Crassostrea virginica

Invasion History

First Non-native North American Tidal Record: 1869
First Non-native West Coast Tidal Record: 1869
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Crassostrea virginica is native to the northwest Atlantic from the Gulf of St. Lawrence to Panama and Venezuela (Bousfield 1960; Abbot 1974: Morris 1975). It is characteristic of shallow, estuarine habitats, and in the southern part of its range, occurs in the intertidal. Beginning in the 19th century, extensive transplants were made to West Coast waters, from British Columbia to southern California, to Hawaii, and to European waters. The Eastern Oyster is established in Hawaii, and a tiny remnant survives in British Columbia, but the vast majority of these introductions were unsuccessful (Carlton 1979; Carlton and Mann 1996; Ruesink et al. 2005; Carlton and Eldredge 2009). Some hatchery-based aquaculture of C. gigas continues on the West Coast and elsewhere (Conte 1996; Rueness et al. 2005).

North American Invasion History:

Invasion History on the West Coast:

The California Gold Rush and rapid population growth around San Francisco Bay and elsewhere on the West Coast created a demand for oysters, and led to rapid overfishing of the native Olympia Oyster (Ostrea lurida). With the coming of the transcontinental railroad, it was possible to ship live Eastern Oysters across the US for consumption and for planting. The first planting was made in San Francisco Bay in 1869. Oysters were shipped to San Francisco Bay from New York (Long Island Sound) and New Jersey in varying quantities, increasing to a peak of 1 million pounds per year by 1900, and then decreasing. Most were shipped to market, but many were placed in the Bay first for 'freshening' and holding. Sometime between 1900 and 1915, growth rates and the quality of oysters raised in San Francisco Bay began to decrease, and production in California shifted to other bays. Oyster culture began in Tomales Bay in 1875, Humboldt Bay in 1896, and Elkhorn Slough in 1923 (Barrett 1963; Carlton 1979). In the late 19th century, oyster plantings occurred in many West Coast bays, including: Yaquina Bay, Oregon in 1872; Willapa Bay, Washington (WA) in 1874; Grays Harbor, WA in 1900; Puget Sound, WA in the 1870s and 1880s; and the Strait of Georgia, British Columbia in 1883 (Carlton 1979). In California, there were some renewed efforts at planting C. virginica in the 1930s, by the California Department of Fish and Game, including Anaheim Bay, Mugu Lagoon, and Morro Bay, but these were not sustained, and did not result in reproduction (Barrett 1963; Carlton 1979).

Rare spawning and settlement was seen in San Francisco Bay (1880-1890s), Yaquina Bay (1899-1903), Willapa Bay (1914-1940s), and the Strait of Georgia (1893, 1911), but the West Coast fisheries were dependent on seed from the East Coast. By the 1920s-1930s, West Coast oyster production had largely shifted to C. gigas, but C. virginica was still cultured, using hatchery stock, in some California bays (Conte 1996; Coan and Valentich-Scott, in Carlton 2007). A small population of C. virginica persisted in the Nasselle River, Willapa Bay, WA into the 1970s (Carlton 1979), but is now extinct (Carlton 1992). The only established reproducing population of C. virginica on the West Coast is in the Nicomekl River, flowing into Boundary Bay, near Langley, British Columbia, which was initially stocked in 1895 (Quayle 1969; Carlton 1979; Carlton 1992; Gillespie 2007). 

