Magallana gigas

Invasion History

First Non-native North American Tidal Record: 1902
First Non-native West Coast Tidal Record: 1902
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Magallana angulata gigas is native to the northwest Pacific including Russia, China, and Korea. Its native range may extend south and west into the Philippines and Indonesia, to Borneo and Sumatra, and west to Pakistan (Carriker and Gaffney 1996). However, the presence of several closely related species and the morphological variation of M. gigas make the boundaries of its range difficult to assess. It is the most widely transplanted shellfish in the world, introduced to at least 52 countries (Food and Agricultural Organization 1998; Ruesink et al. 2005). It is now the world's most widely cultivated oyster. It has established breeding populations in the northeast Pacific (US-Canada), southwest Pacific (Australia-New Zealand), northeast Atlantic-Mediterranean (Europe), southwest Atlantic (Argentina-Brazil), and Indian Ocean (South Africa). It is also successfully cultured using hatcheries and imported spat in many places where conditions are unsuitable for breeding, and has been introduced unsuccessfully to many regions (Food and Agricultural Organization 1998; Ruesink et al. 2005). A repeated pattern in different regions has been for M. gigas to go from being largely confined to culture areas, with only sporadic and limited reproduction, to becoming a major biomass component and ecosystem engineer. This process, which has taken 3-10 decades, has occurred in British Columbia and Washington State (Quayle 1969; Klinger et al. 2006; Kelly et al. 2008; Padilla 2010), the North Sea in Europe (Diederich 2005; Beukema and Dekker 2011), the Atlantic coast of Patagonia (Escapa 2004), Hawaii (Carlton and Eldredge 2009), and Australia (Krassoi et al. 2008). The transition from cultured hatchery-dependent populations, to feral self-sustaining populations complicates the assignment of dates of invasion.

North American Invasion History:

Invasion History on the West Coast:

In North America, Magallana gigas was first introduced to Puget Sound, Washington (WA) in 1902, following overfishing of the native Olympic Oyster (Ostrea lurida) and unsuccessful stocking of M. virginica (Eastern Oyster). Early transplants were unsuccessful due to mortality in shipping, but after numerous subsequent imports, large-scale cultivation was underway in Washington State by 1928 (Chew 1979). In British Columbia, imports began in 1912, but large-scale natural spawning was not seen until 1932 (Quayle 1969). Fairly regular settlement of M. gigas spat, outside areas of cultivation, now occurs from Pendrell Sound, British Columbia, to Willapa Bay, WA (Quayle 1969; Ruesink et al. 2005). This species is now the basis of the West Coast oyster industry, with commercial culture taking place from southern British Columbia to Morro Bay, California (CA) (Chew 1979; Quayle 1969; Conte 1996). However, these operations were largely dependent on imported seed from Japan, and later (1970s onward) on hatchery-reared spat (Barrett 1963; Quayle 1969; Carlton 1979; Conte 1996). South of Willapa Bay, natural spawnings of M. gigas were rare (Span 1978; Carlton 1979; Boyd et al. 2000; Coan et al. 2000), but hatchery-based oyster aquaculture operations occur in several Oregon bays and south to Morro Bay, CA (Carlton 1979; Conte 1996), the Pacific Coast of Baja California (Rodriguez and Ibarra-Obando 2008), and the Gulf of California (Arizpe 1996).

In California, since 2000, there have been collections of 'wild' M. gigas in San Francisco Bay and southern California estuaries (Andy Chang, personal communication; Ruiz et al. unpublished data; Cohen et al. 2002; de Rivera et al. 2005; Burnaford et al. 2011; Goodwin et al. 2011). At least some of the San Francisco Bay occurrences have resulted from the breeding of illegally planted M. gigas (Andy Chang, personal communication). Transport of oysters in ship fouling or larvae in ballast water are also possible vectors. There is some evidence for multiple cohorts of oysters in San Francisco Bay, but at this time we consider the establishment of M. gigas to be uncertain.

Invasion History on the East Coast:

Magallana gigas attracted some attention in the mid-20th century because of its large size and rapid growth. A bushel of Pacific Oysters was planted in Barnegat Bay, New Jersey, but failed to grow, and died within two years. A number of illegal and government plantings were made in estuaries from Delaware to Maine from the 1930s to the 1980s, but settlement of larvae and establishment of Pacific Oysters was not observed (Dean 1979; Hickey 1979; Andrews 1980). There was particular interest in Maine, because of the limited existing Eastern Oyster (M. virginica) stocks there. Plantings were made in 1949 in Blue Hill Bay and in the 1970s in Damariscotta River and Goose Pond, a lagoon of Penobscot Bay (Dean 1979; Shatkin et al. 1997). However, we are not aware of more recent introduction attempts.

