic Invasions 17: In press

Herms, Daniel A. McCullough, Deborah G. (2014) Emerald Ash Borer Invasion of North America: History, Biology, Ecology, Impacts, and Management, Annual Review of Entomology 59: 13–30

Hoagland, K. Elaine (1986) Unsolved problems and promising approaches in the study of Corbicula, American Malacological Bulletin, Special Edition <missing volume>: 203-209

Holthuis, L. B. (1978) A collection of decapod crustacea from Sumbs, Lesser Sunda Islands , Zoologische Verhandelingen 162(1): 3-55

Hornbach, Daniel J. (1992) Life history traits of a riverine population of the Asian clam Corbicula fluminea, American Midland Naturalist 127: 248-257

Horwitz, Richard J. (1986) Fishes of the Delaware estuary in Pennsylvania., In: Majundar, S.K., Brenner, F. J., Rhoads, A. F.(Eds.) Endangered and Threatened Species Programs in Pennsylvania.. , Philadelphia. Pp. 177-201

Hoves, Mark; Neves, Richard (1991) Distribution and life history of the James Spinymussel., Endangered Species Technical Bulletin 16(3): 9

Howlett, D.; Baker, R. (1999) Corbicula fluminea (Muller): New to U.K., Journal of Conchology 36(6): 83

Hughes, Rofer N. (18785) Feeding behavior of the sessile gastropod Tispsycha tulipa, Journal of Molluscan Studies 51: 326-330

Ilarri, Martina I. and 5 authors (2012) Associated macorbenthos with the invasvie Asian clam Corbicula fluminea, Journal of Sea Research 72: 113-120

Ilarri, Martina I.; Sousa, Ronaldo (2012) A handbook of global freshwater invasive species, Earthscan, New York NY. Pp. 174-183

Invasive Species Specialist Group (International Union for the Conservation of Nature) 2022 100 of the World's Worst Invasive Alien Species-Oreochromis mossambicus. http://www.iucngisd.org/gisd/100_worst.php



Isom, Billy D. (1986) Historical Review of Asiatic clam (Corbicula) invasion and biofouling of waters and industries in the Americas, American Malacological Bulletin, Special Edition <missing volume>: 1-5

Jassby, Alan (2008) Phytoplankton in the upper San Francisco estuary: Recent biomass trends, their causes and their trophic significance., San Francisco Estuary and Watershed Science 2007(2): 1-24

Jassby, Alan D.; Cloern, James E.; Cole, Brian E. (2002) Annual primary production: Patterns and mechanisms of change in a nutrient-rich tidal ecosystem., Limnology and Oceanography 47(3): 698-712.

Jordan, Robert A.; Sutton, Charles E. (1984) Oligohaline benthic invertebrate communities at two Chesapeake Bay power plants, Estuaries 7(3): 192-212

Karatayev, Alexander Y. and 5 authors (2012) Exotic molluscs in the Great Lakes host epizootically important trematodes, Journal of Shellfish Research 31: 885-894

Karatayev, Alexander Y.;Howells, Robert G.; Burlakova, Lyubov E.; Sewell, Brian D. (2005) History of spread and current distribution of Corbicula fluminea (Müller) in Texas, Journal of Shellfish Research 24(2): 553-559

Keillor, Phillip (1993) Using filtration and induced infiltration intakes to exclude organsims from water supply systems, Engineering notes- University of Wisconsin Sea Grant 4: 1-14

Keller, Reuben P.; Drake, John M.; Lodge, David M. (2007) Fecundity as a basis for risk assessment of nonindigenous freshwater molluscs, Conservation Biology 21(1): 191-200

Kennedy, Victor S.; Huekelem, Laurie van (1985) Gametogenesis and larval production in a population of the introduced Asiatic clam, Corbicula sp., (Bivalvia: Corbiculidae), in Maryland., Biological Bulletin 168: 50-60

Killgore, K. Jack; Morgan, Raymond P. II; Rybicki, Nancy B. (1989) Distribution and abundance of fishes associated with submersed aquatic plants in the Potomac River, North American Journal of Fisheries Management 9: 101-111

Kimmerer, Wim (2005) Long-term changes in apparent uptake of silica in the San Francisco estuary., Limnology and Oceanography 50(3): 793-798

Kirkendale, Lisa; Clare, Jeremy (2008) The Asiatic Clam (Corbicula fluminea) ‘rediscovered’ on Vancouver Island, Victoria Naturalist 65(3): 12-16

