Invasion History

First Non-native North American Tidal Record: 1933
First Non-native West Coast Tidal Record: 1933
First Non-native East/Gulf Coast Tidal Record: 1970

General Invasion History:

Styela clava is native to the Northwest Pacific, from Shanghai to the Sea of Okhotsk and the southeastern Bering Sea (Benyaminson, in Golikov et al. 1976; Nishikawa 1991). It has become widely distributed in coastal waters, through ship fouling and transport with oysters (Crassostrea gigas; Buizer 1980; Cohen and Carlton 1995; Lutzen 1999). The morphology of this tunicate, with its great attachment strength, and streamlined body, is well adapted for transport on the hulls of moving ships (Murray et al. 2012). In 1933, it was first collected in the Northeast Pacific in Newport Beach California, and has subsequently spread south to San Diego (in 1959), and north to San Francisco Bay (in 1949), Coos Bay Oregon (in 1993) and Vancouver Island, British Columbia (in 1995; Carlton and Cohen 1995). On the East Coast of North America, it was first collected in 1970 in Beverly, Massachusetts and Portsmouth, New Hampshire, and has subsequently spread south to Chincoteague, Virginia and north to Prince Edward Island and Nova Scotia (Berman et al. 1992; U.S. National Museum of Natural History 2002-2012; Locke et al. 2007; Paul Fofonoff, personal observations 2013; Moore et al. 2014). In the northeast Atlantic, it was recorded in Plymouth, England in 1953, and has subsequently spread east, along the southern North Sea to the Limfjord and Kattegat in Denmark and south to Portugal and a Mediterranean lagoon (Buizer 1999; Lutzen 1999; Davis and Davis 2005; Davis and Davis 2007). In 1972, it invaded Port Philip Bay, Australia, and in 1977, Sydney Harbour (Hewitt et al. 1999). In 2005, it was discovered in New Zealand (Biosecurity New Zealand 2005a; Biosecurity New Zealand 2005b).

North American Invasion History:

Invasion History on the West Coast:

Styela clava was first collected on the West Coast in Newport, California in 1933 (Carlton 1979) and in Elkhorn Slough in 1935 (Carlton 1979; Lambert and Lambert 1998; Wasson et al. 2001). Styela clava was found at Point Richmond, in San Francisco Bay, in 1949, King Harbor (Redondo Beach) and Mission Bay (San Diego) in 1971, and San Diego Bay in 1972 (Carlton 1979; Cohen and Carlton 1995; Lambert and Lambert 1998). It was also collected in 1972 in Morro Bay, but has not been seen there since (Carlton 1979; Needles 2007). Surprisingly, although it has been established for 50 years in San Francisco Bay, it has not been reported from nearby Tomales Bay (Fairey et al. 2002), and is a recent invader in Bodega Harbor and Humboldt Bay (reported in 2000, Boyd et al. 2002).

North of California, Styela clava was first reported in Coos Bay, Oregon in 1993 (Cohen and Carlton 1995), and Vancouver Island, British Columbia and Drayton Harbor, in Blaine, Washington (on Georgia Straits) in 1998 (Lambert 2003). In 2001, it was collected in Neah Bay, Juan de Fuca Straits, on the Olympic Peninsula, Washington (deRivera et al. 2005), and in 2005 in the Hood Canal, part of Puget Sound (Center for Aquatic Resource Studies 2009).

Invasion History on the East Coast:

In the Northwest Atlantic, Styela clava was first collected in 1970 in Beverly, Massachusetts and in Portsmouth Harbor, New Hampshire (Berman et al., 1992). South of Cape Cod, the earliest record appears to be from 1973, on Long Island (Carlton, 1989). It was established in Narragansett Bay by 1976 (Fofonoff, personal observation; Plough 1978), and is now found throughout southern New England (Whitlatch and Osman 2000; MIT Sea Grant 2003; Carman et al. 2007; Carman et al. 2009). In 1998 it was found in Kennebunkport and Boothbay Harbor, Maine (Whitlatch and Osman 2000), and in 2000, in Portland Harbor, Casco Bay, Maine. In 1998, S. clava appeared in the Brudenell River, Prince Edward Island, in the Gulf of St. Lawrence, Canada, where it has caused serious problems by fouling cultured mussels and the lines and racks used in their rearing (Locke et al. 2007). This tunicate has spread to several estuaries on Prince Edward Island (Ramsay et al. 2008; Arsenault et al. 2009). In 2012-2013, it was found at several sites in the Canso Canal, between the mainland of Nova Scotia and Cape Breton Island, and on the Atlantic coast of Nova Scotia, in Halifax and Lunenburg harbors (Moore et al. 2014). 

The southern extent of the range of S. clava is poorly known. It has been found in a New Jersey coastal inlet ~50 km north of Cape May (McDermott, personal communication, 1999). In November 2003, several S. clava were found washed ashore on Cape Henlopen, at the mouth of Delaware Bay (Fofonoff, unpublished data). In the Chesapeake Bay region, in 1994, a single specimen of S. clava (Catalog number 20711) was collected in Chincoteague VA (U.S. National Museum of Natural History 2002; Linda Cole, personal communication). We have not collected this species in Chesapeake Bay proper, but in 2013 we found established populations in the Indian River Inlet, Delaware; Ocean City Inlet, Maryland; and Chincoteague Inlet, Virginia (Paul Fofonoff and Kristen Larson, personal observations).

