Invasion History

First Non-native North American Tidal Record: 1949
First Non-native West Coast Tidal Record: 1949
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Molgula manhattensis is native to the East and Gulf coasts of the United States. It was described from Manhattan Island, NY by De Kay in 1843 (De Kay 1843), but it was collected earlier by Gould (in 1841, near Boston), and earlier still by Couthouy (1838) in Massachusetts (Van Name 1912). In the northwest Atlantic, M. manhattensis occurs from Maine to Texas (Van Name 1921; Van Name 1945; Gosner 1978; McDougall 1943; US Museum of Natural History 2009). It was reported from several locations on the East Coast, from Boston to North Carolina in the 19th century (Gould 1841; DeKay 1843; Perkins 1871; Gould 1870; Verrill and Smith 1873). By the early 20th century, it was known over its present range on the Atlantic coast of North America (Van Name 1921). Some authors considered it synonymous with Ascidia tubifera and Molgula socialis, which gives it a wide European range, from Portugal to the White Sea (Van Name 1945; Carlton 2001 personal communication), while others consider it a distinct introduced species, confined to scattered European ports (Monniot 1969). Currently, M. tubifera is considered synonymous with M. manhattensis, while M. socialis has been found to be genetically distinct and presumably native to the northeast Atlantic (Haydar et al. 2011, Shenkar et al. 2011).

Recent genetic analysis supports M. manhattensis' native status in the northwest Atlantic, and introduced status in Japan, San Francisco Bay, and the Black Sea (Haydar et al. 2011). The genetic diversity of European populations was much lower than that of the northwest Atlantic M. manhattensis, but the presence of some unique genotypes led Haydar et al. (2010) to consider this species cryptogenic in Europe. However, this could be explained by under-sampling of American populations, or by the extinction of American source populations (James T. Carlton; April Blakeslee, personal communications). The data presented by Haydar et al. (2010) appear to us to be consistent with introduced status in Europe. In European waters, treating it separately from M. socialis, M. manhattensis ranges from Bergen, Norway to Galicia (northern Spain), and a few locations in the Mediterranean Sea, such as the Venice Lagoon (Monniot 1969; Dybern 1969; Vazquez and Urgorri 1992). 

Molgula manhattensis has been introduced to the Pacific coast of North America, from Newport Beach, California (Lambert and Lambert 1998) to British Columbia (Cohen and Carlton 1995; Cohen et al. 1998; Lambert 2003). It is also introduced to Japan (Asakura 1992), Australia (Hewitt et al. 1999), and Argentina (Orensanz et al. 2002). It is tolerant of a wide range of temperatures, salinity, and pollution levels (Van Name 1945).

North American Invasion History:

Invasion History on the West Coast:

Molgula manhattensis was first discovered in Tomales Bay, California in 1949 (Hedgepeth 1952, cited by Carlton 1979) and in San Francisco Bay in the 1950s (Hopkins 1968, cited by Cohen and Carlton 1995), where it is now abundant and widespread (Carlton 1979; Cohen and Carlton 1995; Cohen et al. 2005). Possible vectors include ship hull fouling or Eastern Oyster (Crassostrea virginica) aquaculture (Carlton 1979).

South of San Francisco Bay, M. manhattensis was collected in Long Beach Harbor and Newport Bay in 1984 (Lambert and Lambert 1998). Newport Bay marks the current southern limit for this tunicate on the West Coast (Lambert and Lambert 1998; Lambert and Lambert 2003; Ruiz et al. unpublished data). It has been found in most of the bays and harbors between San Francisco Bay and Los Angeles, including Elkhorn Slough (in 1998, Wasson et al. 2001), Morro Bay (in 2005, Needles 2007), Santa Barbara (in 2001, Fairey et al. 2002), Ventura (in 1994, Lambert and Lambert 1998), Channel Islands Harbor (in 2001, Fairey et al. 2002), and Marina del Rey (in 1995, Lambert and Lambert 1998).