Kreiser, Brian R.; Mitton, Jeffry B. (1995) The evolution of cold tolerance in Corbicula fluminea (Bivalvia: Corbiculidae), The Nautilus 109(4): 11-112

Lee, Byeong-Gweon; Lee, Jung-Suk; Luoma, Samuel N. (2006) Comparison of selenium bioaccumulation in the clams Corbicula fluminea and Potamocorbula amurensis: a bioenergetic modeling approach, Environmental Toxicology and Chemistry 25(7): 1933-1940

Lee, Taehwan; Siripattrawan, Sirirat; Ituarte, Cristian F.; and Foighil, Diarmaid O´ (2005) Invasion of the clonal clams: Corbicula lineages in the New World., American Malacological Bulletin 20: 113-122

Leff, Laura G.; Burch, Jarrett H.; McArthur, J. Vaun (1990) Spatial distribution, seston removal, and potential competitive interactions of the bivalves Corbicula fluminea and Elliptio complanata, in a coastal plain stream, Freshwater Biology 24(409-416): <missing location>

Leland, Harry V., Scudder, Barbara C. (1990) Trace elements in Corbicula fluminea from the San Joaquin River, California, The Science of the Total Environment <missing volume>: 641-672

Leland, Harry V.; Fend, Steven V. (1998) Benthic invertebrate distributions in the San Joaquin River, California, in relation to physical and chemical factors., Canadian Journal of Fisheries and Aquatic Science 55: 1051-1067

Lippson, Alice Jane; Lippson, Robert L. (1997) <missing title>, Johns Hopkins University Press, Baltimore. Pp. <missing location>

López-López, Eugenia; Sedeño-Díaz, J. Elías; Vega, Perla Tapia; Oliveros, Eloiza (2010) Invasive mollusks Tarebia granifera Lamarck, 1822 and Corbicula fluminea Müller, 1774 in the Tuxpam and Tecolutla rivers, Mexico: spatial and seasonal distribution patterns, Aquatic Invasions 5(S1): 435-450

Lopez, Cary B. and 6 authors (2006) Ecological values of shallow-water habitats: implications for the restoration of disturbed ecosystems, Ecosystems 9: 422-440

Lorenz, Stefan; Pusch, Martin T. (2013) Filtration activity of invasive mussel species under wave disturbance conditions, Biological Invasions Published online: <missing location>

Lucas, Lisa V.; Cloern, James E.; Thompson, Janet K.; Monsen, Nancy E. (2002) Functional variability of habitats within the Sacramento-San Joaquin Delta: Restoration implications, Ecological Applications 12(5): 1528-1547

Lucy, Frances E.; Karatayev, Alexander Y.; Burlakova, Lyubov E. (2012) Predictions for the spread, population density, and impacts of Corbicula fluminea in Ireland, Aquatic Invasions 7: published online

MacKenzie, Richard Ames; Bruland, Gregory L. (2012) Nekton communities in Hawaiian coastal wetlands: the distribution and abundance of introduced fish species, Estuaries and Coasts 35: 212-226

Mangan, Brian P. (2002) Range expansion of the Asiatic clam, Corbicula fluminea, into the North Branch of the Susquehanna River., Journal of the Pennsylvania Academy of Science 76(1): 40-42

Marescaux, Jonathan; Pigneur, Lise-Marie; Van Doninck, Karine (2010) New records of Corbicula clams in French rivers, Aquatic Invasions 5: S35-S39

Matthews, Milton A.; McMahon, Robert F. (1999) Effects of temperature and temperature acclimation on survival of zebra mussels (Dreissena polymorpha) and Asian clams (Corbicula fluminea) under extreme hypoxia, Journal of Molluscan Studies 65: 317-325

McCann, James A.; Arkin, Lori; Williams, James D. (1996) <missing title>, University of Florida, Center for Aquatic Plants, Gainesville. Pp. unpaged

McCrady, Ellen Joy (1990) <missing title>, M. S. Thesis, University of Texas at Arlington, Arlington, Texas. Pp. <missing location>

McDowell, W. G.; Benson, A. J.; Byers, J. E. (2014) Climate controls the distribution of a widespread invasive species: implications for future range expansion, Freshwater Biology 59: 847-857

McMahon, Robert F. (1983) Ecology of an invasive pest bivalve, Corbicula., In: (Eds.) The Mollusca, Ecology. Vol. 6.. , New York. Pp. 505-561