Invasion History Elsewhere in the World:

In Europe, Styela clava was first collected during 1953 in Plymouth, England, where it was mistakenly described as a new species, S. mammiculata (Carlsile 1954 cited by Lutzen 1999). It spread east along the English Channel (record in 1957 from Southampton) and across to France (record in 1968 from Dieppe) into the North Sea (record in 1974 from Den Helder, Netherlands) and to Denmark by 1991 (Buizer 1980; Lutzen 1999). In Ireland, it was first collected in Cork Harbor in 1971, and has spread north to a marina near Dublin on the Irish Sea, and to Kerry on the Atlantic Coast (Minchin et al. 2006). Styela clava spread south along the Atlantic coast of Europe, reaching Brest and Roscoff in Brittany in 1973 (Lutzen 1999) and Arcachon, France on the Bay of Biscay (Davis and Davis 2005). It now occurs at many locations on the Biscay and Atlantic coasts of Spain, with a current southern limit at Cascais, Portugal (Davis and Davis 2005). In 2006, it was found in a lagoon (Bassin de Thau) in Sete, France on the Mediterranean Sea. This basin is used intensively for oyster culture (Davis and Davis 2008). In 2004, S. clava was reported in Romanian waters of the Black Sea, as a 'casual' fouling organism, presumably not established (Micu and Micu 2004, cited by Skolka and Preda 2010). In 2012, an established population was found in the Sea of Marmara, Turkey, connecting the Black and Mediterranean Seas (Cinar 2016).

In 1972, Styela clava was found in Port Phillip Bay, Victoria, Australia, where it has become quite abundant. In 1977, it was found in Sydney Harbour, New South Wales. However, S. clava was not found in a 2000 port survey (Australian Museum Business Services 2002). In 2005, 
S. clava was found in several locations in New Zealand, including Auckland, Littleton, and Christchurch (Biosecurity New Zealand 2005a).


Description

Styela clava is a solitary tunicate with a leathery, but thin, and bumpy tunic. Its body is cylindrical or club-shaped and narrows posteriorly to a stalk that is anchored to the substrata by a disk shaped holdfast. The wrinkled or creased looking stalk is often 20-50% of the total body length. Styela clava can grow up to 150 mm in length. Colors can range from yellowish to reddish to brownish. The oral and atrial siphons are located close together and are directed anteriorly. Both siphons have four lobes and appear striped with alternating dark and light brownish to purplish bands (Kott 1985; Nishikawa 1991; Lambert 2003).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Chordata
Subphylum:   Tunicata
Class:   Ascidiacea
Order:   Stolidobranchia
Family:   Styelidae
Genus:   Styela
Species:   clava

Synonyms

Bostryorchis clava (Redikorzev, 1916)
Styela barnharti (Van Name, 1945)
Styela mammiculata (Carlisle, 1954)

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).

In all part of its native and introduced range, S. clava is more frequently reported from anthropogenic stuctures than from natural surfaces, (Simkanin et al. 2012). Dock floats are especially favored habitats, probably because their motion provides rapid water exchange, and a fresh supply of food-laden water (Glasby 2001). Other colonized man-made structures include pilings, piers, aquaculture structures, and boat hulls (Carman et al. 2010; Davidson et al. 2010; Simkanin et al. 2012). Natural habitats include rocky reefs, bivalve colonies, seaweeds, (White and Orr 2011; Simkanin et al. 2012).

Food:

Phytoplankton

Consumers:

snails, fish, crabs, urchins

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)-2Field data, Denmark (Lutzing 1998)
Maximum Temperature (ºC)26.6Field data, Thau Lagoon, France (Davis and Davis 2009)
Minimum Salinity (‰)18Experimental data; animals from Denmark, transferred stepwise (Lutzen 1998).
Maximum Salinity (‰)35Field data (Lutzen 1998, experimental data (Sims 1984, highest tested)
Minimum Reproductive Temperature15Field data, Davis and Davis 2007; Wong et al. 2011
Maximum Length (mm)90 Kott 1985; Nishikawa 1991; Lambert 2003
Broad Temperature RangeNoneCold temperate-Warm temperate
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Economic Impacts

Fisheries: The largest economic impacts of Styela clava have been seen in Prince Edward Island, Canada, where dense populations of tunicates have fouled cages used for fish aquaculture, and ropes, racks, and other gear used in mussel culture (Locke et al. 2007; Arsenault et al. 2009). The tunicates compete with cultured mussels, reducing mussel harvests by ~50%. The estimated economic impact is 34-88 million $CAN per year (Coulatti et al. 2006). Another negative potential impact of S. clava and other tunicates is that when they foul aquaculture gear and boats, they can retain and transport viable cells and cysts of toxic phytoplankton (Rosa et al. 2013). On the positive side, S. clava is a popular food item in Korea. Styela clava and S. plicata are extensively cultured on long lines in Korea and Japan (Lambert et al. 2016). Fisheries for S. clava, in New England, for the Korean market appear feasible (Karney and Rhee 2009).

