Invasion History

First Non-native North American Tidal Record: 1905
First Non-native West Coast Tidal Record: 1905
First Non-native East/Gulf Coast Tidal Record: 2005

General Invasion History:

Amathia verticillata is an erect, soft-bodied bryozoan, first described from the Mediterranean Sea (Naples, Italy) and now widespread in tropical, subtropical, and warm-temperate waters around the world (Winston 1977; Gordon and Mawatari 1992) We consider it cryptogenic in the tropical-subtropical Atlantic from North Carolina and Bermuda to Santos, Brazil, (Osburn 1914; Marcus 1937; Osburn 1940; Maturo 1958; Winston 1982, but see Farrapeira, 2011) and the coast of west Africa (Senegal and Gana, (Cook 1968), Pacific coast of Central America (Soule et al. 1980, cited by Carlton and Cohen 1995), Arabian Sea and Bay of Bengal waters of India (Robertson 1921, cited by Cohen and Carlton 199); Karande 1968, cited by Winston 1977; Rao and Ganapati 1978), and southern China, in the South China Sea (Huang 2001). 
 
The origin of this bryozoan is unknown, but it is a documented invader in California (1905, Cohen and Carlton 1995), Hawaii (before 1921, Carlton and Eldredge 2007), New Zealand (Gordon and Mawatari 1992), Australia (before 1975, Keough and Ross 1999), Japan (early 1900s, Asakura 1992); South Korea (Je et al. 1988), the Azores (2008, Amat and Tempera 2009), Madeira (Wirtz and Canning-Clode 2009), and the Canary Islands (2010, Canning-Clode et al. 2013; Minchin 2012), and Taiwan (Minchin et al. 2016). Its larval period is brief (Hyman 1959), so that fouling and ballast water (as fragments of adult colonies or attached to bits of flotsam) are likely vectors for its introduction. A recent review documents the probable introduction of this bryozoan to the Mediterranean Sea, where it was first discovered in the early 19th century but is largely restricted to harbors and marinas (Marchini et al. 2015). Galil and Gevli (2014) argued for a Caribbean origin of A. verticillata, because of its strong association with corals, rocks, and seagrasses there, and the presence of a specialized nudibranch predator there (Okenia zoobotryon).

North American Invasion History:

Invasion History on the West Coast:

Robertson (1905, cited by Cohen and Carlton 1995) observed dense growths of Amathia verticillata in San Diego Bay, where it is still abundant in eelgrass beds (Williams 2007). It is found in Mazatlan, Mexico (Alvarez-Leon and Banta 1984), and the Gulf of California (in 1958, Soule 1963). The pattern of its spread along the California coast is unknown, because most of the records in smaller harbors and bays are quite recent. It was found in Mission Bay, La Jolla, and Newport Bay by 1963 (Soule 1963), and Los Angeles Harbor 'in recent years' (Soule et al. 1980, cited by Cohen and Carlton 1995). In 1993, it appeared in Redwood Creek, South San Francisco Bay (Cohen and Carlton 1995), but has not been seen since (James T. Carlton, personal communication 2013). In a 2001 survey, it was found in small harbors in southern California, including Oceanside Harbor, Channel Islands Harbor, and Avalon Harbor, on Catalina Island (Fairey et al. 2002).

Invasion History on the East Coast:

Amathia verticillata was collected in the Dry Tortugas, Florida, as early as 1908 (Osburn 1914), although it was earlier collected in Bermuda in 1876 (Maturo and Schopf, 1968). It has been treated as native, from North Carolina to Venezuela, by most writers (Osburn 1914; Osburn 1940; Weiss 1948; Maturo 1957; Winston 1982; Pederson and Peterson 2002). However, Carlton and Ruckelshaus (1997) and Gossett et al. (2004) list it as an introduction to Florida and the Gulf of Mexico. We consider it cryptogenic here. A colony found in 2005 at the restoration shipyard of Mystic Seaport, Connecticut, on Long Island Sound (James T. Carlton, personal communication), was an introduction by boat fouling. It was collected in the Mystic River again in 2010 (James T. Carlton, personal communication). Establishment of this warm-water bryozoan in Long Island Sound is unlikely until further warming occurs. It was collected in the Mystic River again in 2010 (James T. Carlton, personal communication). Additional occurrences have been seen from 2014 to 2016, from Wachapreague and Kiptopeake, Virginia and from New Bedford, Massachusetts. Establishment of this warm-water bryozoan, north of Cape Hatteras, is unlikely until further warming occurs.

