Invasion History

First Non-native North American Tidal Record: 1998
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1998

General Invasion History:

Magallana ariakensis is thought to be native to the coasts of China, but was introduced very early to Ariake Bay, in southern Japan, from which it was first described (Zhou and Allen 2003). Under the name Crassostrea rivularis, it has been reported from India and Pakistan, and possibly Malaysia and Borneo. However, its occurrence outside China and Japan is uncertain (National Research Council 2003; Zhou and Allen 2003; Reece et al. 2008) and because of its confused taxonomy, the full extent of its native range is unknown. This species was inadvertently introduced to the US West Coast with shipments of M. gigas (Pacific Oyster) from Japan. In the 1990s, culture of M. ariakensis on the West Coast was very localized, and wild populations were not reported (Langdon and Robinson 1996). This oyster attracted attention as a possible replacement or supplement for oyster populations affected by disease, first briefly in France (Cochennec et al. 1998; Goulletquer et al. 2002), and then to a much greater degree on the East Coast of the US, in Chesapeake Bay and North Carolina. In each of these, risks associated with the new oyster were perceived to exceed the potential benefits (Hallerman et al. 2001; National Research Council 2003; Burreson et al. 2004; Bushek et al. 2008; Xaio et al. 2011) and the decision was made against the introduction (Fahrenthold 2009).

North American Invasion History:

Invasion History on the West Coast:

Magallana ariakensis appeared in Oregon hatcheries with shipments of M. gigas from Kumamoto Province, Japan, by the 1970s and created interest as an oyster that would have marketable qualities in the summer, when the dominant commercial Pacific Oyster (M. gigas) has soft flesh due to spawning. This oyster, then known as M. rivularis, was planted in Tillamook Bay and Yaquina Bay, Oregon (Breese and Malouf 1977). These plantings apparently did not result in established populations. Another planting of M. ariakensis in Oakland Bay, Puget Sound, Washington, in 1981, did spawn, presumably because of low temperatures (Perdue and Erickson 1984), but a population did not become established. In the 1990s, culture of on the West Coast was still very localized, and wild populations were not reported (Langdon and Robinson 1996).

Invasion History on the East Coast:

On the East Coast of North America, the decline of Eastern Oyster (Magallana virginica) populations, and reduced recruitment, due in part to the diseases MSX (Haplosporidum nelsoni) and 'Dermo' (Perkinsus marinus) led to an interest in importing exotic, disease-resistant oysters. This interest was strongest in Virginia, because of the near-eradication of native oysters by disease (Burreson and Mann 1994; Calvo et al. 1999; Leffler et al. 2002). Experiments with Pacific Oysters (M. gigas) suggested that this species is poorly adapted to the high summer temperatures and low salinities of many East Coast estuaries (Burreson and Mann 1994; DeBrosse and Allen 1996; Calvo et al. 1999). Starting in 1998, trials were carried out with M. ariakensis stock obtained from Oregon hatcheries. Successful small-scale trials led to extensive stocking of triploid oysters, up to several hundred thousand per year, which provided some economic benefit to Virginia watermen. Environmental and fisheries concerns led to an extensive program of research on many aspects of the biology of M. ariakensis. Possible options included the release of fertile diploid oysters, or expanded aquaculture of triploids. However, the risks, including the presence of a new parasite, Bonamia sp. in high salinity waters, the low genetic diversity of the cultured stock, the risk of triploid reversion, and the high vulnerability of M. ariakensis to predators, were perceived to outweigh the benefits. The Army Corps of Engineers and officials of the states of Maryland and Virginia decided to prohibit introduction of diploid M. ariakensis, end cultivation of triploids in open waters, and instead transfer resources to restoration of the native Eastern Oyster (Fahrenthold 2009; Wheeler 2009). The decision does permit continued research on culture of M. ariakensis in closed systems (Wheeler 2009).

Invasion History Elsewhere in the World:

In 1994, Magallana ariakensis was imported to hatcheries in France to test its potential as a replacement for M. gigas, in the event of a catastrophic disease outbreak. However, the discovery of a Bonamia-like parasite in these oysters led to the abandonment of work with this species (Cochennec et al. 1998).


Description

Magallana ariakensis resembles other oysters in having unequal valves and an irregular shape. The right (lower) valve is thinner, flatter, and smaller than the left. 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 flat and not strongly rippled, as in M. gigas (Langdon and Robinson 1996). Colors of the lamellae on the outer surface vary from gray and yellowish brown to purple, while the inner surface of the valves is smooth and grayish white, with purple on the edges. The muscle scar on the inner surface of the valves is large and purplish (Coan et al. 2000). Substantial geographical variations were noted in different areas of Japan and China (Zhou and Allen 2003). Magallana ariakensis is reported to grow to 200-240 mm (Carriker and Gaffney 1996).

