Invasion History

First Non-native North American Tidal Record: 1874
First Non-native West Coast Tidal Record: 1874
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Mya arenaria's current native range is from subarctic Labrador, Canada to Cape Hatteras, North Carolina and sporadically to South Carolina (Abbott 1974; Gosner 1978; Carlton 2023a). Records of M. arenaria in the Northwest Pacific, from the Yellow Sea, China to the Bering Sea (Zenekevich 1963; Golikov et al. 1976) are now referred to the very similar M. japonica, which requires genetic identification.  Two specimens of M. japonica have been identified in Haida Gwaii, British Columbia, but the extent of this species on the West Coast is unknown (Zhang et al. 2018). Based on the fossil record, Mya arenaria originated in the North Pacific Ocean, possibly around Japan, during the Miocene period and soon colonized the Atlantic, reaching the European coast in the late Pliocene, but then dying out during most of its range in the Pleistocene. In Europe, the West Coast, and Alaska, it is absent for prehistoric human shell middens, disregarding some probable misidentifications (Carlton 1979).The surviving populations were on the East Coast of North America, and the East Coast of Asia (Vermeij 1989; Strasser 1999). Mya arenaria appears to be extinct in the Arctic Ocean, though determining its present distribution is complicated by occurrence of subfossil shells and other species of Mya and related genera (Bernard 1979, James T. Carlton, personal communication). Humans have re-introduced M. arenaria to much of its former range, and beyond. Vikings may have transported this clam to Scandinavia as early as the 13th century, and later shipping and food introductions may have moved it to most of the European coast, from the Barents Sea to the Iberian Peninsula (Petersen 1992; Strasser 1999). It is also established in a few estuaries along the Mediterranean Sea (Zenetos et al. 2003) and in the Black Sea (Gomiou et al. 2002). Softshell Clams were apparently introduced to the West Coast with plantings of Eastern Oysters (Crassostrea virginica) by 1874, and were soon deliberately transplanted as food as far north as Alaska (Carlton 1979; Powers 2006). Recent genetic studies support the recent (post-Pleistocene) introduction of Mya arenaria to Europe and the West Coast of North America (Cross et al. 2016; Lasota et al. 2016).

North American Invasion History:

Invasion History on the West Coast:

Mya arenaria was first reported on the West Coast in San Francisco Bay, California in 1874 (as M. hemphilli, Newcomb 1874, cited by Carlton 1979). It rapidly became abundant and widespread in the Bay, supporting fisheries, as early as the 1880s, and spreading as far upstream as Collinsville and Sherman Lake in the Delta (Cohen and Carlton 1995). Some early introductions to other estuaries, such as Coos Bay, Oregon (OR) (~1875, Dall 1897, cited by Carlton 1979) may have also occurred with oyster plantings, but M. arenaria rapidly became a desirable food item, and was planted deliberately. Early plantings occurred in the Siuslaw River, OR; Willapa Bay, Washington (WA) (in 1884, Stearns 1885, cited by Carlton 1979); Grays Harbor, WA (in 1888, Collins 1892, cited by Palacios et al. 2000); Puget Sound, WA (in 1888, introduced from Willapa Bay, Smith 1896, cited by Carlton 1979); the San Juan Islands (Smith 1896, cited by Carlton 1979); Vancouver Island, British Columbia (BC) (Departure Bay, Strait of Georgia; Taylor 1895, cited by Carlton 1979) and Clayoquot Sound, BC (Newcomb 1893, cited by Carlton 1979). In the 20th century, government and individual plantings occurred in many smaller estuaries from California to British Columbia. In California, populations were established from Bolinas Lagoon to Humboldt Bay and Crescent City by 1920-1922, mostly by state stocking (Weymouth 1920, Bonnot 1940, cited Carlton 1979). In Oregon and Washington, first reports of established populations in smaller estuaries are often later (1917-1950s, Edmondson 1922 and Marriage 1953, cited by Carlton 1979), but this may reflect less sampling in this region.

North of Vancouver Island, BC, M. arenaria was collected in the Queen Charlotte Islands in Massett Inlet in 1939 (Carl and Guiguet 1972; Carlton 1979); Prince Rupert in 1955 (Quayle 1960, cited by Carlton 1979); and Ketchikan, Alaska (AK) in 1946 (Hanna 1966; Carlton 1979). As mentioned above, the history of M. arenaria in Alaska is complicated by the presence of subfossil shells of this species and by the occurrence of similar related species. However, excluding some dubious records, it was present at Hooper Bay, AK (61.5°N) by 1924 (Baxter, personal communication, cited by Carlton 1979), and is common in Bristol Bay (58°N) and Norton Sound (64°N) (Bernard 1979), where it may have been present by 1905. Drift shells have been reported as far north as Kotzebue Sound (67ºN; Bernard 1979). Populations are well established south of the Aleutians, in Prince William Sound (Feder and Paul 1974, cited by Carlton 1979; Powers 2006), Kachemak Bay (1999, Hines and Ruiz 2001), and Kodiak Island (Nybakken 1969, cited by Carlton 1979). These northern occurrences probably represent individual, undocumented introductions, rather than long-range larval dispersal (Carlton 1979).

While M. arenaria has been an extremely successful invader, north of San Francisco Bay, it has not become established in several locations to the south. It was introduced to Santa Cruz, California in 1881 (Stearns 1881, cited by Carlton 1979), and to Morro Bay in 1915 (Heath 1916, cited by Carlton 1979), but both stockings failed. Mya arenaria stocked in Elkhorn Slough, before 1911, may have survived for a while, but by the 1990s, it was locally extinct (Wasson et al. 2001). In San Francisco Bay (Nichols and Thompson 1985; Poulton et al. 2004), Grays Harbor, WA (Palacios et al. 2000) and probably elsewhere, the Softshell Clam has undergone great fluctuations in abundance. In South San Francisco Bay, Nichols and Thompson (1985b) considered it to be an ‘irruptive species, appearing in abundance only one year during a 10-year period’. In the upper estuary of San Francisco Bay, from San Pablo to Suisun Bays, M. arenaria shows great spatial patchiness, as well as temporal variation. During dry periods, when salinities are high, it extends its range into Suisun Bay, but disappears from the upper reaches during flood periods (Nichols and Thompson 1985a). In Grays Harbor, soon after introduction in 1895-1897, a massive population explosion took place, followed by catastrophic mortality, leaving extensive 'death assemblages' of shells (Palacios et al. 2000). In general though, M. arenaria has been notably successful in establishing populations in San Francisco Bay and northward. It is capable of surviving and reproducing at lower salinities than native West Coast bivalves and is often the dominant (or only) marine bivalve in upper estuaries. It also tends to occur higher in the intertidal zone than native clams. Consequently, its invasion success may have been due to filling an unoccupied niche (Carlton 1979).
 