Invasion History on the Gulf Coast:

Invasion History in Hawaii:

Robertson (1921) reported having received specimens of A. verticillata from Honolulu Harbor (Robertson 1921, cited by Carlton and Eldredge 2009). It was present in Pearl Harbor by 1921, and in Kaneohe Bay by 1939 (Carlton and Eldredge, 2009), and is also known from Maui (Coles et al. 2004). Recent reports have A. verticillata rapidly becoming abundant at Palmyra Island in the Line Islands, about 2000 km south of Hawaii (Knapp et al. 2011). 

Invasion History Elsewhere in the World:

Amathia verticillata is a recent introduction in the Indian Ocean, the South China Sea, and Japan, appearing in the early 1900s, and first collected in Tokyo Bay in 1963 (Asakura 1992; Jebakumar et al. 2017). This bryozoan was newly recorded on the south coast of South Korea, where it became a pest fouling cultured pearl oysters (Je et al. 1988). However, Otani (2006) lists it as a cryptogenic species in Japan. It was first collected in Auckland Harbor, New Zealand about 1960 and has since been found in several harbors on the North Island (Gordon and Mawatari 1992).  Amathia verticillata was collected in New South Wales before 1975 (Russ and Wake 1975, cited by Keough and Ross 1999) and in South and Western Australia before 1982 (Bock 1982, cited by Keough and Ross 1999; Huisman et al. 2008). 
 
Amathia verticillata was first reported from Europe in Cadiz, Spain, in 1807, and described from Naples in 1822 Marchini et al. (2015) has described this species as 'pseudo-indigenous', probably introduced very early, but still largely confined to harbors and marinas, and preferring artificial substrates (Marchini et al. 2015). Early Mediterranean records, from west to east include Malaga, Spain, 1823, Algiers, Algeria 1848, Livorno, Italy 1848, Naples, Italy 1822 and Alexandria, Egypt 1828 (Marchini et al. 2015). Locally in the Mediterranean, A. verticillata rapidly colonize marinas, and provide habitat for other introduced species (Marchini et al. 2015). A. verticillata is a recent invader in the Azores, where it appeared in marinas in 2008 on three of the islands (Amat and Tempera 2009), and Madeira, where it was first collected in May of 2009 (Wirtz and Canning-Clode 2009). 
 
In the Eastern Pacific, Amathia verticillata was first collected in Los Frailes Bay in the Gulf of California in 1958,), Baja California and at Mazatlán, Sinaloa (Soule 1963). This bryozoan was collected further south in Oaxaca in 2017 and 2018 (Humara-Gil, et al. 2019). It was first reported from the Pacific coast of South America in 2014, in the Salinas Yacht Club, Ecuador (Priscilla Martinez, personal communication, February 2015, cited by McCann et al. 2015).


Description

Amathia verticillata is an erect, un-calcified bryozoan, with colonies made of irregularly thick transparent, branching or triple-quadruple branching stolons with zooids attached. Colonies can resemble ‘stringy, gelatinous noodles or sauerkraut up to 2 m long’ (Gordon and Mawatari 1992). The zooids are ovoid, about 380 X 580 µm in size, and occur in clusters on either side of the thick transparent stolon. The polypides have eight tentacles and large gizzards. The zooids brood whitish embryos (description from Winston 1982; Gordon and Mawatari 1992; Vieira et al. 2014). Colonies vary greatly in size, 400-700 mm in Brazil (Farrapeira et al. 2010) to 2200 mm (Amat et al. 2011). 

