Invasion History

First Non-native North American Tidal Record: 1947
First Non-native West Coast Tidal Record: 1947
First Non-native East/Gulf Coast Tidal Record: 1975

General Invasion History:

The origin of Botryllus schlosseri is currently unknown. In the Northeast Atlantic, it ranges from Norway to the Mediterranean (Hayward and Ryland 1991; Rinkevich et al. 1995; Ben-Shlomo et al. 2001; Lejeusne et al. 2010). On the coast of Asia, it ranges from Peter the Great Bay, Russia to Hong Kong and southern China (Nishikawa 1991; Huang 2001). It is now widespread on temperate coasts on both sides of the North and South Atlantic and Pacific, and oceanic islands such as Bermuda, the Azores, and New Zealand. It is a common component of fouling communities and has likely been spread largely by shipping and aquaculture transfers. A recent genetic analysis indicates that 'B. schlosseri' is a complex of at least five cryptic species (A-E), but only one, clade A, had a widespread distribution in the Northeast and Northwest Atlantic, and Northeast Pacific. The other clades had restricted distributions in Europe (Bock et al. 2012). However, Yund et al. (2015) have identified at least one subclade which appears to be native to the Northwest Atlantic. This subclade is most genetically diverse in that region, and 9 of its 12 haplotypes are unique. The divergence between the prevailing Northwest Atlantic haplotype and those from European waters is too great to be accounted for by evolution (Yund et al. 2015). We cannot exclude the occurrence of European genotypes or cryptic species in East Coast waters, and so we will treat B. schlosseri as a single cryptogenic species. A recent analysis of Clade A (Nydam et al. 2017) suggests a Pacific origin for Clade A, with highest genetic diversity in specimens from the Northeast Pacific. However these populations are introduced (Carlton 1979). Northwest Pacific populations were not included in Nydam et al.'s (2017) study, and could be a possible source of Clade A.

North American Invasion History:

Invasion History on the West Coast:

Botryllus schlosseri was first collected on the West Coast at Mare Island Naval Yard, in San Francisco Bay, California in 1947 (Carlton 1979) and has since spread to the north and south through ship-fouling, and aquaculture transfers. It was reported in San Diego, CA in 1965 (Lambert and Lambert 1998), and reached Humboldt Bay, CA by the 1970s and Coos Bay, OR by 1978 (Boyd et al. 2002, Carlton 1989). In surveys conducted between 1994 and 2003 it was collected in 16 embayments between San Francisco and Mexico (Lambert and Lambert 1998; Wasson et al. 2001; Cohen et al. 2002; Fairey et al. 2002; deRivera et al. 2005). In 2013, it was also reported in Morro Bay, CA (Smithsonian surveys). Further south, it was reported from Ensenada, Mexico in 2000 (Lambert and Lambert 2003) and Bahia San Quintin, Mexico in 2005 (Rodriguez and Ibarra-Obando 2008). It has been found in several areas north of San Francisco Bay including Tomales Bay (in 2001, Fairey et al. 2002), Bodega Harbor (in 1997, Stoner et al. 2002), San Juan Islands, WA (in the late 1960s or early 1970s, Lambert et al. 1987 cited in Lambert and Lambert, 1998), Vancouver Island, British Columbia (prior to 1998; Lambert and Lambert, 1998) and at its current northern limit, Sitka, AK in 2000 (Ruiz et al., unpublished data).  Zwahleb et al. (2022), consider records from Puget Sound before the 1990s to be doubtful, but found this tunicate tp be increasingly abundant after 1999.  Puget Sound populations have a low genetic diversity, indicative of recnet establishment (Zwahleb et al. (2022),

Invasion History on the East Coast:

Recent genetic analysis suggests that Botryllus schlosseri includes genotypes native to the Northwest Atlantic (Yund et al. 2015), although the presence of genotypes or cryptic species from elsewhere in the world cannot be excluded at this time. Consequently, we will treat it as a cryptogenic species in the Gulf of Mexico and and most of the East Coast. It should be noted, though that it has apparently recently expanded its range into Atlantic Canada, most likely by human transport, so that we will treat it as introduced in those bioregions (NA-ET1, NA-S2, NA-S3, comprising the Atlantic coast of Newfoundland and Nova Scotia, and the Gulf of St. Lawrence).

While we consider B. schlosseri cryptogenic in the Gulf of Maine and southward, it apparently a recent invader on the Atlantic coast of the Maritime Provinces, and the Gulf of St. Lawrence. Many of these populations have been identified as having Northwest Atlantic genotypes (Yund et al. 2015), so they may have been introduced from harbors futher south along the East Coast. Botryllus schlosseri is now abundant on the Atlantic coast of Nova Scotia at Lunenburg, and other locations, and in the Bras d'Or Lakes estuary on Cape Breton Island (Locke et al. 200; Sephton et al. 2011). It was collected in the Gulf of St. Lawrence by Hooper (cited by Callahan et al. 2010), on the west coast of Newfoundland in 1975 , but its occurrence in the Gulf was not widely published until it was found on Prince Edward Island in the Gulf of St. Lawrence by 2001 (Locke et al. 2007). In 2011, it was found on floating docks in Conception Bay, on the outer Atlantic coast of Newfoundland, but was removed as a control measure (Robinson 12/8/2011).