Invasion History Elsewhere in the World:

Mya arenaria is present in European fossil deposits from the Pliocene, but is believed to have become extinct in the late Pliocene, and is absent from prehistoric shell middens (Strasser 1999; Behrends et al. 2005). Several specimens in a Danish sand-dune deposit were estimated to date from1245-1295 CE, using C14 dating (Petersen et al. 1992) indicating a possible introduction by Vikings returning from North America. The Softshell Clam appeared very early in Western Europe, with a report from the Netherlands in 1765 (Baster 1765, cited by Wolff 2005). Generally, from Atlantic France to the western Baltic, the date of invasion is unknown, but is believed to be between 1500 and 1700 (Strasser 1999; Goulletquer et al. 2002). Charles Lyell, examining rocks near Stockholm, Sweden, in 1834, noted that this bivalve was abundant in the Baltic, but was absent in fossils, unlike other common Baltic mollusks (Lyell 1835, cited by Munthe 1894). By 1900, it was established along the west and south coasts of Sweden (Swedish Environmental Protection Agency 2006). It is now found in the inner Baltic, including the Gulfs of Bothnia, Riga, and Finland (Strasser 1999; Leppakoski and Olenin 2000; Swedish Environmental Protection Agency 2006; Zaiko et al. 2011; Olenin and Leppakoski 2012). Mya arenaria may have been originally transported with ballast stones or as food, and after its initial introduction, was probably extensively planted as seafood (Strasser 1999). European populations show a low level of genetic diversity, and a relatively homogenous population, indicating rapid gene flow and geographical expansion (Lasota et al. 2004).

In the farther reaches of Europe, Mya arenaria was found on the east coast of Iceland in 1958 (Oskarssen 1961, cited by Strasser 1999). It is established and is absent from the fossil record on the island (Simonarson and Leifsdottir 2009). It is reported to occur all along the coast of Norway, and is abundant in the Oslofjord (Winther and Gray 1985, cited by Strasser 1985), but specific records are not available for the western and northern coast (Strasser 1985). This clam is established in the White Sea (Maximovich and Guerassimova 2003), and to the Barents Sea, east of Svyaty Nos, on the Kola Peninsula (Zenkevich 1963; Galkin 1998). It was present in the White Sea, at least as early as 1963 (Zenkevich 1963; Russanova 1963, cited by Maximovich and Guerassimova 2003).

In the southern part of its European range, M. arenaria is believed to have become established in French waters by 1700 (Goulletquer et al. 2002). However, the first definite record on the Iberian Peninsula was in the Ria de Aveiro, Portugal in 1997 (Conde et al. 2012b). An early record (1988) from the Lima estuary was a misidentification, but M. arenaria is now established in the Lima (2010) and Tagus (2007) estuaries, Portugal (Conde et al. 2009; Conde 2012a). The distribution of the Softshell Clam in the Mediterranean Sea is spotty. Established populations were discovered in two French lagoons, Berre and Vaine in 1976 (Zenetos et al. 2003), and in the Gulf of Saronicos, Greece, in 1984 (Zenetos et al. 2005). Isolated collections were made in Sicily and the Adriatic (Zenetos et al. 2003; not established, Occhipinti-Ambrogi et al. 2011). Mya arenaria was found in the Black Sea in Romania in 1966, and became abundant enough to be regarded as a pest (Gomiou et al. 2002, Skolka and Preda 2010). It has spread to the Sea of Marmara (in 1996, Albayrak and Balcis 1996, cited by Albayrak 2011), and to the Sea of Azov (Savchuck 1980, cited by Zaitsev and Ozturk 2001).


Description

Mya arenaria is a bivalve with a thin, elongate, elliptical shell, gaping at the anterior and posterior ends even when closed. The pallial sinus is deep and somewhat V-shaped. The hinge is asymmetrical, with a long, tongue-shaped chondrophore in the left valve, and a heart-shaped pit on the right. The shell is chalky white with a thin dull-brown or yellowish periostracum. The typical maximum size is 75-100 mm, with a record length of 163 mm. It usually borrows in soft muddy to sandy sediments in shallow waters and intertidal mud flats (Abbott 1974; Gosner 1978; Coan et al. 2000; Coan and Valentich-Scott 2007). In large specimens, the siphon may extend for as much as 200 mm to reach the surface (Newell and Hidu 1986).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Mollusca
Class:   Bivalvia
Subclass:   Heterodonta
Order:   Myoida
Superfamily:   Myoidea
Family:   Myidae
Genus:   Mya
Species:   arenaria

Synonyms

Mya acuta (Say, 1822)
Mya acuta mercenaria (Say, 1822)
Mya alba (Agassiz, 1839)
Mya arenaria corbuloides (Comfort, 1938)
Mya communis (Megerle von Mühlfeld, 1811)
Mya corpulenta (Conrad, 1845)
Mya declivis (Pennant, 1777)
Mya elongata (Locard, 1866)
Mya hemphilli (Newcomb, 1874)
Mya lata (J. Sowerby, 1815)
Mya oonogai (Makiyama, 1935)
Mya subovata (Woodward, 1833)
Mya subtruncata (Woodward, 1833)
Sphenia ovoidea (Carpenter, 1864)
Mya paternalis (Matsumoto, 1930)

Potentially Misidentified Species

Mya japonica
Mya japonica (Japanese Softshell Clam) has been found to be a genetically distinct species, occuring from the Yellow Sea, China, to the Bering Sea, Russia (Golikov et al. 1976; Bernard 1979; Zhang et al. 2018). Morphological differences are small, but M. japonica has a taller shell, with a more rounded posterior end, rougher submarginal wrinkles, and a more impressed pallial line. However, morphological variability is high. Genetic analysis and spermatozoon morphology indicates that the two species diverged 4.1-12.5 Myr ago (Zhang et al. 2018). Two specimens of M. japonica have been collected in British Columbia, the first record of this species in the Eastern Pacific (Zhang et al. 2018).