Around Chesapeake Bay, interest in M. gigas intensified as the native M. virginica  declined due to overfishing and disease (MSX- Haplosporidium nelsoni, Dermo- Perkinsus marinus). Magallana gigas was considered to be more disease-resistant than the Eastern Oyster, and was considered as a potential replacement, especially in Virginia, where oyster losses were greatest (Andrews 1980; DuPaul 1992). Numerous culture experiments were undertaken with diploid and triploid (sterile) M. gigas in order to assess the disease resistance of the Pacific Oyster and its adaptability to the Chesapeake Bay environment. Experiments in quarantined flumes indicated that M. gigas had lower prevalence and intensity of P. marinus and H. nelsoni infections (Barber 1996; Barber and Mann 1994; Chu et al. 1996; Krantz 1992). Plantings of sterile triploid M. gigas in Chesapeake Bay, Virginia, and North Carolina indicated that this oyster grew well at high salinities, but performed poorly at low salinities (Calvo et al. 1999; Grabowski et al. 2004). Benefits of a disease-resistant oyster would include restoration of the oyster-reef environment and of a filter-feeding biomass in at least part of Chesapeake Bay, as well as revival of oystering (Gottlieb and Schweighofer 1996; Lipton et al. 1992; Mann et al. 1991). Although M. gigas showed strong disease resistance, trials in Chesapeake Bay suggested that this oyster was not well-adapted to the local environment. In quarantined flumes, M. gigas had high non-disease mortality in summer (Barber and Mann 1994), and heavy Polydora spp. infestations (Mann and Burreson 1994; DeBrosse and Allen 1996). By 1998-2000, research interests in Virginia and North Carolina had shifted to M. ariakensis, which demonstrated better growth and survival under Chesapeake Bay conditions (Hallerman et al. 2001; National Research Council 2003).

Invasion History on the Gulf Coast:

At least one unsuccessful attempt was made to introduce M. gigas to the Gulf Coast. Kavanaugh (1941) reported very briefly that 'Japanese oysters' in Louisiana, showed 'amazingly serious infestation' by spionid polychaetes (Polydora spp.), and that native oysters were not seriously affected. This is the only report that we have of this oyster in the Gulf of Mexico.

Invasion History in Hawaii:

A small shipment of M. gigas was planted at Mokapu, Oahu on Kaneohe Bay in 1926, but did not become established. Larger plantings were made at Pearl Harbor in 1938, and in Kaneohe Bay (2 million spat planted) in 1939. Pacific Oysters are now established in Pearl Harbor and abundant in Kaneohe Bay (Coles et al. 1999; Coles et al. 2002; Carlton and Eldredge 2009 - 2000 oysters planted, established).

Invasion History Elsewhere in the World:

In the northeastern Atlantic, Magallana gigas was imported to Marennes, France in small quantities in 1966. This was followed by a disease epizootic in M. angulata (Portuguese Oyster), which was then the predominant commercial species (itself imported to supplant the overfished Ostrea edulis or the European Flat Oyster). Consequently, large imports of M. gigas were made to replace the lost M. angulata stocks (Grizel and Héral 1991). In the United Kingdom, laboratory stocks were imported in 1965 and 1972, and the experimental field plantings of lab-reared spat, in 1967 and 1973. Spawning and recruitment were rare in British waters, owing to low water temperatures (Walne and Helm 1979). Extensive plantings of M. gigas were made in the Atlantic waters of Europe in the 1970s, from Spain to Ireland, and east to Germany and Denmark (Ruesink et al. 2005; Minchin 2007; Troost 2010; Wrange et al. 2010). This culture was largely hatchery-based, but natural spawning and settlement were seen in the 1970s and 1980s in many locations, particularly the Wadden Sea area of the Netherlands, Germany and Denmark (Reise 1998; Gittenberger et al. 2010; Troost 2010; Wrange et al. 2010), where extensive oyster beds were replacing mussel beds by the year 2000. The occurrence of successful spawning and massive recruitment in northern Europe in recent decades has been attributed in part to climate change (Troost 2010; Wrange et al. 2010; Thomas et al. 2016). Successful spawning and apparent establishment took place by 2007 in Espevik, Norway (60⁰N). Established populations also occur in Denmark and Sweden, along the Kattegatt, at the mouth of the Baltic (Wrange et al. 2010) and along the Atlantic coast of France, Portugal and Spain (Grizel and Héral 1991, Ruesink et al. 2005). Hatchery and wild M. gigas populations along the Atlantic coasts of Europe, from Germany (Sylt) to southern France (Arcachon), show little genetic differentiation, being strongly determined by hatchery and aquaculture practices (Meistertzheim et al. 2013).