North of San Francisco and Tomales Bays, it was found in Coos Bay, Oregon (in 1974, Carlton 1989; Wonham and Carlton 2005), Humboldt Bay, California (in 1996, Boyd et al. 2002), Willapa Bay, Washington (in 2000; Cohen et al. 2001), Puget Sound, Washington (in 1998; Cohen et al. 1998), Vancouver Island, British Columbia (in 1998, Lambert 2003) and Prince Rupert, British Columbia (in 2009, Murray et al. 2011). In 2010, it was discovered heavily fouling a barge in Yaquina Bay, Oregon, its first record in that estuary (Estabrook et al. 2010). Its current northern limit is Prince Rupert, British Columbia (Murray et al. 2011).

Invasion History Elsewhere in the World:

One specimen of Molgula manhattensis was collected in 1973 in Panama Bay, on the Pacific side of the Panama Canal (USNM 19556, US Museum of Natural History 2007) but we know of no other records from the Canal area. In the Northwest Pacific, it was first recorded in Japan in Hiroshima Harbor in 1972 and in Tokyo Bay in 1975 (Asakura 1991; Nishikawa 1991). It now ranges from Golden Horn Bay, Sea of Japan, near Vladivostok, Russia (in 1999, Zvyagintsev et al. 2003) to Guangdong (Canton), China (Huang 2001). In the Southwest Pacific, M. manhattensis was first recorded in Australia in Port Phillip Bay, Victoria, in 1967 (Kott 1985, Keough and Ross 1999). By 1976, it had become established in Moreton Bay, in Queensland, Australia (Kott 1985). In the Southwest Atlantic, it is established in Mar del Plata, Argentina (in 1945, Orensanz et al. 2002).

In European waters, M. manhattensis ranges from Bergen, Norway to Galicia, Spain and a few locations in the Mediterranean Sea, such as the Venice Lagoon (Monniot 1969; Dybern 1969; Vazquez and Urgorri 1992). It has also been identified by molecular analysis from Germany (Sylt, Wadden Sea), the Netherlands (Grevelingen and Delfzijl), Belgium (Oostende) and France (Le Havre) (Haydar et al. 2011).


Description

Molgula manhattensis is a solitary ascidian species with a globular-like body with some lateral compression. Large specimens of M. manhattensis can measure 20-50 mm in length and height. The tunic is firm, tough and moderately thick. Some areas on the tunic may develop into small hair-like projections called papillae which frequently catch and accumulate particles or organisms to its tunic and siphons. Molgula manhattensis is typically greenish-olive or yellowish-green. The divergent siphons are fairly close together at the anterior end. The oral siphon is shorter and stouter with six lobes around the opening. The atrial siphon has four lobes around the somewhat square opening and the length of the siphon can be up to half the body length. Molgula manhattensis usually attaches a small area of its ventral surface to a substrate, although when found in dense clusters, attachment areas become variable (Van Name 1945, Lambert 2003; Gretchen Lambert, personal communication 2012).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Chordata
Subphylum:   Tunicata
Class:   Ascidiacea
Order:   Stolidobranchia
Family:   Molgulidae
Genus:   Molgula
Species:   manhattensis

Synonyms

Ascidia amphora (Agassiz, 1843)
Ascidia manhattensis (DeKay, 1843)
Ascidia rustica (Couthouy, 1828)
Ascidia tubifera (Oersted, 1844)
Caesira manhattensis (Hartmeyer, 1909)
Caesira sordida (Van Name, 1912)
Gymnocystis ampulloides (Giard, 1872)
Gymnocystis manhattensis (Huntsman, 1922)
Molgula ampulloides (Van Beneden, 1846)
Molgula coepiformis (Herdman & Sorby, 1882)
Molgula macrosiphonica (Kupfer, 1872)
Molgula platei (Arenback, 1928)
Molgula sordida (Stimpson, 1852)

Potentially Misidentified Species

Ecology

General:

Life History- A solitary tunicate is ovoid, elongate or vase-like in shape, with two openings or siphons. Most solitary tunicates attach to substrates by their side or base, but some attach with a conspicuous stalk. They are sessile filter feeders with two siphons, an oral and an atrial siphon. Water is pumped in through the oral siphon, where phytoplankton and detritus is filtered by the gills, and passed on mucus strings to the stomach and intestines. Waste is then expelled in the outgoing atrial water.