Botryllus schlosseri was first recorded in North America in about 1838 simultaneously from both Boston, Massachusetts (at a meeting of January 17, 1838, and thus almost certainly based upon 1837 collections; Couthouy 1838), and New York City (in a publication dated January 1, 1839, and thus based upon material collected no later than 1838) (James T. Carlton, personal communication). It was later described as B. gouldii from New York circa 1870 (Verrill 1871). Later accounts gave wider ranges: Portland, Maine southward to New Jersey. The current northern range includes the Gulf of Maine from Massachusetts Bay to Cobscook Bay, Maine (Yund and Feldgarden 1992; Blezard 1999; Dijkstra et al. 2007; MIT Sea Grant 2009); Passamaquoddy Bay, New Brunswick (LeGresley et al. 2008); the Atlantic coast of Nova Scotia at Lunenburg; the Bras d'Or lakes estuary on Cape Breton Island (Locke et al. 2007); and Placentia Bay, Newfoundland (Callahan et al. 2010). It was collected in the Gulf of St. Lawrence by Hooper, on the west coast of Newfoundland (in 1975, cited by Callahan et al. 2010), but its occurrence in the Gulf was not widely publicized until it was found on Prince Edward Island in the Gulf of St. Lawrence by 2001 (Locke et al. 2007).

It is now found as far south as Florida, but records between New Jersey and Florida are spotty. It has been reported in Beaufort and other areas of North Carolina (Van Name 1945, Sutherland and Karlson 1977, USNM 15481, U.S. Museum of Natural History 2003) and Chesapeake Bay (Gosner 1978). Plough (1978) collected it off Sapelo Island, GA and it was reported by Mook (1983) from the Indian River Lagoon, FL, but was not found in recent late summer surveys (e.g. Ruiz et al. unpublished data), but this species may grow best in winter and spring at lower latitudes.

In Chesapeake Bay, B. schlosseri was not reported in the course of an extensive survey of Chesapeake benthos in 1915-1922 (Cowles 1930) or in shoreline surveys near Norfolk, VA by Ferguson et al. (1949). However, it was reported as fouling the dredge, “Chinook”, in Hampton Roads, VA in 1923 (Visscher 1928). It was listed as a ‘rare species’ in deeper waters of the lower Bay by Wass (1965). A colony was collected at Virginia Institute of Marine Science (Gloucester Point, VA) in 1962 (Calder 1972), and it was common to abundant on piers in Norfolk, VA in 1964-1965 (Calder and Brehmer 1967). 'For nearly twenty years, it was a rare inhabitant of deep waters near the mouth of Chesapeake Bay’, as indicated in the checklist by Wass (1963). During the drought years of the mid 1960’s, B. schlosseri suddenly appeared at Gloucester Point on oyster trays and eventually erupted to cover nearly all tufts of eelgrass (Zostera marina) and widgeongrass (Ruppia maritima) inhabiting shallow waters in the lower York River and Mobjack Bay, VA. In 1973 Andrews reported: ‘It is a fast-growing pernicious pest on trays in the cool months of spring and fall but barely survives hot summers in Virginia...B. schlosseri was not vigorous in the wet year of 1971, but it was still present on trays of oysters in the spring of 1972. After Agnes it disappeared and no trace has been found at any fouling stations. It may not recover its distribution of the 1960's until another series of droughts occurs'. Botryllus schlosseri was abundant on fouling panels in Lynnhaven Bay, VA in 1977 (Otsuka and Dauer 1980) and common on settling plates (1994-95) in all of the major lower Bay regions sampled except Norfolk, VA (15-18ppt), and not at all sites. Botryllus schlosseri is much more common on spring-early summer plates than summer-fall plates (Ruiz et al. unpublished data). The spread of B. schlosseri in the lower Bay region in the 1960’s suggests a recent introduction, either from Europe or from further north along the East Coast. However, this pattern could also represent population fluctuations in response to long-term salinity or temperature changes.

Very few records are known from the Atlantic Coast south of Cape Hatteras, NC. Aside from the Pearse et al. record from Beaufort, NC cited by Van Name (1945), it was reported from Beaufort in 1971 by Sutherland and Karlson (1977) and collected off North Carolina in 1981 (USNM 15481, U.S. Museum of Natural History 2003). Plough (1978) collected it off Sapelo Island, GA. It was reported by Mook (1983) from the Indian River Lagoon, FL, but was not found in later surveys (e.g. Ruiz et al. unpublished data). However, it was not found on SERC plates deployed in summer months (June-Sept) in Charleston, SC and Jacksonville, FL harbors, or the Indian River Lagoon, FL (Ruiz et al. unpublished data). As noted for the Gulf, this species may grow best in winter and spring at lower latitudes.

Invasion History on the Gulf Coast:

Botryllus schlosseri was collected in the Gulf of Mexico in 1887 off Cedar Key, FL (USNMNH 6993, US. National Museum of Natural History 2003; Van Name 1921; Van Name 1945), and was collected from the Tortugas northwest to St. Andrew Sound, FL. Most USNMNH collections were made between December and April, though two were made in June. This temperate species may be most abundant in winter months in the Gulf. Botryllus schlosseri was never found on SERC settling plates deployed in this region during the summer (Ruiz et al. unpublished data).

Invasion History Elsewhere in the World:

Botryllus schlosseri has been introduced to many remote parts of the world by shipping. It was reported as absent from Bermuda by Van Name (1921), but was listed as being present in Bermuda by Berrill (1932). It was first collected in the Azores Islands in 1971 (Morton and Britton 2000). In the Southern Hemisphere, it is known from Chile (in 1948, Valdivia et al. 2005; Ben-Shlomo et al. 2010), Argentina (in 1964, Orensanz et al. 2002; Ben-Shlomo et al. 2010), South Africa (in 1955, Monniot et al. 2001), Australia (in 1905, Kott 1985), and New Zealand (in 1922, Cranfield et al. 1998).


