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

M130	Chesapeake Bay		Economic Impact	Industry
Corbicula fluminea caused fouling of nuclear and conventional power plants, by clogging water pumps and condensers, including Potomac River Steam Electric Station, Alexandria VA, and the 12th Street Generating Plant, Richmond VA. This resulted in reduced efficiency, decreased output, and outages due to time required for cleaning (Diaz 1974; Potter and Liden 1986).
M130	Chesapeake Bay		Ecological Impact	Herbivory
In 1980, the biomass of C. fluminea in the tidal fresh Potomac River was estimated to be sufficient to filter all of the phytoplankton from one stretch (Rosier Bluff to Hatton Point, MD; River Km 160-165) every 3-4 days (Cohen et al. 1984). Cerco and Noel (2010) estimated filtering rates for bivalves (Corbicula and Rangia) in the oligohaline waters of Chesapeake Bay and its tributaries. Corbicula comprised <1-60% of the filter-feeding biomass in the major tributaries and upper Bay, being most abundant in the Potomac. The two species together removed 14% to 40% of the carbon load, 11% to 23% of the nitrogen load, and 37% to 84% of the phosphorus load from the water column (Cerco and Noel 2010).
M130	Chesapeake Bay		Ecological Impact	Competition
In the nontidal James River, Corbicula fluminea was thought to have virtually eliminated the native unionid Pleurobema collina (James River Spiny mussel), which formerly ranged from Richmond to the headwaters and is now confined to a few headwater streams. Abundances of Fusconaia masoni (Atlantic Pigtoe), Alismidonta undulata (Triangle Floater) and Strophitus undulatus (Squawfoot) may have been seriously reduced, but Elliptio complanata appeared to have been unaffected (Clarke 1986). Later studies have stressed the effects of habitat disturbance (siltation, stream modification, pollution) in the decline of P. collina in the James River Basin (Howe and Neves 1991). The importance of C. fluminea in the decline of native mussels is unresolved.
M130	Chesapeake Bay		Ecological Impact	Habitat Change
Changes possibly caused by C. fluminea's high filtering rates in the Potomac include: increased light penetration resulting in regrowth of native submerged aquatic vegetation, decreased down-bay transport of phosphates, disappearance of blooms of the blue-green alga Microcystis, and changes in sediment composition due to deposition of pseudofeces (Phelps 1994). 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.
M130	Chesapeake Bay		Ecological Impact	Trophic Cascade
Corbicula fluminea's invasion, especially in the Potomac River, has resulted in major changes in local foodwebs in tidal fresh and oligohaine regions, and has apparently shifted much of the flow of nutrients and energy from the water column to the benthos, affecting both the primary producers (phytoplankton and submerged vascular plants), and higher level predators, such as fishes and waterfowl. Changes attributed to C. fluminea include bay transport of phosphates on particles, disappearance of blooms of the blue-green alga Microcystis, and changes in sediment composition due to deposition of pseudofeces (Phelps 1994).
M130	Chesapeake Bay		Ecological Impact	Food/Prey
A wide variety of fishes are known to eat C. fluminea, but many of these species are not native to the Chesapeake region. Among possible native predators are suckers (Catostomidae); and White Catfish and Bullheads (Amieurus spp.) Fishes of the same families are listed by McMahon (1983) as predators of C. fluminea. Corbicula fluminea is an important food source for the endangered Shortnose Sturgeon Acipenser brevirostris and rare Atlantic Sturgeon A. oxyrhynchus in the Delaware River (Horwitz 1986); and probably in the Chesapeake as well. Five species of Chesapeake Bay ducks, Aix sponsa (Wood Duck), Anas clypeata (Northern Shoveler), Anas acuta (Pintail), Anas platyrhychos (Mallard), and Anas rubripes (American Black Duck) were found to be feeding on C. fluminea during 1973-76 (Perry 1981). Several additional species of diving ducks, known to feed on molluscs, including the Ring-Neck Duck (Athya collariformis), Bufflehead (Bucephala albeola), and Canvasback (Athya vallisnerae), also increased in the freshwater tidal Potomac during the height of the Corbicula invasion (Phelps 1994).
M130	Chesapeake Bay		Economic Impact	Fisheries
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), including increased catches of Micropterus salmoides (Largemouth Bass) by sportsmen (Phelps 1994).
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).
S120	Savannah River		Ecological Impact	Herbivory
Filtration by Corbicula fluminea resulted in significant reductions in suspended organic matter in a Savannah River tributary. The invading clam filtered at much higher rates than the native unionid mussel Elliptio complanata, but the presence of the clam had no effect on the mussel's growth rates (Leff et al. 1990).
P260	Columbia River		Ecological Impact	Herbivory
In reservoirs, Corbicula preferentually filtered diatoms, but in the tidal river, near Portland, they preferred flagellates, but in both environments, they avoided cyanobacteria (Bolam et al. 2019).
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).
OR	Oregon		Ecological Impact	Herbivory
In reservoirs, Corbicula preferentually filtered diatoms, but in the tidal river, near Portland, they preferred flagellates, but in both environments, they avoided cyanobacteria (Bolam et al. 2019).

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DOI: 10.1111/ddi.13666

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