Magallana gigas, imported from Japan, was first introduced to the Mediterranean Sea by 1964, in the Thau Lagoon, near Sete, France, again as a replacement for declining stocks of Ostrea edulis and M. angulata. It soon was widely cultured in the Mediterranean from Morocco to Israel (Ruesink et al. 2005). This oyster appears to be, at least locally, established in coastal lagoons and estuaries in Tunisia, France, Italy, Greece, and Turkey (Zenetos et al. 2003; Ruesink et al. 2005; Zenetos et al. 2005; Albayrak 2011; Antit et al. 2011). The status of M. gigas in the Black Sea is uncertain - it is known mostly as single individuals near ports and oyster farms (Skarlato and Sarobogov 1972, cited by Zoloterev 1996; Gomiou et al. 2002; Skolka and Preda 2010).

Magallana gigas has been widely cultured in the Southern Hemisphere, beginning in 1947 in Tasmania, Australia (Nell 2001), in 1950 in South Africa (Robinson et al. 2005), and in 1977 in Chile (Castilla et al. 2005). In Chile, Pacific Oysters remain confined to aquaculture facilities, possibly because of low water temperatures (Castilla et al. 2005). However, breeding populations quickly developed in Tasmania and by the 1960s in mainland Australia (Nell 2001), and locally by 2001 on the southern coast of South Africa (Robinson et al. 2005). In Argentina, a failed aquaculture operation led to established populations on the Patagonian coast (Orensanz et al. 2002; Escapa 2004). Surprisingly, spat and adults of M. gigas were identified by molecular means in cultures of native oysters (M. brasiliana and M. rhizophorae) in Brazil, at latitudes between 27 and 29⁰S (Melo et al. 2010). Some populations of M. gigas in New Zealand (1st record 1961, Cranfield et al. 1998) are believed to have resulted in transport by shipping, and are not associated with known aquaculture operations (Krassoi et al. 2008). Pacific Oysters have been widely introduced to tropical and subtropical regions and islands (e.g., Puerto Rico, Virgin Islands, Madeira, Guam, Tonga, Fiji, Belize, Malaysia), but with the exception of Hawaii, these introductions have not resulted in established populations or successful hatchery-based aquaculture (Ruesink et al. 2005). Carrasco and Baron (2010) concluded that M. gigas could establish populations in regions with mean sea surface temperature ranging from 14 to 28.9⁰C for the warmest month and from -1.9 to 19.8⁰C for the coldest month of the year. The Pacific Oyster's occurrence in slightly warmer water in Brazil may have been due to unintentional selection of oysters in Brazilian shellfish laboratories (Melo et al. 2010).

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Description

Magallana gigas resembles other oysters in having unequal valves and an irregular shape. The shape of the shell varies greatly with the growth environment. For instance, on hard substrate the shell can be rounded, domed and fluted; on soft substrate it can be flatter and less ridged; and when crowded the shell is often narrower (Quayle 1969). The right (lower) valve may be deeply cupped. Both valves are covered with concentric growth layers (lamellae) on the outer surface, but with fewer and stronger ridges on the left (upper) side. The edges of the lamellae are strongly rippled into spines and ridges (Coan and Valentich-Scott 2007; Langdon and Robinson 1996). Shells can be white to off-white to gray, sometimes with brown or purple on the ridges. The interior of the shell is smooth and white, with a purple muscle scar (Quayle 1969; Coan et al. 2000). Magallana gigas matures at about 80 mm, but is reported to occasionally grow to 400-450 mm (Carriker and Gaffney 1996). The larvae are illustrated by Quayle (1969). 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 (Quayle 1969).

Magallana gigas is a genetically diverse species. In different parts of Japan, different strains are cultivated with different growth patterns and ecological preferences. The most widely planted form is the Miyagi strain, from the central Pacific coast of Japan which is large and fast-growing (Quayle 1969). In addition, many closely related species are found in the Northwest and Indo-West Pacific regions. Magallana angulata (Portuguese Oyster), introduced to Europe in the 16th century, is very closely related (Ó'Foighil et al. 1998; Huvet et al. 2004; Lapegue et al. 2004; Reece et al. 2008).

The genus name Magallana has been proposed for Pacific members of the genus Crassostrea, based on genetic divergence between Pacific and Atlantic oysters of the genus (Salvi et al. 2014; Salvi and Mariottini 2020). Bayne and 23 co-authors disagreed with the proposed name changes, based on the limited scope of the genetic analysis, the absence of morphological differentiation, and the inconveninece of changing thename of an economically important species (Bayne et al. 2017). A further genetic analysis by Salvi and Mariottini (2020) owed that the Indo-Pacific and western Atlantic 'Crassotrea' clustered in two separate groups, justifying the use of the name Magallana for the Indo-Pacific species (James T. Carlton, personal communication).