Invasion History in Hawaii:

Crassostrea virginica is established in Pearl Harbor, Oahu. Small plantings of C. virginica were made in 1866, although the currently established population may have resulted from a more extensive stocking (38,000 oysters) in 1895. The population has fluctuated due to pollution and disease, but reefs and settled oysters are still abundant (Coles et al. 1999; Carlton and Eldredge 2009). Smaller stockings in Kaneohe Bay, Oahu (1921-1926) did not result in breeding populations (Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

In France, shipments of C. virginica were made, beginning in 1861, to Le Havre, Ile Vaast-la-Hogue, and Arcachon. A French author wrote that the 'American Oyster is little appreciated in France', and its rearing was abandoned (Bouchon-Brandeley 1877, cited by Carlton and Mann 1996). However, numerous other introductions to European waters were made to replace depleted stocks of the European Oyster (Ostrea edulis). Introductions to English waters began in 1871 and in Denmark in 1880 (Utting and Spencer 1992; Carlton and Mann 1996). Large shipments of Eastern Oysters for planting in Europe continued until the 1930s (Carlton and Mann 1996; Ruesink et al. 2005; Wolff and Reise 2002). Introductions to the Netherlands and Ireland occurred as late as 1939, and there may have been many unreported introductions in European waters (Carlton and Mann 1996).

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Description

Crassostrea virginica is white to lead-gray in color, and its shell shows concentric layers of growth. The shell is rough and heavy, and generally narrow near the hinge (umbo) and widening away from the hinge, but this can vary according to conditions of growth. The lower (right) valve is somewhat cupped. The upper (left) valve is smaller and fatter, and advances forward as the shell grows. The interior of the shell is dull white, but the muscle scar is deep violet. Adult oysters are 150-250 mm in size. Description from: Quayle 1969; Morris 1975; Gosner 1978; Coan et al. 2000; Coan and Valentich-Scott, in Carlton 2007.

Larvae are described and illustrated by Chanley and Andrews (1971). Early veligers are nearly circular, but late larvae of this and other oysters are distinguished by the asymmetrical umbo. They settle at a length of about 300 µm (Chanley and Andrews 1971).

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Taxonomy

Ecology

General:

Crassostrea virginica, like other oysters, is a protandric hermaphrodite, maturing first as a male and then becoming female in subsequent seasons. Females release eggs, and males release sperm, into the water column, where fertilization occurs. The fertilized egg develops first into a ciliated trochophore larva, and then into a shelled veliger larva. The larva feeds on phytoplankton, and grows, eventually developing a foot and becoming a pediveliger, competent for settlement. In laboratory culture, larval settlement occurred at about 11-30 days at 19 to 33⁰C (Shumway 1996). Adult oysters grow up to 250 mm (Abbott 1974; Morris 1975).

Crassostrea virginica is characteristic of estuarine waters in eastern North America. This oyster grows at salinities of 5-42 PSU, and can tolerate brief exposures to salinities as low as 2 PSU (Shumway 1996). In Chesapeake Bay, extensive intertidal beds are common. Successful reproduction occurs at 19-36⁰C, and 12.5 to 35 PSU (Shumway 1996). It tolerates a wide variety of water temperatures, from -1.8 to 36⁰C, while air temperatures up to 42⁰C can be tolerated during low tide exposures (Shumway 1996).

Food:

Phytoplankton

Consumers:

Crabs; Fishes, Gastropods; Parasites; Humans

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Mangroves	None
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	-2	Field range, water temperature (Shumway 1996).
Maximum Temperature (ºC)	36	Field range, water temperature. Air temperatures of 46-49 C can be tolerated during low-tide exposure. (Shumway 1996).
Minimum Salinity (‰)	5	Field range. But 2 ppt can be tolerated for up to a month (Shumway 1996).
Maximum Salinity (‰)	42	Field range (Shumway 1996).
Minimum Reproductive Temperature	18	Thresholds vary greatly among populations. (Shumway 1996).
Maximum Reproductive Temperature	30	Upper limits vary greatly among populations. (Shumway 1996).
Minimum Reproductive Salinity	12.5	Minimum for larval development, experimental (Shumway 1996).
Maximum Reproductive Salinity	35	Maximum for larval development, experimental. (Shumway 1996).
Minimum Duration	10	Larval duration at 30-33 C, experimental (Shumway 1996).
Maximum Duration	30	Larval duration at 19 C, experimental (Shumway 1996).
Minimum Width (mm)	250	Morris 1975
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Successful invasions of Crassostrea virginica have been rare and limited (Pearl Harbor, Hawaii; Nicomekl River, British Columbia), and impacts have not been reported from the oysters themselves. However, oysters and their reefs constitute a major marine habitat, and oyster transfers have constituted one of the major vectors of introductions in the marine environment (Galtsoff 1932; Carlton 1979; Cohen and Carlton 1995; Eno et al. 1997; Reise et al. 1999; Wolff and Reise 2002).