Shipping and Industry: Styela clava can grow in very dense populations on boat hulls, docks, fishing gear, and aquaculture equipment.

Ecological Impacts

Competition: Styela clava typically grows in dense populations, dominating large patches in fouling communities. Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes from southern California harbors (Lambert and Lambert 1998). It covered 18.5% of fouling plate surfaces in one-year old assemblages near Avery Point, Connecticut, Long Island Sound (Altman and Whitlatch 2007). It can reduce the growth rate of co-occurring species through its intense filtration of the water column (Osman and Whitlatch 2000). In English waters, the population growth of S. clava was paralleled by a decrease in Ciona intestinalis (Lutzen 1999). Styela clava grows in very dense populations in Port Phillip Bay, Australia, apparently displacing other fouling organisms. Because of its high filtering rate, it was expected to compete for food with other suspension-feeding taxa (Currie et al. 1999); however, this effect was not seen in the experiments conducted by Ross et al. (2007).

Habitat Change: The dense aggregations frequently formed by Styela clava exclude other organisms but also provide secondary substrate for other fouling organisms. The tunics of S. clava are frequently fouled by colonial tunicates. In experimental studies, soft sediment fauna from Port Phillip Bay, Australia, showed idiosyncratic responses to increased Styela density, with crustaceans and tanaids, decreasing, while the deposit-feeding bivalve Laternula rostrata increased (Ross et al. 2007).

Predation: Styela clava can inhibit settlement of co-occurring fouling organisms through predation on planktonic larvae, as a result of high filtering rates (Osman and Whitlatch 2000).

Food/Prey: Newly settled S. clava in Long Island Sound are prone to intense predation by the snails Astyris lunata (= Mitrella lunata) and Costoanachis translirata (= Anachis lafresnayi of authors) (Osman and Whitlatch 2000; Whitlatch and Osman 2009). Older juveniles are subject to predation by fishes, mostly Tautog (Tautoga onitis) and Cunner (Tautogolabrus adspersus) (Osman and Whitlatch 2000).