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Taxonomy

Ecology

General:

Magallana gigas like other oysters, is a protandric hermaphrodite, maturing first as a male, and then often 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 16 to 30⁰C (Quayle 1969; His et al. 1989). Gonads can develop in M. gigas at 80 mm (National Research Council 2003). Adult M. gigas feed on phytoplankton of 6-32 um with ~100% retention efficiency, but are less efficient with smaller organisms (Nielsen et al. 2016). Adult oysters are reported to grow to 450 mm, although 300 mm in length is a more typical maximum (Quayle 1969; Carriker and Gaffney 1996).

Magallana gigas is characteristic of protected coastal waters in China and Japan. This oyster normally grows at salinities of 23-28 PSU, and can tolerate brief exposures to salinities as low as 5-10 PSU (Nell and Holliday 1988; Carriker and Gaffney 1996; Gray and Langdon 2018). It tolerates a very wide temperature range, from -1.8 to 35⁰C, although temperatures over 30⁰C are stressful (Shpigel et al. 1992; Carrasco and Barón 2010). Settlement and survival are best at sites at sites portected from wave exposure (Teschke et al. 2020).

Food:

Phytoplankton

Consumers:

Crabs, Fishes, Starfish, Humans

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Oyster Reef	None
General Habitat	Coarse Woody Debris	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	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)	-1.8	Based on geographical range (Carrasco and Baron 2010).
Maximum Temperature (ºC)	35	Crassostrea gigas (Pacific Oysters) shows signs of metabolic stress at 30 C (Shpigel et al. 1992; Gray and Langdon 2018).
Minimum Salinity (‰)	5	Substantial growth and reproduction occurs only above 20 ppt. (His et al. 1989; Mann et al. 1991; Nell and Holliday 1988; Gray and Langdon 2018).
Maximum Salinity (‰)	41	Successful aquaculture, Nell and Holliday 1988
Minimum Reproductive Temperature	16	Field and experimental data (His 1991; Mann et al. 1991)
Maximum Reproductive Temperature	30	Field and experimental data (His 1991; Mann et al. 1991)
Minimum Reproductive Salinity	15	Experimental conditions for larval growth (Nell and Holliday 1988). Optimum salinities for reproduction and larval growth are 20-30 ppt (His et al. 1989; Mann et al. 1991; Nell and Holliday 1988).
Maximum Reproductive Salinity	40	Experimental conditions for larval growth (Nell and Holliday 1988). Optimum salinities for reproduction and larval growth are 20-30 ppt (His et al. 1989; Mann et al. 1991; Nell and Holliday 1988).
Minimum Duration	11	Larval Period (His et al. 1989)
Maximum Duration	30	Larval Period (His et al. 1989)
Minimum Length (mm)	80	Carriker and Gaffney (1996)
Maximum Length (mm)	450	Carriker and Gaffney (1996)
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Magallana gigas is the world's most widely cultivated and eaten shellfish (Carriker and Gaffney 1996; Ruesink et al. 2005), but it is also a highly successful invader, and a powerful ecosystem engineer, creating complex reefs, replacing native shellfish, and altering estuarine foodwebs through suspension-feeding (Herbert et al. 2016).

Economic Impacts

Fisheries - Magallana gigas is the most widely cultivated and harvested shellfish species in the world, introduced to at least 52 countries (Carriker and Gaffney 1996; Ruesink et al. 2005). Among the more notable introductions have been those to the west coast of North America (Chew 1979; Quayle 1969), European waters (Grizel and Héral 1991; Walne and Helm 1979), and Australia (Nell 2001). The disease resistance of this oyster, its adaptability to a wide range of environments, the long development of culture techniques, and its large size are among the reasons for its widespread introduction (Quayle 1969; Andrews 1980; Mann et al. 1991; Ruesink et al. 2005). Profitable culture in natural waters is possible, using hatcheries or imported seed, even in regions where M. gigas cannot breed successfully in the wild, such as California and Pacific Mexico (Arizpe 1996; Conte 1996; Ruesink et al. 2005).

Disadvantages include bland flavor compared to other species, including Ostrea eduis and M. virginica (DuPaul 1992), and risks to native oyster populations, including competition, hybridization, and introductions of associated organisms (parasites, fouling species and oyster predators) (Galtsoff 1932; Grizel and Héral 1991; Mann et al. 1991; Ruesink et al. 2005). In the Wadden Sea area of northern Europe, settlement of M. gigas has covered valuable beds of mussels (Mytilus edulis) and cockles (Cerastoderma edule), and interfered with the use of fishnets (Troost 2010). In Willapa Bay and Grays Harbor, Washington (WA), the pesticide Carbaryl is used to kill mud shrimps which burrow in oyster beds, creating general environmental concerns, as well as killing other fisheries species, such as Dungeness Crabs (Metacarcinus magister) and English Sole (Parophrys vetulus).