Solitary ascidians are hermaphroditic, meaning that both eggs and sperm are released to the atrial chamber. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but many species have a partial block to self-fertilization. Depending on the species, eggs may be externally or internally fertilized. In external fertilizers, eggs and sperm are released through the atrial siphon into the surrounding water column were fertilization takes place. In internal fertilizers, eggs are brooded and fertilized within the atrial chamber and then released into the water column upon hatching. Fertilized eggs hatch into a tadpole larva with a muscular tail, notochord, eyespots, and a set of adhesive papillae. The lecithotrophic (non-feeding, yolk-dependent) larva swims briefly before settlement. Swimming periods are usually less than a day and some larvae settle immediately after release, but the larval period can be longer at lower temperatures. Once settled, the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Barnes 1983).

Food:

Phytoplankton, detritus

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatGrass BedNone
General HabitatCoarse Woody DebrisNone
General HabitatUnstructured BottomNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone


Tolerances and Life History Parameters

Minimum Salinity (‰)10Field data- Cory 1967; Wass 1972
Maximum Salinity (‰)35Field data; Probably tolerates higher salinities
Minimum Duration0Larval period, Graves 1933
Maximum Duration0Larval period, Graves 1933
Maximum Length (mm)25Antero-posterior, large specimens, rarely reach 35 mm (Van Name 1945)
Minimum Height (mm)25Dorso-ventral. large specimens, rarely reach 35 mm (Van Name 1945)
Broad Temperature RangeNoneCold-temperate-Warm-Temperate
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Economic Impacts

Fisheries: Molgula manhattensis is regarded as a serious fouler of oyster cultch (empty shells, placed to encourage spat settlement) and living oysters (Andrews 1953; Andrews 1973).

Shipping and Industry: Molgula manhattensis is an important ship-fouling organism (Lippson and Lippson 1984; Millar 1971; Visscher 1927).

Ecological Impacts

Competition: In Chesapeake Bay, where it is native, Molgula manhattensis is capable of rapidly settling on and overgrowing most other fouling community organisms (Andrews 1953; Calder and Brehmer 1967; Otsuka and Dauer 1982). It frequently attains 100% cover on settling plates (Otsuka and Dauer 1982; Ruiz et al. unpublished data). Where introduced, extremely dense populations have been noted in San Francisco Bay, clogging bottom trawls (Ganssle 1968, cited by Cohen and Carlton 1995). Communities of M. manhattensis and Pinuauy crocea occupied considerable space on fouling plates in the winter of 1980-1981, but were displaced by heavy settlement of mussels in spring (Okamura 1986). When another tunicate, Ciona intestinalis was removed in experimental trials, M. manhattensis covered up to 92% of fouling plates (Blum et al. 2007). Complete cover of M. manhattensis was observed at some sites and times in Newport and Alamitos Bays, California (Lambert and Lambert 2003), suggestive of competition.

Habitat Change: The rapid growth of Molgula manhattensis means that it can quickly cover fouled surfaces with multiple layers of tunicates ~10-20 mm deep. This is likely to inhibit settlement of many other organisms (Osman and Whitlatch 1995). Although, hydroids, Polydora spp., Corophium spp. (and related genera), and bryozoans do settle on its tunic (Otsuka and Dauer 1982), and the presence of M. manhattensis did not affect recruitment on adjacent bare surfaces (Osman and Whitlatch 1995).