Description

Botryllus schlosseri is a colonial tunicate that can form colonies as large as 25 mm x 150 mm and up to 2 mm thick. Color is highly variable within and among colonies, and can be yellow, dark purple, red, brown or black. Colonies consist of many radial clusters of 5-20 zooids forming star-shaped systems. Individual zooids can be 1.75 mm - 5 mm long. There are 12-16 oral tentacles around the oral siphon and 7-9 stigmatal rows. The stomach has 8-10 folds and is somewhat spirally deflected with a prominent pyloric caecum. On each side of peribranchial cavity are 1-4 ovaries which are located above prominent testis. Tailed larvae, about 400 µm long, with eight ampullae, may be present in the cavity (Van Name 1945; Nishikawa 1991; Lambert and Lambert 1998).

A recent genetic analysis indicates 'Botryllus schlosseri' is probably a complex of cryptic species. Five clades (A-E) were identified in Europe, but only one of these (A) was found at sites in the Northwest Atlantic and Northeast Pacific. The other clades had restricted distributions on the coasts of Europe (Bock et al. 2012). However, Yund et al. (2015) have found that clade A is comprised of several subclades, at least one (BS2) of which is native to the Northwest Atlantic.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Chordata
Subphylum:   Tunicata
Class:   Ascidiacea
Order:   Stolidobranchia
Family:   Styelidae
Genus:   Botryllus
Species:   schlosseri

Synonyms

Alcyonium borlasii (Turton, 1814)
Alcyonium schlosseri (Pallas, 1766)
Aplidium verrucosum (Dalyell, 1839)
Botryllus aurolineatus (Giard, 1872)
Botryllus badius (Alder & Hancock, 1912)
Botryllus bivittatus (Milne-Edwards, 1841)
Botryllus calendula (Giard, 1872)
Botryllus calyculatus (Alder & Hancock, 1912)
Botryllus castaneus (Alder & Hancock, 1848)
Botryllus gemmeus (Savigny, 1816)
Botryllus gouldii (Verrill, 1871)
Botryllus marionis (Giard, 1872)
Botryllus miniatus (Alder & Hancock, 1912)
Botryllus minutus (Savigny, 1816)
Botryllus namei (Hartmeyer and Michaelsen, 1928)
Botryllus polycyclus (Savigny, 1816)
Botryllus pruinosus (Giard, 1872)
Botryllus rubens (Alder and Hancock, 1848)
Botryllus rubigo (Giard, 1872)
Botryllus smaragdus (Milne-Edwards, 1841)
Botryllus stellatus (Gaertner, 1774)
Botryllus violaceus (Milne-Edwards, 1841)
Botryllus violatinctus (Hartmeyer, 1909)
Botryllus virescens (Alder and Hancock, 1848)
Botryllus morio (Giard, 1873)

Potentially Misidentified Species

Botrylloides diegensis
possibly native to northeast Pacific

Botrylloides nigrum
None

Botrylloides violaceus
northwest Pacific species, widely introduced

Botryllus planus
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 Botryllus have small zooids, arranged in systems, and fully embedded in a mass of tunic material. Each zooid has an oral siphon and an atrial canal, an 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 the atrial cavity. Once mature, they 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). At Millport Scotland, B. schlosseri had only one generation per year, while at Plymouth, England, there were at least two (Millar 1954).

In all part of its native and introduced range, B. schlosseri 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).

Food:

Phytoplankton

Consumers:

Crabs, snails, urchins, starfish

Competitors:

Other fouling organisms

Trophic Status:

Suspension Feeder

SusFed

Habitats

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

Life History


Tolerances and Life History Parameters

Minimum Temperature (ºC)-1Field Data- Calder 1966; Ruiz et al. unpublished data; Whitlatch et al. 1995
Maximum Temperature (ºC)30Brunetti 1974; Brunetti et al. 1980; Brunetti et al. 1984. Temperature tolerances vary with acclimation and geographical location. For B. schlosseri, from Lynn Harbor MA, acclimated at 17 C, the median lethal 24 h temperature (LT50) was 29.4 C, but signficantly lower (28.3) for this species from Bodega Bay CA (Sorte et al. 2013).
Minimum Salinity (‰)14Experimental and Field data- Brunetti et al. 1980 (Venice Lagoon); Epelbaum et al. 2009; Ruiz et al. unpublished data; Experimental data, Dijkstra and Harris 2007 (New Hampshire)
Maximum Salinity (‰)44Experimental data- Brunetti et al. 1980
Minimum Duration0Larva, laboratory (Brunetti and Geghi 1982)
Maximum Duration1Larva, laboratory. Typical value, ~50% settlement is 0.04 days (Brunetti and Geghi 1982)
Broad Temperature RangeNoneCold temperate-Subtropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Economic Impacts

Shipping and Industry: The colonial ascidian Botryllus schlosseri is a common fouling organism in temperate harbors and on ships (Visscher 1927; Milkman 1967) and has been called 'a fast-growing pernicious pest' on oyster trays in lower Chesapeake Bay, but is absent or rare on natural oyster beds, presumably because of siltation (Andrews 1973). However, it is an important experimental organism in embryology, genetics, and immunology (Milkman 1967; Rinkevich et al. 1995).