Predator-Parasite Vectors- Eastern Oyster transplants have transferred many organisms from the East Coast of North America to the West Coast, and to the coasts of northern Europe. It is striking that many East Coast natives have become established in both regions. The Atlantic Oyster Drill (Urosalpinx cinerea) an important oyster predator, the Common Atlantic Slipper Shell (Crepidula fornicata) a fouling gastropod, and the infaunal polychaetes Streblospio benedicti and Clymenella torquata are all introduced both on the West Coast and in Europe, and now create problems for the culture of introduced Pacific Oysters (C. gigas), as well as affecting native species (Quayle 1969; Carlton 1979; Cohen and Carlton 1995; Eno et al. 1997; Reise et al. 1999; Wolff and Reise 2002).

The introduction of C. virginica introduced the parasite Perkinsus marinus, the causative agent of 'Dermo' disease to Hawaii. However, this parasite has not been reported to affect native fauna (Kern et al. 1973; Carlton and Eldredge 2009). Transfers of Eastern Oysters within their native range, from Chesapeake Bay and more southern regions to northern estuaries have extended the range of P. marinus northward along the coast to Maine (Ford 1996; Reece et al. 2001). Another inadvertent transfer, within the native range occurred, when Haplosporidium nelsoni (MSX parasite), caused massive mortalities of Eastern Oysters in Chesapeake Bay. Subsequent oyster transfers from the Gulf of Mexico are believed to have introduced the parasitic barnacle Loxothylacus panopei, which infects mud crabs (Eurypanopeus depressus; Rhithropanopeus harrisii), to Chesapeake Bay (Van Engel et al. 1966; Hines et al. 1997).

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Regional Impacts

NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in central California waters, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes [Alitta (=Neanthes, Nereis), Streblospio benedicti], slippershells (Crepidula convexa, C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Cohen and Carlton 1995; Wasson et al. 2001; de Rivera et al. 2005).
P090	San Francisco Bay		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in West Coast o Bay waters, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes [Alitta (=Neanthes succinea, Nereis), Streblospio benedicti], slippershells (Crepidula convexa, C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), tube-dwelling amphipods (Ampelisca, Ampithoe, Monocorophium spp.) and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Cohen and Carlton 1995).
P080	Monterey Bay		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in Elkhorn Slough, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes (Streblospio benedicti), Atlantic Oyster Drill (Urosalpinx cinerea), tube-dwelling amphipods (Ampelisca, Ampithoe, Monocorophium spp.) and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Wasson et al. 2001; de Rivera et al. 2005).
P110	Tomales Bay		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in central California waters, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes [Alitta (=Neanthes, Nereis), Streblospio benedicti], Atlantic Oyster Drill (Urosalpinx cinerea), tube-dwelling amphipods (Ampelisca, Ampithoe, Monocorophium spp.) and tunicates (Brtryllus schlosseri, Molgula manhattensis) (Carlton 1979; de Rivera et al. 2005).
NEP-IV	Puget Sound to Northern California		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established on the Pacific Northwest outer coast, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes [Alitta (=Neanthes, Nereis), Streblospio benedicti], slippershells (Crepidula convexa, C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Boyd et al. 2002; Cohen et al. 2001; Wonham and Carlton 2005).
P130	Humboldt Bay		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in Humboldt Bay, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes (Streblospio benedicti), slippershells (Crepidula convexa, C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Boyd et al. 2002; Wonham and Carlton 2005).
CA	California		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in central California waters, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes [Alitta (=Neanthes, Nereis), Streblospio benedicti], slippershells (Crepidula convexa, C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Cohen and Carlton 1995; Wasson et al. 2001; de Rivera et al. 2005)., While C. virginica has not become established in San Francisco Bay waters, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes [Alitta (=Neanthes succinea, Nereis), Streblospio benedicti], slippershells (Crepidula convexa, C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), tube-dwelling amphipods (Ampelisca, Ampithoe, Monocorophium spp.) and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Cohen and Carlton 1995)., While C. virginica has not become established in Elkhorn Slough, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes (Streblospio benedicti), Atlantic Oyster Drill (Urosalpinx cinerea), tube-dwelling amphipods (Ampelisca, Ampithoe, Monocorophium spp.) and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Wasson et al. 2001; de Rivera et al. 2005)., While C. virginica has not become established in central California waters, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes [Alitta (=Neanthes, Nereis), Streblospio benedicti], Atlantic Oyster Drill (Urosalpinx cinerea), tube-dwelling amphipods (Ampelisca, Ampithoe, Monocorophium spp.) and tunicates (Brtryllus schlosseri, Molgula manhattensis) (Carlton 1979; de Rivera et al. 2005)., While C. virginica has not become established in Humboldt Bay, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Cliona sp. (boring sponges) and other sponges, polychaetes (Streblospio benedicti), slippershells (Crepidula convexa, C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Boyd et al. 2002; Wonham and Carlton 2005).

