Invasion History

First Non-native North American Tidal Record: 1966
First Non-native West Coast Tidal Record: 1966
First Non-native East/Gulf Coast Tidal Record: 1978

General Invasion History:

Botrylloides violaceus was first described in Japan in 1927. It is native to the Northwest Pacific from northern Japan to southern Korea and northern China (Nishikawa 1991; Rho and Lee 1991; Rho and Park 1998; Rho et al. 2000).  Botrylloides violaceus is now widely introduced, being found in the Northeast Pacific, Northwest Atlantic, and parts of the Northeast Atlantic.

North American Invasion History:

Invasion History on the West Coast:

Botrylloides violaceus was first collected on the West coast in 1966 in Santa Barbara Harbor (Lambert, personal communication) and in San Francisco Bay in 1973 (Cohen and Carlton 1995). Subsequently, it was found in Bahia San Quintin, Mexico (Rodriguez and Ibarra-Obando 2008), Ensenada, Mexico (2000, Lambert and Lambert 2003), San Diego Bay, California (1994, Lambert, personal communication), Coos Bay, Oregon (1978, Carlton, unpublished data), Willapa Bay, Washington (1980, Cohen and Carlton 1995), Puget Sound, Washington (1998, Cohen et al. 1998), Prince William Sound, Alaska (1999, Lambert and Sanamyan 2001) and Kachemak Bay, Alaska (1999, Ruiz et al. 2006).

Invasion History on the East Coast:

On the East Coast, the history of the invasion of Botrylloides violaceus was complicated by confusions with B. diegensis. Botrylloides diegensis, possibly native to the northeast Pacific, was introduced to Eel Pond, adjacent to Woods Hole Harbor MA, by a scientist, who wanted a supply of experimental animals around 1972. He reported successful overwintering and reproduction of these animals (Carlton 1989). Consequently, later occurrences of Botrylloides sp. in the NW Atlantic were attributed to this species (Berman et al. 1992). Subsequent examination of Botrylloides sp. along the Northeast Coast of North America indicates that all the specimens found were actually B. violaceus (Gretchen Lambert, personal communication 2000), but the fate of the transplanted B. diegensis in the Eel Pond is unknown. Since B. violaceus is presumed to have been a recent (late 1970s) and local invader to the West Coast, it is unlikely that this species was introduced to Eel Pond instead of the reported B. diegensis. A separate invasion, around 1980 is more likely for B. violaceus.

Owing to the confusion with B. diegensis, the date of first introduction and pattern of early spread in the Northwest Atlantic is obscure. Whitlach et al. (1995) found B. violaceus in 1980 in Long Island Sound and by 1981-1982, B. violaceus had spread to Great Bay, New Hampshire, and to the Damariscotta River, Maine (1982, on oyster-culture nets, Dijkstra et al. 2007). By the mid 1990s it had reached Penobscot Bay (Whitlach and Osman 2001). In 2004, B. violaceus was found fouling aquaculture facilities in Savage Harbour, Prince Edward Island, Canada (Locke et al. 2007).

We are unsure of its range in estuaries south of Long Island Sound. However, in 2002, Gretchen Lambert confirmed our preliminary identifications of B. violaceus from lower Chesapeake Bay, collected in 2000 and 2001. Specimens were found on settling plates on both the Eastern (Kiptopeke/VA/Chesapeake Bay) and Western shores (Norfolk/VA/Little Creek (Hampton Roads); Poquoson/VA/Hampton Roads; Belle Isle Marina/VA/Rappahannock River; Amoco Refinery, Yorktown/VA/York River) (Ruiz et al., unpublished data). In July 2013, we found B. violaceus in the Indian River Inlet, Delaware, and the Chincoteague Inlet, Virginia, on the Atlantic coast of the DelMarVa peninsula (Paul Fofonoff, and Kristen Larson, personal observations). We are unsure of its invasion south of the Chesapeake Bay. Whether it is established here is unknown.

Invasion History Elsewhere in the World:

In the Northeast Atlantic, B. violaceus was first collected in 1993, in the Lagoon of Venice, in the Mediterranean Sea (Zaniolo et al. 1998). It has also been found in the Western Scheldt estuary, in the Netherlands, in 2000 (Gretchen Lambert, personal communication 2001) and in British waters in 2005 (MarLin 2006). In 2009, it was collected in three locations in Spain, Nazaré and Bueu on the Atlantic Ocean and Santander on the/Bay of Biscay (El Nagar et al. 2010).

Botrylloides violaceus has also been reported from the coast of Queensland, Australia (Kott 1985; Kott 1998), however, these reports may refer to another species (Gretchen Lambert, pers. comm.).


Description

Botrylloides violaceus is a colonial tunicate that can vary in color, ranging from purple, light lavender, red, yellow, orange and brown. In all cases the colony is entirely one color. Botrylloides violaceus colonies are encrusting, usually 2 – 3 mm in thickness (Saito et al. 1981) and can be large, up to 200 mm x 20 mm.  The tunic is soft, easily torn and the zooids are easily freed from the tunic (Lambert 2003). Zooids are arranged in ladder-like chains, with several common cloacal openings. Between chains of zooids the tunic surface is sometimes elevated. Zooids have 10-11 rows of stigmata and 9-12 stomach folds. Ova (reproductive cells) are located dorsal-posterior to testis, consisting of up to 16 follicles. The larvae of B. violaceus are incubated in the tunic. They are nourished by the tunic vascular system and continue to grow even after the adult zooid dies. The larvae are particularly large (up to 3 mm in length) with 24-32 lateral ampullae (Saito et al. 1981; Nishikawa 1991; Lambert 2003). Fully developed larvae are released from the incubatory pouch via the common cloacal openings (Saito et al. 1981).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Chordata
Subphylum:   Tunicata
Class:   Ascidiacea
Order:   Stolidobranchia
Family:   Styelidae
Genus:   Botrylloides
Species:   violaceus

Synonyms

Botrylloides violaceum (Oka, 1927)
Botryllus aurantius (Oka, 1927)
Botrylloides aurantium (Oka, 1927)
Botrylloides lateritium (Beniaminson, 1975)
Botrylloides carnosum (Oka, 1927)

Potentially Misidentified Species

Botrylloides diegensis
This name, for a botryllid possibly native to the West Coast, was widely used during early studies of B. violaceus' invasion on the East Coast of North America (Berman et al. 1992; Whitlach and Osman 1995; Whitlach et al. 1995; Osman and Whitlach 2000).

Botrylloides firmus
Records of B. violaceus from Australia (Kott 1985) were re-identified as B. firmus (Kott 2005).

Botryllus leachii
Northeast Atlantic species, used for B. violaceus on the East Coast by Myers (1990).