Ecological Impacts

Competition: Bancroft (1903) listed hydroids, bryozoans, and 'worms' as competitors for space with B. schlosseri at Woods Hole, MA and Naples, Italy. Botryllus schlosseri is abundant and occasionally dominant in spring-early summer fouling communities in the lower Chesapeake Bay (Andrews 1973; Ruiz et al. unpublished data). When fouling plates in Lynnhaven Bay, VA were covered with wire mesh to exclude larger predators, B. schlosseri became the dominant fouling organism during December-March (Otsuka and Dauer 1982). In Long Island Sound, NY, recruitment of other fouling organisms, including native Spirorbis spp., was reduced in the vicinity of B. schlosseri colonies (Osman and Whitlatch 1995), and on colonial tunicates including B. schlosseri in Portsmouth Harbor, NH (Dijkstra et al. 2007). However, in Humboldt Bay (CA), fouling plate experiments (Nelson 2009) found that colonial tunicates (Botryllus schlosseri and Botrylloides violaceus), growing in sheets, were able to quickly occupy space on fouling plates, but did not decrease recruitment or species richness. In Langstone Harbor, Hampshire, England, Schmidt and Warner (1986) found that B. schlosseri paired with Trididemnum tenerum or Botrylloides leachii, on plexiglass panels resulted in stand-offs, colonies touching without overgrowth. However, B. schlosseri and the other species were always overgrown when paired with Diplosoma listerianum. In experiments in Wells Harbor, Maine, Botryllus schlosseri grew rapidly on some artificial substrates (rubber and metal), outcompeting native organisms, but grew more slowly on natural substrates (shell, marble, slate) (Tyrell and Byers 2007). In Bodega Harbor, California, Botryllus schlosseri was one of the eight most abundant fouling organisms both in 1969-1971 and in 2005-2009 (Sorte and Stachowicz 2011). Spawning periods and abundance of species in this group appeared to be favored by a 1?C increase in average temperatures at this site over a 30-year period (Sorte and Stachowicz 2011).

Botryllus schlosseri, Botrylloides violaceus, 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 (Wong and Vercaemer 2012). Negative effects on eelgrass are likely to be widespread.

Habitat Change:

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).