Ecological Impacts

Competition - Introductions of new oyster species are often motivated by the decline of the previously dominant oyster due to overfishing or disease, but in some cases they have led to further damage to the remaining populations. Introductions of M. angulata (Portuguese Oyster) in France coincided with the decline of the native Ostrea edulis (European Flat Oyster) in the 19th century (Galtsoff 1932); the replacement of M. angulata by M. gigas in the 1970's seems to have largely been a consequence of a disease of unknown origin (Grizel and Héral 1991). In Australia, competition with M. gigas is considered a threat to the native Saccostrea commercialis (Sydney Rock Oyster) (Mann et al. 1991; Nell 2001). On the West Coast of North America, competition between M. gigas and the native Olympia Oyster (Ostrea lurida) is limited since M. gigas tends to settle, and is cultivated in intertidal areas, while the native oyster tends to grow in lower intertidal and subtidal areas. However, where they do overlap, M. gigas grows much faster, and has a higher filtration rate (Ruesink et al. 2005).

Magallana gigas also competes for space and food with bivalves other than oysters, such as Mytilus edulis (Blue Mussel) and Cerastoderma edule (Common Cockle). In the Wadden Sea (southern North Sea) of Netherlands-Germany-Denmark, M. gigas has been settling on intertidal mussel and cockle beds (Reise 1998; Diederich 2005).

Habitat Change - Both cultivated populations of M. gigas and naturally settled reefs can make large structural changes in littoral communities. These changes are greatest in soft-bottom habitats, such as Willapa Bay, WA; Bahia Anagada, Argentina; and the Wadden Sea (southern North Sea) of Netherlands-Germany-Denmark, which include vast intertidal mudflats. Cultivation takes place on man-made structures, while natural beds result from settlement on mussel beds, logs, or other isolated hard substrates (Escapa 2004; Ruesink et al. 2005; Ruesink et al. 2006; Troost 2010). Cultivated and natural beds create large, complex structures, with extensive hard substrate for organisms to settle on, and lots of nooks and crannies providing shelter for native and introduced mobile organisms (Escapa 2004; Diederich 2005; Hosack et al. 2006; Ruesink et al. 2006; Gittenberger et al. 2010; Markert et al. 2010; LeJart and Hily 2011). While Pacific Oysters settle on and cover mussel beds, they also provide substrate for mussel settlement, and can result in increased biodiversity in their invaded habitats (Markert et al. 2010; LeJart and Hily 2011). On hard substrates, such as rocky shores, impacts of M. gigas are less dramatic (Ruesink et al. 2005). However, intertidal oysters provide a light-colored substrate, cooler than exposed dark rocks, and favoring the survival of limpets (Lottia sp.) at high tide (Padilla 2010), and also increase habitat for barnacle settlement (Bourne 1979, cited by Ruesink et al. 2005). The deposits of pseudofeces can also increase the diversity and abundance of deposit feeders (LeJart and Hily 2011).

Invasions by M. gigas do have negative impacts on habitats. Their high filtration rates result in the deposition of partially digested pseudofeces, which can accumulate around the oyster beds, creating anoxic zones in the sediment, limiting infauna and adversely affecting eelgrass beds (Kelly et al. 2008; Troost 2010). The large accumulations of shell which M. gigas creates in the intertidal zone have a negative effect on the native oyster (O. lurida) by attracting large numbers of settling larvae to the intertidal zone, where their survival is poor, acting as a recruitment sink (Ruesink et al. 2005).

Parasite-Predator vector - In many regions of the world, parasites, epifauna, and predators have been imported with shipments of M. gigas. Known parasites of M. gigas which are now established on the Pacific coast of North America, or in France, include three viruses, three bacterial diseases, three protistans (other than haplosporidians) (Marteilia refringens, Marteilioides chungmuensis and Mikrocytos mackini), the copepod Mytilicola orientalis, and at least one disease of unknown etiology (Mann et al. 1991). The first imports of M. gigas to France coincided with a viral epizootic which largely wiped out the then-dominant commercial oyster M. angulata (Portuguese Oyster), but the origin of this disease is unknown (Grizel and Héral 1991). Many species of macro-organisms have been introduced to, or transferred locally in European and West Coast waters with M. gigas, these include macroalgae (eg. Sargassum muticum), flatworms (Pseudostylochus ostreophagus), snails (e.g. Pteropurpura inornata, Japanese Oyster Drill), clams, bryozoans (Schizoporella japonica), and tunicates (Perophora japonica, Styela clava). Some of these species have had negative impacts on oysters and surrounding communities (Cohen and Carlton 1995; Grizel and Héral 1991; Mann et al. 1991; Cohen et al. 1998; Reise et al. 1999; Goulletquer et al. 2002).