Botryllus schlosseri
None

Ecology

General:

Life History- A colonial (or compound) tunicate consists of many zooids, bearing most or all of the organs of a solitary tunicate, but modified to varying degrees for colonial life. Colonial tunicates of the genera Botrylloides have small zooids, usually not organized in systems, and fully embedded in a mass of tunic material. Each zooid has an oral siphon and an atrial canal, opening to a shared cloacal chamber. Water is pumped into the oral siphon, through finely meshed ciliated gills on the pharynx, where phytoplankton and detritus is filtered, and passed on mucus strings to the stomach and intestines. Excess waste is expelled in the outgoing atrial water (Van Name 1945; Barnes 1983).

Colonial tunicates reproduce both asexually, by budding, and sexually, from fertilized eggs developing into larvae. Buds can form from the body wall of the zooid. Colonies vary in size and can range from small clusters of zooids to huge spreading masses. The zooids are hermaphroditic, with eggs and sperm being produced by a single individual. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but many species have a partial block to self-fertilization. Eggs are internally fertilized and embryos are incubated in a brood pouch. Once they are mature, 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 can 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 (Van Name 1945; Barnes 1983).

In all part of its native and introduced range, B. violaceus 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, and eelgrass (White and Orr 2011; Simkanin et al. 2012; Wong and Vercaemer 2012; Carman et al. 2016). Predation may limit or slow the spread of B. violaceus to natural habitats (Simkanin et al. 2013). In habitats in Cape Cod and Massachusetts Bays, settlement was much higher on floating docks, than in eelgrass or rocky habitats (Wagstaff 2017). In experiments with controlled injections of larvae, increased propagule pressure resulted in increased recruitment, as did unoccupied vs occupied space, and plates on floating docks versus those on natural rocks (Simkanin et al. 2017).

Food:

Phytoplankton, detritus

Consumers:

fish, snails, crabs, urchins, starfish

Competitors:

Colonial tunicates, bryozoans

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatCoarse Woody DebrisNone
General HabitatGrass BedNone
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)-0.6Field, based on coldest site in geographical range, Boston MA (Zerebecki and Sorte 2011)
Maximum Temperature (ºC)27.4Experimental, 24 h 50% survival, using animals acclimated at 17 C from Lynn Harbor MA (Sorte et al. 2013). For Bodega Bay CA animals, acclimated at 12 C, the median lethal temperature was 25. 3 C (Zerebecki and Sorte 2011)
Minimum Salinity (‰)16Experimental data, Dijkstra and Harris 2007; Epelbaum et al. 2009
Maximum Salinity (‰)38highest tested, Epelbaum et al. 2009
Minimum Reproductive Salinity16Some colonies metamorphosed at salinities at 16-35 PSU, but showed reduced growth, and their colonies had fewer zooids than those at higher salinities (Lambert et al. 2016).
Minimum Duration0.2Larval period (Saito et al. 1981)
Maximum Duration0.4Larval period (Saito et al. 1981)
Broad Temperature RangeNoneCold temperate-Warm Temperate
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Economic Impacts

Shipping and Industry: The colonial ascidian Botrylloides violaceus is becoming an abundant component of ship and dock fouling communities in the Northeast and Northwest Atlantic, and the Northeast Pacific (Berman et al. 1992; Cohen and Carlton 1995; Lambert et al. 1998; Ruiz et al. 2000; Lambert and Lambert 2003; Dijkstra et al. 2007). It has been found in aquaculture operations in the Gulf of St. Lawrence, Canada but no negative impacts have been reported yet (Ramsay et al. 2008). Due to its abundance, wide distribution, and frequent dominance, this ascidian is likely to have substantial impacts on shipping, aquaculture, and fisheries.

Ecological Impacts

Competition: Botrylloides violaceus frequently displaces other fouling organisms, including native and introduced tunicates, bryozoans, barnacles, and mussels through competition for space and food. Evidence of this was found during experiments with fouling plates in New England waters (Myers 1990; Osman and Whitlatch 1995; Stachowicz et al. 1999; Osman and Whitlatch 2000; Stachowicz et al. 2002a,b; Osman and Whitlatch 2004; Bullard et al. 2004; Agius 2007; Altman and Whitlatch 2007; Rajbanshi and Pederson 2007; Dijkstra and Harris 2009). B.violaceus, in conjunction with other introduced tunicates, is abundant in harbors of Long Island Sound but its abundance decreases sharply in outer coast locations where colonies are susceptible to predation by fishes and gastropods. In experiments, B. violaceus was found to be most successful in communities of low diversity (Stachowicz et al. 1999; Stachowicz et al. 2002a) and in years with high water temperatures in spring (Stachowicz et al. 2002b). On the West Coast, B. violaceus is a recent invader of San Diego Bay, CA. At two locations in California (Mission Bay, 1997; Los Angeles Harbor, 2000) it has formed extensive areas of 100% cover, indicating strong competitive ability (Lambert and Lambert 2003). B. violaceus was one of several invasive fouling species which showed increased growth (% coverage) at temperatures above the ambient temperature of 13.5C in Bodega Harbor, California, while the native species Distaplia occidentalis showed reduced survival (Sorte et al. 2010). Modelling and field observations indicate that increasing water temperatures in the Gulf of Maine will increase the season for asexual reproduction of B. violaceus, as well as the frequency of sexual reproduction (Dijkstra et al. 2017).

Botrylloides violaceus, Botryllus schlosseri, and a native sponge Halichrondria panicea were found to adversely affect native eelgrass Zostera marina in southeastern Nova Scotia by fouling the leaves of the grass, and reducing the availability of light. Fouling increased the mortality of the plants. The violet morph of B. violaceus had a more negative effect than the lighter-colored tunicates, which transmitted more light through their bodies (Wong and Vercaemer 2012). Negative effects on eelgrass are likely to be widespread.

Habitat Change: By 2003-2006, tunicates, including B. violaceus, replaced the mussel Mytilus edulis as the dominant species in fouling communities in Portsmouth Harbor, New Hampshire (Dijkstra and Harris 2009). A major functional habitat change occured because while mussels (e.g. M. edulis) provided a year-round structure for other organisms to settle upon, in what is known as secondary settlement, most tunicates do not provide this secondary settlement structure type. However, B. violaceus dies off seasonally which creates large areas of bare substrate for organisms to colonize (Dijkstra and Harris 2009). Based on experiments on fouling of eelgrass plants in Nova Scotia, the spread of Botrylloides violaceus and Botryllus schlosseri is likely to have an adverse impact on eelgrass beds, increasing mortality of the plants and decreasing their productivity (Wong and Vercaemer 2012).