References

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

Bancila. Raluca I.; Skolka, Marius; Ivanova, Petya; Surugiu, Victor; Stefanova, Kremena; Todorova. Valentina; Zenetos, Argyro (2022) Alien species of the Romanian and Bulgarian Black Sea coast: state of knowledge, uncertainties, and needs for future research, Aquatic Invasions 17: Published online

Barrett, Elinore M. (1963) The California Oyster Industry., California Department of Fish and Game Fish Bulletin 123: 1-103

Bousfield, E. L. (1960) Canadian Atlantic Sea Shells, In: (Eds.) . , Ottawa. Pp. <missing location>

Boyd, Milton J.; Mulligan, Tim J; Shaughnessy, Frank J. (2002) <missing title>, California Department of Fish and Game, Sacramento. Pp. 1-118

California Department of Fish and Wildlife (2014) Introduced Aquatic Species in California Bays and Harbors, 2011 Survey, California Department of Fish and Wildlife, Sacramento CA. Pp. 1-36

California Department of Health Services (2007) <missing title>, California Department of Health Services, Sacramento CA. Pp. <missing location>

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>

Carlton, James T. (1992) Introduced marine and estuarine mollusks of North America: An end-of-the-20th-century perspective., Journal of Shellfish Research 11(2): 489-505

Carlton, James T. (Ed.) (2007) The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon Fourth Edition, Completely Revised and Expanded, University of California Press, Berkeley. Pp. <missing location>

Carlton, James T.; Eldredge, Lucius (2009) Marine bioinvasions of Hawaii: The introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian archipelago., Bishop Museum Bulletin in Cultural and Environmental Studies 4: 1-202

Carlton, James T.; Mann, Roger (1996) Transfers and world-wide introductions., In: Kennedy, V. S., Newell, R. I. E., Eble, A. F.(Eds.) The Eastern Oyster. , College Park. Pp. 691-706

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

Chanley, Paul; Andrews, J. D. (1971) Aids for identification of bivalve larvae of Virginia, Malacologia 11(1): 45-119

Coan, Eugene V.; Valentich-Scott, Paul (2007) The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon, University of California Press, Berkeley CA. Pp. 807-859

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>

Coen, Loren D.; Bishop, Melanie J. (2015) The ecology, evolution, impacts and management of host-parasite interactions of marine molluscs, Journal of Invertebrate Pathology 131: 177-211