However, the associate of M. gigas which has had the largest ecological and economic impact is probably the protist Haplosporidium nelsoni, which infects the Pacific Oyster with minimal symptoms, but produces the symptoms of the MSX disease, with high mortality, in the Eastern Oyster (M. virginica) (Friedman 1996; Burreson et al. 2000). From 1958 to the present, outbreaks of this disease have caused high mortality in Chesapeake and Delaware Bays, and elsewhere on the East Coast of North America. It seems likely that one of the many early unofficial introductions of M. gigas to the East Coast may have introduced H. nelsoni, although transport of oysters in fouling or spores in ballast water cannot be excluded (Burreson et al. 2000).

In a sort of reverse-parasite vector role, Magallana gigas, together with the Common Atlantic Slipper Shell Crepidula fornicata and other filter feeders, such as the Softshell Clam (Mya arenaria) were found to affect transmission of native parasites (the trematode Himasthla elongata) of the Common Cockle (Cerastoderma edule) and the Blue Mussel (Mytilus edulis), by filtering out the metacercariae, without becoming infected themselves. The effect of these invaders was to reduce the parasite load of the native bivalves (Thieltges et al. 2008; Thieltges et al. 2009).

While it is a highly desired seafood item, Magallana gigas is also an ecosystem engineer and poses a challenge to managers of marine protected areas. This oyster can interfere with native mussel fisheries, create reefs which can obstruct navigation, litter beaches with 'razor-sharp' shells, and drastically effect native marine communities. Regional planning and risk assessment is desirable for oyster culture operations in new areas. Environmental measurements can be used to determine the risk of reproduction of cultured oysters. One option is to require use of triploid oysters in culture, to limit reproduction, but reversion of triploids can occur. Heavy settlement of 'wild' oysters can interfere with culture operations by fouling equipment and cultured oysters. Dredging has been used to eliminate 'wild' oysters, but the habitat damage is considerable. In some areas, hand collection is sufficient to maintain oyster-free zones (Herbert et al. 2016).