Regional Impacts

NWP-4aNoneEcological ImpactCompetition
Molgula manhattensis comprised 42% of the macrofouling on one wharf with highly polluted water, in Golden Horn Bay, Vladivostok, Russia, but was a minor component at two other sites (Koryakova et al. 2002).
P090San Francisco BayEcological ImpactCompetition
Extremely dense populations have been noted in San Francisco Bay, clogging bottom trawls (Ganssle 1968, cited by Cohen and Carlton 1995). Communities of Molgula manhattensis and Pinuauy crocea occupied considerable space on fouling plates in the winter of 1980/81, but were displaced by heavy settlement of mussels in Spring (Okamura 1986). When another tunicate, Ciona intestinalis was removed from experimental trials, M. manhattensis covered up to 92% of fouling plates (Blum et al. 2007).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
Extremely dense populations have been noted in San Francisco Bay, clogging bottom trawls (Ganssle 1968, cited by Cohen and Carlton 1995). Communities of Molgula manhattensis and Pinuauy crocea occupied considerable space on fouling plates in the winter of 1980/81, but were displaced by heavy settlement of mussels in Spring (Okamura 1986). When another tunicate, Ciona intestinalis was removed from experimental trials, M. manhattensis covered up to 92% of fouling plates (Blum et al. 2007).
NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactCompetition
Complete cover of Molgula manhattensis was observed at some sites and times in Newport and Alamitos Bays (Lambert and Lambert 2003), suggestive of competition.
P040Newport BayEcological ImpactCompetition
Complete cover of Molgula manhattensis was observed at some sites and times in Newport and Alamitos Bays (Lambert and Lambert 2003), suggestive of competition.
P050San Pedro BayEcological ImpactCompetition
Complete cover of Molgula manhattensis was observed at some sites and times in Newport and Alamitos Bays (Lambert and Lambert 2003), suggestive of competition.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEA-II None 1762 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 1838 Native Estab
MED-VII None 1969 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 1842 Native Estab
CAR-VII Cape Hatteras to Mid-East Florida 1852 Native Estab
NEP-V Northern California to Mid Channel Islands 1949 Def Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1898 Native Estab
NEP-VI Pt. Conception to Southern Baja California 1984 Def Estab
NEP-IV Puget Sound to Northern California 1974 Def Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1998 Def Estab
NEA-V None 1969 Def Estab
NWP-3b None 1972 Def Estab
NEA-IV None 1845 Def Estab
AUS-VIII None 1967 Def Estab
AUS-XII None 1976 Def Estab
MED-II None 1969 Def Estab
B-III None 1969 Def Estab
SA-I None 1962 Def Estab
NWP-4a None 1992 Def Estab
M130 Chesapeake Bay 1887 Native Estab
M090 Delaware Bay 0 Native Estab
M080 New Jersey Inland Bays 1869 Native Estab
M060 Hudson River/Raritan Bay 1843 Native Estab
M040 Long Island Sound 1871 Native Estab
S190 Indian River 0 Native Estab
G130 Pensacola Bay 0 Native Estab
M020 Narragansett Bay 0 Native Estab
M010 Buzzards Bay 1873 Native Estab
G260 Galveston Bay 0 Native Estab
P050 San Pedro Bay 1984 Def Estab
P270 Willapa Bay 2000 Def Estab
P170 Coos Bay 1974 Def Estab
G070 Tampa Bay 0 Native Estab
S180 St. Johns River 0 Native Estab
P040 Newport Bay 1984 Def Estab
P060 Santa Monica Bay 1995 Def Estab
P064 _CDA_P064 (Ventura) 1994 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 2001 Def Estab
