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).
P210	Yaquina Bay		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in Yaquina 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), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979).
P270	Willapa Bay		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in Willapa Bay, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including sponges, polychaetes (Streblospio benedicti), slippershells (C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Cohen et al. 2001; Wonham and Carlton 2005).
NEP-III	Alaskan panhandle to N. of Puget Sound		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in British Columbia and Puget sound waters, exept for a tiny population in boundary 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 [Alitta (=Neanthes, Nereis), Clymenellla torquata, Streblospio benedicti], slippershells (Crepidula convexa, C. plana, C. fornicata), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Quayle 1969; Carlton 1979; Cohen et al. 2001; de Rivera et al. 2005; Gillespie et al. 2007).
NEA-II	None		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in northern European waters, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including Streblospio benedicti, slippershells (Crepidula fornicata), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Molgula manhattensis) (Eno et al. 1997; Reise et al. 1999; Wolff and Reise 2002).
SP-XXI	None		Ecological Impact	Parasite/Predator Vector
Crassostrea virginica, stocked in Pearl Harbor, apparently carried the protistan parasite Perkinsus marinus, which caused 'Dermo' disease and massive mortalities in Eastern Oysters in the harbor in 1972. However, there are no reports of this parasite affecting other invertebrates at that time (Kern et al. 1973; Carlton and Eldredge 2009).
WA	Washington		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in Willapa Bay, Eastern Oyster introductions have been a probable/possible vector for many fouling organisms and predators including sponges, polychaetes (Streblospio benedicti), slippershells (C. plana), Atlantic Oyster Drill (Urosalpinx cinerea), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979; Cohen et al. 2001; Wonham and Carlton 2005).
OR	Oregon		Ecological Impact	Parasite/Predator Vector
While C. virginica has not become established in Yaquina 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), and tunicates (Botryllus schlosseri, Molgula manhattensis) (Carlton 1979).
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).
HI	Hawaii		Ecological Impact	Parasite/Predator Vector
Crassostrea virginica, stocked in Pearl Harbor, apparently carried the protistan parasite Perkinsus marinus, which caused 'Dermo' disease and massive mortalities in Eastern Oysters in the harbor in 1972. However, there are no reports of this parasite affecting other invertebrates at that time (Kern et al. 1973; Carlton and Eldredge 2009).

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