Cohen, Andrew N. and 22 authors (2001) <missing title>, Washington State Department of Natural Resources, Olympia. 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, Andrew; and 16 authors. (1998) <missing title>, Washington State Department of Natural Resources, Olympia, Washington. Pp. 1-37

Coles, S. L.; DeFelice, R. C.; Eldredge, L. G.; Carlton, J. T. (1999b) Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands., Marine Biology 135(1): 147-158

Conte, Fred S. (1996) California oyster culture., California Aquaculture A-7: 1-7

Cronin, L. Eugene (1986) Chesapeake fisheries and resource stress in the 19th century, Journal of the Washington Academy of Sciences 76(3): 188-198

de Rivera, Catherine, and 27 authors (2005) Broad-scale non-indigenous species monitoring along the West Coast in National Marine Sanctuaries and National Estuarine Research Reserves report to National Fish and Wildlife Foundation, National Fish and Wildlife Foundation, Washington, D.C.. Pp. <missing location>

do Amaral, Vanessa Simao; Simone, Luiz Ricardo L. (2014) Revision of genus Crassostrea (Bivalvia: Ostreidae) of Brazil, Journal of the Marine Biological Association of the United Kingdom 94(4): 811-836

Eldredge, L. G.; Smith, C. M. (2001) Introduced marine species of Hawaii, Bishop Museum Technical Report 21: 1-60

Eldredge, L.G. (1994) Perspectives in aquatic exotic species management in the Pacific Islands Vol. I. Introductions of commercially significant aquatic organisms to the Pacific islands, South Pacific Commission. Inshore Fisheries Research Project, Technical Document 7: 1-127

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

Ewers, Christine; Normant-Saremba, Monika; Keirsebelik, Heleen; Schoelynck, Jonas (2023) The temporal abundance-distribution relationship in a global invader sheds light on species distribution mechanisms, Aquatic Invasions 18(2): 179-197
179–197. https:// doi.org/10.3391/ai.2023.18.2.105548

Ford, Susan E. (1985) Chronic infections of Haplosporidium nelsoni (MSX) in the oyster Crassostrea virginica, Journal of Invertebrate Pathology 45: 94-107

Ford, Susan E. (1996) Range extension by the oyster parasite Perkinsus marinus into the northeastern United States: Response to climate change?, Journal of Shellfish Research 15(1): 45-46

Galtsoff, Paul S. (1932) Introduction of Japanese Oysters into the United States, United States Department of Commerce, Bureau of Fisheries, Fisheries Circular 12: 1-16

Gillespie, Graham E. (2007) Distribution of non-indigenous intertidal species on the Pacific Coast of Canada, Nippon Suisan Gakkaishi 73(6): 1133-1137

Gould, Augustus A. (1870) <missing title>, Wright and Potter, State Printers, Boston. Pp. <missing location>

Hines, Anson H.; Alvarez, Fernando; Reed, Sherry A. (1997) Introduced and native populations of a marine parasitic castrator: Variation in prevalence of the rhizocephalan Loxothylacus panopaei in xanthid crabs, Bulletin of Marine Science 61(2): 197-214

Ignacio, B. L; Absher, A. M.; Lazoski, C.; Sole-Cava, A. M. (2000) Genetic evidence of the presence of two species of Crassostrea on the coast of Brazil, Marine Biology 136: 987-991

Kennedy, Victor S. (1995) Ecological role of the eastern oyster, Crassostrea virginica, with remarks on disease., Journal of Shellfish Research 15(1): 177-183

Kern, Frederick G.; Sullivan, L. Cecelia; Takata, Michio (1973) Labyrinthomyxa-like organisms associated with mass mortalities of oysters, Crassostrea virginica, from Hawaii, Proceedings of the National Shellfisheries Association 63: 43-46