Mya truncata
Mya truncata is native to the Arctic Ocean. The extent of its range into the temperate Atlantic and Pacific has been obscured by its similarity to M. arenaria (Carlton 1979; Strasser 1999; Zhang et al. 2018). Genetic analysis suggests that it is a species complex (Zhang et al. 2018

Mya uzenensis
Mya uzenensis (Siberian Softshell Clam) is native to Alaska and northeast Russia (Zhang et al. 2018).

Ecology

General:

Mya arenaria is a large bivalve which inhabits gravelly to muddy bottoms, from the mid-intertidal to about 100 m depth, though they are rare below 9-10 m. In regions with large tidal ranges, they are most-abundant in intertidal mudflats (Gosner 1978; Newell and Hidu 1986). They require temperatures above 12-15°C for spawning, but do not tolerate temperatures above 28°C for prolonged periods (Newell and Hidu 1986). Mya arenaria is unusually tolerant of low salinities, and can be acclimated to feed at 3 PSU (Castagna and Chanley 1973). In estuaries such as Chesapeake Bay and brackish seas such as the Baltic, Softshell Clams can be abundant at salinities as low as 4-5 PSU, while at marine salinities (25-25 PSU), predation may reduce their abundance in subtidal waters (Newell and Hidu 1986; Carlton 1979). Sexes are usually separate, but there is a low incidence of hermaphroditism. Size appears more important than age in determining maturity. Maturity occurs at about 20 mm length, while market size is about 50 mm. Market size is reached in about 1.5 years in Connecticut, 3-6 years in Maine, 5 years in New Brunswick and the White Sea (Sadykhova 1979; Newell and Hidu 1986).

Mya arenaria usually spawns twice a year in spring and fall, mostly in the southern part of its range (Connecticut, Rhode Island, but also in Oslofjord, Norway and southern England), but once a year mostly further north (White Sea- Russia, Maine, New Brunswick, Ireland, Sweden, Wadden Sea, but also the Black Sea) Spawning usually occurs at 10-25ºC, but the temperature range is quite variable (Sadykhova 1979; Newell and Hidu 1986; Strasser 1999; Cross et al. 2012). Reported fecundity ranges from about 100,000 to 3 million eggs (Newell and Hidu 1986). The eggs and sperm are released through the exhaling siphon. Fertilized eggs develop into trochophore larvae within 9 hours and a few hours later they grow their first shell (called 'D-shaped', or 'straight-hinged). The larvae swim and feed on phytoplankton, using a ciliated velum. At about 12-20 days, and 175-230 μm, they develop a ciliated foot, and begin to investigate substrates for settlement (Chanley and Andrews 1971; Newell and Hidu 1986). At the end of this pediveliger stage, the velum is lost and the larvae settle, moving by crawling, and attaching to grains of sand or sediment, seaweeds or surfaces, using byssal threads. As the clams grow, they burrow deeper, the siphons elongate, and the byssus glands atrophy. At about 5 mm size, clams are called 'seed'. As they grow, they tend to move shoreward (Newell and Hidu 1986). Mortality is very high for larvae and seed clams, but once clams reach adult size, a life span of 10 years is typical, with some specimens living for 20 years (Strasser 1999).

Softshell clams are suspension feeders and can burrow up to 20 cm (in large specimens), with their siphon protruding above the surface. They draw water through the incurrent siphon, to the gills, where food particles are trapped in mucus and carried by cilia to the mouth to be ingested. Particles which are too large or inedible, or simply too dense for ingestion, are rejected by the labial palps as pseudofeces. Diatoms and flagellates are optimal food, but clams can obtain some nutrition from suspended detritus. Feeding rates are influenced by temperature, salinity, and food quality. Filtration and assimilation drops to very low levels below 3ºC. These clams are able to feed in water with considerable quantities of suspended silt and are able to sort cells for silt particles before ingestion (Newell and Hidu 1986). Larvae and newly settled spat are very vulnerable to predation. Small clams are eaten by fishes, crabs, clam worms (Nereidae), moon snails (Naticidae), birds, etc. When clams reach ~60 mm in length, they are less vulnerable to predation (Newell and Hidu 1986).

Food:

Phytoplankton

Consumers:

crabs, fishes, birds, humans

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatGrass BedNone
General HabitatUnstructured BottomNone
General HabitatOyster ReefNone
General HabitatSalt-brackish marshNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Tidal RangeMid IntertidalNone
Vertical HabitatEndobenthicNone

Life History


Tolerances and Life History Parameters

Minimum Temperature (ºC)0Based on range (Abbott 1974).
Maximum Temperature (ºC)32.5Experimental, 24 hr LC 50 (Kennedy and Mihursky 1971).
Minimum Salinity (‰)3Experimental, acclimation (Castagna and Chanley 1973)
Maximum Salinity (‰)35Based on field occurences (Castagna and Chanley 1973)
Minimum Reproductive Temperature4Season and temperature of spawning is highly variable- a Massachusetts population spawned at 4-6 C (Brousseau 1979, cited by Strasser 1999), while larvae in the laboratory developed poorly below 8 C (Stickney 1979, cited by Strasser 1999).
Maximum Reproductive Temperature23Upper limit for optimal development in the laboratory (Stickney 1964, cited by Strasser 1999).
Minimum Reproductive Salinity10Stickney (1965), cited by Castagna and Chanley (1973)
Maximum Reproductive Salinity35Stickney (1965), cited by Castagna and Chanley (1973)
Minimum Duration10Larval duration, laboratory- Loosanoff and Davies 1963, cited by Strasser 1999
Maximum Duration35Larval duration, laboratory- Loosanoff and Davies 1963, cited by Strasser 1999
Minimum Length (mm)20Minimum size at first sexual maturity (Newell and Hidu 1986)
Maximum Length (mm)163But more usually, up to 100 mm (Abbott 1974; Gosner 1978)
Broad Temperature RangeNonePolar-Warm temperate
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Mya arenaria is an important shellfish species in its native range, from Atlantic Canada to Chesapeake Bay, supporting both commercial and recreational fisheries. On the West Coast of the US, it supported commercial fisheries in San Francisco Bay and elsewhere historically, but is now mainly taken by recreational clammers (Cohen and Carlton 1995). Surprisingly, it is apparently rarely eaten in Europe, and may be more frequently used as bait (Eno et al. 1997; Strasser 1999). Where it is abundant, it is an important suspension-feeder, grazing phytoplankton, and an important food item for fishes, invertebrates, and birds (Nichols and Thompson 1985a; Zaiko et al. 2011) It is also a potential competitor with native bivalves (Moller 1986; Conde et al. 2011).

