Corbicula Fluminea

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Phylum: Mollusca Corbicu la fluminea Class: Bivalvia, Heterodonta Order: Veneroida

The Asian clam Family: Corbiculidae

Taxonomy: Corbicula fluminea was originally General Morphology: Bivalve mollusks are described as one of three species in the bilaterally symmetrical with two lateral valves genus Tellina (Araujo et al. 1993). The or shells that are hinged dorsally and taxonomy of this group was based on shell surround a mantle, head, foot and viscera morphology and color, which are variable (see Plate 393B, Coan and Valentich-Scott characters (Sousa et al. 2008a), and has lead 2007). The Veneroida is a large and diverse to the synonymization of several previously bivalve heterodont order that is characterized described species that were found to display by well developed hinge teeth. There are 22 intraspecific variation (Araujo et al. et al. local families, and members of the 1993). Corbicula fluminea was synonymized Corbiculidae have an elongated hinge and with C. fluviatilis by Prashad in 1929, C. serrate teeth laterally, C. fluminea is common manilensis in 1933 (Talvera and Faustino in in freshwater habitats and has a conspicuous Britton and Morton 1979) and C. leana by triangular shape with low commarginal ridges Morton (1977). This synonymization lead to (see Plate 398B, 415E, Coan and Valentich- two Corbicula species with widespread Scott 2007). distribution: C. fluminea (a freshwater Body: species) and C. fluminalis (an estuarine Color: species) (Araujo et al. et al. 1993 but see Interior: The ligament is thick and Sousa et al. 2008a). These species can be strong, and is entirely external and posterior further distinguished by geographic location to the beaks (Fig. 2). and characters of biology and morphology, as Exterior: all North American populations were Byssus: A small byssus is present determined to be only C. fluminea (see Britton only in first year (Morton 1979a) (not figured). and Morton 1979; Morton 1982; Araujo et al. Gills: et al. 1993). Shell: The overrall shell shape is triangular (Coan and Valentich-Scott 2007) to Description subcircular. Both valves are similar and Size: Individuals range in size from less than regular, and the shell is thick. 2.5– 6.4 cm in length, and are rarely over 5.0 Interior: The adductor muscle scars cm (Ward and Whipple 1963). Individuals are approximately equal (Coan and Valentich- over 40 mm in length are often found in canal Scott 2007) (Fig. 3). The pallial line is bottoms (Eng 1979). The illustrated incomplete anteriorly (Corbiculidae, Britton specimen (from the Columbia River) is 14 mm and Morton 1979). in length. Mean wet weight for C. fluminea Exterior: The exterior sculpture has from the Potomac River were estimated for heavy concentric ridges and faint growth rings year classes 1, 2, 3, and 4 to be 0.48, 1.47, (Britton and Morton 1979) (Fig. 1). 4.30, and 10.37 grams, respectively (Cohen Hinge: The hinge is very elongate, et al. 1984). with anterior and posterior serrate lateral Color: Tan exterior; the shell interior is white, teeth (Corbiculidae, Burch 1975) (Fig. 3). smooth, polished, and is sometimes with There are three hinge teeth on each valve, purple markings (Britton and Morton 1979). A which are "divergent, pseudocardinals" periostracum is present and thick (Coan and (Clarke1981) (Fig. 3). A long row of serrate Valentich-Scott 2007), particularly in canal hinge teeth are present both anteriorly and specimens; specimens from earth-lined posteriorly (Fig. 3) (Corbiculidae, Burch habitats may lack periostracum (Eng 1979). 1975). (Note: Ward and Whipple 1963 lists