Recent molecular work has confirmed that Amathia verticillata is a single global species (Nascimento et al. 2021). A recent genetic analysis of the family Vesiculariidae supports a synonymy of the genera Amathia, Bowerbankia, and Zoobotryon (Waeschenbach et al. 2015). 

Nascimento KB, Migotto AE, Fehlauer-Ale KH (2021). Molecular data suggest the worldwide introduction of the bryozoan Amathia verticillata (Ctenostomata, Vesiculariidae).  Marine Biology 168:33, 9 pp.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Bryozoa
Class:   Gymnolaemata
Order:   Ctenostomata
Suborder:   Carnosa
Superfamily:   Vesicularioidea
Family:   Vesiculariidae
Genus:   Amathia
Species:   verticillata

Synonyms

Amathia goodei (Verrill, 1901)
Hydra verticillata (Della Chaije, 1828)
Zoobotryon pellucidus (Ehrenberg, 1839)
Amathia verticillata (Waeschenbach et al., 2015)

Potentially Misidentified Species

Ecology

General:

Life History- Amathia verticillata is an erect, uncalcified bryozoan, composed of many individual zooids. The zooids feed by extending the ciliated tentacles of the lophophore as a funnel, creating a current, and driving food particles into their mouths. The food is guided along the tentacles and through the pharynx by the cilia. Larger food particles can be moved or captured by flicking or contracting the tentacles. The zooids are hermaphroditic, and produce large yolky eggs, which hatch into lecithotrophic larvae, which are planktonic for short periods (less than 1 day). Larvae settle on a substrate and metamorphose into the first zooid of a colony, an ancestrula (Barnes 1983). 
 
Ecology- This bryozoan is found on wood, rock, vegetation (including eelgrass and mangroves), shells, and man-made structures, including vessel hulls (Winston 1982; Gordon and Mawatari 1992; Minchin 2012). In an investigation of chemical compounds affecting feeding on A. verticillata, a Brazilian population had alkaloids that deterred feeding, while Florida populations had variable resistance to feeding. Dos Santos et al. (2017) suggest that the success of this bryozoan is due to factors other than toxicity. 

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatGrass BedNone
General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatCanalsNone
General HabitatMangrovesNone
General HabitatVessel HullNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone

Life History


Tolerances and Life History Parameters

Minimum Salinity (‰)15Nair et al. 1992
Maximum Salinity (‰)40Field salinity (Shark Bay, Western Australia) (Wyatt et al. 2005)
Minimum Length (mm)100Colony length, Azores (Amat et al. 2009, on marina floats)
Maximum Height (mm)2,200Colony length, Azores (Amat et al. 2009, on marina floats)
Broad Temperature RangeNoneWarm temperate-Tropical
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Economic Impacts

Shipping/Boating- Amathia verticillata is known from ship fouling (Woods Hole Oceanographic Institution 1952) and is reported to cause fouling problems to boats and marinas in southern California (Johnson et al. 2006).

Fisheries- Amathia verticillata causes major problems by clogging shrimp-fishing gear in Galveston Bay, Texas (Gossett et al. 2004). It was reported as a pest species fouling cultured pearl oysters, Pinctada fuca, on the south coast of Korea (Je et al. 1988).

Ecological Impacts

Habitat Change- Amathia verticillata grows epiphytically on the leaves of eelgrass (Zostera marina), killing plants in San Diego Bay, which creates gaps in the seagrass canopy, leaving bare patches of sediment available for colonization by other algal species (Williams 2007).