Economic Impacts

Fisheries- Mya arenaria is an important commercial fisheries species, eaten steamed or fried in eastern North America. Intertidal populations in New England and the Maritimes are harvested with rakes, forks, or hoes, while subtidal populations in Chesapeake and Delaware Bays are taken with hydraulic dredges (Newell and Hidu 1986). In San Francisco Bay, they supported a commercial fishery from the 1880s to 1948, but the fishery steadily declined by 1926 and ended by 1948 (Cohen and Carlton 1995). Elsewhere on the West Coast, the fishery has been mostly recreational, as indicated by state agency websites. Native clams and the Japanese Littleneck (Venerupis philippinarum) are often preferred to Softshell Clams. However, at least one culture operation is taking place in Skagit Bay, WA (Washington Department of Fish and Wildlife 2012, http://wdfw.wa.gov/fishing/shellfish/clams/eastern_softshell.html). In Europe, it is not frequently eaten, and does not support commercial fisheries. Web searches for it under the English name 'Sand Gaper' and 'fisheries', 'fishing', etc., turned up only references to using it as bait on a recreational basis (e.g., http://www.ukmarinesac.org.uk/activities/bait-collection/bc1_1.htm). In the Black Sea, it became very abundant about 4-5 years after its original discovery. When large masses of clams washed ashore, they were fed to chickens (Gomiou et al. 2002). Similarly, in the Sea of Marmars, Turkey, it is of 'no commercial importance', except as a food for larger fishes (Ozturk 2002).

Aesthetic- Soon after its invasion in the Black Sea, by the 1970s, masses of decaying M. arenaria began washing ashore, attracting masses of seagulls (Gomiou et al. 2002). Mass early occurrences and mortalities are also known from Grays Harbor, WA in the late 1800s, though aesthetic impacts were not reported (Palacios et al. 2000).

Ecological Impacts

Herbivory- When abundant, Mya arenaria is a significant herbivore in estuaries, because of its large size and powerful filtration, and its ability to survive in low salinities and wide tidal ranges, where large native bivalves are often rare. Estimated feeding rates of M. arenaria in the southwestern Baltic Sea, off Germany, indicate that this clam can filter the entire water column once or several times a day, depending on water depth (Forster and Zettler 2004). Large biomasses in San Francisco Bay (Nichols and Thompson 1985a; Nichols and Thompson 1985b), the Skagerrak (Moller 1986), the Baltic (Bubinas and Vaitonis 2003; Forster and Zettler 2004; Obolewski and Piesik 2005; Zaiko et al. 2011), and Black Sea (Gomiou et al. 2002) imply significant feeding rates.

Competition- Introduced populations of Mya arenaria in several locations are believed to have reduced or partially replaced native bivalves, including Macoma nasuta (Bent-Nose Macoma) in San Francisco Bay (Cohen and Carlton 1995), Macoma balthica in the Baltic Sea (Obolewski and Piesik 2005), Lentidium mediterraneum in the Black Sea (Skolka and Preda 2010), and Cerastoderma edule (Edible Cockle) in the Skagerrak, Sweden (Moller 1986). In the case of C. edule, competition was reciprocal, with one species or the other having heavy recruitment in some years, and inhibiting recruitment of the other (Moller 1986).

Food/Prey- Mya arenaria, when abundant, has been an important prey organism for clam worms (Nereidae), predatory snails, shrimps, crabs, fishes, ducks, and shorebirds in invaded regions (Carlton 1979; Sadykhova 1979; Ozturk 2002; Bubinas and Vaitonis 2003; Cloern et al. 2007; Skolka and Preda 2010). Because it tolerates low salinities and wide tidal ranges better than many native clams, it has the potential to increase the food supply for predators in estuaries.

Habitat Change- Mya arenaria, as a powerful burrower and filterer, has the potential to alter habitats and sediment characteristics through bioturbation and deposition of peudofeces and also through suspension feeding, increasing water clarity, and light penetration (Obolewski and Piesik 2005; Queiros et al. 2011; Zaiko et al. 2011). Introduced populations of Mya arenaria have often gone through boom-and bust phases, leaving 'death assemblages' of empty shells, providing habitat for many other benthic organisms (Strasser 1999; Palacios et al. 2000).

Trophic Cascade- During periods of exceptional abundance, Mya arenaria may have effects throughout the food web, affecting phytoplankton abundance, and in turn, zooplankton, mysids, and fish recruitment. This may have happened in 1976-1977 in Suisun Bay, California (Nichols and Thompson 1985b; Cohen and Carlton 1995). High abundances of Mya arenaria during 'boom' periods, or its empty shells during 'busts,' can affect the abundance of predators with implications for other benthic organisms. For example, high abundances of M. arenaria shells supported elevated abundances of juvenile Dungeness Crabs (Metacarcinus magister) in Grays Harbor, WA which could lead to increased predation on other benthic organisms (Palacios et al. 2000).