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12744 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] two teeth in each valve, which is a possible the basis of electrophoresis, that specimens error.) The beaks are subcentral (Ward and from 5 major U.S. populations (California, Whipple 1963), heavy, and inflated (Britton Texas, Arkansas, Tennessee and South and Morton 1979). The umbones are often Carolina) are all of the same species. There eroded, especially in acidic waters (Britton is, however, a great deal of variation in shell and Morton 1979). shape, sculpture, etc. Although C. fluminea Eyes: has been synonymized with C. manilensis, Foot: The foot is rather large (Clarke 1981) there are still a few questions about its (not figured). reproductive habits and longevity (Morton Siphons: Siphons are of similar size and 1979a). Corbicula fluminalis, may be larger shape (Britton and Morton 1979) (not than C. fluminea, could live longer (to 8 figured). years), and seems to occupy river mouths Burrow: rather than streams (Britton and Morton 1979) (see taxonomy). Possible Misidentifications The Corbiculidae, of which Corbicula Ecological Information is the only North American genus (Burch Range: Type locality is the Artemovka River 1975), are noted for a heavy shell, strong in Russia (Kantor et al. 2009) and is endemic concentric sculpture, a hinge ligament that is to southeast Asia (Morton 1979b). Corbicula internal, and with a peg-shaped chondrophore fluminea is a widespread species with a on the left valve (Coan and Valentich-Scott history of invasion that is well documented; it 2007). Corbicula fluminea is characterized by is a non-native invasive species and, where a thick, triangular shell with commarginal present, leads to considerable ecological and ridges and dark brown-black periostracum. economic impacts in many aquatic systems There are three other freshwater bivalve (e.g., hydrology, biogeochemistry, families in the Pacific Northwest. The biodiversity, Sousa et al. 2008a; Sousa et al. Unionidae are represented by two genera: 2008b; Crespo et al. 2015 and references the monotypic Gonidea (angulata) has a therein). It has been transported (e.g., via smooth but irregular elongate subtriangular ballast water, as bait, etc.; Sousa et al. shell, with a distinctive high sharp ridge. 2008a) to North America in (first half of 20th Anodonta, the second genus, has a thin century) and South America in the 1960–70s, smooth elliptical inflated shell, sometimes Europe in the 1980s (see Fig. 1, Crespo et al. winged at the posterior end. It has fine 2015). This species was introduced to North parallel ridges (not deep heavy ones as in America from southeast Asia in the 1930s Corbiculidae) and its hinges lack teeth. The and is now present in all major U.S. drainages third family, the Sphaeriidae, belongs to the (in the Pacific and Atlantic) below 40° latitude same superfamily (Sphaeriacea) as the (Britton and Morton 1979; McMahon 1982). Corbiculidae (Burch 1975). There are several Its range in the Pacific Northwest includes genera, including Sphaerium, Musculium, and Washington, Idaho, Oregon to northern Pisidium. Sphaeridae can have regular California and it also occurs in the Imperial valves, subcentral beaks and radial sculpture Valley, California, and in Arizona. There were as do Corbiculidae. The lateral hinge teeth in likely two “epicenters” of human mediated Sphaeriidae are smooth, however, not serrate introductions in North America; the first to the as in Corbicula. northwest from southeast Asia and the A great number of species of second a population in the Ohio River (see Corbicula have been named worldwide, many Fig. 1, McMahon 1982). On the eastern coast of them superfluous. The taxonomy of the of North America, C. fluminea was first genus remains unclear, and there is still observed in Potomac River in 1977 (Cohen et uncertainty as to how many species have al. 1984; Phelps 1994). Outside of North been introduced into North America from Asia America, established populations were and elsewhere (Britton and Morton 1979). It is reported in the Negro River Basin in possible that only C. fluminea exists here: Patagonia, Argentina, which is the M.H. Smith et al. 1979 have suggested, on southernmost reach if the species distribution