Magallana ariakensis appears to be part of a species complex, including the closely related M. honkongensis and M. nippona. It is less closely related, but still quite similar to at least nine other Magallana species from Indo-West-Pacific waters. A recent genetic study indicates that populations of M. ariakensis from northern and southern China constitute two separate sibling species. Hatchery stock in the US, including that of the Virginia Institute of Marine Science, used for Chesapeake Bay and North Carolina trials, include both species, although they are dominated by the northern genotype (Zhang et al. 2005). Molecular tools are needed for accurate identification of the species in this group (Reece et al. 2008).

The genus name Magallana was proposed for Pacific members of the genus Crassostrea, based on genetic divergence between Pacific and Atlantic oysters of the genus (Salvi et al. 2014). 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). Currently, the genus name Magallana is largely used in European literature. A recent article (Salvi et al. 2020) provides sufficient phylogenetic justification for adopting the name Magallana for the northwest Pacific oysters formerly included in 'Crassostrea.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Mollusca
Class:   Bivalvia
Subclass:   Pteriomorphia
Order:   Ostreoida
Family:   Ostreidae
Genus:   Crassostrea
Species:   ariakensis

Synonyms

Crassostrea rivularis (Gould, 1861)
Ostrea ariakensis (Fujita, 1913)
Ostrea discoidea (Gould, 1850)
Ostrea rivularis (Gould, 1861)
Mageallana ariakensis (Salvi & Marriotini, 2016)
Crassostrea ariakensis ((Fujita), 1913)

Potentially Misidentified Species

Magallana angulata
None

Magallana belcheri
None

Magallana gigas
None

Magallana gryphoides
None

Magallana iredalei
None

Magallana madrasensis
None

Magallana nippona
Japan

Magallana plicatula
None

Magallanahongkongensis
China

Magallanasikamea
None

Ecology

General:

Magallana ariakensis, like other oysters, is a protandric hermaphrodite, maturing first as a male, and often becoming female in subsequent seasons. Females release eggs, and male 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 14 days at 28⁰C (Breese and Malouf 1977; Langdon and Robinson 1996; National Research Council 2003). Gonads can develop in M. ariakensis at 2-3 months old and 40-60 mm (National Research Council 2003). Adult oysters are reported to grow to 200-240 mm in length (Carriker and Gaffney 1996).

Magallana ariakensis is characteristic of estuarine habitats in China and Japan, usually occurring in muddy intertidal zones. This oyster normally grows at salinities of 10-30 PSU, and can tolerate brief exposures to salinities as low as 2 PSU (Carriker and Gaffney 1996; Zhou and Allen 2003).

Food:

Phytoplankton

Consumers:

Crabs, snails

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatUnstructured BottomNone
General HabitatOyster ReefNone
General HabitatRockyNone
General HabitatMangrovesNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)2Field observations, China (Zhou and Allen 2003)
Maximum Temperature (ºC)35Field observations, China (Zhou and Allen 2003)
Minimum Salinity (‰)6Field observations, Virginia, very little mortality (Calvo et a1. 2001)
Maximum Salinity (‰)35Field observations, Virginia, very little mortality (Calvo et a1. 2001)
Minimum Reproductive Temperature20Experimental (Breese and Maloof 1977; Langdon and Robinson 1996)
Maximum Reproductive Temperature30Experimental (Breese and Malouf 1977; Langdon and Robinson 1996)
Minimum Reproductive Salinity15Experimental (Breese and Malouf 1977; Langdon and Robinson 1996)
Maximum Reproductive Salinity30Experimental (Breese and Malouf 1977; Langdon and Robinson 1996)
Minimum Duration14Experimental (28 C, Breese and Malouf 1977; Langdon and Robinson 1996)
Maximum Length (mm)240Carikker and Gaffney 1996
Broad Temperature RangeNoneCold temperate-Subtropical
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Economic Impacts

Magallana ariakensis supports regional fisheries in China, southern Japan, and probably elsewhere throughout its native range (Hallerman et al. 2001; Wu and Pan 2000). This oyster was accidentally introduced to the West Coast of North America with stocks of M. gigas (Pacific Oyster). It has been cultured at a few hatcheries in Washington and Oregon since the 1970s. Its advantages over the more widely cultured M. gigas include better quality in summer, superior flavor, and wider temperature and salinity tolerances (Langdon and Robinson 1996). However, this oyster has not successfully been introduced to the wild outside Asia, so the impacts listed below are potential and have not been realized.

Fisheries- In 1994, Magallana ariakensis was imported to hatcheries in France to test its potential as a replacement for M. gigas, in the event of a catastrophic disease outbreak. However, the discovery of a Bonamia-like parasite in these oysters led to the abandonment of work with this species (Cochennec et al. 1998).