Regional Impacts

NA-ET3Cape Cod to Cape HatterasEconomic ImpactFisheries
Botryllus schlosseri was 'a fast-growing pernicious pest' on oyster trays in lower Chesapeake Bay, but is absent or rare on natural oyster beds, presumably because of siltation (Andrews 1973). Botryllus schlosseri was found fouling aqaculture gear at nine sites, and cultured Bay Scallops (Argopecten irradians) at one site, of 26 aquaculture sites surveyed on Marthas Vineyard (Carman et al. 2010). This tunicate was also reported at aquaculture sites in New York State (Carman et al. 2010).
M130Chesapeake BayEconomic ImpactFisheries
Botryllus schlosseri was 'a fast-growing pernicious pest' on oyster trays in lower Chesapeake Bay, but is absent or rare on natural oyster beds, presumably because of siltation (Andrews 1973).
NA-ET3Cape Cod to Cape HatterasEconomic ImpactShipping/Boating
Botryllus schlosseri (Golden Star Tunicate) was a common fouling organism in a survey of ship fouling among ships in northeastern US ports (Visscher 1927).
NA-ET3Cape Cod to Cape HatterasEcological ImpactCompetition
Bancroft (1903) listed hydroids, bryozoans, and 'worms' as competitors for space with B. schlosseri at Woods Hole and Naples. B. schlosseri is abundant and occasionally dominant in spring-early summer fouling communities in lower Chesapeake Bay (Andrews 1973; Ruiz et al. unpublished data). When fouling plates in Lynnhaven Bay were covered with wire mesh to exclude larger predators, B. schlosseri became dominant during December-March (Otsuka and Dauer 1982). Recruitment of other fouling organisms including native Spirorbis spp. was reduced in the vicinity of B. schlosseri colonies in Long Island Sound (Osman and Whitlatch 1995).
M010Buzzards BayEcological ImpactCompetition
Bancroft (1903) listed hydroids, bryozoans, and 'worms' as competitors for space with B. schlosseri at Woods Hole and Naples.
M130Chesapeake BayEcological ImpactCompetition
B. schlosseri is abundant and occasionally dominant in spring-early summer fouling communities in the lower Bay (Andrews 1973; Ruiz et al. unpublished data). When fouling plates in Lynnhaven Bay were covered with wire mesh to exclude larger predators, B. schlosseri became dominant during December-March (Otsuka and Dauer 1982).
M040Long Island SoundEcological ImpactCompetition
Recruitment of other fouling organisms including native Spirorbis spp. was reduced in the vicinity of B. schlosseri colonies in Long Island Sound (Osman and Whitlatch 1995). Recruitment of other introduced fouling organisms (Diplosoma listerianum; Botrylloides spp.) was reduced in the vicinity of B. schlosseri colonies in Long Island Sound (Osman and Whitlatch 1995).
NEA-IINoneEcological ImpactCompetition
In Langstone Harbor, Hampshire, England, Schmidt and Warner (1986) found that B. schlosseri paired with Trididemnum tenerurum or Botrylloides leachii, on Plexiglass panels resulted in stand-offs, colonies touching without overgrowth. However, B. schlosseri and the other species were always overgrown when paired with Diplosoma listerianum.
NA-ET2Bay of Fundy to Cape CodEcological ImpactCompetition
Recruitment of fouling organisms was reduced on colonial organisms incluiding B. schlosseri in Portsmouth Harbor, NH (Dijkstra et al. 2007). In experiments in Wells Harbor, Maine, Botryllus schlosseri 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).
N120Wells BayEcological ImpactCompetition
In experiments in Wells Harbor, Maine, Botryllus schlosseri 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).
NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactCompetition
Large monospecific colonies of Botyrllus schlosseri were noted at King Harbor and Port Hueneme, indicating strong competition (Lambert and Lambert 2003).
P060Santa Monica BayEcological ImpactCompetition
Large monospecific colonies of Botyrllus schlosseri were noted at King Harbor, indicating strong competition (Lambert and Lambert 2003).
P062_CDA_P062 (Calleguas)Ecological ImpactCompetition
Large monospecific colonies of Botyrllus schlosseri were noted at King Harbor and Port Hueneme, indicating strong competition (Lambert and Lambert 2003).
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 quickly occupy space on fouling plates, but did not decrease recruitment or species richness.
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 quickly occupy space on fouling plates, but did not decrease recruitment or species richness.
N195_CDA_N195 (Cape Cod)Economic ImpactFisheries
Botryllus schlosseri was found fouling aqaculture gear at nine sites, and cultured Bay Scallops (Argopecten irradians) at one site, of 26 aquaculture sites surveyed on Marthas Vineyard (Carman et al. 2010).
NA-ET2Bay of Fundy to Cape CodEconomic ImpactFisheries
Botryllus schlosseri was reportedly fouling aquaculture sites in Maine (Carman et al. 2010).
NA-S3NoneEconomic ImpactFisheries
Botrylloides violaceus was reportedly fouling mussel (Mytilus edulis a)quaculture sites in Prince Edward Island (Carman et al. 2010; Arens et al. 2011). Fouling by B. schlosseri did not affect cultured mussels through competion for phytoplankton, unlike Ciona intestinalis or Styela clava (Comeau et al. 2015). However, fouling by Botryllus schlosseri and Botrylloides violaceus had little effect on mussel growth and production (Arens et al. 2011). High-pressure water spraying reduced fouling of mussels. However, further studies showed that frequent spraying could affect mussel production, and could increase the spread of B. schlosseri through fragmentation (Paetzold et al. 2012).
P112_CDA_P112 (Bodega Bay)Ecological ImpactCompetition
Botryllus sclosseri was one of the eight most abundant species in Bodega Harbor both in 1969-1971 and 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).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
Botryllus sclosseri was one of the eight most abundant species in Bodega Harbor both in 1969-1971 and 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). Heavy recruitment of B. schlosseri can affect subsequent community development. When recruitment was artifically enhanced (at Santa Cruz), communities remained dominated by B. schlosseri for several months, compared to controls (Sams and Keough 2012).
WA-IVNoneEcological ImpactHabitat Change
Fouling of the eelgrass Zostera capensis in Langebaans Lagoon by B. schlosseri is a concern, (Griffiths et al. 2009), but the present extent of the problem is unclear.
N130Great BayEcological ImpactCompetition
Recruitment of fouling organisms was reduced on colonial organisms incluiding B. schlosseri in Portsmouth Harbor NH (Dijkstra et al. 2007).
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactCompetition
The native eelgrass Zostera marina was adversely affected by fouling by Botryllus schlosseri. However, fouling by a native sponge, Halichondria panicea, produced a greater reduction of chlorophyll than Botryllus schlosseri or any of the morphs of B. violaceus (Wong and Vercaemer 2012).
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactHabitat Change
The spread of introduced fouling organisms (B. schlosseri and B. violaceus) 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).
AUS-VIIINoneEcological ImpactCompetition
Heavy recruitment of B. schlosseri can affect subsequent community development. However, impacts varied by site and season (Sams and Keough 2012).
P080Monterey BayEcological ImpactCompetition
Heavy recruitment of B. schlosseri can affect subsequent community development. When recruitment was artifically enhanced (at Santa Cruz), communities remained dominated by B. schloserris for several months, compared to controls.(Sams and Keough 2012).
P080Monterey BayEcological ImpactHabitat Change
Fouling plates (at Santa Cruz) with enhanced B. schlosseri recruitment also had increased densites of Balanus crenatus and an unidentified sponge (Sams and Keoguh 2012).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactHabitat Change
i>B. schlosseri recruitment also had increased densites of Balanus crenatus and an unidentified sponge (Sams and Keoguh 2012).
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 2020).
OROregonEcological 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).
CACaliforniaEcological ImpactCompetition
Botryllus sclosseri was one of the eight most abundant species in Bodega Harbor both in 1969-1971 and 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). Heavy recruitment of B. schlosseri can affect subsequent community development. When recruitment was artifically enhanced (at Santa Cruz), communities remained dominated by B. schlosseri for several months, compared to controls (Sams and Keough 2012)., In fouling plate experiments in Humboldt Bay, Nelson (2009) found that colonial tunicates (Botryllus schlosseri and Botrylloides violaceus), growing in sheets, were able to quickly occupy space on fouling plates, but did not decrease recruitment or species richness., Large monospecific colonies of Botyrllus schlosseri were noted at King Harbor, indicating strong competition (Lambert and Lambert 2003)., Large monospecific colonies of Botyrllus schlosseri were noted at King Harbor and Port Hueneme, indicating strong competition (Lambert and Lambert 2003)., Heavy recruitment of B. schlosseri can affect subsequent community development. When recruitment was artifically enhanced (at Santa Cruz), communities remained dominated by B. schloserris for several months, compared to controls.(Sams and Keough 2012)., Botryllus sclosseri was one of the eight most abundant species in Bodega Harbor both in 1969-1971 and 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).
CACaliforniaEcological ImpactHabitat Change

i>B. schlosseri recruitment also had increased densities of Balanus crenatus and an unidentified sponge (Sams and Keough 2012)., Fouling plates (at Santa Cruz) with enhanced B. schlosseri recruitment also had increased densites of Balanus crenatus and an unidentified sponge (Sams and Keough 2012).