McMahon, Robert F. (2000) Nonindigenous freshwater organisms: Vectors, biology, impacts, Lewis Publishers, Boca Raton FL. Pp. 315-341

Mellink, E.; Ferreira-Bartrina, V. (2000) On the wildlife of wetlands of the Mexican portion of the Rio Colorado Delta, Southern California Academy of Sciences 99(3): 115-127

Mills, Edward L.; Leach, Joseph H.; Carlton, James T.; Secor, Carol L. (1993) Exotic species in the Great Lakes: a history of biotic crises and anthropogenic introductions., Journal of Great Lakes Research 19(1): 1-54

Mills, Edward L.; Scheuerell, Mark D.; Carlton, James T.; Strayer, David (1997) Biological invasions in the Hudson River: an inventory and historical analysis., New York State Museum Circular 57: 1-51

Minchin, Dan (2014) The distribution of the Asian clam Corbicula fluminea and its potential to spread in Ireland, Management of Biological Invasions 5: in press

Modesto, Vanessa and 5 authors (2013) Spatial and temporal dynamics of Corbicula fluminea (Muller, 1774) in relation to environmental variables in the Mondego estuary (Portugal), Journal of Molluscan Studies 79: 302-309

Montagna, Paul A.; Estevez, Ernest D.; Palmer, Terry A.; Flannery, Michael S. (2008) Meta-analysis of the relationship between salinity and molluscs in tidal river estuaries of southwest Florida, U.S.A, American Malacological Bulletin 24(1): 101-115

Morgan, D. E.; Keser, M.; Swenarton, J. T.; Foertch, J. F. (2003) Population dynamics of the Asiatic clam, Corbicula fluminea in the lower Connecticut River: establishing a foothold in New England., Journal of Shellfish Research 22(1): 193-203

Morton, Brian (1986) Corbicula in Asia- an updated synthesis, American Malacological Bulletin, Special Edition <missing volume>(Special Edition): 113-124

Morton, Brian; Tong, K. Y. (1985) The salinity tolerance of Corbicula fluminea (Bivalvia: Corbiculoidea) from Hong Kong, Malacological Review 18: 91-95

Müller, Oliver; Baur, Bruno (2011) Survival of the invasive clam Corbicula fluminea (Müller) in response to winter water temperature, Malacologia 53(2): 367-371

Nakano, Daisuke; Strayer, David L. (2014) Biofouling animals in fresh water: biology, impacts, and ecosystem engineering, Frontiers in Ecology and the Environment 12(3): 167: 175

Nelson, S. Mark; McNabb, Cal (1994) New record of Asiatic clam in Colorado, Journal of Freshwater Ecology 9(1): 79

Nichols, Barry L.; Domermuth, Robert B. (1981) Apppearance of the Asiatic clam, Corbicula fluminea, in the Susquehanna River, Proceedings of the Pennsylvania Academy of Science 55: 181-182

Nichols, S. J.; Black, M. G. (1994) Identification of larvae: The zebra mussel (Dreissena polymorpha), quagga mussel (Dreissena rostriformis bugensis), and Asian clam (Corbicula fluminea), Canadian Journal of Zoology 72: 406-417

Novais, Adriana; Souza, Allan T.; Ilarri, Martina ; Pascoal, Cláudia; Sousa, Ronaldo (2015) Facilitation in the low intertidal: effects of an invasive species on the structure of an estuarine macrozoobenthic assemblage, Marine Ecology Progress Series 522: 157-167

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

Ohlmeyer, Garrett (2017) Roseau cane-killing bug found in Terrebonne, Lafourche, Houma Today Published online: <missing location>

Okawa, Takuya; Kurita, Yoshihisa; Kanno, Kazuki; Koyama, Akihiko; Onikura, Norio (2016) Molecular analysis of the distributions of the invasive Asian clam, Corbicula fluminea (O.F. Müller, 1774), and threatened native clam, C. leana Prime, 1867, on Kyushu Island, Japan, BioInvasions Records 5: In press

Oscoz, Javier; Tomás, Pedro; Durán, Concha (2009) Review and new records of non-indigenous freshwater invertebrates in the Ebro River basin (Northeast Spain), Aquatic Invasions 5(3): 263-284

Owen, Debra A., Cahoon, Lawrence B. (1991) An investigation into the use of exotic and native bivalves as indicators of eutrophication-induced hypoxia, Journal of the Elisha Mitchell Scientific Society 107(2): 71-74