Regional Impacts

NEP-VNorthern California to Mid Channel IslandsEconomic ImpactFisheries
By the 1880s, M. arenaria supported a commercial fishery of 500-900 tons per year in San Francisco Bay, but this declined to 100 tons per year by 1916 to 1926, and ended after 1948, due to overharvesting, pollution, and possible preference for Venerupis phillipinarum (Japanese Littleneck). However, recreational harvests continue to the present (Cohen and Carlton 1995). Extensive plantings were carried out along the California coast by individuals and the California Department of Fish and Game (Weymouth 1920; Bonnot 1940, both cited by Carlton 1979). Recreational clamming probably occurs in many other estuaries where clams are common.
NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
Mya arenaria may have replaced Macoma nasuta in clam beds in San Francisco Bay (Cohen and Carlton 1995).
P090San Francisco BayEconomic ImpactFisheries
By the 1880s, M. arenaria supported a commercial fishery of 500-900 tons per year in San Francisco Bay, but this declined to 100 tons per year by 1916 to 1926, and ended after 1948, due to overharvesting, pollution, and possible preference for Venerupis phillipinarum (Japanese Littleneck). However, recreational harvests continue to the present (Cohen and Carlton 1995).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactHerbivory
Mya arenaria, when abundant, has had significant impact as a filter-feeder. During periods of high salinity, it has been one of several filter-feeders contributing to low phytoplankton biomass in Suisun Bay (Nichols and Thompson 1985a).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactFood/Prey
Mya arenaria is an important prey organism for ducks, shorebirds, flounders, skates, rays, and native crabs and shrimps (Carlton 1979; Cloern et al. 2007).
P090San Francisco BayEcological ImpactCompetition
Mya arenaria may have replaced Macoma nasuta in clam beds in San Francisco Bay (Cohen and Carlton 1995).
P090San Francisco BayEcological ImpactHerbivory
Mya arenaria, when abundant, has had significant impact as a filter-feeder. During periods of high salinity in 1976-1977, it has been one of several filter-feeders contributing to low phytoplankton biomass in Suisun Bay (Nichols and Thompson 1985a).
P090San Francisco BayEcological ImpactFood/Prey
Mya arenaria is an important prey organism for ducks, shorebirds, flounders, skates, rays, and native crabs and shrimps (Carlton 1979; Cloern et al. 2007).
NEP-IVPuget Sound to Northern CaliforniaEconomic ImpactFisheries
In Humboldt Bay. 'It is taken for bait and food by sport clammers.' (Boyd et al. 2002). Recreational clamming for M. arenaria is also popular in Oregon. According to the Oregon Division of Fish and Wildlife, this clam is present in nearly every Oregon estuary (http://www.dfw.state.or.us/mrp/shellfish/bayclams/dig_softshell.asp). In Washington, they are less popular than Butter Clams (Saxidomus gigantea) or Littlenecks (Leukoma staminea- Pacific Littleneck; Venerupis philippinarum- Japanese Littleneck) (Washington Department of Fish and Wildlife 2012, http://wdfw.wa.gov/fishing/shellfish/clams/eastern_softshell.html).
P090San Francisco BayEcological ImpactTrophic Cascade
During a drought in 1976-1977 in Suisun Bay, a high abundance of Mya arenaria and other marine filter-feeders may have contributed to a low phytoplankton abundance, which in turn contributed to low zooplankton abundance and a scarcity of the omnivorous Neomysis mercedis, an important food for juvenile fishes. This, in turn, may have led to decreased recruitment of Morone saxatilis (Striped Bass), an economically important introduced gamefish (Nichols and Thompson 1985b; Cohen and Carlton 1995).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactTrophic Cascade
During a drought in 1976-1977 in Suisun Bay, a high abundance of Mya arenaria and other marine filter-feeders may have contributed to a low phytoplankton abundance, which in turn contributed to low zooplankton abundance and a scarcity of the omnivorous Neomysis mercedis, an important food for juvenile fishes. This, in turn, may have led to decreased recruitment of Morone saxatilis (Striped Bass), an economically important introduced gamefish (Nichols and Thompson 1985b; Cohen and Carlton 1995).
NEP-IVPuget Sound to Northern CaliforniaEcological ImpactHabitat Change
In Grays Harbor WA, large shell deposits provide a highly favorable habitat for settling Dungeness Crab (Metacarcinus magister) juveniles. High densities of these crabs may, in turn, limit the recruitment of M. arenaria).
CACaliforniaEcological ImpactCompetition
Mya arenaria may have replaced Macoma nasuta in clam beds in San Francisco Bay (Cohen and Carlton 1995)., Mya arenaria may have replaced Macoma nasuta in clam beds in San Francisco Bay (Cohen and Carlton 1995).
CACaliforniaEcological ImpactFood/Prey
Mya arenaria is an important prey organism for ducks, shorebirds, flounders, skates, rays, and native crabs and shrimps (Carlton 1979; Cloern et al. 2007)., Mya arenaria is an important prey organism for ducks, shorebirds, flounders, skates, rays, and native crabs and shrimps (Carlton 1979; Cloern et al. 2007).
CACaliforniaEcological ImpactHerbivory
Mya arenaria, when abundant, has had significant impact as a filter-feeder. During periods of high salinity, it has been one of several filter-feeders contributing to low phytoplankton biomass in Suisun Bay (Nichols and Thompson 1985a)., Mya arenaria, when abundant, has had significant impact as a filter-feeder. During periods of high salinity in 1976-1977, it has been one of several filter-feeders contributing to low phytoplankton biomass in Suisun Bay (Nichols and Thompson 1985a).
CACaliforniaEcological ImpactTrophic Cascade
During a drought in 1976-1977 in Suisun Bay, a high abundance of Mya arenaria and other marine filter-feeders may have contributed to a low phytoplankton abundance, which in turn contributed to low zooplankton abundance and a scarcity of the omnivorous Neomysis mercedis, an important food for juvenile fishes. This, in turn, may have led to decreased recruitment of Morone saxatilis (Striped Bass), an economically important introduced gamefish (Nichols and Thompson 1985b; Cohen and Carlton 1995)., During a drought in 1976-1977 in Suisun Bay, a high abundance of Mya arenaria and other marine filter-feeders may have contributed to a low phytoplankton abundance, which in turn contributed to low zooplankton abundance and a scarcity of the omnivorous Neomysis mercedis, an important food for juvenile fishes. This, in turn, may have led to decreased recruitment of Morone saxatilis (Striped Bass), an economically important introduced gamefish (Nichols and Thompson 1985b; Cohen and Carlton 1995).