Hiebert, T.C. 2015. Corbicula fluminea. In: Oregon Estuarine Invertebrates: Rudys' Illustrated Guide to Common Species, 3rd ed. T.C. Hiebert, B.A. Butler and A.L. Shanks (eds.). University of Oregon Libraries and Oregon Institute of Marine Biology, Charleston, OR. to date worldwide (Molina et al. 2015) and the Rosa et al. 2015; caffeine, Aguirre-Martinez et Iberian Peninsula (Spain and Portugal, Araujo al. 2015; Oliveira et al. 2015; metals Bonnail et al. 1993). Potentially high genetic diversity et al. 2016; Oliveria et al. 2016). exists within populations, which is observed Salinity: Considered a freshwater species, molecularly (e.g., Hongze Lake, China, Li et but can survive a gradual increase of salinity, al. 1994) and morphologically (a dark morph i.e., up to salinity of 22 for 80 days. Salinity observed in southwestern United States, range reported for C. fluminea is 10–14 Nichols and Black 1994) (Sousa et al. 2008a). (Crespo et al. 2015). In shock tests, however, Local Distribution: Local distribution mortality was 50% within 10 days of salinity includes the Columbia River system (from over 10, and individual sodium uptake was which the illustrated specimen originated), measured 50% sodium at salinity of 4.5 Siuslaw River at Florence, and possibly in the (Evans et al. 1979). The closely related Umpqua River (Carlton 1979). species C. fluminalis, is differentiated from C. Habitat: An opportunist, C. fluminea can live fluminea by some authors as an estuarine, in quiet or fast moving water, in streams, where C. fluminea is exclusively freshwater rivers, canals, lakes and reservoirs. It can (Araujo et al. et al. 1993 but see Sousa et al. utilize either an "r" or a "k" reproductive 2008a). strategy, and its only limiting factor seems to Temperature: Individuals cannot survive be space (Britton and Morton1979). It has cold temperatures (i.e., one severe winter can expanded geographically very successfully in kill a population, Horning and Keup 1964). 50 years, especially into irrigation canals in Short warm water periods may be a limiting northern California, where it is considered a factor in the northern range (Eng 1979). The pest. In canals, it finds a nursery in the distribution of this species seems to be limited incrustation formed of corophiid amphipod by low temperatures (Crespo et al. 2015; tubes and colonial hydroids on the concrete Gama et al. 2016), as individuals are walls. Locally, the densely packed individuals negatively effected by high temperatures accumulate as a solid layer (sometimes up to (temperature range 2–34.8˚C, Crespo et al. 92 cm thick) composed of clams, fine silt, and 2015). Growth occurs only at temperatures mucus. Individuals become so dominant that over 14°C, and spawning takes place only at they have the capacity to change their 13–16 °C. (California, Eng 1979). environment radically. In the Columbia River Tidal Level: Usually a shallow water or near- system, they live in stream bottoms. shore species. Prolonged exposure to air can cause death by Associates: Individuals become very accumulation of metabolic wastes (rather than dominant and outcompete native bivalves desiccation as the clam shell can close within whose communities it can live (Morton tightly). Mean survival is 26.8 days at 20 °C, 1979a; Crespo et al. 2015). In canals, the at high relative humidity (r.h.); 13.9 days at 20 amphipod Corophium spinicorne provides °C., at low r.h.; 8.3 days at 30 °C., at high r.h.; tubes that harbor recently settled clams; the 6.7 days at 30 °C., at low r.h (McMahon colonial hydrozoan Cordylophora lacustris 1979). Mortality rate increases with increased also provides habitat (Eng 1979). The turbidity (e.g., 150 nephelometric turbidity oligochaete, Chaetogaster limnaei, can units, Avelar et al. 2015). Bioturbation from sometimes be found within the mantle cavity C. fluminea increases soluble and labile of C. fluminea (Eng 1976). phosphorus within sediment (up to 64 mm Abundance: Often more than 1,000 depths, Chen et al. 2016). Individuals prefer individuals per square meter (McMahon 2000; oxygenated, sandy sediment (Crespo et al. Majdi et al. 2014). Maximum densities in fall, 2015). Corbicula fluminea is a common winter, summer were up to 4,500 clams per bioindicator (e.g., lead contamination, Clarke ~1 square meter in the Delta-Mendota Canal, et al 1979; potassium, Daum et al. 1979; California (Eng 1979); mean density range Sousa et al. 2008a) and experiment species was 173–2,990 clams per square meter in a (e.g., Baudrimont et al. 1997; Basack et al. river near Charlottesville, Virginia (Hornback 1998; Cataldo et al. 2001; Miller et al. 2005; 1992); and up to 525 individuals per square Vale et al. 2014; Erdogan and Erdogan 2015;