Ecological Impacts

On the East Coast, wild Magallana ariakensis existed only as experimentally planted sterile triploids. These oysters were monitored for reversion to diploid (potentially fertile) condition (Calvo et al. 1999; Allen 2000; Luckenbach 1999). Their interactions with native biota were studied, since little was known of this oyster even in its native habitat. In April 2009, the decision was made to end experiments with triploid M. ariakensis in open Chesapeake Bay waters (Fahrenthold 2009; Wheeler 2009).

Food/Prey- Magallana virginica's formerly abundant larvae and spat, as well as adult pseudofeces production and adult biomass, supported a diverse and dynamic oyster-reef foodweb. The extent to which M. ariakensis could perform this role was investigated in mesocosm experiments (Luckenbach 1999). Experiments indicated that M. ariakensis was much more sensitive than M. virginica to blooms of the toxic dinoflagellates Prorocentrum minimum and Karlodinium veneficium (Brownlee et al. 2008).

The faster growth of M. ariakensis is accompanied by a thinner shell. In laboratory experiments, this oyster was more vulnerable to predation by Blue Crabs (Callinectes sapidus) than M. virginica (Bishop and Peterson 2006). In additional experiments, M. ariakensis of varying sizes were more vulnerable than M. virginica to a range of predators, including three species of mud crabs (Xanthidae) and two species of polyclad flatworms, as well as M. sapidus. The native oyster increased its shell strength in response to exudates from predators, but M. ariakensis did not show this response (Newell et al. 2007). However, in experiments, M. ariakensis and M. virginica did not differ in vulnerability to predation by Cownose Rays, a powerful predator on bivalves in Chesapeake Bay (Fisher et al. 2011).

Parasitism- One of the most serious risks of a M. ariakensis introduction is potential transmission of a new disease to Magallana virginica, the native Eastern Oyster. Since the Magallana ariakensis stock introduced in Virginia were raised for several generations in the laboratory, and closely observed, risks of protozoan or bacterial diseases are virtually absent, but a larger, though still low, risk remains from viral diseases (Burreson, in Hallerman et al. 2001). However, several diseases are known from cultured and wild stocks of M. ariakensis. In its native range, along the coast of Guangdong Province, China, oyster populations have been subject to mortalities each year from February to May, affecting about 90% of the population. The causative organism is an intracellular Rickettsia-like prokaryote, infecting the gills, digestive glands, and mantle (Wu and Pan 2000).

In France, mortalities were noted in quarantined M. ariakensis. These were caused by a Bonamia-like parasite affecting the gills and digestive gland. This type of parasite had not previously been seen in oysters of the genus Magallana (Cochennec et al. 1998). In 2003, infections of a Bonamia parasite, resulting in substantial mortality, were seen in triploid M. ariakensis planted in Morehead Sound, North Carolina. The parasite was genetically most similar to known Bonamia species from Australia and New Zealand (Burreson et al. 2004; Bishop et al. 2006). The Bonamia parasite requires high salinities and high temperatures (above 20 ppt and 20⁰C, Audemard et al. 2008). In 2004, it was found at very low prevalence in triploid M. ariakensis at two sites in Chesapeake Bay (Schott et al. 2008).

Hybridization- Magallana ariakensis and M. virginica gametes do not form viable hybrids. Instead, gametes of the two species, released into the water, fuse, but fail to develop. If fertile M. ariakensis were introduced, this could inhibit their recruitment, but conversely, a rapidly growing population of this oyster could adversely affect the native M. virginica. This 'gamete sink' appeared to favor the native oyster, but the magnitude and outcomes of these effects are impossible to predict (Bushek et al. 2008).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NWP-3a None 1913 Native Estab
NWP-2 None 0 Native Estab
NWP-4a None 0 Native Estab
CAR-VII Cape Hatteras to Mid-East Florida 1999 Def Unk
NA-ET3 Cape Cod to Cape Hatteras 1998 Def Unk
NEA-V None 1994 Def Unk
M130 Chesapeake Bay 1998 Def Unk
M128 _CDA_M128 (Eastern Lower Delmarva) 1998 Def Unk
S040 New River 1999 Def Unk
S030 Bogue Sound 2001 Def Unk
S020 Pamlico Sound 2001 Def Unk
NEP-IV Puget Sound to Northern California 1977 Def Failed
P240 Tillamook Bay 1977 Def Failed
P210 Yaquina Bay 1977 Def Failed
NEP-III Alaskan panhandle to N. of Puget Sound 1981 Def Failed
P290 Puget Sound 1981 Def Failed
S010 Albemarle Sound 1999 Def Unk
S045 _CDA_S045 (New) 2001 Def Unk

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude

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