Regional Impacts

NEP-VIPt. Conception to Southern Baja CaliforniaEconomic ImpactShipping/Boating
Amathia verticillata is reported to cause fouling problems to boats and marinas in southern California (Johnson et al. 2006).
NWP-3aNoneEconomic ImpactFisheries
Amathia verticillata fouls cultured pearl oysters (Pinctada fucaon the south coast of Korea (Je et al. 1988).
CAR-INorthern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern FloridaEconomic ImpactFisheries
Amathia verticillata causes major problems by clogging shrimp-fishing gear in Galveston Bay, Texas (Gossett et al. 2004).
NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactCompetition
Amathia verticillata grows epiphytically on the leaves of eelgrass (Zostera marina) in San Diego Bay, killing them (Williams 2007).
NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactHabitat Change
Amathia verticillata kills eelgrass plants in San Diego Bay, creating gaps in the seagrass canopy and bare patches of sediment, which are then colonized by adult plants (Williams 2007).
P020San Diego BayEcological ImpactCompetition
Amathia verticillata grows epiphytically on the leaves of eelgrass (Zostera marina) in San Diego Bay, killing them (Williams 2007).
P020San Diego BayEcological ImpactHabitat Change
Amathia verticillata kills eelgrass plants in San Diego Bay, creating gaps in the seagrass canopy and bare patches of sediment, which are then colonized by adult plants (Williams 2007).
CAR-INorthern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern FloridaEcological ImpactHabitat Change
Drifting mats of detached Amathia verticillata form an important habitat for a wide variety of invertebrate taxa on muddy bottoms of Biloxi Bay, Gulf of Mexico (Pederson and Peterson 2002).
G170West Mississippi SoundEcological ImpactHabitat Change
Drifting mats of detached Amathia verticillata form an important habitat for a wide variety of invertebrate taxa on muddy bottoms of Biloxi Bay, Gulf of Mexico (Pederson and Peterson 2002).
MED-VNoneEconomic ImpactIndustry
Amathia verticillata has clogged power plant intakes in Israel (Gordon 2018).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
CAR-VII Cape Hatteras to Mid-East Florida 1958 Crypto Estab
NA-ET4 Bermuda 0 Crypto Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1908 Crypto Estab
CAR-IV None 1940 Crypto Estab
MED-II None 1848 Def Estab
MED-III None 1822 Def Estab
MED-VII None 1898 Def Estab
CIO-I None 1968 Def Estab
CIO-II None 1905 Def Estab
NWP-3b None 0 Def Estab
SA-II None 1937 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1905 Def Estab
NEP-VII None 1958 Def Estab
NEP-VIII None 1958 Def Estab
SEP-H None 1914 Def Unk
NEP-V Northern California to Mid Channel Islands 1980 Def Estab
AUS-X None 1975 Def Estab
AUS-VII None 1977 Def Estab
NZ-IV None 1960 Def Estab
SP-XXI None 1921 Def Estab
CAR-III None 0 Crypto Estab
WA-I None 1973 Def Estab
WA-II None 1968 Crypto Estab
CAR-II None 0 Crypto Estab
MED-V None 1828 Def Estab
AUS-V None 0 Def Estab
AUS-III None 2002 Def Estab
S190 Indian River 0 Crypto Estab
G260 Galveston Bay 0 Crypto Estab
P050 San Pedro Bay 1980 Def Estab