P062 _CDA_P062 (Calleguas) 2001 Def Estab
P080 Monterey Bay 1998 Def Estab
P090 San Francisco Bay 1955 Def Estab
P110 Tomales Bay 1949 Def Estab
P130 Humboldt Bay 1996 Def Estab
P290 Puget Sound 1998 Def Estab
P294 _CDA_P294 (Nooksack) 2003 Def Estab
G310 Corpus Christi Bay 0 Native Estab
S080 Charleston Harbor 1898 Native Estab
SEP-H None 1973 Def Unk
P070 Morro Bay 2005 Def Estab
P093 _CDA_P093 (San Pablo Bay) 1955 Def Estab
M100 Delaware Inland Bays 0 Native Estab
N170 Massachusetts Bay 1838 Native Estab
N180 Cape Cod Bay 0 Native Estab
NWP-2 None 2001 Def Estab
NWP-3a None 2001 Def Estab
AR-V None 1969 Def Estab
G108 _CDA_G108 (St. Andrew-St. Joseph Bays) 0 Native Estab
G170 West Mississippi Sound 0 Native Estab
M050 Great South Bay 1843 Native Estab
M070 Barnegat Bay 0 Native Estab
N130 Great Bay 0 Native Estab
MED-IX None 1971 Def Estab
MED-VI None 1995 Def Estab
P210 Yaquina Bay 2010 Def Unk
S030 Bogue Sound 0 Native Estab
PAN_PAC Panama Pacific Coast 1973 Def Unk
B-I None 0 Def Estab
NZ-IV None 2018 Def Unk
B-II None 1919 Def Estab
AR-IV None 2018 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
4868 Couthuoy 1838 1838 1838-01-01 Nahant Native 42.4264 -70.9194
4869 MIT Sea Grant 2003 2003 2000-01-01 Boston Native 42.3750 -70.7500
4870 MIT Sea Grant 2003 2003 2000-01-01 Annisquam (Gloucester) Native 42.6583 -70.6778
4871 MIT Sea grant 2003 2003 2000-01-01 Gloucester Native 42.5958 -70.6694
4872 MIT Sea grant 2003 2003 2000-01-01 Salem Native 42.5236 -70.8667
4873 MIT Sea grant 2003 2003 2000-01-01 Boston Native 42.3417 -70.9667
4874 MIT Sea grant 2003 2003 2000-01-01 Plymouth Native 42.0333 -70.4167
4876 MIT Sea grant 2003 2003 2000-01-01 Barnstable Native 41.7167 -70.2667
4877 Verrill and Smith 1873 1873 1872-01-01 Martha's Vineyard Native 41.4417 -70.7750
4878 MIT Sea Grant 2003 2003 2000-01-01 New Bedford-Woods Hole-Vineyard Haven Native 41.5167 -70.6833
4881 MIT Sea Grant 2003 2003 2000-01-01 Narragansett Bay Native 41.6011 -71.3225
4882 MIT Sea Grant 2003 2003 2000-01-01 Fall River Native 41.6833 -71.2167
4883 Perkins 1871 None 1871-01-01 New Haven Native 41.0833 -73.0000
4884 DeKay 1843 1843 1843-01-01 Manhatten Native 40.5833 -74.1500
4885 DeKay 1843 1843 1843-01-01 Long Island Native 40.6903 -73.1019
4886 Perkins 1869 None 1869-01-01 Great Egg Bay Native 39.3007 -74.6329
4890 Richards 1938; Loveland and Shafto 1984 1938 1984-01-01 Barnegat Bay Native 39.7581 -74.1919
4891 Van Name 1912; Van Name 1945 1912 1912-01-01 oyster beds/DE-NJ Native 39.0500 -75.1500
4892 Dean and Hurd 1980 1980 1980-01-01 Lewes Native 38.7744 -75.1397
4893 Maurer 1977 1977 1977-01-01 Indian River-Rehoboth Bays Native 38.6614 -75.0989
4894 Van Name 1912 1912 9999-01-01 Patapsco River mouth Native 39.1736 -76.4436
4895 Ruiz et al., unpublished data 1995 1995-08-01 Annapolis Native 38.9583 -76.4456
4896 Abbe 1987 1987 9999-01-01 Calvert Cliffs Nuclear Power Plant Native 38.5250 -76.5144
4897 Cory 1967 1967 9999-01-01 Eagle Harbor-Solomons Island Native 38.3214 -76.4586
4898 Humphries et al. 1985 1985 9999-01-01 Cambridge Native 38.6344 -76.3269
4899 Frey 1946 1946 9999-01-01 Cobb Island-Point Lookout Native 39.0767 -76.5942
4900 Andrews 1956; Wass 1963; Andrews 1973 1956 9999-01-01 Gloucester Point-Norfolk/VA/Hampton Roads Native 37.2539 -76.4972
4901 Larsen 1985 1985 9999-01-01 Wreck Shoal-Newport News Native 36.9414 -76.4439
4902 Ferguson and Jones 1949; Otsuka and Dauer 1982 1949 9999-01-01 Virginia Beach Native 36.8886 -76.0764