Herbivory: Experiments by Byrne and Stachowicz (2009) indicate that B. violaceus has a lower filtration rate than the possibly native B. diegensis in Bodega Harbor, California. Similar results were obtained for other exotic/native species pairs. It is suggested that the cumulative effect of increased invasions in fouling filter-feeding communities may increase seasonal consistency of filtration, due to spreading out of recruitment times, rather than increased rates.

Food/Prey: Gastropods (Costoanachis avara, C. translirata), crabs, starfish, and fishes (Tautogolabrus adspersus, Tautoga onitis) may prey upon newly settled (less than one week old) B. violaceus in Long Island Sound, NY. This predation may restrict the tunicate's distribution, especially in open coast areas (Osman and Whitlatch 2004).

Regional Impacts

NA-ET3Cape Cod to Cape HatterasEcological ImpactCompetition
Botrylloides violaceus is a strong competitor for space with other fouling organisms (Myers 1990; Osman and Whitlatch 1995; Stachowicz et al. 1999; Stachowicz et al. 2002a; Stachowicz et al. 2002b; Bullard et al. 2004; Agius 2007; Altman and Whitlach 2007).
NA-ET3Cape Cod to Cape HatterasEcological ImpactFood/Prey
Botrylloides violaceus is preyed on by gastropods (Costanachis avara, C. translirata), by crabs, starfish, and by fishes (Tautogolabrus adspersa, Tautoga onitis), which resitrict the tunicate's distribution, especially in open coast areas (Osman and Whitlatch 2004).
NA-ET2Bay of Fundy to Cape CodEcological ImpactCompetition
Botrylloides violaceus is a strong competitor for space with other fouling organisms including the native mussel Mytilus edulis and the introduced tunicate Diplosoma listerianum in Boston Harbor (Rajbanshi and Pederson 2007).  Botylloides violaceus was the most abundant colonial tunicate on fouling plates in Portsmouth Harbor in 1984-1985 (Berman et al. 1992) and in 2003-2005, partially replacing B. schlosseri, the previous dominant colonial ascidian (Dijkstra et al. 2007). In Portsmouth Harbor, by 2003-2006, B. violaceus, along with Didemnum vexillum, replaced the mussel M. edulis (1979-1982) as dominant species in fouling communities (Dijkstra and Harris 2009).
M010Buzzards BayEcological ImpactCompetition
Botrylloides violaceus is a strong competitor for space with other fouling organisms (Myers 1990; Agius 2007).
M040Long Island SoundEcological ImpactCompetition
Botrylloides violaceus is a strong competitor for space with other fouling organisms (Osman and Whitlatch 1995; Stachowicz et al. 1999; Stachowicz et al. 2002a; Stachowicz et al. 2002b; Bullard et al. 2004; Altman and Whitlatch 2007).
M040Long Island SoundEcological ImpactFood/Prey
Newly setteld (less than one week old) Botrylloides violaceus is preyed on by gastropods (Costanachis avara, C. translirata) and by fishes (Tautogolabrus adspersa, Tautoga onitis), which may restrict the tunicate's distribution, especially in open coast areas (Osman and Whitlatch 2000).
N170Massachusetts BayEcological ImpactCompetition
Botrylloides violaceus is a strong competitor for space with other fouling organisms including the native mussel Mytilus edulis and the introduced tunicate Diplosoma listerianum in Boston Harbor (Agius 2007; Rajbanshi and Pederson 2007).
NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactCompetition
The colonial tunicate Botrylloides violaceus is a recent invader of San Diego Bay. At two locations, in Mission Bay, in 1997, and in Los Angeles Harbor in 2000, it formed extensive areas of 100% cover, indicating strong competitive ability (Lambert and Lambert 2003).
P030Mission BayEcological ImpactCompetition
The colonial tunicate Botrylloides violaceus is a recent invader of San Diego Bay. At one location in Mission Bay, in 1997 (South Shore Boat Ramp) and one in Los Angeles Harbor in 2000, it formed extensive areas of 100% cover, indicating strong competittive ability (Lambert and Lambert 2003).
P050San Pedro BayEcological ImpactCompetition
The colonial tunicate Botrylloides violaceus is a recent invader of San Diego Bay. At two locations, in Mission Bay, in 1997, and in Los Angeles Harbor (Watchorn Marina) in 2000, it formed extensive areas of 100% cover, indicating strong compeition (Lambert and Lambert 2003).
NA-S3NoneEconomic ImpactFisheries
Botrylloides violaceus was seen overgrowing mussel lines on Prince Edward Island (Gittenberger 2009). High-pressure water spraying reduced fouling of mussels. However, fouling by Botryllus schlosseri and Botrylloides violaceus had little effect on mussel growth and production (Arens et al. 2011). The abundance of B. violaceus was much smaller than that of B. schlosseri, so impacts were smaller (Paetzold et al. 2012)/
N130Great BayEcological ImpactCompetition
B. violaceus was the most abundant colonial tunicate on fouling plates in Portsmouth Harbor in 2003-2005, partially replacing B. schlosseri, the previous dominant colonial tunicate (Dijkstra et al. 2007). In Portsmouth Harbor, by 2003-2006, B. violaceus and to a lesser extent, D. vexillum replaced the mussel M. edulis (1979-1982) as the dominant species in fouling communities (Dijkstra and Harris 2009).
N120Wells BayEcological ImpactCompetition
In experiments in Wells Harbor, Maine, Botrylloides violaceus grew rapidly on some artficial substrates (rubber and metal), outcompeting native organisms, but grew more slowly on natural substrates (shell, marble, slate) (Tyrell and Byers 2007).
P130Humboldt BayEcological ImpactCompetition
In fouling plate experiments in Humboldt Bay (Nelson 2009), found that colonial tunicates (Botryllus schlosseri and Botrylloides violaceus), growing in sheets, were able to quicly occupy space on fouling plates, but did not decrease recruitment or species richness.
NEP-IVPuget Sound to Northern CaliforniaEcological ImpactCompetition
In fouling plate experiments in Humboldt Bay, (Nelson 2009) found that colonial tunicates (Botryllus schlosseri and Botrylloides violaceus), growing in sheets, were able to quicly occupy space on fouling plates, but did not decrease recruitment or species richness.
NA-ET2Bay of Fundy to Cape CodEcological ImpactHabitat Change
In Portsmouth Harbor, by 2003-2006, B. violaceus and to a lesser extent, D. vexillum replaced the mussel M. edulis (1979-1982) as dominant species in fouling communities (Dijkstra and Harris 2009). A major functional change is that while mussel shells provided a year-round structure on the substrate, available to settlement by other organisms, colonial tunicates are more resistant to secondary settlement, and die off seasonally, creating large areas of bare substrate which can be colonized by other organisms (Dijkstra and Harris 2009).
N130Great BayEcological ImpactHabitat Change
In Portsmouth Harbor, by 2003-2006, B. violaceus and to a lesser extent, D. vexillum replaced the mussel M. edulis (1979-1982) as dominant species in fouling communities (Dijkstra and Harris 2009). A major functional change is that while mussel shells provide structure, available for settlement and colonization by other organisms, the colonial tunicates are more resistant to secondary settlement, and die off seasonally, creating large areas of bare substrate which can be colonized by other organisms (Dijkstra and Harris 2009).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