P020 San Diego Bay 1905 Def Estab
S180 St. Johns River 0 Crypto Estab
P030 Mission Bay 1963 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 2001 Def Estab
P058 _CDA_P058 (San Pedro Channel Islands) 2001 Def Estab
P040 Newport Bay 1959 Def Estab
P062 _CDA_P062 (Calleguas) 2001 Def Estab
P090 San Francisco Bay 1993 Def Estab
P022 _CDA_P022 (San Diego) 1963 Def Estab
S200 Biscayne Bay 2004 Crypto Estab
G090 Apalachee Bay 0 Crypto Estab
NWP-2 None 0 Crypto Estab
CAR-V None 0 Crypto Estab
NA-ET3 Cape Cod to Cape Hatteras 2005 Def Unk
M040 Long Island Sound 2005 Def Unk
SP-XVI None 2007 Def Estab
NEA-VI None 2008 Def Estab
NWP-3a None 1986 Def Estab
AUS-II None 2008 Def Estab
SP-XII None 2010 Def Estab
SA-III None 2009 Def Estab
M020 Narragansett Bay 2011 Def Unk
G074 _CDA_G074 (Crystal-Pithlachascotee) 0 Crypto Estab
G310 Corpus Christi Bay 0 Crypto Estab
S140 St. Catherines/Sapelo Sounds 0 Crypto Estab
S206 _CDA_S206 (Vero Beach) 1908 Crypto Estab
NEA-V None 1807 Def Estab
AUS-XII None 2011 Def Estab
G170 West Mississippi Sound 0 Crypto Estab
S030 Bogue Sound 1957 Crypto Estab
M128 _CDA_M128 (Eastern Lower Delmarva) 2014 Def Unk
SEP-Z None 2015 Def Estab
MED-I None 1825 Def Estab
MED-IV None 1934 Def Estab
MED-VI None 1969 Def Estab
M010 Buzzards Bay 2016 Def Unk
PAN_PAC Panama Pacific Coast 1914 Crypto Estab
SA-I None 2013 Def Unk
AUS-VIII None 1975 Def Estab
EAS-VI None 2014 Def Estab
SA-IV None 0 Def Estab
WA-IV None 2016 Def Estab
SEP-I None 2014 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
8136 Soule 1963 1959 1959-01-01 Newport Harbor Def 33.6084 -117.9092
8138 Soule 1963 1963 1963-01-01 La Jolla Def 32.8512 -117.2731
8139 Soule 1963 1963 1963-01-01 Mission Bay Def 32.7792 -117.2342
8140 Robertson 1905, cited by Cohen and Carlton 1995 1905 1905-01-01 San Diego Bay Def 32.7184 -117.1945
8141 Soule 1963 1958 1958-01-01 Los Frailes Bay Def 23.3669 -109.4239
8142 Alvarez-Leon and Banta 1984 1958 1958-01-01 Bahia de los Olas Altas Def 23.2000 -106.4500
8143 US National Museum of Natural History 2007 1914 9999-01-01 Taboga Island Crypto 8.7833 -79.5500
8144 Robertson 1921, cited by Cohen and Carlton 1995 1921 9999-01-01 Honolulu Harbor Def 21.3074 -157.8689
8145 Carlton and Eldredge 2009 1939 1939-01-01 Kaneohe Bay Def 21.4628 -157.8103
8146 Coles et al. 2004 2003 2003-01-01 Kahului Harbor Def 20.8962 -156.4723
8147 Knapp et al. 2011 2007 2007-01-01 Palmyra Island Def 5.8811 -162.0725
8148 Cuffy 2011 2010 2010-01-01 Midway Atoll Def 28.0000 -177.0000
8151 Bogue Sound None 9999-01-01 Beaufort Crypto 34.7163 -76.6646
8152 Ruiz et al. unpublished data 2002 2002-01-01 Jacksonville Crypto 30.3369 -81.6614
8153 Winston 1982 1982 1982-01-01 Haulover Canal, Indian River Lagoon Crypto 28.7430 -80.7473
8154 Winston 1982 1982 9999-01-01 Sebastian Grass Flats, Indian River Lagoon Crypto 27.8603 -80.4473
8155 Winston 1982 1982 1982-01-01 None Crypto 27.3000 -80.2562
8156 Ruiz et al. unpublished data 2004 2004-01-01 Biscayne Bay Crypto 25.5658 -80.2164
8157 Joseph and Nichy 1955, cited by Winston 1977 1955 1955-01-01 Apalachee Bay Crypto 30.0372 -84.1708
8158 US National Museum of Natural History 2012, 1914 1914-05-23 Ensenada De Cajon, Cape San Antonio Crypto 21.9136 -84.8967