4903 Kott 1985 1985 1852-01-01 Charleston Native 32.7764 -79.9311
4904 US National Museum of Natural History 2002 9999-01-01 Morehead City Native 34.6922 -76.6725
4905 Ruiz et al., unpublished data None 9999-01-01 Jacksonville Native 30.4011 -81.4011
4907 Lambert and Lambert 1998 1994 1984-01-01 Newport Beach Def 33.6083 -117.9083
4908 Lambert and Lambert 2003 1994 1994-10-01 Costa Mesa Def 33.6411 -117.9178
4910 Lambert and Lambert 2003 1996 1984-01-01 Long Beach Marina Def 33.7497 -118.1172
4911 Lambert and Lambert 2003 1984 9999-01-01 Long Beach Def 33.7669 -118.1883
4912 Lambert and Lambert 2003 1998 9999-01-01 Impound Marina, Long Beach Def 33.7669 -118.1883
4913 Lambert and Lambert 2003 1997 9999-01-01 Watchorn Marina, Long Beach Def 33.7669 -118.1883
4914 Lambert and Lambert 2003 1995 9999-01-01 Venice Def 33.9908 -118.4592
4916 Lambert and Lambert 2003 1998 9999-01-01 Island Packers, Ventura Def 34.2783 -119.2922
4917 Lambert and Lambert 2003 1994 9999-01-01 Pierpoint Yacht Club, Ventura Def 34.2783 -119.2922
4921 de Rivera et al. 2005 2003 9999-01-01 Marshall Def 38.1606 -122.8931
4922 de Rivera et al. 2005 2003 9999-01-01 Nick's Cove Def 38.1997 -122.9200
4923 de Rivera et al. 2005 2003 9999-01-01 Sacramento Landing Def 38.1497 -122.9053
4924 de Rivera et al. 2005 2003 9999-01-01 Shell Beach Def 38.1175 -122.8733
4925 de Rivera et al. 2005 2003 9999-01-01 Thomas Station Def 38.1697 -122.9089
4926 Hedgepeth 1962, cited by Carlton 1979 1950 9999-01-01 None Def 37.7083 -122.2792
4927 Hopkins 1968 1968 9999-01-01 South San Francisco Bay Def 37.6750 -122.3803
4928 Hopkins 1968 1968 9999-01-01 San Pablo Bay Def 38.0667 -122.3833
4929 Hopkins 1968 1968 9999-01-01 Martinez Def 38.0833 -122.2500
4940 Cohen et al. 2005 2004 9999-01-01 San Leandro Marina Def 37.6978 -122.1900
4943 de Rivera et al. 2005 2003 2003-01-01 Santa Cruz Def 36.9575 -122.0106
4945 Cohen et al. 2001 2000 2000-01-01 Port of Bay Center Def 46.6283 -123.9525
4946 Cohen et al. 2001 2000 2000-01-01 Stackpole Slough,Willapa Bay Def 46.6058 -124.0433
4947 Cohen et al. 2001 2000 2000-01-01 Round Island Def 46.3953 -123.9533
4948 Calrton 1989 1974 1974-01-01 Coos Bay Def 43.3731 -124.2067
4949 Carlton 1989 1986 1986-01-01 Charleston/OR/Inner Boast Basin Def 43.3403 -124.3289
4951 Cohen et al. 1998 1998 1998-01-01 Cape Flattery Def 47.2151 -123.1007
4952 de Rivera et al. 2005 2003 9999-01-01 La Conner Marina Def 48.3003 -122.4458
4954 Lambert 2003 1998 9999-01-01 French Creek Marina Def 49.0000 -123.3333
4956 Kott 1976 cited in Kott 1985 1976 9999-01-01 Yarra River Def -37.8500 144.9000
4958 Kott 1976 cited in Kott 1985 1976 9999-01-01 Brisbane River Def -27.4000 153.1500
4960 Tokioka and Kado 1972 cited in Kott 1985 1972 9999-01-01 Takehara City Def 34.3286 132.6344
4961 Nakauchi and Kajihara 1981 cited in Kott 1985 1981 9999-01-01 Tokyo Harbor Def 34.0667 134.4667
4963 Zvyagintsev 2003 1999 1999-01-01 Harbor of Vladivostok Def 43.0833 132.0000
4964 Zvyagintsev 2003 2003 1999-01-01 Russkii Island Def 43.2500 131.7870
4965 USNM 13433 1977 1977-08-18 Off South Carolina Native 32.7000 -78.8000
4970 USNM 6906 1915 2015-12-10 Chesapeake Bay, Severn River, Off Mouth Of River Native 39.0200 -76.3750
4971 USNM 6908 1915 2015-12-03 VA, Chesapeake Bay Native 37.2200 -76.0750
4972 USNM 7167 1916 2016-04-25 Annapolis Native 38.6708 -76.4183
4973 USNM 10505 1938 1938-05-06 Galveston Native 29.0833 -94.6583
4974 USNM 10507 1940 1940-04-05 Daytona Beach Native 28.9583 -80.7917