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Regional Impacts

P110	Tomales Bay		Economic Impact	Fisheries
Commercial oyster operations, using M. gigas began in Tomales Bay in 1928, and continue to the present. Oyster culture here was intially dependent on seed imported from Japan, but now uses seed produced in US hatcheries (Barrett 1963; Conte 1996).
NEP-V	Northern California to Mid Channel Islands		Economic Impact	Fisheries
Commercial oyster operations, using M. gigas began in Tomales Bay in 1928, and continue to the present. Major locations of oyster rearing included Morro Bay, Elkhorn Slough, Drakes Estero, and Tomales Bay (Barrett 1963; Carlton 1979; Conte 1996). Culture of M. gigas continues in Morro Bay, Drakes Estero and Tomales Bay (Conte 1996). In San Francisco Bay, commercial Pacfiic Oyster rearing occurred form 1932 to 1939. Oyster culture here was intially dependent on seed imported from Japan, but now uses seed produced in US hatcheries (Barrett 1963, Conte 1996). California Pacific Oyster growers produced 1.5 million pounds of shucked meat in 1995. About 90% of Calfornia's production occurred in Drakes Estero and Humboldt Bays (Conte 1996).
P100	Drakes Estero		Economic Impact	Fisheries
Commercial culture of M. gigas began in Drakes Estero in 1932 and continues to the present. Oyster culture here was intially dependent on seed imported from Japan, but now uses seed produced in US hatcheries (Barrett 1963, Conte 1996). Drakes Estero is one of the two most important oyster-growing sites in California About 90% of production occurred in Drakes Estero and Humboldt Bays (Conte 1996).
P080	Monterey Bay		Economic Impact	Fisheries
Culture of M. gigas continued in Elkhorn Slough from 1929 to the 1980s (Barrett 1963; Conte 1996; Wasson et al. 2001)
P070	Morro Bay		Economic Impact	Fisheries
Culture of M. gigas in Morro Bay started in 1932 and continues to the present (Barrett 1963; Conte 1996; Morro Bay National Estuary Program 2005 http://www.mbnep.org/index.php).
P090	San Francisco Bay		Economic Impact	Fisheries
Commercial rearing of M. gigas took place in San Francisco Bay from 1932 to 1939, when the company involved went out of business (Barrett 1963).
P112	_CDA_P112 (Bodega Bay)		Economic Impact	Fisheries
Commercial rearing of M. gigas occurred in San Francisco Bay from 1932 to 1938 (Barrett 1963, cited by Carlton 1979)
NEP-VI	Pt. Conception to Southern Baja California		Economic Impact	Fisheries
Substantial aquaculture operations for M. gigas occur in Bahia San Quitin, Baja California, Mexico (Rodriguez and Ibarra-Obando 2008).
P130	Humboldt Bay		Economic Impact	Fisheries
Magallana gigas is reared in extensive aquaculture operations in Humboldt Bay. These began in 1953 and continue to the present. About 90% of Calfornia's production occurred in Drakes Estero and Humboldt Bays (Conte 1996).
NEP-IV	Puget Sound to Northern California		Economic Impact	Fisheries
Willapa Bay and Grays Harbor are major oyster-growing areas, producing more than 10% of the US oyster crop, through intensively managed culture (Feldman et al. 2000; Ruesink et al. 2006). A negative impact of this aquaculture operation is the use of the pesticide carbaryl to kill the mud shrimps Neotrypaea californiensis and Upogebia pugettensis, which interfere with oyster culture by burrowing and suspending sediment. The pesticide also kills juvenile Dungeness Crabs (Metacarcinus magister), English sole (Parophrys vetulus), and other commerical and sport fishery species, as well as raising general environmental concerns (Feldman et al. 2000). In Oregon, aquaculture of M. gigas began in 1906 in Yaquina Bay, and 1940-1948 in Netarts, Tillamook, Winchester, and Coos Bays (Carlton 1979), and continues to the present day (Oregon Department of Fish and Wildlife 2011, http://www.dfw.state.or.us/mrp/shellfish/bayclams/about_oysters.asp; Oregon Department of State Lands 2011, http://www.oregon.gov/DSL/SSNERR/docs/EFS/EFS34aquaculture.pdf?ga=t).
NEP-IV	Puget Sound to Northern California		Ecological Impact	Habitat Change
Intensive oyster production has greatly altered Willapa Bay. Most of the production takes place in the intertidal zone, which was formerly mudflat. The native Olympic Oyster, O. lurida, now rare, was primarily subtidal. Oyster growth in the interitdal zone has created large areas of hard, stuctured habitat, which supports greatly increased densities of epibenthic invertebrates, incluiding mussels, scaleworms, and tube-dwelling amphipods (Ruesink et al. 2005; Ruesink et al. 2006; Hosack et al. 2006). However, the large accumulations of shell which M. gigas creates in the intertidal zone has a negetive effect on the native oyster by attracting large numbers of settling larvae of O. lurida, in the interitdal zone, where their survival is poor, acting as a recuriment sink (Ruesink et al. 2005)
NEP-IV	Puget Sound to Northern California		Ecological Impact	Herbivory
The greatly increased oyster biomass has resulted in an increase in filtration rate of about 25%, from 0.8 to 1.3% of the bay's volume (Ruesink et al. 2006). This is an underestimate, since it is based on harvested biomass, and excludes feral populations of M. gigas.
NEP-IV	Puget Sound to Northern California		Ecological Impact	Competition
Competition between the introduced Pacific Oyster (Magallana gigas) and the native Olympia Oyster (Ostrea lurida) is expected to be minimal, since M. gigas tends to settle, and is cultivated in intertidal areas, while the native oyster tends to grow in the lower intertidal and subtidal areas. However, where they do overlap, M. gigas grows much faster, and has a higher filtration rate (Ruesink et al. 2005). Competition for space occurs when M. gigas displaces native Eelgrass (Zostera marina), in culture operations (Wagner et al. 2012).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- Although M. gigas has not become definitely established in central California, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including Pteropurpura (=Ocinebrellus) inornata (Japanese Oyster Drill) in Tomales Bay, the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum and Styela clava (Carlton 1979; Cohen and Carlton 1995; Wasson et al. 2001; de Rivera et al. 2005).