Regional Impacts

NEP-VNorthern California to Mid Channel IslandsEconomic ImpactFisheries
Common oyster fouling organism (Carlton 1979)
NEP-VNorthern California to Mid Channel IslandsEconomic ImpactShipping/Boating
Common hull and dock fouling oganism (Carlton 1979)
P090San Francisco BayEconomic ImpactShipping/Boating
Common dock and hull fouling organism.
M040Long Island SoundEcological ImpactCompetition
Styela clava is frequently abundant and often the dominant fouling organism in harbors. It covered 18.5% of fouling plate surface in one-year old assemblages near Avery Point, CT (Altman and Whitlatch 2007). It can reduce the growth rate of co-occurring species through its intense filtration of the water column (Osman and Whitlatch 2000).
NA-S3NoneEconomic ImpactFisheries
Fouling of aquaculture cages (Arsenault et al. 2009). Competition with cultured mollusks for food and space. Reduces mussel harvests by ~50%, estimated impact 34-88 miliion $CAN per year (Coulatti et al. 2006).
NA-ET3Cape Cod to Cape HatterasEcological ImpactCompetition
Styela clava is frequently abundant and often the dominant fouling organism in harbors. It covered 18.5% of fouling plate surfaces in one-year old assemblages near Avery Point, Connecticut, Long Island Sound (Altman and Whitlatch 2007). It can reduce the growth rate of co-occurring species through its intense filtration of the water column (Osman and Whitlatch 2000). Styela clava is abundant in harbors with varying water quaility, including those with poor, fair and good quality (Carman et al. 2007).
NA-ET2Bay of Fundy to Cape CodEconomic ImpactFisheries
Farming Styela clava for the Korean market appears feasible (Karney and Rhee 2009).
NA-S3NoneEcological ImpactCompetition
Competition with cultured mussels for food and space (Coulatti et al. 2006; Gittenberger 2009; Comeau et al. 2015).
NA-ET3Cape Cod to Cape HatterasEcological ImpactPredation
Styela clava can inhibit settlement of co-occurring fouling organisms through predation on planktonic larvae, as a result of high filtering rates (Osman and Whitlatch 2000).
AUS-VIIINoneEcological ImpactHabitat Change
In experimental studies with various densities of Styela clava, fauna from Port Phillip Bay showed idiosyncratic responses to increased Styela density, with crustaceans, and tanaids, as a group decreasing, while the deposit-feeding bivalve Laternula rostrata increased. These changes appeared to be due to subtle changes in hydrodynamics, sediment quality, emigration or altered survival in Styela rather than community-wide effects (Ross et al. 2007).
NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P020San Diego BayEcological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P030Mission BayEcological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P023_CDA_P023 (San Louis Rey-Escondido)Ecological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P027_CDA_P027 (Aliso-San Onofre)Ecological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P040Newport BayEcological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P050San Pedro BayEcological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P060Santa Monica BayEcological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P062_CDA_P062 (Calleguas)Ecological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P064_CDA_P064 (Ventura)Ecological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P065_CDA_P065 (Santa Barbara Channel)Ecological ImpactCompetition
Styela clava is abundant in Southern California harbors, often forming dense patches with 100% cover (Lambert and Lambert 1998; Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
M040Long Island SoundEcological ImpactPredation
Styela clava can inhibit settlement of co-occurring fouling organisms through predation on planktonic larvae, as a result of high filtering rates (Osman and Whitlatch 2000).
M040Long Island SoundEcological ImpactFood/Prey
Newly settled Styela clava in Long Island Sound are prone to intense predation by the snails Astyris lunata (= Mitrella lunata) and Costoanachis translirata (= Anachis lafresnayi of authors) (Osman and Whitlatch 2000; Whitlatch and Osman 2009). Older juveniles are subject to predation by fishes, mostly Tautog (Tautoga onitis) and Cunner (Tautogolabrus adspersus; Osman and Whitlatch 2000).
M010Buzzards BayEcological ImpactCompetition
Styela clava is abundant in harbors with poor and fair as well as good water quality (Carman et al. 2007).
N185_CDA_N185 (Cape Cod)Ecological ImpactCompetition