Botrylloides violaceus was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5⁰C above the ambient temperature in Bodega Harbor (13.5⁰C), while tha native Distaplia occidentalis showed reduced survival (Sorte et al. 2010). Botrylloides violaceus was one of a group of 7 non-native species, most of which were rare or absent in 1970-1971 species, but were among the 8 most abundant species in 2006. Spawning periods and abundance of species in this group appeared to be favored by a 1⁰C increase in average temperatures at this site over a 30-year period (Sorte and Stachowicz 2011).
P112_CDA_P112 (Bodega Bay)Ecological ImpactCompetition
Botrylloides violaceus was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5⁰C above the ambient temperature in Bodega Harbor (13.5⁰C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010). Botrylloides violaceus was one of a group of 7 non-native species, most of which were rare or absent in 1969-1971 surveys, but were among the 8 most abundant species in 2006. Spawning periods and abundance of species in this group appeared to be favored by a 1⁰C increase in average temperatures at this site over a 30-year period (Sorte and Stachowicz 2011).
NA-ET3Cape Cod to Cape HatterasEconomic ImpactFisheries
Botrylloides violaceus was found fouling aquaculture gear at 18 sites, and cultured Bay Scallops (Argopecten 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 New York State and Rhode Island (Carman et al. 2010).
NA-ET2Bay of Fundy to Cape CodEconomic ImpactFisheries
Botrylloides violaceus was reportedly fouling aquaculture sites in Maine (Carman et al. 2010; Bullard et al. 2015)
N195_CDA_N195 (Cape Cod)Economic ImpactFisheries
Botrylloides violaceus was found fouling aquaculture gear at 18 sites, and cultured Bay Scallops (Argopecten irradians) at two sites, of 26 aquaculture sites surveyed on Marthas Vineyard (Carman et al. 2010).
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactCompetition
The native eelgrass Zostera marina was adversely affected by fouling by Botrylloides violaceus. The burgundy colored morph had a greater effect than that of orange or cream-colored colonies, as indicated by lower chlorophyll concentrations in the leaf, and leaf mortality. However, fouling by a native sponge, Halichondria panicea, produced a greater reduction of chlorphyll that any of the morphs of B. violaceus, or Botryllus schlosseri (Wong and Vercaemer 2012).
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactHabitat Change
The spread of introduced fouling organisms (B. violaceus and B. schlosseri) to eelgrass beds is considered likely to reduce the primary productivity and the extent of grass beds in Nova Scotia waters (Wong and Vercaemer 2012).
NEA-IINoneEcological ImpactCompetition
Botrylloides violaceus appeared to compete and outgrow Botryllus schlosseri on fouling plates in some Netherlands estuaries, where salinity was above 30 PSU, but was rare or absent in estuaries with lower salinities (14-29 PSU) (Gittenberger and Moons 2011).
NEA-IINoneEconomic ImpactShipping/Boating
Fouling impacts fave been reported for the British Isles (Minchin et al. 2013).
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).
N070Damariscotta RiverEconomic ImpactFisheries
Botrylloides violaceus was reportedly fouling aquaculture sites on the Damariscotta River (Bullard et al. 2015)
NEP-IVPuget Sound to Northern CaliforniaEcological ImpactFood/Prey
In feeding trials, the native crabs Hemigrapsus oregonensis, the flatworm Eurylepta leoparda and the nudibranch Hermissenda crassicornis fed heavily on the native tuinicate Distaplia occidentalis but at much lower rates on the non-native Botryllus schlosseri and Botrylloides violaceus) (Kincaid and de Rivera 2020).
P170Coos BayEcological ImpactFood/Prey
In feeding trials, the native crabs Hemigrapsus oregonensis, the flatworm Eurylepta leoparda and the nudibranch Hermissenda crassicornis fed heavily on the native tuinicate Distaplia occidentalis but at much lower rates on the non-native Botryllus schlosseri and Botrylloides violaceus) (Kincaid and de Rivera 2021).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEP-V Northern California to Mid Channel Islands 1973 Def Estab
NWP-4a None 0 Native Estab
NEP-IV Puget Sound to Northern California 1979 Def Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1998 Def Estab
NEP-II Alaska south of Aluetians to the Alaskan panhandle 1999 Def Unk
NA-ET3 Cape Cod to Cape Hatteras 1980 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 1981 Def Estab
MED-VII None 1993 Def Estab
NWP-3a None 0 Native Estab
NEP-VI Pt. Conception to Southern Baja California 1966 Def Estab
NEA-II None 2000 Def Estab
NA-S3 None 2004 Def Estab
M010 Buzzards Bay 1998 Def Estab
P050 San Pedro Bay 1994 Def Estab
P170 Coos Bay 1978 Def Estab
P130 Humboldt Bay 1992 Def Estab
P270 Willapa Bay 1979 Def Estab
M060 Hudson River/Raritan Bay 2003 Def Estab
M020 Narragansett Bay 2000 Def Estab
M040 Long Island Sound 1980 Def Estab
M130 Chesapeake Bay 2000 Def Estab
N130 Great Bay 1981 Def Estab
P020 San Diego Bay 1994 Def Estab
NEA-III None 2005 Def Estab
P030 Mission Bay 1994 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 1995 Def Estab
P040 Newport Bay 1997 Def Estab
P060 Santa Monica Bay 1998 Def Estab
P064 _CDA_P064 (Ventura) 1996 Def Estab
P062 _CDA_P062 (Calleguas) 1997 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 1966 Def Estab
P080 Monterey Bay 1998 Def Estab
P090 San Francisco Bay 1973 Def Estab
P110 Tomales Bay 2001 Def Estab
P112 _CDA_P112 (Bodega Bay) 2000 Def Estab
P286 _CDA_P286 (Crescent-Hoko) 2003 Def Estab
P290 Puget Sound 1998 Def Estab
P297 _CDA_P297 (Strait of Georgia) 1998 Def Estab
M030 Gardiners Bay 2003 Def Estab
M013 _CDA_M013 (Cape Cod) 1998 Def Estab
N185 _CDA_N185 (Cape Cod) 2000 Def Estab
N180 Cape Cod Bay 1998 Def Estab
N170 Massachusetts Bay 2000 Def Estab
N165 _CDA_N165 (Charles) 1998 Def Estab
N140 Hampton Harbor 2003 Def Estab
N135 _CDA_N135 (Piscataqua-Salmon Falls) 1998 Def Estab
N125 _CDA_N125 (Piscataqua-Salmon Falls) 1998 Def Estab
N116 _CDA_N116 (Piscataqua-Salmon Falls) 1998 Def Estab
N100 Casco Bay 1998 Def Estab
N070 Damariscotta River 1978 Def Estab
N050 Penobscot Bay 1998 Def Estab
N010 Passamaquoddy Bay 2004 Def Estab
NWP-4b None 0 Native Estab
NWP-3b None 0 Native Estab
P180 Umpqua River 1986 Def Estab
N195 _CDA_N195 (Cape Cod) 2003 Def Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 2007 Def Estab
P027 _CDA_P027 (Aliso-San Onofre) 2001 Def Estab
P058 _CDA_P058 (San Pedro Channel Islands) 2001 Def Estab
P070 Morro Bay 2001 Def Estab
M023 _CDA_M023 (Narragansett) 2007 Def Estab
N120 Wells Bay 2004 Def Estab
NEA-V None 2009 Def Estab
P210 Yaquina Bay 2010 Def Estab
N080 Sheepscot Bay 1998 Def Estab
P292 _CDA_P292 (San Juan Islands) 2005 Def Estab
P288 _CDA_P288 (Dungeness-Elwha) 2005 Def Estab
AUS-XII None 2003 Def Unk
N040 Blue Hill Bay 2009 Def Estab
NEP-VII None 2012 Def Estab
NEP-VIII None 2012 Def Estab
WA-I None 2007 Def Estab
P293 _CDA_P293 (Strait of Georgia) 2007 Def Estab
M080 New Jersey Inland Bays 2013 Def Estab
M100 Delaware Inland Bays 2013 Def Estab
M120 Chincoteague Bay 2013 Def Estab
NEA-VI None 2009 Def Estab
NEA-IV None 2005 Def Estab
M050 Great South Bay 2013 Def Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 0 Def Unk
G300 Aransas Bay 0 Def Unk
G102 _CDA_G102 (Apalachicola Bay) 2011 Def Unk
G074 _CDA_G074 (Crystal-Pithlachascotee) 2012 Def Unk
G010 Florida Bay 2012 Def Unk
S200 Biscayne Bay 2015 Def Unk
AR-IV None 2018 Def Unk