4976 USNM 17633 1986 1986-06-01 Yacht Harbor And Redwood City Def 37.0000 -122.0000
4979 Monniot 1969 1969 9999-01-01 Germany, Baltic Sea Def 54.5000 12.0000
4980 Monniot 1969 1969 9999-01-01 Harbor Ostende Def 51.2167 2.9167
4981 Monniot 1969 1969 9999-01-01 Arcachon Def 44.6500 -1.1670
4982 Monniot 1969 1969 9999-01-01 Venice Def 45.5800 12.5600
4983 Monniot 1969 1969 9999-01-01 Brittany Def 49.0000 -5.0000
4984 Vazquez and Urgorri 1992 1992 9999-01-01 Punta Nudillo Def 43.4583 -8.2333
4985 Vazquez and Urgorri 1992 1992 9999-01-01 A Grana Def 43.4794 -8.2581
5075 Huang (ed) 2001 2001 9999-01-01 Lushun Def 38.8000 121.2667
5076 Huang (ed) 2001 2001 9999-01-01 Tanggu Def 39.0211 117.6469
5077 Huang (ed) 2001 2001 9999-01-01 Penglai Def 37.8086 120.7508
5078 Huang (ed) 2001 2001 9999-01-01 Yantai Def 37.5333 121.4000
5079 Huang (ed) 2001 2001 9999-01-01 Qingdao Def 36.0814 120.3367
5080 Huang (ed) 2001 2001 9999-01-01 Zhifu Bay Def 37.5667 121.3833
5081 Huang (ed) 2001 2001 9999-01-01 Liangyungang Def 34.5833 119.1667
5082 Huang (ed) 2001 2001 9999-01-01 Luoyuan Bay Def 26.0833 119.2833
5083 Huang (ed) 2001 2001 9999-01-01 Xiamen Harbor Def 24.4600 118.0789
5084 Huang (ed) 2001 2001 9999-01-01 Dongshan Bay Def 23.7006 117.4311
5085 Huang (ed) 2001 2001 9999-01-01 Shantou Harbor Def 23.3667 116.6667
5086 Huang (ed) 2001 2001 9999-01-01 Daya Bay Def 21.6206 111.1272
5087 Orensanz et al. 2002 1945 1945-01-01 Mar del Plata Harbor Def -38.0000 -57.0000
5088 Altman and Whitlatch 2007 2001 2001-06-15 Avery Point Native 41.3208 -72.0444
5089 Altman and Whitlatch 2007 2001 2001-06-15 Bushy Point Rock Native 41.3053 -72.0550
5229 Dragovich and Kelly 1964 1961 9999-01-01 Tampa Bay Native 27.6883 -82.5725
5230 Dybern 1969 1969 9999-01-01 Bergen Def 60.2658 5.2525
5232 SERC Records None 9999-01-01 Puget Sound Def 47.0500 -122.9000
5233 SERC Records None 9999-01-01 Great Bay Native 43.1000 -70.7670
5749 US National Museum of Natural History 2008) None 1898-02-18 Drum Bay Native 29.9044 -89.2737
5751 US National Museum of Natural History 2008 1940 1940-04-04 Cape Canaveral Native 28.6250 -80.5000
5752 US National Museum of Natural History 2008 None 1969-01-01 Punta Gorda Native 26.9298 -82.0454
5753 US National Museum of Natural History 2008 1969 1969-01-01 Punta Gorda Native 26.9298 -82.0454
5754 US National Museum of Natural History 2008 1968 1968-03-18 Alligator Harbor Native 29.9072 -84.3705
6784 Haydar et al. 2011 2010 2010-01-01 None Def 40.0000 25.0000
6785 Monniot 1969 1969 969-01-01 Sigean Def 43.0333 2.9833
6786 Minchin et al. 2007 1998 1998-01-01 Irish Sea Def 53.0000 -6.0000
6787 Haydar 2010 None 9999-01-01 Black Sea Def 43.0000 28.0000
6788 Haydar et al. 2010 None 9999-01-01 Ise of Sylt Def 54.9000 8.3333
6789 Haydar 2010 None 9999-01-01 North Sea Def 51.7333 3.9833
6790 Haydar 2010 None 9999-01-01 le Havre Def 49.5000 0.1333
6874 Murray et al. 2011 2009 2009-08-01 Prince Rupert Def 54.3167 -130.3333
767413 Ruiz et al., 2015 2012 2012-08-17 Tomales- Shell Beach, Bodega Bay, California, USA Def 38.1163 -122.8713
767768 Ruiz et al., 2015 2013 2013-07-26 Pier 32 Marina, San Diego Bay, CA, California, USA Def 32.6516 -117.1077
767805 Ruiz et al., 2015 2011 2011-09-15 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9117 -122.3494
767815 Ruiz et al., 2015 2011 2012-09-20 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6979 -122.1912
767826 Ruiz et al., 2015 2011 2011-09-20 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8067 -122.4432