Styela clava is abundant in harbors with poor and fair as well as good water quality (Carman et al. 2007).
NEA-IINoneEcological ImpactCompetition
Populations increased 'explosively' in the Eastern Scheldt, presumably displacing native species (Buizer 1980). In English waters, the population growth of S. clava was paralelled by a decrease in Ciona intestinalis (Lutzen 1999).
NA-ET3Cape Cod to Cape HatterasEcological ImpactFood/Prey
Newly settled Styela clava in Long Island Sound are prone to intense predation by the snails Astyris lunata (= Mitrella lunata) and Costoanachis translirata (= Anachis lafresnayi of authors) (Osman and Whitlatch 2000; Whitlatch and Osman 2009). Older juveniles are subject to predation by fishes, mostly Tautog (Tautoga onitis) and Cunner (Tautogolabrus adspersus; Osman and Whitlach 2000).
AUS-VIIINoneEcological ImpactCompetition
Styela clava grows in very dense populations in Port Phillip Bay, apparently displacing other fouling organisms. Because of its high filtering rate, it was expected to compete for food with other suspension-feeding taxa (Currie et al. 1999). This was not seen in experiments conducted by Ross et al. (2007).
N195_CDA_N195 (Cape Cod)Economic ImpactFisheries
Styela clava was found fouling aquaculture gear at 21 sites, and cultured Bay Scallops (Aequipecten irradians) at two sites, of 26 aquaculture sites surveyed on Marthas Vineyard (Carman et al. 2010). This tunicate was also reported at aquaculture sites in Rhode Island (Carman et al. 2010).
NA-ET3Cape Cod to Cape HatterasEconomic ImpactFisheries
Styela clava was found fouling aquaculture gear at 21 sites, and cultured Bay Scallops (Aequipecten irradians) at two sites, of 26 aquaculture sites surveyed on Marthas Vineyard (Carman et al. 2010). Styela clava was also reported to be fouling aquaculture sites in Rhode Island and New York state (Carman et al. 2010).
N195_CDA_N195 (Cape Cod)Ecological ImpactCompetition
Styela clava is abundant in harbors with poor and fair as well as good water quality (Carman et al. 2007)
NEA-IINoneEconomic ImpactShipping/Boating
Fouling impacts fave been reported for the British Isles (Minchin et al. 2013).
SA-INoneEcological ImpactHabitat Change
Styela clavad facilitated the invasion of the seaweed Undaria pinnatifida by providing an attachment site for sporelings (Pereyra et al. 2015;Pereyra et al. 2017)
NEA-IIINoneEconomic ImpactFisheries
Fouling of cultured mussels by a variety of non-native tunicates was reported beginning in 2013, and was a serious problem by 2016 (Palanisamy et al. 2018).
NZ-IVNoneEconomic ImpactFisheries
The impact of Styela clava on Greenshell Mussel (Perna canaliculus) aquaculture in New Zealand was estimated at NZ $23.9 million (Soliman and Inglis 2018).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET2 Bay of Fundy to Cape Cod 1970 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 1973 Def Estab
NEP-IV Puget Sound to Northern California 1993 Def Estab
NEP-V Northern California to Mid Channel Islands 1935 Def Estab
NA-S3 None 1998 Def Estab
NEA-II None 1957 Def Estab
AUS-VIII None 1972 Def Estab
B-II None 1991 Def Estab
NEA-III None 1953 Def Estab
NWP-3b None 0 Native Estab
NWP-4a None 0 Native Estab
NWP-3a None 0 Native Estab
NEA-IV None 1971 Def Estab
NWP-5 None 0 Native Estab
NWP-4b None 0 Native Estab
NEP-VI Pt. Conception to Southern Baja California 1933 Def Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1995 Def Estab
AUS-X None 1977 Def Estab
NEA-V None 1980 Def Estab
NZ-IV None 2002 Def Estab
P050 San Pedro Bay 1994 Def Estab
P130 Humboldt Bay 2000 Def Estab
M010 Buzzards Bay 1986 Def Estab
M040 Long Island Sound 1988 Def Estab
P020 San Diego Bay 1972 Def Estab
N130 Great Bay 1970 Def Estab
M020 Narragansett Bay 1976 Def Estab
M080 New Jersey Inland Bays 1998 Def Estab
M120 Chincoteague Bay 1994 Def Estab
M090 Delaware Bay 2003 Def Estab
M030 Gardiners Bay 1973 Def Estab
N185 _CDA_N185 (Cape Cod) 1998 Def Estab
N180 Cape Cod Bay 1998 Def Estab
N170 Massachusetts Bay 1970 Def Estab
N116 _CDA_N116 (Piscataqua-Salmon Falls) 1998 Def Estab
N100 Casco Bay 2000 Def Estab
N070 Damariscotta River 2007 Def Estab
P030 Mission Bay 1971 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 1994 Def Estab
P040 Newport Bay 1933 Def Estab
P060 Santa Monica Bay 1971 Def Estab
P062 _CDA_P062 (Calleguas) 1994 Def Estab
P064 _CDA_P064 (Ventura) 1994 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 1994 Def Estab
P070 Morro Bay 1972 Def Unk