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
4451 Lambert and Lambert 2003 2000 2000-08-01 Ensenada Def 31.8667 -116.6167
4453 Lambert and Lambert 2003 1995 1995-05-01 Harbor Island, San Diego Def 32.7253 -117.2064
4454 Lambert and Lambert 2003 1995 1995-05-01 Fiddlers Cove, San Diego Def 32.6519 -117.2342
4457 Lambert and Lambert 2003 1995 1995-05-01 Dana Landing Def 32.7675 -117.2365
4458 Lambert and Lambert 2003 1998 1998-05-01 Bahia Point, San Diego Def 32.7756 -117.2467
4459 Lambert and Lambert 2003 1997 1997-05-01 Mission Bay Yacht Club, San Diego Def 32.7778 -117.2489
4461 Lambert and Lambert 2003 1996 1996-10-01 Oceanside Def 33.2078 -117.3950
4462 Lambert and Lambert 2003 1997 1997-05-01 Fun Zone, Newport Def 33.6084 -117.9092
4463 Lambert and Lambert 2003 1995 1995-05-01 Impound Marina, Long Beach Def 33.7639 -118.2444
4464 Lambert and Lambert 2003 1998 1998-05-01 Long Beach Marina Def 33.7545 -118.1290
4465 Lambert and Lambert 2003 1994 1994-10-01 Watchorn Marina, Long Beach Def 33.7203 -118.2764
4467 Lambert and Lambert 2003 1998 1998-05-01 Santa Monica Def 33.9722 -118.4522
4468 Lambert and Lambert 2003 1997 1997-05-01 Jack's Landing Def 34.1636 -119.2228
4469 Lambert and Lambert 2003 1997 1997-05-01 Anacapa Island Marina Def 34.1731 -119.2269
4471 Lambert and Lambert 2003 1997 1997-05-01 Island Packers, Ventura Def 34.2495 -119.2648
4472 Lambert and Lambert 2003 1966 1966-01-01 Santa Barbara Def 34.4072 -119.6887
4474 Wasson et al. 2000; de Rivera et al. 2005 1998 1998-01-01 Moss Landing Def 36.8002 -121.7872
4476 Wasson et al. 2000; de Rivera et al. 2005 1998 1998-01-01 South Marsh trail, Moss Landing Def 36.8181 -121.7396
4478 de Rivera et al. 2005a 2003 2003-08-01 Santa Cruz Def 36.9658 -122.0016
4481 Cohen et al. 2005 2004 2004-05-24 San Leandro Marina Def 37.6966 -122.1932
4482 Cohen et al. 2005 2004 2004-05-24 Coast Guard Island, Oakland Def 37.7812 -122.2458
4484 Cohen et al. 2005 2004 2004-05-26 Richmond Marina Def 37.9139 -122.3542
4485 Cohen et al. 2005 2004 2004-05-27 Pete's Harbor Def 37.5006 -122.2242
4488 Cohen et al. 2005 2004 2004-05-24 Fruitvale Bridge, Alameda Def 37.7690 -122.3906
4490 de Rivera et al. 2005a 2003 2003-08-01 Clark Def 38.1810 -122.9105
4491 de Rivera et al. 2005a 2003 2003-08-01 Marshall Def 38.1497 -122.8885
4492 de Rivera et al. 2005a 2003 2003-08-01 Miller Park Def 38.1996 -122.9217
4493 de Rivera et al. 2005a 2003 2003-08-01 Nick's Cove Def 38.2010 -122.9228
4494 de Rivera et al. 2005a 2003 2003-08-01 Sacramento Landing Def 38.1504 -122.9058
4495 de Rivera et al. 2005a 2003 2003-08-01 Shell Beach Def 38.1147 -122.8694
4496 de Rivera et al. 2005a 2003 2003-08-01 Thomas Station Def 38.1287 -122.8654
4497 de Rivera et al. 2005a 2003 2003-08-01 Tomales Beach Def 38.1756 -122.9234
4499 de Rivera et al. 2005a 2003 2003-08-01 Porto Bodega Marina Def 38.3344 -123.0526
4500 de Rivera et al. 2005 2003 2003-08-01 Mason's Marina Def 38.3321 -123.0588
4501 deRivera et al. 2005 2001 2001-09-18 Humboldt Bay Def 40.8074 -124.1666
4502 Carlton, unpublished data; de Rivera et al. 2005a 1979 1979-01-01 Charleston Boat Basin Def 43.3465 -124.3267
4503 de Rivera et al. 2005a 2003 2003-08-01 Empire Pier Def 43.3933 -124.2812
4504 de Rivera et al. 2005a 2003 2003-08-01 Port of Coos Bay Citrus Dock Def 43.3823 -124.2193
4505 de Rivera et al. 2005a 2003 2003-08-01 Valino Island Def 43.3167 -124.3214
4506 Carlton 1979; Cohen et al, 2001 2001 1979-01-01 Nahcotta Small Boat Basin Def 46.5007 -124.0307
4507 Cohen et al. 2001 2000 2000-05-01 Wilson Point Def 46.6494 -123.9527
4508 Cohen et al. 2001 2000 2000-05-22 Stackpole Slough, Willapa Bay Def 46.6059 -124.0432
4509 de Rivera et al. 2005 2003 2003-08-01 Makah Marina Def 48.3677 -124.6116
4510 Cohen et al. 1998 1998 1998-09-01 Seabeck Marina Def 47.6462 -122.8277
4511 Cohen et al. 1998 1998 1998-09-10 Port Townsend Def 48.1170 -122.7605
4512 Cohen et al. 1998 1998 1998-09-09 Grapeview (Fairharbor Marina) Def 47.3318 -122.8351
4513 Cohen et al. 1998 1998 1998-09-08 Edmonds (Port of Edmonds Marina) Def 47.8084 -122.3899
4514 Cohen et al. 2001 2000 2000-05-19 Taylor Shellfish Rafts Def 47.1529 -122.9644
4515 Cohen et al. 1998 19998 1998-09-08 Des Moines (City Marina) Def 47.3993 -122.3299
4516 Cohen et al. 1998 1998 1998-09-11 Deception Pass Marina Def 48.4057 -122.6463
4517 Cohen et al. 1998 1998 1998-09-10 Brownsville Marina Def 47.6557 -122.6165
4518 Cohen et al. 1998 1998 1998-09-11 Anacortes (Cap Sante Boat Haven) Def 48.5110 -122.6094
4519 Cohen et al. 1998 1998 1998-09-11 Blaine (Blaine Marina) Def 48.9940 -122.7596