8159 Gossett et al. 2004) None 9999-01-01 Galveston Bay Crypto 29.5697 -94.9367
8160 US Museum of Natural History 2012) 1959 1959-01-01 Clearwater Crypto 27.9659 -82.8001
8162 US National Museum of Natural History 2012 None 9999-01-01 Corpus Christi Bay Crypto 27.7722 -97.2747
8163 Prezant et al. 2002 1992 1992-01-01 St. Catherines Island Crypto 31.6569 -81.1515
8164 US National Museum of Natural History 2012 1982 1982-12-15 off Florida Crypto 26.7669 -82.1011
8165 Creary 2003 None 9999-01-01 Kingston Harbour Crypto 17.9539 -76.8037
8166 Montoya-Cadavid et al. 2007 None 9999-01-01 Gulf of Morrosquilllo Crypto 9.5833 -75.6667
8167 Osburn 1940 1940 1940-01-01 Ensenada Crypto 17.9314 -66.9246
8168 Farrapeira 201 2006 2006-11-01 Port of Natal, Def -5.7503 -35.2042
8169 Kelmo et al. 2004 1995 1995-01-01 Praio do Forte, Bahia Def -12.5492 -37.9903
8170 Farrapeira et al. 2008, cited by Farrapeira 2010 2008 2008-01-01 Porto de Suape Def -8.3958 -34.9678
8171 Winston 1984; Winston 2004 1980 9999-01-01 None Crypto 16.8029 -88.0821
8172 Junqueira et al. 2004, cited by Farrapeira 2010 2004 9999-01-01 Sepetiba Def -22.9833 -43.7000
8173 Marcus 1937; Vieira et al. 2008 1937 1937-01-01 Guaruja Def -23.9936 -46.2564
8174 Marcus 1937 1937 1937-01-01 Santos Def -23.9370 -46.3251
8175 Cook 1968 1968 1968-01-01 Tema Crypto 5.6667 0.0000
8177 d'Hondt 1975 1973 1973-01-01 Goree Crypto 14.6669 -17.3983
8178 Canning-Clode et al., in prep.) 2010 2010-01-01 Canary Islands Def 28.1000 -15.4000
8179 Wirtz and Canning-Clode 2009 2009 2009-05-28 Quinta do Lorde harbor, Madeira Def 32.7417 -16.7111
8180 Amat and Tempera 2009 2008 2008-08-02 Marina of Horta, Faial Def 38.5796 -28.7164
8181 Amat and Tempera 2009 2008 2008-08-31 Lajo de Pico (lagoon) Def 38.4700 -28.4000
8182 Amat and Tempera 2009 2008 2008-09-29 Marina of Villa Franca do Campo, Sao Miguel Def 37.8097 -25.2142
8183 Carrada and Sacchi 1966, cited by Winston 1977 1828 1828-01-01 Bay of Naples Def 40.7353 14.2753
8184 Carrada and Sacchi 1966, cited by Winston 1977 1966 1966-01-01 Stagno of Tortoli, Sardinia Def 39.9433 9.6703
8185 Antit et al. 2011 2009 2009-01-01 Tunis Harbor Def 36.8000 10.1833
8186 Neviani 1937; Gautier 1958, cited by Winston 1977 1937 1937-01-01 Lagoon of Venice Def 45.4131 12.2972
8189 Ardizzone and Riggio, cited by Minchin 2012 1981 1981-01-01 Palermo Def 38.1167 13.3667
8190 Abdel Salam and Ramadan 2008 1939 1939-01-01 Alexandria Def 31.1980 29.9192
8191 Guerra-Garcia et al. 2011 2009 2009-01-01 Cadiz Def 36.5333 -6.2833
8195 Karande 1968, cited by Winston 1977 1968 1968-01-01 Mumbai Crypto 18.9750 72.8258
8196 Robertson 1921, cited by Cohen and Carlton 1995 1921 1921-01-01 Chennai (Madras) Crypto 13.0839 80.2700
8197 Rao and Ganapati 1978 1978 1978-01-01 Visakhapatnam Crypto 17.6884 80.2188
8198 Huang 2001 None 9999-01-01 Hainan Island Crypto 19.2000 109.7000
8199 Huang 2001) None 9999-01-01 Guangdong Province Crypto 23.0000 113.5000
8200 Je et al. 1988 1988 1988-01-01 Chungmu Def 34.8333 128.4167
8201 Asakura 1992 1925 9999-01-01 Tateyama Def 35.0000 139.8667