P080 Monterey Bay 1935 Def Estab
P090 San Francisco Bay 1949 Def Estab
P112 _CDA_P112 (Bodega Bay) 2000 Def Estab
P286 _CDA_P286 (Crescent-Hoko) 2001 Def Estab
P297 _CDA_P297 (Strait of Georgia) 1998 Def Estab
P290 Puget Sound 2005 Def Estab
P170 Coos Bay 1993 Def Estab
N195 _CDA_N195 (Cape Cod) 1986 Def Estab
N190 Waquoit Bay 2003 Def Estab
P027 _CDA_P027 (Aliso-San Onofre) 1994 Def Estab
MED-II None 2006 Def Estab
P056 _CDA_P056 (Los Angeles) 1971 Def Estab
M023 _CDA_M023 (Narragansett) 2007 Def Estab
N140 Hampton Harbor 2003 Def Estab
N080 Sheepscot Bay 1998 Def Estab
N050 Penobscot Bay 2007 Def Estab
N165 _CDA_N165 (Charles) 2009 Def Estab
MED-IX None 2004 Def Unk
M100 Delaware Inland Bays 2013 Def Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 2012 Def Estab
N165 _CDA_N165 (Charles) 2004 Def Estab
SA-I None 2013 Def Estab
MED-VIII None 2012 Def Estab
B-III None 207 Def Unk
B-I None 2013 Def Estab
NEP-II Alaska south of Aluetians to the Alaskan panhandle 1975 Crypto Estab
AR-V None 2013 Def Estab
M060 Hudson River/Raritan Bay 2019 Def Estab
MED-VII None 2021 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
3940 Fofonoff, personal observation None 2003-12-01 Cape Henlopen Def 38.8032 -75.0946
3941 McDermott, personal communication 1998 1998-01-01 0 mi NE of Cape May/ Def 38.9350 -74.9064
3942 MIT Sea Grant 2003 2003 2003-08-07 South Jamesport Def 40.9364 -72.5778
3943 Carlton 1989 1972 1972-01-01 Long Island Sound Def 41.0833 -73.0000
3945 MIT Sea Grant 2003 2003 2003-08-08 Stamford Def 41.0533 -73.5392
3946 MIT Sea Grant 2003 2003 2003-08-08 Milford Yacht Club, Milford/ Def 41.2222 -73.0569
3947 MIT Sea Grant 2003 2003 2003-08-07 Brewer Yacht Yard, Mystic Def 41.3542 -71.9669
3948 Whitlach and Osman 2000 1997 1997-08-01 Groton Def 41.3500 -72.0789
3993 MIT Sea Grant 2003 2000 2000-08-16 Warwick Def 41.6839 -71.3917
3994 MIT Sea Grant 2003 2003 2003-08-06 Allen Harbor, North Kingstown Def 41.6237 -71.4126
3995 Plough 1978, MITSea Grant 2003 1978 1978-01-01 Wickford Marina, Wickford Def 41.5739 -71.4619
3996 MIT Sea Grant 2003 2000 2000-08-15 Colt State Park, Bristol Def 41.6697 -71.3014
3997 MIT Sea Grant 2003 2000 2000-08-15 Coasters Harbor Island, Newport Def 41.5107 -71.3270
3998 MIT Sea Grant 2003 2004 2004-04-01 Beavertail Point Def 41.4490 -71.3995
3999 Whitlach and Osman 2000 1998 1998-08-01 Sakonnet Point Def 41.4542 -71.1953
4000 MIT Sea Grant 2003 2000 2000-08-06 Westport River Def 41.5125 -71.0894
4001 MIT Sea Grant 2003 2000 2000-08-11 Massachusetts Maritime Academy, Bourne Def 41.7394 -70.6244
4002 MIT Sea Grant 2003 2000 2000-08-11 New Bedford Def 41.6778 -70.9167
4003 MIT Sea Grant 2003 1986 1986-01-01 Woods Hole Def 41.5257 -70.6703
4004 Whitlach and Osman 2000 1998 1998-08-01 Hyannis Def 41.6528 -70.2833
4005 Whitlach and Osman 2000 1998 1998-08-01 Barnstable Def 41.7167 -70.2667
4006 MIT Sea Grant 2003 2000 2000-08-10 Sandwich Def 41.7650 -70.4750
4007 Whitlach and Osman 2000 1998 1998-08-01 Plymouth Def 41.9750 -70.6667
4008 MIT Sea Grant 2003 2000 2000-08-09 Quincy Def 42.2528 -71.0028
4009 MIT Sea Grant 2003 2000 2000-08-07 Rowes Wharf, Boston Def 42.3569 -71.0414
4010 Berman et al. 1992 None 1970-01-01 Beverly Def 42.5583 -70.8806
4011 MIT Sea Grant 2003 2000 2000-08-08 Gloucester State Pier Def 42.5958 -70.6694
4012 MIT Sea Grant 2003 2003 2003-08-03 Hampton State Pier Def 42.9375 -70.8394
4013 Berman et al. 1992 1985 1985-01-01 Portsmouth Def 43.0717 -70.7631
4014 Whitlach and Osman 2000 1998 1998-08-01 Kennebunkport Def 43.3617 -70.4772
4015 MIT Sea Grant 2003 2003 2003-08-04 Portland Yacht Services, Portland/ Def 43.6575 -70.2411
4016 Whitlach and Osman 2000 1998 1998-01-01 Boothbay Harbor Def 43.8522 -69.6286
4017 Fisheries and Oceans Canada 2002 1996 1996-01-01 Brudenell River (Gulf of St.Lawrence) Def 46.1833 -62.5500
4114 Lambert and Lambert 2003 2000 2000-08-01 Ensenada Def 31.8667 -116.6167
4116 Abbott and Johnson 1972; Carlton 1979 1971 1971-01-01 San Diego Def 32.7792 -117.2333
4117 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-08-27 Oceanside Def 33.2078 -117.3942
4118 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-09-18 Dana Point Def 33.4597 -117.6947
4120 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-09-04 Long Beach Def 33.7489 -118.2229
4121 Carlton 1979; Cohen et al. 2002 1971 1971-01-01 Redondo Beach Def 33.8464 -118.3969
4122 Lambert and Lambert 1998; Cohen et al. 2002 1994 1994-10-01 Port Hueneme Def 34.1481 -119.2019