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12744 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] meter in the Negro River Basin (Molina et al. that have settled to the benthos and exhibit 2015). secondary settlement by riding currents (i.e., not true larvae) (Brink 2001). Larvae are Life-History Information released (as juveniles or late pediveligers, Reproduction: The reproductive organs are Kraemer and Galloway 1986) into the composed of an anterior testis and posterior plankton, and this species is the only ovary, which both discharge into a common freshwater bivalve that does this (Eng 1979; gonoduct (Britton and Morton 1979). (For Crespo et al. 2015). Both primary and oocyte and sperm morphology see, Fig. 4 secondary growth lines give larval shell a Kraemer and Galloway 1986.) Early “pleated appearance” (Nichols and Black reproduction and high fecundity contribute to 1994). (For larval identification key for C. the success of this species (see Range). fluminea, Driessena polymorpha, and D. Both a protogynous and a protandric rosteriformis bugensis see Nichols and Black consecutive hermaphrodite (Britton and 1994). Morton 1979), it can also function as a Juvenile: Like other bivalve species (e.g. simultaneous hermaphrodite (Crespo et al. Mytilus edulis, Macoma balthica), post-larval 2015), making it highly successful (i.e., juvenile) C. fluminea may exploit two reproductively (Sousa et al. 2008a). dispersal periods. The initial larval dispersal Individuals are also capable of androgenetic and settlement into the benthos, is followed self-fertilization (Crespo et al. 2015). by potential dispersal from the benthos by Corbicula fluminea incubates its young for thread drifting on water currents via mucous about one month in the inner demibranch of threads by juveniles (i.e., “thread drifting”, each ctenidium (Morton 1979a) i.e., the Martel and Chia 1991). Clams produce these branchial water tubes (see Fig. 1, Sousa et al. mucous threads (see Fig. 2, Prezant and 2008a). There are two reproductive peaks in Chalermwat 1984) in response to water California: April to May and August to current stimuli via ctenidial mucocytes September and fall and spring peaks were (Prezant and Chalermwat 1984). Juvenile reported in Arkansas, the former lasting settlement flexibility greatly aids in dispersal longer than the latter in Arkansas (Kraemer potential. and Galloway 1986). (In Kentucky, a cooler Longevity: Two to 4 years (Britton and climate, there was only but one peak per Morton 1979) or one to 5 years (Sousa et al. year.) Cleavage is via a coeloblastula that is 2008a), with large animals in canal bottoms 175 µm in diameter at 24 hours post living longest. Two to 3 years were estimated fertilization and development proceeds with for individuals collected near Charlottesville, cone-shaped gastrulae by 30 hr (175–180 Virginia (Hornback 1992). µm); trochophore larvae (180 µm in length); Growth Rate: Rapid growth is one of the veliger larvae at 24–48 hrs (190–250 µm). successful strategies of C. fluminea (Sousa et Larva: Larvae are brooded until they are 210 al. 2008a). Clams primarily grow from March µm and released as spat with very straight to October, when temperatures are over umbo (see Fig. 4, Brink 2001) (see 14°C. (Eng 1979). In central California, Reproduction). Pediveligers develop after outside canals, clams grow to a mean of 12 3–5 days (230 µm) at which point the velum is mm the first season, and 15 mm the second shed metamorphosis occurs into a straight- (Heinsohn 1958), with two growth rings hinge juvenile (see Fig. 11 and for full formed each year (Britton and Morton 1979). description of development see Kraemer and Growth varies with depth, with specimens at 8 Galloway 1986 and for list of life-history m deep are longer than those at 12 m (Abbott characters, see Table 1, Sousa et al. 2008a). 1979). Growth of individuals collected in a Larvae have a sloping posterior shoulder with river near Charlottesville, Virginia was 11.1 rounded posterior end that is slightly shorter mm in the first year, 17.6 mm in the second than the anterior end. “Larvae” collected in year (Hornback 1992). plankton are likely thread drifting (see Food: A non-selective filter feeder Macoma balthica, Nutricola tantilla, Hiatella (Boltovskoy et al. 1995) that also pedal feeds. arctica, descriptions in this guide) juveniles Diet consists primarily of diatoms (Hanna