CACaliforniaEconomic ImpactFisheries
By the 1880s, M. arenaria supported a commercial fishery of 500-900 tons per year in San Francisco Bay, but this declined to 100 tons per year by 1916 to 1926, and ended after 1948, due to overharvesting, pollution, and possible preference for Venerupis phillipinarum (Japanese Littleneck). However, recreational harvests continue to the present (Cohen and Carlton 1995). Extensive plantings were carried out along the California coast by individuals and the California Department of Fish and Game (Weymouth 1920; Bonnot 1940, both cited by Carlton 1979). Recreational clamming probably occurs in many other estuaries where clams are common., By the 1880s, M. arenaria supported a commercial fishery of 500-900 tons per year in San Francisco Bay, but this declined to 100 tons per year by 1916 to 1926, and ended after 1948, due to overharvesting, pollution, and possible preference for Venerupis phillipinarum (Japanese Littleneck). However, recreational harvests continue to the present (Cohen and Carlton 1995).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P095 _CDA_P095 (Tomales-Drakes Bay) 1922 Non-native Established
P116 _CDA_P116 (Big Navaro-Garcia) 1920 Non-native Established
P117 _CDA_P117 (Mattole) 1920 Non-native Established
P120 Eel River 1920 Non-native Established
P135 _CDA_P135 (Mad-Redwood) 1920 Non-native Established
P143 _CDA_P143 (Smith) 1920 Non-native Established
P105 _CDA_P105 (Tomales-Drakes Bay) 1919 Non-native Established
P100 Drakes Estero 1919 Non-native Established
P130 Humboldt Bay 1917 Non-native Established
P110 Tomales Bay 1916 Non-native Established
P112 _CDA_P112 (Bodega Bay) 1916 Non-native Established
P070 Morro Bay 1915 Non-native Failed
P093 _CDA_P093 (San Pablo Bay) 1895 Non-native Established
P080 Monterey Bay 1881 Non-native Extinct
NEP-IV Puget Sound to Northern California 1875 Non-native Established
P090 San Francisco Bay 1874 Non-native Established
NEP-V Northern California to Mid Channel Islands 1874 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
697173 ISS 2000-2002 Survey Data 2001 2001-09-19 Tomales Bay Infaunal 01 Non-native 38.2062 -122.9381
697278 Introduced Species Study 2005 2005-11-14 Cal Maritime Academy/Vallejo Non-native 38.0661 -122.2299
697884 Bonnot 1932 1932 Suisun Bay - Martinez Non-native 38.0287 -122.1333
697982 Stearns 1881 1881 Santa Cruz, Monterey Bay Non-native 36.9552 -122.0106
697997 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-24 Fruitvale Bridge, San Francisco Bay Non-native 37.7690 -122.2296
698184 Markmann 1986 1986 Suisun Bay off Middle Point near Nichols (Station D8) Non-native 38.0599 -121.9900
698450 Boyd et al. 2002 (Humboldt Bay Report) 2002 Samoa Channel HB, St. 13 Non-native 40.8195 -124.1733
699261 Introduced Species Study 2010 2010-06-02 Oakland Inner Harbor - Shipping cranes Non-native 37.7947 -122.3095
699262 Introduced Species Study 2005 2005-06-07 Oakland Inner Harbor - Shipping cranes Non-native 37.7947 -122.3095
699299 Introduced Species Study 2005 2005-10-20 San Pablo Bay Pumphouse Non-native 38.0446 -122.4326
699842 Introduced Species Study 2005 2005-06-09 Paradise Area Non-native 37.9062 -122.4768
699868 Introduced Species Study 2010 2010-05-31 Redwood Creek - Marina Non-native 37.5021 -122.2130
699988 Introduced Species Study 2005 2005-06-09 McNears Beach Non-native 37.9962 -122.4556
699989 Introduced Species Study 2010 2010-06-12 McNears Beach Non-native 37.9962 -122.4556
700056 Boyd et al. 2002 (Humboldt Bay Report) 2002 Southport Landing Non-native 40.6952 -124.2494
700316 Introduced Species Study 2010 2010-06-03 Berkeley Flats/Berkeley Pier Non-native 37.8600 -122.3256
700355 Bonnot 1932 1932 San Pablo Bay - Napa River Non-native 38.1402 -122.2764
700509 Introduced Species Study 2005 2005-06-08 Sea Plane Lagoon Non-native 37.7761 -122.2998
700679 Boyd et al. 2002 (Humboldt Bay Report) 2002 Jacoby Creek Non-native 40.8435 -124.0838
700934 Bonnot 1932 1932 South San Francisco Bay Non-native 37.5457 -122.1645
701078 Stearns 1881 1881 San Francisco Bay Non-native 37.8494 -122.3681
701079 Newcomb 1874; Stearns 1881, 1900. 1874 Alameda County shoreline Non-native 37.8494 -122.3681
701384 Boyd et al. 2002 (Humboldt Bay Report) 2002 Mad River Slough Channel St. 11 Non-native 40.8273 -124.1649
701424 Introduced Species Study 2010 2010-06-30 Hercules Wharf Non-native 38.0231 -122.2928
701425 Introduced Species Study 2005 2005-10-19 Hercules Wharf Non-native 38.0231 -122.2928
701519 Markmann 1986 1986 Sherman Lake near Antioch (Station D11) Non-native 38.0422 -121.7995
702568 Introduced Species Study 2005 2005-07-08 Richmond Marina Non-native 37.9137 -122.3504
702780 Bonnot 1932 1932 Central San Francisco Bay Non-native 37.8595 -122.3884
702897 Boyd et al. 2002 (Humboldt Bay Report) 2002 Bracut Non-native 40.8313 -124.0845
703291 Introduced Species Study 2005 2005-11-15 China Camp Non-native 38.0025 -122.4617
703292 Introduced Species Study 2010 2010-06-12 China Camp Non-native 38.0025 -122.4617
703510 ISS 2000-2002 Survey Data 2001 2001-09-19 Tomales Bay Infaunal 02 Non-native 38.2067 -122.9392
703595 Introduced Species Study 2005 2005-06-10 Toll Plaza Non-native 37.8266 -122.3166
703783 Introduced Species Study 2010 2010-06-13 Hayward Landing Non-native 37.6447 -122.1543
703788 Introduced Species Study 2005 2005-06-10 Hayward Landing Non-native 37.6447 -122.1543
704271 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-23 Brisbane Lagoon, San Francisco Bay Non-native 37.6862 -122.3906
704690 California Department of Fish and Game 1916 1915 Morro Bay Non-native 35.3500 -120.8500
716873 MacGinitie 1935 1927 Elkhorn Slough Non-native 36.8091 -121.7860
716876 Markmann 1986 1986 Sacramento River above Point Sacramento (Station D4) Non-native 38.0622 -121.8179
716878 Thompson and Nichols 1984 1975 Sand Point, Palo Alto Non-native 37.4630 -122.1011
716880 Robinson et al. 2011 2005 China Camp Non-native 38.0008 -122.4616
716881 Bonnot 1940, and A.G. Smith Collection, both cited in Carlton 1979a 1922 Bolinas Lagoon Non-native 37.9183 -122.6811
716882 Weymouth 1920 1919 Drakes Estero Non-native 38.0474 -122.9422