767837 Ruiz et al., 2015 2011 2011-09-14 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5880 -122.3160
767847 Ruiz et al., 2015 2011 2011-09-16 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9724 -122.4796
767859 Ruiz et al., 2015 2011 2011-09-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6725 -122.3864
767863 Ruiz et al., 2015 2011 2011-09-27 Petaluma Marina, San Francisco Bay, CA, California, USA Def 38.2304 -122.6136
767872 Ruiz et al., 2015 2011 2011-09-13 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.8046 -122.3985
767882 Ruiz et al., 2015 2011 2012-09-15 Berkeley Marina, San Francisco Bay, CA, California, USA Def 37.8758 -122.3181
767893 Ruiz et al., 2015 2011 2012-09-19 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8609 -122.4853
767910 Ruiz et al., 2015 2011 2011-09-21 South Beach Harbor, San Francisco Bay, CA, California, USA Def 37.7797 -122.3871
767924 Ruiz et al., 2015 2011 2011-09-20 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7947 -122.2822
767935 Ruiz et al., 2015 2011 2011-09-22 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
767954 Ruiz et al., 2015 2011 2011-09-12 Paradise Cay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9156 -122.4769
767970 Ruiz et al., 2015 2011 2011-09-12 Corinthian Yacht Club, San Francisco Bay, CA, California, USA Def 37.8103 -122.3228
767992 Ruiz et al., 2015 2012 2012-08-24 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9134 -122.3523
768011 Ruiz et al., 2015 2012 2012-08-23 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8609 -122.4853
768044 Ruiz et al., 2015 2012 2012-08-27 Port of San Francisco Pier 31, San Francisco Bay, CA, California, USA Def 37.8078 -122.4060
768068 Ruiz et al., 2015 2012 2012-09-11 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
768091 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6633 -122.3817
768115 Ruiz et al., 2015 2012 2012-08-29 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3174
768136 Ruiz et al., 2015 2012 2012-09-04 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.5023 -122.2130
768156 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9736 -122.4802
768181 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Def 37.7987 -122.3228
768201 Ruiz et al., 2015 2012 2012-09-07 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7940 -122.2787
768211 Ruiz et al., 2015 2012 2012-08-31 Glen Cove Marina, San Francisco Bay, CA, California, USA Def 38.0663 -122.2130
768238 Ruiz et al., 2015 2012 2012-09-13 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6962 -122.1919
768256 Ruiz et al., 2015 2012 2012-09-12 Emeryville, San Francisco Bay, CA, California, USA Def 37.8396 -122.3133
768282 Ruiz et al., 2015 2013 2013-08-15 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7656 -122.2858
768303 Ruiz et al., 2015 2013 2013-08-20 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3163
768321 Ruiz et al., 2015 2013 2013-08-22 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7926 -122.2746
768343 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9723 -122.4829
768363 Ruiz et al., 2015 2013 2013-08-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6639 -122.3821
768386 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.5024 -122.2134
768407 Ruiz et al., 2015 2013 2013-08-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9138 -122.3522
768437 Ruiz et al., 2015 2013 2013-08-21 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6980 -122.1908

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