4520 Lambert, unpublished, cited by Cohen 2005 1993 1993-01-01 French Creek, Vancouver Island Def 49.3500 -124.3500
4521 Lambert, unpublished, cited by Cohen 2005 1993 1993-01-01 Maple Bay, Vancouver Island Def 48.8167 -123.6167
4522 Ruiz et al., unpublished data 2003 2003-01-01 Ketchikan Def 55.3428 -131.6486
4523 Ruiz et al., unpublished data 2001 2003-01-01 Galankin Island Def 57.0311 -135.3267
4524 Lambert and Sanamyan 2001 2000 2000-03-24 Sitka Sea Farms Def 57.0539 -135.3472
4527 Ruiz et al., unpublished data 2000 2000-08-01 Norfolk Def 36.9124 -76.1848
4528 Ruiz et al., unpublished data 2000 2000-08-01 Poquoson Def 37.1840 -76.4223
4529 Ruiz et al., unpublished data 2000 2000-08-01 Belle Isle Marina Def 37.0964 -76.2920
4530 MIT Sea Grant 2003 2003 2003-09-09 South Street Seaport, New York City Def 40.7065 -74.0032
4531 MIT Sea Grant 2003 2003 2003-08-08 Brewer Yacht Haven Marine Center, Stamford Def 41.0534 -73.5387
4533 MIT Sea Grant 2003 1998 1998-01-01 Bridgeport Def 41.1634 -73.1754
4535 MIT Sea Grant 2003 2003 2003-08-07 Brewer Yacht Yard, Mystic Def 41.3334 -71.9759
4536 Whitlach and Osman 1995 1980 1980-01-01 Avery Point (Groton) Def 41.3154 -72.0634
4537 Whitlach and Osman 1995 2003 2003-08-07 Stirling Yacht Harbor, Greenport, Long Island Def 41.1004 -72.3548
4538 MIT Sea Grant 2006 2004 2004-04-10 Beavertail State Park Def 41.4490 -71.3995
4539 MIT Sea Grant 2003 2000 2000-08-15 Coasters Harbor Island Def 41.5107 -71.3270
4540 MIT Sea Grant 2003 2003 2003-08-06 Newport Shipyard Def 41.4901 -71.3217
4541 (1998, Whitlatch and Osman 2000 1998 1998-01-01 Jamestown Def 41.4971 -71.3673
4542 MIT Sea Grant 2003 2000 2000-08-15 Wickford Marina, Wickford Def 41.5754 -71.4423
4543 MIT Sea Grant 2003 2000 2000-08-17 Narragansett Bay T-wharf, Prudence Island Def 41.5882 -71.3245
4544 MIT Sea Grant 2003 2000 2000-08-16 North Kingston Def 41.6237 -71.4126
4545 MIT Sea Grant 2003 2000 2000-08-15 Roger Williams University Dock, Bristol Def 41.6484 -71.2609
4546 MIT Sea Grant 2003 2000 2000-08-16 Warwick Cove Marina Def 41.6839 -71.3917
4547 MIT Sea Grant 2003 2003 2003-08-14 Edgewood Yacht Club, Cranston Def 41.7765 -71.3884
4548 MIT Sea Grant 2003 2000 2000-08-11 Fall River Def 41.7062 -71.1620
4549 Whitlatch and Osman 2000 1998 1998-01-01 Tiverton Def 41.6246 -71.2064
4550 Whitlatch and Osman 2000 1998 1998-08-01 Sakonnet Point Def 41.4542 -71.1953
4551 Whitlatch and Osman 2000 1998 1998-08-01 Westport Def 41.5125 -71.0894
4553 MIT Sea Grant 2003 2003 2000-08-11 Massachusetts Maritime Academy, Bourne Def 41.7396 -70.6239
4554 Whitlatch and Osman 2000 1998 1998-08-01 West Falmouth Def 41.6057 -70.6495
4555 Whitlatch and Osman 2000 1998 1998-08-01 Hyannis Def 41.6315 -70.2870
4556 Whitlatch and Osman 2000 1998 1998-08-01 Chatham Def 41.7437 -69.9559
4558 Whitlatch and Osman 2000 1998 1998-08-01 Barnstable Def 41.7167 -70.2667
4559 MIT Sea Grant 2003 2000 2000-08-19 Sandwich Marina Def 41.7650 -70.4750
4560 Whitlatch and Osman 2000 1998 1998-08-01 Sagamore Def 41.7701 -70.5284
4561 Whitlatch and Osman 2000 1998 1998-08-01 Manomet Point, Plymouth Def 41.9268 -70.5389
4562 MIT Sea Grant 2003 2000 2000-08-09 Plymouth Town Wharf Def 41.9623 -70.6662
4563 MIT Sea Grant 2003 2000 2000-08-09 Duxbury Town Pier Def 42.0001 -70.6578
4564 MIT Sea Grant 2003 2000 2000-08-09 MWRA Quincy (sewage plant) Def 42.2918 -70.9745
4565 MIT Sea Grant 2003 2000 2000-08-07 Deer Island, Boston Def 42.3518 -70.9606
4566 MIT Sea Grant 2003 2000 2000-08-07 Rowes Wharf, Boston Def 42.3570 -71.0409
4567 MIT Sea Grant 2003 2000 2000-08-08 Hawthorne Cove Marina, Def 42.5220 -70.8823
4568 MIT Sea Grant 2003 2000 2000-08-08 Tucks Point Marina, Beverly Def 42.5676 -70.7787
4569 MIT Sea Grant 2003 2000 2000-08-08 Cape Ann Marina, Gloucester Def 42.6209 -70.6912
4570 MIT Sea Grant 2003 2003 2003-08-03 Hampton State Pier Def 42.9375 -70.8394
4571 Berman et al. 1989 1980 1980-08-01 Fox Point Def 43.1212 -70.8589
4572 Blezard 1999 1998 1998-08-01 Newcastle Def 43.0723 -70.7162
4573 Whitlach and Osman 2000 1998 1998-08-01 Kennebunkport Def 43.3618 -70.4767
4574 Whitlach and Osman 2000 1998 1998-08-01 Cape Porpoise Def 43.3634 -70.4320
4576 MIT Sea Grant 2003 2003 2003-08-04 Port Harbor Marine, South Portland Def 43.6414 -70.2414
4578 MIT Sea Grant 2003 2003 2003-08-04 Brewer South Freeport Marina Def 43.8204 -70.1053
4579 Whitlatch and Osman 2000 1998 1998-08-01 Boothbay Harbor Def 43.8465 -69.6348
4580 USGS Woods Hole Science Center 2006 2002 2002-10-07 Thrumcap Island Def 43.8203 -69.5497