716883 California Department of Fish and Game 1916 1916 Tomales Bay Non-native 38.1285 -122.8730
716884 Packard 1918 1916 Bodega Harbor Non-native 38.3235 -123.0478
716885 Weymouth 1920 1919 Navarro River mouth Non-native 39.1921 -123.7611
716887 Weymouth 1920 1919 Eel River mouth Non-native 40.6415 -124.3123
716888 California Department of Fish and Game 1917; Weymouth 1920 1917 Humboldt Bay Non-native 40.7498 -124.2095
716890 Weymouth 1920 1919 Stone Lagoon Non-native 41.2448 -124.0925
716892 Weymouth 1920 1919 Big Lagoon Non-native 41.1752 -124.1147
716893 Weymouth 1920 1919 Lake Earl Non-native 41.8257 -124.1887
716894 Monroe et al. 1975; C.D. Snow (Oregon Fish Commission), pers. comm. 1977, in Carlton 1979 1975 Smith River Delta Non-native 41.9310 -124.1985
759942 Cooper 1886 1886 San Francisco Bay Non-native 37.8494 -122.3681
759943 Weymouth 1920 1919 Humboldt Bay Non-native 40.7498 -124.2095
759944 Weymouth 1920 1919 Ten Mile River mouth Non-native
759945 Weymouth 1920 1919 Big River mouth Non-native 39.3022 -123.7934
759946 Weymouth 1920 1919 Bodega Harbor Non-native 38.3235 -123.0478
759947 Weymouth 1920 1919 Estero Americano Non-native 38.3081 -122.9845
759948 Weymouth 1920 1919 Tomales Bay Non-native 38.1285 -122.8730
759949 Weymouth 1920 1919 Abbotts Lagoon Non-native 38.1142 -122.9527
759950 Weymouth 1920 1919 San Francisco Bay Non-native 37.8494 -122.3681
759951 A.G. Smith Collection, cited in Carlton 1979a 1924 Big Lagoon Non-native 41.1708 -124.1272
759952 MacGinitie 1935 1935 Elkhorn Slough at Highway 1 Bridge (Station 7) Non-native 36.8093 -121.7848
759953 MacGinitie 1935 1935 Old Salinas River between Sandholdt Road Bridge and mouth of Elkhorn Slough (Station 8) Non-native 36.8029 -121.7859
759954 Baily 1932 1932 Lake Merritt Non-native 37.8025 -122.2578
759955 Bonnot 1932 1932 Tomales Bay Non-native 38.1285 -122.8730
759956 Bonnot 1932 1932 Bodega Bay Non-native 38.3262 -123.0495
759957 Bonnot 1940 1940 Humboldt Bay Non-native 40.7498 -124.2095
759958 Bonnot 1940 1940 Bodega Bay Non-native 38.3262 -123.0495
759959 Bonnot 1940 1940 Tomales Bay Non-native 38.1285 -122.8730
759960 Bonnot 1940 1940 Bolinas Bay [sic] Non-native 37.9183 -122.6811
759961 Bonnot 1940 1940 San Francisco Bay Non-native 37.8494 -122.3681
759962 Bonnot 1940 1940 Elkhorn Slough Non-native 36.8086 -121.7856
759963 Graham and Gay 1945 1941 Fruitvale Avenue Bridge Non-native 37.7689 -122.2296
759964 Ganssle 1966 1963 San Pablo Bay Non-native 38.0600 -122.3900
759965 Ganssle 1966 1964 San Pablo Bay Non-native 38.0600 -122.3900
759966 Standing et al. 1975 1975 Bodega Harbor Non-native 38.3235 -123.0478
759967 Markmann 1986 1986 Suisun Bay off Bulls Head Point (Station D6) Non-native 38.0420 -122.1220
759968 Markmann 1986 1986 Grizzly Bay at Dolphin (Station D7) Non-native 38.1171 -122.0396
759969 Markmann 1986 1986 Honker Bay near Wheeler Point (Station D9 Non-native 38.0792 -121.9303
759970 Barnhart et al. 1992 1992 Humboldt Bay Non-native 40.7498 -124.2095
759971 McDonald 1969a 1965 Arcata (Humboldt Bay) Non-native 40.8500 -124.1000
759972 McDonald 1969a 1965 Millerton Marsh, Tomales Bay Non-native 38.1072 -122.8411
759973 McDonald 1969a 1965 Elkhorn Slough General Location Non-native 36.8086 -121.7856
759974 Wicksten 1978 1978 Coyote Point Non-native 37.5922 -122.3210
759975 Cohen and Chapman 2005 2005 2005-11-27 Dolphin # 11 Non-native 38.0530 -122.3307
759976 Chapman and Dorman 1975 1971 1971-02-20 Pinole Point Non-native 38.0133 -122.3659
759977 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 1 (Deep) Non-native 38.0125 -122.3928
759978 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 1 (Deep) Non-native 38.0125 -122.3928
759979 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 1 (Deep) Non-native 38.0125 -122.3928
759980 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 1 (Deep) Non-native 38.0125 -122.3928
759981 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 1 (Deep) Non-native 38.0125 -122.3928
759982 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 1 (Deep) Non-native 38.0125 -122.3928
759983 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 1 (Deep) Non-native 38.0125 -122.3928
759984 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 11 (16-18') Non-native 38.0169 -122.4258
759985 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 11 (16-18') Non-native 38.0169 -122.4258
759986 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 11 (16-18') Non-native 38.0169 -122.4258
759987 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 11 (16-18') Non-native 38.0169 -122.4258
759988 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 2 (Deep) Non-native 38.0378 -122.3619
759989 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 2 (Deep) Non-native 38.0378 -122.3619
759990 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 2 (Deep) Non-native 38.0378 -122.3619
759991 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 2 (Deep) Non-native 38.0378 -122.3619
759992 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 2 (Deep) Non-native 38.0378 -122.3619
759993 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 2 (Deep) Non-native 38.0378 -122.3619
759994 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
759995 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
759996 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
759997 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
759998 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
759999 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
760000 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
760001 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
760002 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
760003 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 12 (16-18') Non-native 38.0506 -122.3625
760004 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 13 (16-18') Non-native 38.0522 -122.1778
760005 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 13 (16-18') Non-native 38.0522 -122.1778
760006 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 13 (16-18') Non-native 38.0522 -122.1778
760007 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 13 (16-18') Non-native 38.0522 -122.1778