Pearce, Timothy A.; Evans, Ryan (2008) Freshwater Mollusca of Plummers Island, Maryland, Bulletin of the Biological Society of Washington 15(1): 20-30

Pérez-Quintero, Juan Carlos (2008) Revision of the distribution of Corbicula fluminea (Müller 1744) in the Iberian Peninsula., Aquatic Invasions 3(3): 355-358

Perry, Matthew C. (1981) Asiatic Clam (Corbicula manilensis) and other foods used by waterfowl in the James River, Virginia, Estuaries 4(3): 229-233

Perry, Matthew C.; Deller, Amy S. (1996) Review of factors affecting the distribution and abundance of waterfowl in shallow-water habitats of Chesapeake Bay, Estuaries 19(2A): 272-276

Pestana, L. B.; Dias, G. M.; Marques, A. C . (2021) Spatial and temporal diversity of non-native biofouling species associated with marinas in two Angolan bays, African Journal of Marine Science 42(4): 413-422

Peterson, Heather A.; Vayssieres, Marc (2010) Benthic assemblage variability in the upper San Francisco estuary: A 27-year retrospective, San Francisco Estuary and Watershed Science <missing volume>: published online

Pfenninger, M.; Reihardt, F.; Streit, B. (2002) Evidence for cryptic hybridization between different evolutionary lineages of the invasive clam genus Corbicula (Verneroidea, Bivalvia)., Journal of Evolutionary Biology 15: 818-829.

Phelps, Harriette L. (1994) The Asiatic clam (Corbicula fluminea) invasion and system-level ecological change in the Potomac River estuary near Washington, D.C., Estuaries 17(3): 614-621

Pigneur, Lise-Marie and 5 authors (2011) Phylogeny and androgenesis in the invasive Corbicula clams (Bivalvia, Corbiculidae) in Western Europe, BMC Evolutionary Biology 11(147): published online

Potter, Jeanne M., Liden, Lawrence H. (1986) Corbicula control at the Potomac Steam Electric Power Station, Alexandria, Virginia, American Malacological Bulletin, Special Edition <missing volume>(Special Edition): 53-58

Robertson, D. Ross; Dominguez-Dominguez, Omar; Solís-Guzmán; María Gloria; Kingon, Kelly C (2021b) Origins of isolated populations of an Indo-Pacific damselfish at opposite ends of the Greater Caribbean, Aquatic Invasions 16: 269-280
0, https://doi.org/10. 3391/ai.2021.16.2.04 Received: 13 May 20

Rosa, Ines C. and 5 authors (2011) The Asian clam Corbicula fluminea in the European freshwater-dependent industry: A latent threat or friendly enemy, Ecological Economics 70: 1805-1813

Salmon, Terry and 21 authors 2014-2022 California Fish Website. https://calfish.ucdavis.edu/



Sampaio, Eduardo; Rodil, Ivan F. (2014) Effects of the invasive clam Corbicula fluminea (Muller 1774) on a representative macrobenthic community from two estuaries at different stages of invasion, Limnetica 33(2): 249-262

Santamaria, Luis: Klaassen, Marcel (2002) Waterbird-mediated dispersal of aquatic organisms: an introduction, Acta Oecologia 23: 115-119

Sickel, James B. (1973) A new record of Corbicula manilensis (Philippi) in the southern Atlantic slope region of Georgia, The Nautilus 87(1): 11-12

Simard, M. Anouk and 6 authors (2012) North American range extension of the invasive Asian clam in a St. Lawrence River power station thermal plume, Aquatic Invasions 12(1): 81-89

Siripattrawan, Sirirat; Park, Joon-Ki; O'Foighil, Diarmiad (2000) Two lineages of the invasive clam Corbicula occur in North America, Journal of Molluscan Studies 66: 423-429

Skolka, Marius; Preda, Cristina (2010) Alien invasive species at the Romanian Black Sea coast: present and perspectives, Travaux du Muséum National d’Histoire Naturelle «Grigore Antipa» 53: 443-467

Sousa, and 6 authors. (2007) Genetic and shell morphological variability of the invasive bivalve Corbicula fluminea (Muller, 1774) in two Portuguese estuaries., Estuarine, Coastal and Shelf Science 74: 166-174

Sousa, R.; Antunes, C.; Guilhermino, L. (2008) Ecology of the invasive clam Corbicula fluminea (Muller 1774) in aquatic ecosystems: an overview., Annales de Limnologie 44(2): 85-94