MAMassachusettsEcological ImpactCompetition
Bancroft (1903) listed hydroids, bryozoans, and 'worms' as competitors for space with B. schlosseri at Woods Hole and Naples.
MAMassachusettsEconomic ImpactFisheries
Botryllus schlosseri was found fouling aqaculture gear at nine sites, and cultured Bay Scallops (Argopecten irradians) at one site, of 26 aquaculture sites surveyed on Marthas Vineyard (Carman et al. 2010).
MEMaineEcological ImpactCompetition
In experiments in Wells Harbor, Maine, Botryllus schlosseri 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).
MEMaineEconomic ImpactFisheries
Botrylloides violaceus was reportedly fouling aquaculture sites on the Damariscotta River( Bullard et al. 2015)

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET2 Bay of Fundy to Cape Cod 1838 Crypogenic Established
NA-ET3 Cape Cod to Cape Hatteras 1871 Crypogenic Established
NEP-IV Puget Sound to Northern California 1972 Non-native Established
NEP-V Northern California to Mid Channel Islands 1947 Non-native Established
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1887 Crypogenic Established
NEP-VI Pt. Conception to Southern Baja California 1965 Non-native Established
NEP-III Alaskan panhandle to N. of Puget Sound 1998 Non-native Established
AUS-IV None 1985 Non-native Established
AUS-VIII None 1977 Non-native Established
AUS-X None 1995 Non-native Established
AUS-VII None 1955 Non-native Established
NZ-IV None 1922 Non-native Established
NZ-VI None 1944 Non-native Established
AR-V None 0 Crypogenic Established
NEA-II None 0 Crypogenic Established
NEA-III None 1776 Crypogenic Established
NEA-IV None 0 Crypogenic Established
MED-II None 0 Crypogenic Established
MED-III None 0 Crypogenic Established
MED-V None 0 Crypogenic Established
MED-IX None 0 Crypogenic Established
NWP-4a None 1941 Crypogenic Established
MED-VII None 0 Crypogenic Established
NWP-4b None 1951 Crypogenic Established
NWP-3a None 1991 Crypogenic Established
NWP-2 None 2001 Crypogenic Established
AUS-III None 1985 Non-native Established
AUS-VI None 1985 Non-native Established
AUS-XIII None 1985 Non-native Established
AUS-XII None 1985 Non-native Established
AUS-V None 1985 Non-native Established
MED-VI None 0 Crypogenic Established
NEA-VI None 1971 Non-native Established
CAR-VII Cape Hatteras to Mid-East Florida 1971 Crypogenic Established
MED-I None 0 Crypogenic Established
WA-IV None 2001 Non-native Established
WA-V None 1946 Non-native Established
NA-S3 None 1975 Non-native Established
NWP-3b None 1953 Crypogenic Established
SEP-C None 1948 Non-native Established
P270 Willapa Bay 2000 Non-native Established
M010 Buzzards Bay 1871 Crypogenic Established
M020 Narragansett Bay 1880 Crypogenic Established
M040 Long Island Sound 1874 Crypogenic Established
M080 New Jersey Inland Bays 1929 Crypogenic Established
M130 Chesapeake Bay 1923 Crypogenic Established
S190 Indian River 1981 Crypogenic Established
P050 San Pedro Bay 1994 Non-native Established
P170 Coos Bay 1978 Non-native Established
N130 Great Bay 1979 Crypogenic Established
P180 Umpqua River 1986 Non-native Established
P130 Humboldt Bay 1972 Non-native Established
M060 Hudson River/Raritan Bay 1871 Crypogenic Established
P020 San Diego Bay 1965 Non-native Established
NEA-V None 0 Crypogenic Established
AUS-XV None 1985 Non-native Established
AUS-IX None 1985 Non-native Established
P030 Mission Bay 1994 Non-native Established
P023 _CDA_P023 (San Louis Rey-Escondido) 1995 Non-native Established
P040 Newport Bay 1997 Non-native Established
P060 Santa Monica Bay 1994 Non-native Established
P062 _CDA_P062 (Calleguas) 1995 Non-native Established
P064 _CDA_P064 (Ventura) 1995 Non-native Established
P065 _CDA_P065 (Santa Barbara Channel) 1996 Non-native Established
P058 _CDA_P058 (San Pedro Channel Islands) 2001 Non-native Established
P080 Monterey Bay 1984 Non-native Established
P086 _CDA_P086 (San Francisco Coastal South) 1994 Non-native Established
P090 San Francisco Bay 1947 Non-native Established
P110 Tomales Bay 2000 Non-native Established
P112 _CDA_P112 (Bodega Bay) 1969 Non-native Established
P286 _CDA_P286 (Crescent-Hoko) 2003 Non-native Established
P290 Puget Sound 1998 Non-native Established
G100 Apalachicola Bay 1955 Crypogenic Established
G110 St. Andrew Bay 1995 Crypogenic Established
G080 Suwannee River 1887 Crypogenic Established
G060 Sarasota Bay 1952 Crypogenic Established
G050 Charlotte Harbor 1913 Crypogenic Established
G030 North Ten Thousand Islands 1982 Crypogenic Established
G020 South Ten Thousand Islands 1982 Crypogenic Established
S030 Bogue Sound 1971 Crypogenic Established
S056 _CDA_S056 (Northeast Cape Fear) 1981 Crypogenic Established