760008 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760009 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760010 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760011 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760012 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760013 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760014 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760015 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760016 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760017 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 3 (Deep; off Commodore Jones Point) Non-native 38.0542 -122.1756
760018 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760019 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760020 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760021 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760022 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760023 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760024 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760025 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760026 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760027 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 23 (6-8') Non-native 38.0547 -122.1744
760028 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 14 (16-18') Non-native 38.0597 -122.0764
760029 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 14 (16-18') Non-native 38.0597 -122.0764
760030 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 14 (16-18') Non-native 38.0597 -122.0764
760031 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 14 (16-18') Non-native 38.0597 -122.0764
760032 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 14 (16-18') Non-native 38.0597 -122.0764
760033 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 24 (6-8') Non-native 38.0606 -122.0769
760034 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 24 (6-8') Non-native 38.0606 -122.0769
760035 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 24 (6-8') Non-native 38.0606 -122.0769
760036 Painter 1966b; Hopkins 1986 1963 Suisun Bay, near Point Edith, Station 24 (6-8') Non-native 38.0606 -122.0769
760037 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 33 (Intertidal; Southampton Bay) Non-native 38.0642 -122.1872
760038 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 33 (Intertidal; Southampton Bay) Non-native 38.0642 -122.1872
760039 Painter 1966b; Hopkins 1986 1963 Carquinez Strait, Station 33 (Intertidal; Southampton Bay) Non-native 38.0642 -122.1872
760040 Painter 1966b; Hopkins 1986 1963 Suisun Bay near Ryer Island, Station 26 (6-8') Non-native 38.0697 -122.0047
760041 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760042 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760043 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760044 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760045 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760046 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760047 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760048 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760049 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760050 Painter 1966b; Hopkins 1986 1963 San Pablo Bay West, Station 21 (6-8') Non-native 38.0889 -122.4636
760051 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760052 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760053 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760054 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760055 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760056 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760057 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760058 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760059 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 22 (6-8') Non-native 38.0900 -122.3569
760060 Painter 1966b; Hopkins 1986 1963 Suisun Bay North, Station 5 (Deep) Non-native 38.0972 -122.0669
760061 Painter 1966b; Hopkins 1986 1963 Suisun Bay North, Station 5 (Deep) Non-native 38.0972 -122.0669
760062 Painter 1966b; Hopkins 1986 1963 Suisun Bay North, Station 5 (Deep) Non-native 38.0972 -122.0669
760063 Painter 1966b; Hopkins 1986 1963 Suisun Bay North, Station 15 (16-18') Non-native 38.1000 -122.0528
760064 Painter 1966b; Hopkins 1986 1963 Suisun Bay North, Station 25 (6-8'; in Grizzly Bay) Non-native 38.1161 -122.0397
760065 Painter 1966b; Hopkins 1986 1963 Suisun Bay North, Station 25 (6-8'; in Grizzly Bay) Non-native 38.1161 -122.0397
760066 Painter 1966b; Hopkins 1986 1963 Suisun Bay North, Station 25 (6-8'; in Grizzly Bay) Non-native 38.1161 -122.0397
760067 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 32 (Intertidal) Non-native 38.1261 -122.3544
760068 Painter 1966b; Hopkins 1986 1963 San Pablo Bay East, Station 32 (Intertidal) Non-native 38.1261 -122.3544
760069 Recher 1966 1962 near Mouth of San Francisquito Creek Non-native 37.4658 -122.1156
760070 DeMartini and Lau 1967, in Hopkins 1986 1967 1967-10-28 Offshore of Candlestick Point (416 meters SE) Non-native 37.7069 -122.3703
760071 DeMartini and Lau 1967, in Hopkins 1986 1967 1967-10-26 Seaward side of Candlestick Cove (495 meters due E of shoreline) Non-native 37.7106 -122.3744
760072 Jones 1961; Hopkins 1986 1955 1955-03-06 Point Richmond, Station D Non-native 37.9058 -122.3850
760073 Jones 1961; Hopkins 1986 1955 1955-01-29 Point Richmond, Station C Non-native 37.9036 -122.3853
760074 Jones 1961; Hopkins 1986 1955 1955-03-06 Point Richmond, Station C Non-native 37.9036 -122.3853
760075 Jones 1961; Hopkins 1986 1955 1955-07-09 Point Richmond, Station C Non-native 37.9036 -122.3853
760076 Jones 1961; Hopkins 1986 1955 1955-09-15 Point Richmond, Station C Non-native 37.9036 -122.3853
760077 Jones 1961; Hopkins 1986 1955 1955-11-26 Point Richmond, Station C Non-native 37.9036 -122.3853
760078 Jones 1961; Hopkins 1986 1955 1955-01-15 Point Richmond, Station B Non-native 37.9114 -122.3889
760079 Jones 1961; Hopkins 1986 1955 1955-03-06 Point Richmond, Station B Non-native 37.9114 -122.3889
760080 Jones 1961; Hopkins 1986 1955 1955-05-28 Point Richmond, Station B Non-native 37.9114 -122.3889
760081 Jones 1961; Hopkins 1986 1955 1955-10-22 Point Richmond, Station B Non-native 37.9114 -122.3889
760082 Jones 1961; Hopkins 1986 1955 1955-11-26 Point Richmond, Station B Non-native 37.9114 -122.3889
760083 Jones 1961; Hopkins 1986 1956 1956-01-24 Point Richmond, Station B Non-native 37.9114 -122.3889
760084 Jones 1961; Hopkins 1986 1955 1955-03-06 Point Richmond, Station A Non-native 37.9211 -122.3881

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