Lipton, Douglas; Lavan, Eileen F.; Strand, Ivar E. (1992) Economics of molluscan introductions and transfers: the Chesapeake Bay dilemma, Journal of Shellfish Research 11(2): 511-519

Lohan, Katrina M. Pagenkopp; Hill-Spanik, Kristina M.; Torchin, Mark E.; Aguirre-Macedo, Leopoldina ; Fleischer, Robert C.; Ruiz, Gregory M. (2016) Richness and distribution of tropical oyster parasites in two oceans, Parasitology Published online: <missing location>

Lohan, Katrina M. Pagenkopp; Hill-Spanik, Kristina M.; Torchin, Mark E; Strong, Ellen E.; Fleischer, Robert C.; Ruiz, Gregory M. (2015) Molecular phylogenetics reveals first record and invasion of Saccostrea species in the Caribbean, Marine Biology 162(5): 957-968

Mach, Megan E.; Levings, Colin D.; McDonald, P. Sean; Chan, Kai M. A. (2012) An Atlantic infaunal engineer is established in the Northeast Pacific: Clymenella torquata (Polychaeta: Maldanidae) on the British Columbia and Washington Coasts, Biological Invasions 14: 503-507

Miller, A. Whitman; Ruiz, Gregory M.; Minton, Mark S.; Ambrose, Richard F. (2007) Differentiating successful and failed molluscan invaders in estuarine ecosystems., Marine Ecology Progress Series 332: 41-51

Morris, Percy A. (1975) A field guide to shells of the Atlantic, Houghton-Mifflin, Boston. Pp. <missing location>

Paglia, Ken 4/22/21 Invasive Seaweed Found in Newport Bay. <missing URL>



Quayle, D. B. (1969) Pacific oyster culture in British Columbia, Canadian Fisheries Research Board Bulletin 169: 1-192

Reece, K. S., Bushek, D., Hudson K. L. & Graves, J. E. (2001) Geographic distribution of Perkinsus marinus genetic strains along the Atlantic and Gulf coasts of the USA., Marine Biology 139: 1047-1055

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

Ruesink, Jennifer (2011) Encyclopedia of Biological Invasions, University of California Press, Berkeley and Los Angeles. Pp. 494-499

Ruesink, Jennifer L. and 6 authors (2005) Introduction of non-native oysters: Ecosystem effects and restoration implications., Annual Review of Ecology and Systematics 36: 643-689

Shebley, W. H. (1917) Introduction of food and game fishes into the waters of California., California Fish and Game 3(1): 1-12

Shumway, Sandra E. (1996) The Eastern Oyster, <missing publisher>, <missing place>. Pp. <missing location>

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

Smith, Hugh M. (1895) A review of the history and results of the attempts to acclimatize fish and other water animals in the Pacific states., Bulletin of the U. S. Fish Commission 15: 379-472

Utting, S. D.; Spencer, B. E. (1992) Introductions of marine Bivalvia Mollusca into the United Kingdom, International Council for Exploration of the Marine Science Symposium 194: 84-91

Van Engel, W. A.; Dillon, William A.; Zwerner, David; Eldridge, Dana (1966) Loxothylacus panopaei (Cirripedia, Sacculinidae) an introduced parasite on a xanthid crab in Chesapeake Bay, U.S.A., Crustaceana 10: 110-112

Wasson, Kerstin; Zabin, C. J.; Bedinger, L.; Diaz, M. C.; Pearse J. S. (2001) Biological invasions of estuaries without international shipping: the importance of intraregional transport, Biological Conservation 102: 143-153

Wells, Harry W. (1961) The fauna of oyster beds, with special reference to the salinity factor, Ecological Monographs 31: 239-266

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

Wolff, Wim; Reise, Karsten (2002) Invasive aquatic species of Europe: Distribution, impacts and management., Kluwer Academic Publishers, Dordrecht, Boston, London.. Pp. 193-205

Wonham, Marjorie J.; Carlton, James T. (2005) Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system, Biological Invasions 7: 369-392

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