M120 Chincoteague Bay 1994 Non-native Established
M070 Barnegat Bay 1938 Crypogenic Established
M030 Gardiners Bay 2003 Crypogenic Established
M026 _CDA_M026 (Pawcatuck-Wood) 1873 Crypogenic Established
N195 _CDA_N195 (Cape Cod) 1871 Crypogenic Established
N180 Cape Cod Bay 2000 Crypogenic Established
N170 Massachusetts Bay 1838 Crypogenic Established
N140 Hampton Harbor 2003 Crypogenic Established
N100 Casco Bay 1945 Crypogenic Established
N010 Passamaquoddy Bay 2004 Crypogenic Established
NA-ET4 Bermuda 1932 Non-native Established
N190 Waquoit Bay 1871 Crypogenic Established
N185 _CDA_N185 (Cape Cod) 2003 Crypogenic Established
N070 Damariscotta River 1992 Crypogenic Established
N050 Penobscot Bay 2007 Crypogenic Established
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 2006 Non-native Established
P027 _CDA_P027 (Aliso-San Onofre) 1994 Non-native Established
P093 _CDA_P093 (San Pablo Bay) 1947 Non-native Established
G090 Apalachee Bay 1978 Crypogenic Established
S140 St. Catherines/Sapelo Sounds 1978 Crypogenic Established
MED-IV None 0 Crypogenic Established
N120 Wells Bay 2004 Crypogenic Established
CIO-II None 2008 Non-native Established
RS-3 None 2003 Non-native Established
SA-I None 1962 Non-native Established
SEP-A' None 2008 Non-native Established
N165 _CDA_N165 (Charles) 2009 Crypogenic Established
N060 Muscongus Bay 2009 Crypogenic Established
N040 Blue Hill Bay 2009 Crypogenic Established
NA-S2 None 2011 Non-native Unknown
NEP-VII None 2012 Non-native Established
WA-I None 2007 Non-native Established
M100 Delaware Inland Bays 2013 Crypogenic Established
P288 _CDA_P288 (Dungeness-Elwha) 2012 Non-native Established
N110 Saco Bay 0 Crypogenic Established
M050 Great South Bay 2013 Crypogenic Established
P070 Morro Bay 2013 Non-native Established
NWP-5 None 0 Crypogenic Established
B-II None 0 Crypogenic Established
B-I None 0 Crypogenic Established
AR-IV None 2011 Non-native Established
SA-III None 2002 Non-native Unknown
M023 _CDA_M023 (Narragansett) 2003 Crypogenic Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
767337 Ruiz et al., 2015 2012 2012-08-14 Spud Point North, Bodega Bay, California, USA Non-native 38.3301 -123.0572
767347 Ruiz et al., 2015 2012 2012-08-21 Lucas/Tides, Bodega Bay, California, USA Non-native 38.3284 -123.0445
767355 Ruiz et al., 2015 2012 2012-08-21 Porto Bodega, Bodega Bay, California, USA Non-native 38.3333 -123.0525
767367 Ruiz et al., 2015 2012 2012-08-22 Tomales-Marshall, Bodega Bay, California, USA Non-native 38.1514 -122.8888
767378 Ruiz et al., 2015 2012 2012-08-21 Tomales-Nick's Cove, Bodega Bay, California, USA Non-native 38.1980 -122.9222
767398 Ruiz et al., 2015 2012 2012-08-16 Tomales-SNPS, Bodega Bay, California, USA Non-native 38.1359 -122.8719
767410 Ruiz et al., 2015 2012 2012-08-17 Tomales- Shell Beach, Bodega Bay, California, USA Non-native 38.1163 -122.8713
767424 Ruiz et al., 2015 2013 2013-07-19 SeaWorld Marina, Mission Bay, CA, California, USA Non-native 32.7676 -117.2314
767442 Ruiz et al., 2015 2013 2013-07-23 Marina Village, Mission Bay, CA, California, USA Non-native 32.7605 -117.2364
767459 Ruiz et al., 2015 2013 2013-07-29 Mission Bay Yacht Club, Mission Bay, CA, California, USA Non-native 32.7778 -117.2485
767479 Ruiz et al., 2015 2013 2013-08-04 Bahia Resort Marina, Mission Bay, CA, California, USA Non-native 32.7731 -117.2478
767492 Ruiz et al., 2015 2013 2013-07-31 Campland on the Bay, Mission Bay, CA, California, USA Non-native 32.7936 -117.2234
767510 Ruiz et al., 2015 2013 2013-08-01 Hyatt Resort Marina, Mission Bay, CA, California, USA Non-native 32.7634 -117.2397
767525 Ruiz et al., 2015 2013 2013-08-03 Mission Bay Sport Center, Mission Bay, CA, California, USA Non-native 32.7857 -117.2495
767539 Ruiz et al., 2015 2013 2013-07-30 Hilton Resort Docks, Mission Bay, CA, California, USA Non-native 32.7791 -117.2128
767554 Ruiz et al., 2015 2013 2013-08-02 The Dana Marina, Mission Bay, CA, California, USA Non-native 32.7671 -117.2363
767565 Ruiz et al., 2015 2013 2013-08-05 Paradise Point Resort, Mission Bay, CA, California, USA Non-native 32.7730 -117.2406
767644 Ruiz et al., 2015 2013 2013-09-03 State Park Marina, Morro Bay, CA, California, USA Non-native 35.3459 -120.8423
767679 Ruiz et al., 2015 2013 2013-07-17 Naval Station San Diego, San Diego Bay, CA, California, USA Non-native 32.6867 -117.1333
767693 Ruiz et al., 2015 2013 2013-07-24 NAB ACU-1 Docks, San Diego Bay, CA, California, USA Non-native 32.6786 -117.1615
767717 Ruiz et al., 2015 2013 2013-07-21 Cabrillo Isle Marina, San Diego Bay, CA, California, USA Non-native 32.7272 -117.1995