Sousa, Ronaldo; Gutierrez, Jorge L.; Aldridge, David C. (2009) Non-indigenous invasive bivalves as ecosystem engineers., Biological Invasions 10: 2367-2385

Sousa, Ronaldo; Ilarri, Martina; Souza, Allan T.; Antunes, Carlos; Guilhermino, Lucia (2011) Rapid decline of the greater European peaclam at the periphery of its distribution, Annales de Limnologie 47: 211-219

Sousa, Ronaldo; Novais, Adriana; Costa, Raquel; Strayer, David L. (2014) Invasive bivalves in fresh waters: impacts from individuals to ecosystems and possible control strategies, Hydrobiologia 735: 233-251

Statzner, Bernhard; Bonada, Nuria; Doledec, Sylvain (2008) Biological attributes discriminating invasive from native European stream macroinvertebrates., Biological Invasions 10: 517-530

Strayer, David L. (1999) Effects of alien species on freshwater mollusks in North America, Journal of the North American Benthological Society 18(1): 74-98

Sytsma, Mark D.; Cordell, Jeffrey R.; Chapman, John W.; Draheim, Robyn, C. (2004) <missing title>, Center for Lakes and Reservoirs, Portland State University, Portland OR. Pp. <missing location>

Tavares, M. R.; .Franco, A. C. S. ; . Ventura, C. R. R.; .Santos, L. N. (2021) Geographic distribution of Ophiothela brittle stars (Echinodermata: Ophiuroidea): substrate use plasticity and implications for the silent invasion of O. mirabilis in the Atlantic, Hydrobiologia 848: 2093-2103

Taylor, Ralph W. (1985) Comments on the distribution of freshwater mussels (Unionaceae) of the Potomac River headwaters in West Virginia, Nautilus 99(2-3): 84-87

Thorp, James H.; Covich, Alan P. (2001) <missing title>, Academic Press, San Diego CA. Pp. <missing location>

Trama, Francesco B. (1982) Occurrence of the Asiatic clam Corbicula fluminea in the Raritan River, New Jersey, Nautilus 96(1): 6-8

Trebitz, Anett S. and 5 authors (2010) Status of non-indigenous benthic invertebrates in the Duluth-Superior Harbor and the role of sampling methods in their detection, Journal of Great Lakes Research 36: 747-756

USGS Nonindigenous Aquatic Species Program 2003-2024 Nonindigenous Aquatic Species Database. https://nas.er.usgs.gov/



Vaughn, Caryn C.; Spooner, Daneil E. (2006) Scale dependent associations between native mussels and invasive Corbicula, Hydrobiologia 568: 531-539

Way, Carl M.; Hornbach, Daniel M.; Miller-Way, Christine A.; Payne, Barry S.; Miller, Andrew C. (1990) Dynamics of filter-feeding in Corbicula fluminea (Bivalvia: Corbiculidae), Canadian Journal of Zoology 68: 115-120

Werner, Stefan; Rothhaupt, Karl-Otto (2002) Effects of the invasive bivalve Corbicula fluminea on settling juveniles and other benthic taxa, Journal of the North American Benthological Society 26: 673-680

Wijnhoven, Sander; Hummel, Herman (2011) Patterns in macrozoobenthic assemblages indicate the state of the environment: insights from the Rhine-Meuse estuary, Marine Ecology Progress Series 436: 29-50

Williams, Ernest H. Jr.; Bunkley-Williams,Lucy; ilyestrom, Craig G. L; Ortiz-Corps, Edgardo A. R. (2001) A review of recent introductions of aquatic invertebrates in Puerto Rico and implications for the management of nonindigenous species., Caribbean Journal of Science 37(3-4): 246-251

Wilson, Sarah; Partridge, Valerie (2007) <missing title>, Washington State Department of Ecology, Olympia. Pp. 244

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

Yamada, Sylvia Behrens; Fisher, Jennifer L.; Kosro, P. Michael (2021) Relationship between ocean ecosystem indicators and year class strength of the invasive European green crab (Carcinus maenas), Progress in Oceanography 196(102618): Published online

Zeng, Cong; Tang, Yangxin; Vastrade, Martin; Coughlan, Neil E; Zhang. Ting; Cai, Yongjiu; Van Doninck, Karine; Li, Deliang (2022) Salinity appears to be the main factor shaping spatial COI diversity of Corbicula lineages within the Chinese Yangtze River Basin, Diversity and Distributions <missing volume>: Published online
DOI: 10.1111/ddi.13666

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