4581 Whitlach and Osman 2000 1998 1998-08-01 Castine Def 44.3829 -68.7989
4582 Whitlach and Osman 2000 1998 1998-08-01 Belfast Def 44.4281 -69.0020
4583 Trott 2004 2004 9999-01-01 Eastport Def 44.9095 -67.0555
4584 Locke et al. 2005 2004 2004-12-01 Savage Harbour Def 46.4167 -62.8333
4590 Locke et al. 2007 2007 2007-01-01 Lunenburg Def 44.3781 -64.3097
5922 Rodriguez and Ibarra-Obando 2008 2005 2005-12-01 Bahia San Quintin Def 30.4500 -116.0000
6040 Lu et al. 2007 2005 2005-09-20 Esquimalt Def 48.4325 -123.4325
6767 Callahan et al. 2010 2008 2008-10-01 Belleoram Def 47.5272 -55.4092
6773 IT Sea Grant 2008 2007 2007-07-29 Darling Maine Center Dock Def 43.9401 -69.5737
6774 MIT Sea Grant 2008 2007 2007-07-30 Journey's End Marina, Rockland Def 44.1045 -69.1017
6775 MIT Sea Grant 2008 2007 2008-07-30 Wayfarer Marina, Camden Def 44.2104 -69.0528
6831 White and Orr 2011 2008 2008-01-01 Bamfield, Vancouver Island Def 48.8150 -125.1583
6849 Bock et al. 2011 2011 2011-01-01 St. Peter's Bay Def 46.4178 -62.5817
6850 Bock et al. 2011 2011 2011-01-01 Cardigan River Def 46.2048 -62.5193
6851 Bock et al. 2011 2011 2011-01-01 Aspy Bay Def 46.9333 -60.3992
6852 Bock et al. 2011 2011 2011-01-01 South Bar Def 46.2097 -60.1953
6854 Bock et al. 2011 2011 2011-01-01 Chester Def 44.5500 -64.2992
6855 Bock et al. 2011 2011 2011-01-01 Lockeport Def 43.7000 -65.0992
6856 Bock et al. 2011 2011 2011-01-01 Meteghan Def 44.2017 -66.1422
6873 Martin et al. 2011 2009 2009-09-01 Head Harbour, Campobello Island Def 44.9466 -66.9063
6889 Sephton et al. 2011 2009 2009-01-01 Wedgeport Def 43.7137 -65.9695
6890 Sephton et al. 2011 2009 2009-01-01 Petit de Grat Def 45.5025 -60.9640
6891 Sephton et al. 2011 2009 2009-01-01 Cheticamp, Cape Breton Island Def 46.6268 -61.0160
6892 Sephton et al. 2011 2006 2006-01-01 Big Bras d'Or Def 46.2812 -60.4250
6939 Tovar-Hernandez et al. 2012 2012 2012-01-01 La Paz Def 24.1422 -110.3108
6940 Tovar-Hernandez et al. 2012 2012 2012-01-01 Guaymas Def 27.9183 -110.8989
6941 Tovar-Hernandez et al. 2012 2012 2012-01-01 Topolobampo Def 25.6167 -109.0500
6942 Tovar-Hernández et al. 2012 None 9999-01-01 Mazatlan Def 23.2200 -106.4200
6943 Tovar-Hernández et al. 2012 2012 2012-01-01 Puerto Vallarta, Def 20.6667 -105.2667
7182 Nishikawa 1991 None 9999-01-01 Akkeshi Native 43.0500 144.8500
7183 Nishikawa 1991 None 9999-01-01 Mikawa Bay Native 34.7667 137.0833
7184 Nishikawa 1991 None 9999-01-01 Iwaya, Native 34.7042 135.2178
7185 Nishikawa 1991 None 9999-01-01 Toshijima Island Native 34.5167 136.8830
7186 Huang 2001 None 9999-01-01 Liangyungang Native 34.6000 119.1667
7187 Nishikawa 1991 None 9999-01-01 Jiaozhou Bay Native 36.1699 120.2983
7188 Huang 2001 None 9999-01-01 Yantai Native 37.4000 121.1000
7189 Huang 2001 None 9999-01-01 Penglai Native 37.8167 120.7333
7190 Nishikawa 1991 None 9999-01-01 Maizuru Bay Native 35.4667 135.3833
7191 Nishikawa 1991 None 9999-01-01 Anamizu Bay Native 37.2333 136.9167
7192 Nishikawa 1991 None 9999-01-01 Moura-ko Native 40.9333 140.8500
7193 Nishikawa 1991 None 9999-01-01 Yaemon-misaki Native 42.0790 139.4980
7194 Nishikawa 1991 None 9999-01-01 Esashi Native 41.8667 140.1333
7195 Nishikawa 1991 None 9999-01-01 Ofuyu Native 43.7347 141.3397
7196 Nishikawa 1991 None 9999-01-01 Rebun Island Native 45.3500 141.0167
7197 Rho 1995 None 9999-01-01 Gojedo Island Native 34.8581 128.6183
7198 Rho et al. 2000 None 9999-01-01 Tangsa Native 35.5747 129.4506
7199 Rho et al. 2000 None 9999-01-01 Rho et al. 2000 Native 34.3833 126.4830
7200 Rho 1995 None 9999-01-01 Chindo Island Native 34.9269 128.0319
7203 Zaniolo et al. 1998 1993 1993-01-01 Lagoon of Venice Def 45.4131 12.2972
7204 El Nagar et al. 2010 2009 2009-06-13 Santander Def 43.4620 -3.7940
7205 El Nagar et al. 2010 2009 2009-07-18 Bueu Def 42.3280 -8.7860
7206 El Nagar et al. 2010 2009 2009-07-16 Nazaré Def 39.5840 -9.0750
7207 MarLin 2006 2005 2005-01-01 Queen Anne's Battery marina pontoon, Plymouth Def 50.3714 -4.1424
7208 Gittenberger 2007 2000 2000-01-01 Beskens Def 51.4000 3.5500
7209 Arenas et al. 2006 2004 2004-09-02 Gosport Def 50.7948 -1.1243
7210 Arenas et al. 2006 2004 2004-09-04 Poole Def 50.7167 -1.9833
7211 2004, Arenas et al. 2006 2004 2004-09-06 Exmouth Def 50.6200 -3.4130
7212 Minchin 2007 2006 2006-01-01 Malahide Marina Def 53.4543 -6.1535
7214 Minchin 2007 2006 2006-06-28 Carlingford Marina Def 54.0502 -6.1878
7215 Kerkchof et al. 2007 2004 2004-01-01 Zeebrugge Def 51.3333 3.2000
7216 Kerckhof et al. 2007 2004 2004-01-01 Oostende Def 51.2333 2.9167
7217 Gittenberger et al. 2010 2009 2009-08-01 Schiermonnikoog Def 53.4914 6.2286