767745 Ruiz et al., 2015 2013 2013-07-18 NAB Fiddlers Cove, San Diego Bay, CA, California, USA Non-native 32.6524 -117.1486
767762 Ruiz et al., 2015 2013 2013-07-26 Pier 32 Marina, San Diego Bay, CA, California, USA Non-native 32.6516 -117.1077
767772 Ruiz et al., 2015 2013 2013-07-20 Chula Vista Marina, San Diego Bay, CA, California, USA Non-native 32.6252 -117.1036
767785 Ruiz et al., 2015 2013 2013-07-28 Marriott Marquis and Marina, San Diego Bay, CA, California, USA Non-native 32.7059 -117.1655
767803 Ruiz et al., 2015 2011 2011-09-15 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9117 -122.3494
767814 Ruiz et al., 2015 2011 2012-09-20 San Leandro Marina, San Francisco Bay, CA, California, USA Non-native 37.6979 -122.1912
767824 Ruiz et al., 2015 2011 2011-09-20 San Francisco Marina, San Francisco Bay, CA, California, USA Non-native 37.8067 -122.4432
767836 Ruiz et al., 2015 2011 2011-09-14 Coyote Point Marina, San Francisco Bay, CA, California, USA Non-native 37.5880 -122.3160
767846 Ruiz et al., 2015 2011 2011-09-16 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9724 -122.4796
767857 Ruiz et al., 2015 2011 2011-09-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Non-native 37.6725 -122.3864
767871 Ruiz et al., 2015 2011 2011-09-13 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.8046 -122.3985
767881 Ruiz et al., 2015 2011 2012-09-15 Berkeley Marina, San Francisco Bay, CA, California, USA Non-native 37.8758 -122.3181
767891 Ruiz et al., 2015 2011 2012-09-19 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Non-native 37.8609 -122.4853
767908 Ruiz et al., 2015 2011 2011-09-21 South Beach Harbor, San Francisco Bay, CA, California, USA Non-native 37.7797 -122.3871
767921 Ruiz et al., 2015 2011 2011-09-20 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7947 -122.2822
767933 Ruiz et al., 2015 2011 2011-09-22 Ballena Isle Marina, San Francisco Bay, CA, California, USA Non-native 37.7676 -122.2869
767952 Ruiz et al., 2015 2011 2011-09-12 Paradise Cay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9156 -122.4769
767987 Ruiz et al., 2015 2012 2012-08-24 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9134 -122.3523
768007 Ruiz et al., 2015 2012 2012-08-23 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Non-native 37.8609 -122.4853
768022 Ruiz et al., 2015 2012 2012-08-28 San Francisco Marina, San Francisco Bay, CA, California, USA Non-native 37.8071 -122.4341
768041 Ruiz et al., 2015 2012 2012-08-27 Port of San Francisco Pier 31, San Francisco Bay, CA, California, USA Non-native 37.8078 -122.4060
768063 Ruiz et al., 2015 2012 2012-09-11 Ballena Isle Marina, San Francisco Bay, CA, California, USA Non-native 37.7676 -122.2869
768086 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Non-native 37.6633 -122.3817
768110 Ruiz et al., 2015 2012 2012-08-29 Coyote Point Marina, San Francisco Bay, CA, California, USA Non-native 37.5877 -122.3174
768132 Ruiz et al., 2015 2012 2012-09-04 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.5023 -122.2130
768155 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9736 -122.4802
768176 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Non-native 37.7987 -122.3228
768196 Ruiz et al., 2015 2012 2012-09-07 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7940 -122.2787
768236 Ruiz et al., 2015 2012 2012-09-13 San Leandro Marina, San Francisco Bay, CA, California, USA Non-native 37.6962 -122.1919
768252 Ruiz et al., 2015 2012 2012-09-12 Emeryville, San Francisco Bay, CA, California, USA Non-native 37.8396 -122.3133
768278 Ruiz et al., 2015 2013 2013-08-15 Ballena Isle Marina, San Francisco Bay, CA, California, USA Non-native 37.7656 -122.2858
768298 Ruiz et al., 2015 2013 2013-08-20 Coyote Point Marina, San Francisco Bay, CA, California, USA Non-native 37.5877 -122.3163
768317 Ruiz et al., 2015 2013 2013-08-22 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7926 -122.2746
768340 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9723 -122.4829
768358 Ruiz et al., 2015 2013 2013-08-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Non-native 37.6639 -122.3821
768382 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.5024 -122.2134
768402 Ruiz et al., 2015 2013 2013-08-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Non-native 37.9138 -122.3522
768419 Ruiz et al., 2015 2013 2013-08-12 San Francisco Marina, San Francisco Bay, CA, California, USA Non-native 37.8078 -122.4354
768450 Ruiz et al., 2015 2013 2013-08-16 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Non-native 37.8611 -122.4851

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