P090	San Francisco Bay		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- Although M. gigas has not become definitely established in San Francisco Bay, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including, the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus and Styela clava (Carlton 1979; Cohen and Carlton 1995).
P110	Tomales Bay		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- Although M. gigas has not become definitely established in Tomales Bay, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including Pteropurpura (=Ocinebrellus) inornata (Japanese Oyster Drill), the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum, and Styela clava (Carlton 1979; Cohen and Carlton 1995).
P080	Monterey Bay		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- Although M. gigas has not become definitely established in Elkhorn Slough, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus and Styela clava (Carlton 1979; Wasson et al. 2001; de Rivera et al. 2005)
P070	Morro Bay		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- Although M. gigas has not become definitely established in Morro Bay, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum, and Styela clava (Carlton 1979; Needles 2007)
NEP-IV	Puget Sound to Northern California		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- The introduction of M. gigas has been a possible/probable vector for a number of oyster foulers or predators on the Washington-Oregon-northern California Coast, including Pteropurpura (=Ocinebrellus) inornata (Japanese Oyster Drill) in Willapa Bay, the parasitic copepod Mytilicola orientalis (widespread), the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum and Styela clava (Carlton 1979; Cohen and Carlton 1995; Boyd et al. 2002; Wonham and Carlton 2005).
P130	Humboldt Bay		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- The introduction of M. gigas has been a possible/probable vector for a number of oyster foulers or predators in Humboldt Bay, including the parasitc copepod Mytilicola orientalis (widespread), the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum and Styela clava (Carlton 1979; Boyd et al. 2002).
P100	Drakes Estero		Ecological Impact	Parasite/Predator Vector
None
CA	California		Ecological Impact	Parasite/Predator Vector
Parasite-Predator vector- Although M. gigas has not become definitely established in central California, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including Pteropurpura (=Ocinebrellus) inornata (Japanese Oyster Drill) in Tomales Bay, the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum and Styela clava (Carlton 1979; Cohen and Carlton 1995; Wasson et al. 2001; de Rivera et al. 2005)., Parasite-Predator vector- The introduction of M. gigas has been a possible/probable vector for a number of oyster foulers or predators in Humboldt Bay, including the parasitc copepod Mytilicola orientalis (widespread), the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum and Styela clava (Carlton 1979; Boyd et al. 2002)., Parasite-Predator vector- Although M. gigas has not become definitely established in San Francisco Bay, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including, the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus and Styela clava (Carlton 1979; Cohen and Carlton 1995)., Parasite-Predator vector- Although M. gigas has not become definitely established in Morro Bay, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum, and Styela clava (Carlton 1979; Needles 2007), Parasite-Predator vector- Although M. gigas has not become definitely established in Elkhorn Slough, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus and Styela clava (Carlton 1979; Wasson et al. 2001; de Rivera et al. 2005), nan, Parasite-Predator vector- Although M. gigas has not become definitely established in Tomales Bay, its introduction has been a possible/probable vector for a number of oyster foulers or predators, including Pteropurpura (=Ocinebrellus) inornata (Japanese Oyster Drill), the parasitc copepod Mytilicola orientalis (widespread), the mussel Musculista senhousia, the bryozoan Schizoporella japonica, and the tunicates Botrylloides violaceus, Didemnum vexillum, and Styela clava (Carlton 1979; Cohen and Carlton 1995).
CA	California		Economic Impact	Fisheries
Commercial oyster operations, using M. gigas began in Tomales Bay in 1928, and continue to the present. Major locations of oyster rearing included Morro Bay, Elkhorn Slough, Drakes Estero, and Tomales Bay (Barrett 1963; Carlton 1979; Conte 1996). Culture of M. gigas continues in Morro Bay, Drakes Estero and Tomales Bay (Conte 1996). In San Francisco Bay, commercial Pacfiic Oyster rearing occurred form 1932 to 1939. Oyster culture here was intially dependent on seed imported from Japan, but now uses seed produced in US hatcheries (Barrett 1963, Conte 1996). California Pacific Oyster growers produced 1.5 million pounds of shucked meat in 1995. About 90% of Calfornia's production occurred in Drakes Estero and Humboldt Bays (Conte 1996)., Magallana gigas is reared in extensive aquaculture operations in Humboldt Bay. These began in 1953 and continue to the present. About 90% of Calfornia's production occurred in Drakes Estero and Humboldt Bays (Conte 1996)., Commercial rearing of M. gigas took place in San Francisco Bay from 1932 to 1939, when the company involved went out of business (Barrett 1963)., Culture of M. gigas in Morro Bay started in 1932 and continues to the present (Barrett 1963; Conte 1996; Morro Bay National Estuary Program 2005 http://www.mbnep.org/index.php)., Culture of M. gigas continued in Elkhorn Slough from 1929 to the 1980s (Barrett 1963; Conte 1996; Wasson et al. 2001), Commercial culture of M. gigas began in Drakes Estero in 1932 and continues to the present. Oyster culture here was intially dependent on seed imported from Japan, but now uses seed produced in US hatcheries (Barrett 1963, Conte 1996). Drakes Estero is one of the two most important oyster-growing sites in California About 90% of production occurred in Drakes Estero and Humboldt Bays (Conte 1996)., Commercial oyster operations, using M. gigas began in Tomales Bay in 1928, and continue to the present. Oyster culture here was intially dependent on seed imported from Japan, but now uses seed produced in US hatcheries (Barrett 1963; Conte 1996)., Commercial rearing of M. gigas occurred in San Francisco Bay from 1932 to 1938 (Barrett 1963, cited by Carlton 1979)

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