767324 Ruiz et al., 2015 2012 2012-08-13 Coast Guard, Bodega Bay, California, USA Def 38.3126 -123.0512
767330 Ruiz et al., 2015 2012 2012-08-14 Spud Point South, Bodega Bay, California, USA Def 38.3281 -123.0574
767336 Ruiz et al., 2015 2012 2012-08-14 Spud Point North, Bodega Bay, California, USA Def 38.3301 -123.0572
767346 Ruiz et al., 2015 2012 2012-08-21 Lucas/Tides, Bodega Bay, California, USA Def 38.3284 -123.0445
767354 Ruiz et al., 2015 2012 2012-08-21 Porto Bodega, Bodega Bay, California, USA Def 38.3333 -123.0525
767366 Ruiz et al., 2015 2012 2012-08-22 Tomales-Marshall, Bodega Bay, California, USA Def 38.1514 -122.8888
767377 Ruiz et al., 2015 2012 2012-08-21 Tomales-Nick's Cove, Bodega Bay, California, USA Def 38.1980 -122.9222
767397 Ruiz et al., 2015 2012 2012-08-16 Tomales-SNPS, Bodega Bay, California, USA Def 38.1359 -122.8719
767409 Ruiz et al., 2015 2012 2012-08-17 Tomales- Shell Beach, Bodega Bay, California, USA Def 38.1163 -122.8713
767441 Ruiz et al., 2015 2013 2013-07-23 Marina Village, Mission Bay, CA, California, USA Def 32.7605 -117.2364
767458 Ruiz et al., 2015 2013 2013-07-29 Mission Bay Yacht Club, Mission Bay, CA, California, USA Def 32.7778 -117.2485
767509 Ruiz et al., 2015 2013 2013-08-01 Hyatt Resort Marina, Mission Bay, CA, California, USA Def 32.7634 -117.2397
767580 Ruiz et al., 2015 2013 2013-08-30 201 Main, Morro Bay, CA, California, USA Def 35.3564 -120.8474
767643 Ruiz et al., 2015 2013 2013-09-03 State Park Marina, Morro Bay, CA, California, USA Def 35.3459 -120.8423
767706 Ruiz et al., 2015 2013 2013-07-25 Navy Ammo Dock, Pier Bravo, San Diego Bay, CA, California, USA Def 32.6939 -117.2276
767802 Ruiz et al., 2015 2011 2011-09-15 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9117 -122.3494
767823 Ruiz et al., 2015 2011 2011-09-20 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8067 -122.4432
767835 Ruiz et al., 2015 2011 2011-09-14 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5880 -122.3160
767856 Ruiz et al., 2015 2011 2011-09-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6725 -122.3864
767880 Ruiz et al., 2015 2011 2012-09-15 Berkeley Marina, San Francisco Bay, CA, California, USA Def 37.8758 -122.3181
767890 Ruiz et al., 2015 2011 2012-09-19 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8609 -122.4853
767907 Ruiz et al., 2015 2011 2011-09-21 South Beach Harbor, San Francisco Bay, CA, California, USA Def 37.7797 -122.3871
767920 Ruiz et al., 2015 2011 2011-09-20 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7947 -122.2822
767932 Ruiz et al., 2015 2011 2011-09-22 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
767968 Ruiz et al., 2015 2011 2011-09-12 Corinthian Yacht Club, San Francisco Bay, CA, California, USA Def 37.8103 -122.3228
767986 Ruiz et al., 2015 2012 2012-08-24 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9134 -122.3523
768006 Ruiz et al., 2015 2012 2012-08-23 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8609 -122.4853
768021 Ruiz et al., 2015 2012 2012-08-28 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8071 -122.4341
768040 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
768062 Ruiz et al., 2015 2012 2012-09-11 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
768085 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6633 -122.3817
768109 Ruiz et al., 2015 2012 2012-08-29 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3174
768154 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9736 -122.4802
768175 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Def 37.7987 -122.3228
768251 Ruiz et al., 2015 2012 2012-09-12 Emeryville, San Francisco Bay, CA, California, USA Def 37.8396 -122.3133
768277 Ruiz et al., 2015 2013 2013-08-15 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7656 -122.2858
768297 Ruiz et al., 2015 2013 2013-08-20 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3163
768339 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9723 -122.4829
768357 Ruiz et al., 2015 2013 2013-08-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6639 -122.3821
768381 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.5024 -122.2134
768401 Ruiz et al., 2015 2013 2013-08-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9138 -122.3522
768418 Ruiz et al., 2015 2013 2013-08-12 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8078 -122.4354
768449 Ruiz et al., 2015 2013 2013-08-16 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8611 -122.4851

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