4123 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-01 Ventura Def 34.2495 -119.2648
4124 Lambert and Lambert 1998; Lambert and Lambert 2003 1994 1994-10-01 Santa Barbara Harbor Def 34.4072 -119.6887
4125 Abbot and Johnson 1972, Carlton 1979 1972 1972-01-01 Morro Bay Def 35.3378 -120.8513
4128 Cohen and Carlton 1995, Ruiz et al. unpublished None 1993-01-01 South San Francisco Bay Def 37.5133 -122.2075
4130 Boyd et al. 2002 2000 2000-01-01 Woodley Island Marina/ Def 40.8097 -124.1594
4131 Cohen and Carlton 1995; Wonham et al. 2005 1993 1993-01-01 Coos Bay Def 43.4294 -124.2286
4132 Lambert 2003 2001 2001-08-18 Neah Bay Def 48.3712 -124.6100
4133 Lambert 2003 1998 1998-09-03 Drayton Harbor (near Blaine, WA) Def 48.9959 -122.7638
4134 Center for Aquatic Resource Studies 2006 2005 2005-10-01 Pleasant Harbor [just S of Brinnon, WA] Def 47.6620 -122.9163
4135 Lambert 2003 1993 1993-01-01 French Creek, Vancover Island/ Def 49.3500 -124.3500
4136 Lambert 2003 1998 1998-09-04 Brechin Point, Nanaimo Def 49.1833 -123.9500
4137 Lambert 2003 2000 2000-12-09 Duncan Def 48.8167 -123.6000
4138 Kashin et al. 2003 None 9999-01-01 Pacific Maritime region Native 42.8000 132.9500
4139 Zvyagintsev 1992 1986 1986-09-01 Amur Bay, Sea of Japan (Zvyagintsev 1992) Native 43.3000 131.8000
4140 Golikov, A. N. et al. 1976 1976-01-01 Vladivostok area Native 43.0000 132.5000
4141 Nishikawa 1991 1991 1991-01-01 Rishiri Island, Hokkaido/ Native 45.1667 141.2500
4142 Nishikawa 1991 1991 1991-01-01 Rebun Island Native 45.3644 141.0331
4143 Nishikawa 1991 1991 1991-01-01 Esashi Native 41.8572 140.1231
4144 Nishikawa 1991 1991 1885-01-01 Hakodate, Native 41.7758 140.7367
4145 Nishikawa 1991 1991 1991-01-01 Mutsu Bay Native 41.0800 140.8378
4146 Nishikawa 1991 1991 1991-01-01 Oga Native 39.5500 139.5000
4147 Nishikawa 1991 1991 1991-01-01 Sado Island Native 38.0333 138.2500
4148 Nishikawa 1991 1991 1991-01-01 Wakasa Bay Native 35.7500 135.6667
4149 Nishikawa 1991 1951 1951-01-01 Akkeshi Native 43.0356 144.8525
4150 Nishikawa 1991 1960 1960-01-01 Matsushima Bay Native 38.3667 141.0717
4151 Nishikawa 1991 1953 1953-01-01 Tokyo Bay Native 35.4169 139.7836
4152 Nishikawa 1991 1953 1953-01-01 Sagami Bay Native 35.3333 139.2500
4153 Nishikawa 1991 1991 1991-01-01 Ise Bay Native 34.7167 136.7167
4154 Nishikawa 1991 1906 1906-01-01 Kii Peninsula Native 34.0000 135.7500
4155 Nishikawa 1991 1991 1967-01-01 Nagasaki Native 32.7500 129.8667
4156 Rho 1991 1991 1977-07-08 Naedo Native 36.9833 126.7500
4157 Rho 1991 1977 1977-07-09 Chisep'o Native 34.8256 128.7097
4158 Rho 1991 1984 1984-07-02 Tae'chon Native 36.3467 126.6047
4159 Rho 1991 1982 1982-07-10 Tolsan-do Island/ Native 34.6300 127.7600
4160 Rho 1991 1985 1985-05-26 Sodol Native 37.8000 129.0000
4161 Rho 1991 1985 1985-10-27 P'ohang Native 36.0403 129.3711
4162 Nishikawa 1991 1967 1967-01-01 Yantai Native 37.5333 121.4000
4163 Nishikawa 1991 1967 1967-01-01 Shanghai Native 31.1094 121.3681
4164 Nishikawa 1991 1885 1885-01-01 East China Sea Native 31.0000 125.0000
4165 Nishikawa 1991 1975 1975-01-01 East China Sea Native 31.2667 123.5833
4166 Huang 2001 2001 2001-01-01 Lushun Native 38.8000 121.2667
4167 Huang 2001 2001 2001-01-01 Tanggu, Native 39.0211 117.6469
4168 Huang 2001 2001 2001-01-01 Qingdao Native 36.0814 120.3367
4169 Huang 2001 2001 2001-01-01 Lianyngang Native 34.5997 119.1594
4170 Huang 2001 2001 2001-01-01 Luoyuan Native 26.4856 119.5492
4171 Lutzen 1998 1953 1953-01-01 Plymouth Def 50.4000 -4.1167
4172 Lutzen 1998 1968 1968-09-01 Celtic Sea Def 51.7000 -5.1167
4173 Lutzen 1998 1985 1985-01-01 Heysham Harbor, Irish Sea Def 54.0333 -2.9000
4174 Lutzen 1998 None 9999-01-01 Millport, Isle of Cumbrae Def 55.7500 -4.9500
4175 Lutzen 1998; Minchin et al. 2006 1971 1971-01-01 Marloge Marina, County Cork Def 51.8986 -8.4958
4176 Minchin et al. 2006 2004 2004-09-01 Fenit Harbor Marina Def 52.2808 -9.8675
4177 Minchin et al. 2006 2004 2004-09-01 Dingle Marina, Dingle Def 52.1408 -10.2689
4178 Minchin et al. 2006 2004 2004-08-01 Dun Laoghaire Marina Def 53.2925 -6.1286
4179 Lutzen 1998 1962 1962-01-01 Poole Def 50.7000 -2.0000
4180 Lutzen 1998 1957 1957-01-01 Southampton Def 50.9000 -1.4000
4181 Lutzen 1998 1958 1958-01-01 Portsmouth Def 50.7667 -1.0833
4182 Lutzen 1998 1962 1962-01-01 Langstone Def 50.8333 -0.9833
4183 Lutzen 1998 1962 1962-01-01 Shoreham Def 50.8333 -0.2333
4184 Lutzen 1998 1969 1969-01-01 Dover Def 51.1333 1.3000
4185 Lutzen 1998 1991 1991-01-01 Hals Def 57.0000 10.3167
4186 Lutzen 1998 1986 1986-01-01 Livo Def 56.9000 9.1000