Hiebert, T.C. 2015. Corbicula fluminea. In: Oregon Estuarine Invertebrates: Rudys' Illustrated Guide to Common Species, 3rd ed. T.C. Hiebert, B.A. Butler and A.L. Shanks (eds.). University of Oregon Libraries and Oregon Institute of Marine Biology, Charleston, OR. 1966) and possibly flagellates (i.e., fluminea (Muller, 1774) (Bivalvia: phytoplankton, Foe and Knight 1985). Corbiculidae) exposed to different Corbicula fluminea removes chlorophyll a turbidity conditions. Brazilian Journal from river water with filtration rates that of Biology. 74:509-514. ranged from 13.8–33.9 ml per hour per gram 5. BASACK, S. B., M. L. ONETO, J. S. (wet weight) clam (Cohen et al. 1984). FUCHS, E. J. WOOD, and E. M. Filtration rates measured in three riverine KESTEN. 1998. Esterases of habitats (Tombigbee and Tangipahoa, Corbicula fluminea as biomarkers of Mississippi, and Ouachita, Louisiana) ranged exposure to organophosphorus from 4–145 ml per hour per gram clam (Way pesticides. Bulletin of Environmental et al. 1990). Increased abundance of C. Contamination and Toxicology. fluminea reduced abundance of benthic 61:569-576. bacteria and flagellates (Hakenkamp et al. 6. BAUDRIMONT, M., J. METIVAUD, R. 2001; Majdi et al. 2014). However, large MAURYBRACHET, F. RIBEYRE, and populations do not necessarily effect food A. BOUDOU. 1997. Bioaccumulation consumption by competition (Eng 1979). and metallothionein response in the Predators: Humans use C. fluminea for fish Asiatic clam (Corbicula fluminea) after bait, and occasionally for food. Other experimental exposure to cadmium predators include fish (Robinson and and inorganic mercury. Environmental Wellborn 1988) and waterfowl (Sinclair and Toxicology and Chemistry. 16:2096- Isom 1963). 2105. Behavior: A burrower and pest in dredged 7. BOLTOVSKOY, D., I. IZAGUIRRE, sands that are used to make concrete and N. CORREA. 1995. Feeding (Morton 1979a). selectivity of Corbicula fluminea (Bivalvia) on natural phytoplankton. Bibliography Hydrobiologia. 312:171-182. 8. BONNAIL, E., A. M. SARMIENTO, T. 1. ABBOTT, T. M. 1979. Asiatic clam A. DELVALLS, J. M. NIETO, and I. (Corbicula fluminea) vertical RIBA. 2016. Assessment of metal distributions in Dale Hollow Reservoir, contamination, bioavailability, toxicity Tennessee, p. 111-118. In: and bioaccumulation in extreme Proceedings, First International metallic environments (Iberian Pyrite Corbicula Symposium. J. D. Britton Belt) using Corbicula fluminea. (ed.). The Texas Christian University Science of the Total Environment. Research Foundation, Fort Worth, 544:1031-1044. Texas. 9. BRINK, L. A. 2001. Mollusca: Bivalvia, 2. AGUIRRE-MARTINEZ, G. V., A. T. p. 129-149. In: Identification guide to DELVALLS, and M. LAURA MARTIN- larval marine invertebrates of the DIAZ. 2015. Yes, caffeine, ibuprofen, Pacific Northwest. A. Shanks (ed.). carbamazepine, novobiocin and Oregon State University Press, tamoxifen have an effect on Corbicula Corvallis, OR. fluminea (Muller, 1774). Ecotoxicology 10. BRITTON, J. C., and B. MORTON. and Environmental Safety. 120:142- 1979. Corbicula in North America: the 154. evidence reviewed and evaluated, p. 3. ARAUJO, R., D. MORENO, and M. A. 249-287. In: Proceeings, First RAMOS. 1993. The Asiatic clam International Corbicula Symposium. J. Corbicula fluminea (Muller, 1774) C. Britton (ed.). Texas Christian (Bivalvia, Corbiculidae) in Europe. University Research Foundation, Fort American Malacological Bulletin. Worth, Texas. 10:39-49. 11. BURCH, J. B. 1975. Freshwater 4. AVELAR, W. E. P., F. F. NEVES, and sphaeriacean clams (Mollusca, M. A. S. LAVRADOR. 2014. Modelling Pelecypoda) of North America. the risk of mortality of Corbicula

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