4187 Lutzen 1998 1980 1980-01-01 Mors Def 56.8333 8.7500
4188 Lutzen 1998 1978 1978-01-01 Nissum Bredning Def 56.6333 8.3667
4189 Lutzen 1998 1995 1995-01-01 Esbjerg Def 55.4667 8.4500
4190 Lutzen 1998 1997 1997-10-01 List, Sylt Def 55.0167 8.4333
4191 Lutzen 1998 1999 1998-06-01 Wilhelmshaven Def 53.5167 8.1333
4192 Lutzen 1998 1999 1982-01-01 Oudeschild, Texel Def 53.0333 4.8667
4193 Buizer 1981; Lutzen 1998 1974 1974-01-01 Den Helder Def 52.9667 4.7667
4194 Buizer 1981; Lutzen 1998 1980 1980-01-01 Lake Grevelingen Def 51.7333 3.9833
4195 Buizer 1980; Lutzen 1998 1974 1974-01-01 Oosterschelde Def 51.5500 4.0000
4196 Lutzen 1998 1988 1988-01-01 Zeebrugge Def 51.3333 3.2000
4197 Lutzen 1998 1995 1995-01-01 Ostend Def 51.2167 2.9167
4198 Lutzen 1998 1980 1980-01-01 Dunkerque Def 51.0500 2.3667
4199 Lutzen 1998 1980 1980-01-01 Ambleteuse Def 50.8000 1.6000
4200 Breton et al. 1995; Lutzen 1998 1977 1977-01-01 Le Havre Def 49.5000 0.1333
4201 Lutzen 1998 1971 1971-01-01 Dinard Def 48.6333 -2.0667
4202 Lutzen 1998 1973 1973-01-01 Brest Def 48.4000 -4.4833
4203 Lutzen 1998 1973 1973-01-01 Roscoff Def 48.7333 -3.9833
4204 Lutzen 1998 1968 1968-01-01 Dieppe Def 49.9333 1.0833
4205 Davis and Davis 2005 2004 2004-01-01 Jersey Def 49.2167 -2.1167
4206 Davis and Davis 2005 2004 2004-01-01 Guernsey Def 49.5833 -2.3333
4207 Davis and Davis 2005 2004 2004-01-01 La Rochelle Def 46.1500 -1.1500
4208 Davis and Davis 2005 2004 2004-01-01 Arcachon Def 44.6500 -1.1667
4209 Davis and Davis 2005 2004 2004-01-01 Santander Def 43.4647 -3.8044
4210 Davis and Davis 2005 2004 2004-01-01 Gijon Def 43.5411 -5.6644
4211 Davis and Davis 2005 1981 1981-01-01 Ria de Arosa Def 42.4667 -8.9500
4212 Davis and Davis 2005 1981 1981-01-01 Ria del Eo Def 43.5383 -7.0228
4213 Davis and Davis 2005 1992 1992-01-01 Ferrol Def 43.4833 -8.2333
4214 Davis and Davis 2005 2003 2003-09-01 Leixos Def 41.1903 -8.7033
4215 Davis and Davis 2005 2003 2003-09-01 Cascais Def 38.6917 -9.4155
4216 Davis and Davis 2005 2003 2003-09-01 Bom Successo Def 38.6933 -9.2100
4217 Holmes 1976 1972 1972-12-01 Melbourne Def -37.8500 144.9167
4218 Keough and Ross 1999 1977 1977-01-01 Sydney Def -33.8667 151.2500
4219 Hayward and Morley 2009 2002 2002-09-01 Auckland Def -36.8333 174.8333
4220 Biosecurity New Zealand 2005 2005 2005-10-01 Christchurch Def -43.5333 172.6333
4221 Biosecurity New Zealand 2005 2005 2005-11-01 Tutukaka Def -35.6000 174.5333
4231 Rednikorzev 1916, cited by Sanamyan 2000 1916 1916-01-01 Kurile Islands Native 50.8356 156.5847
4867 Davis and Davis 2008 2005 2005-06-01 Sette Def 43.3833 3.6000
6044 Locke et al. 2007; Arsenault et al. 2009 2002 2002-09-15 Malpeque Bay Def 46.5333 -63.7833
6778 MIT Sea Grant 2008 2007 2007-07-30 Journey's End Marina, Camden Def 44.2104 -69.0528
6779 MIT Sea Grant 2008 2007 2007-07-30 Darling Maine Center Dock, Walpole Def 43.9401 -69.5737
6830 White and Orr 2011 2008 2008-07-15 Bamfield Def 48.8150 -125.1583
767464 Ruiz et al., 2015 2013 2013-07-29 Mission Bay Yacht Club, Mission Bay, CA, California, USA Def 32.7778 -117.2485
767484 Ruiz et al., 2015 2013 2013-08-04 Bahia Resort Marina, Mission Bay, CA, California, USA Def 32.7731 -117.2478
767530 Ruiz et al., 2015 2013 2013-08-03 Mission Bay Sport Center, Mission Bay, CA, California, USA Def 32.7857 -117.2495
767558 Ruiz et al., 2015 2013 2013-08-02 The Dana Marina, Mission Bay, CA, California, USA Def 32.7671 -117.2363
767570 Ruiz et al., 2015 2013 2013-08-05 Paradise Point Resort, Mission Bay, CA, California, USA Def 32.7730 -117.2406
767671 Ruiz et al., 2015 2013 2013-07-16 Naval Base Point Loma, San Diego Bay, CA, California, USA Def 32.6886 -117.2343
767711 Ruiz et al., 2015 2013 2013-07-25 Navy Ammo Dock, Pier Bravo, San Diego Bay, CA, California, USA Def 32.6939 -117.2276
767806 Ruiz et al., 2015 2011 2011-09-15 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9117 -122.3494
767827 Ruiz et al., 2015 2011 2011-09-20 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8067 -122.4432
768069 Ruiz et al., 2015 2012 2012-09-11 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
768092 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6633 -122.3817
768182 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Def 37.7987 -122.3228
768304 Ruiz et al., 2015 2013 2013-08-20 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3163
768364 Ruiz et al., 2015 2013 2013-08-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6639 -122.3821

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