8202 Asakura 1992 1964 9999-01-01 Tokyo Bay Def 35.4167 139.7833
8203 Ohgaki 2007 None 9999-01-01 Tanabe Bay Def 33.7333 135.3833
8204 P. Colin, personal communication, 2010 9999-01-01 Guam Def 13.5000 144.8000
8205 Huisman et al. 2008 2005 2005-01-01 Port Hedland Def -20.3100 118.6001
8206 Wyatt et al. 2005 2002 2002-01-01 Denham Def -25.9269 113.5339
8207 Hewitt et al. unpublished 2005 2005-01-01 Esperance Def -33.8611 121.8919
8208 Brock 1983 1977 1977-01-01 Adelaide Def -34.9289 138.6011
8209 Russ and Wake 1975, cited by Keough and Ross 1999 1975 1975-01-01 New South Wales Def -33.8600 151.2111
8210 Tillbrook 2012 None 9999-01-01 Townsville Motor Boat and Yacht Club Def -19.2586 146.8225
8211 Gordon and Mawatari 1992 1960 1960-01-01 Waitemata Harbour Def -36.8299 174.7378
8212 Cranfield et al. 1998 1998 1998-01-01 Whangarei Harbour Def -35.8000 174.4100
8213 Gordon and Mawatari 1992 1992 1992-01-01 Manakau Def -37.0000 174.6667
767418 Ruiz et al., 2015 2013 2013-07-19 SeaWorld Marina, Mission Bay, CA, California, USA Def 32.7676 -117.2314
767451 Ruiz et al., 2015 2013 2013-07-29 Mission Bay Yacht Club, Mission Bay, CA, California, USA Def 32.7778 -117.2485
767471 Ruiz et al., 2015 2013 2013-08-04 Bahia Resort Marina, Mission Bay, CA, California, USA Def 32.7731 -117.2478
767489 Ruiz et al., 2015 2013 2013-07-31 Campland on the Bay, Mission Bay, CA, California, USA Def 32.7936 -117.2234
767501 Ruiz et al., 2015 2013 2013-08-01 Hyatt Resort Marina, Mission Bay, CA, California, USA Def 32.7634 -117.2397
767518 Ruiz et al., 2015 2013 2013-08-03 Mission Bay Sport Center, Mission Bay, CA, California, USA Def 32.7857 -117.2495
767534 Ruiz et al., 2015 2013 2013-07-30 Hilton Resort Docks, Mission Bay, CA, California, USA Def 32.7791 -117.2128
767547 Ruiz et al., 2015 2013 2013-08-02 The Dana Marina, Mission Bay, CA, California, USA Def 32.7671 -117.2363
767674 Ruiz et al., 2015 2013 2013-07-17 Naval Station San Diego, San Diego Bay, CA, California, USA Def 32.6867 -117.1333
767690 Ruiz et al., 2015 2013 2013-07-24 NAB ACU-1 Docks, San Diego Bay, CA, California, USA Def 32.6786 -117.1615
767714 Ruiz et al., 2015 2013 2013-07-21 Cabrillo Isle Marina, San Diego Bay, CA, California, USA Def 32.7272 -117.1995
767727 Ruiz et al., 2015 2013 2013-07-22 Coronado Cays Marina, San Diego Bay, CA, California, USA Def 32.6257 -117.1309
767740 Ruiz et al., 2015 2013 2013-07-18 NAB Fiddlers Cove, San Diego Bay, CA, California, USA Def 32.6524 -117.1486
767755 Ruiz et al., 2015 2013 2013-07-26 Pier 32 Marina, San Diego Bay, CA, California, USA Def 32.6516 -117.1077
767770 Ruiz et al., 2015 2013 2013-07-20 Chula Vista Marina, San Diego Bay, CA, California, USA Def 32.6252 -117.1036
767780 Ruiz et al., 2015 2013 2013-07-28 Marriott Marquis and Marina, San Diego Bay, CA, California, USA Def 32.7059 -117.1655

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Asakura, A. (1992) Recent introductions of marine benthos into Tokyo Bay (review): Process of invasion into an urban ecosystem with discussion on the factors inducing their successful introduction, Journal of the Natural History Museum and Institute, Chiba 2(1): 1-14

Barnes, Robert D. (1983) Invertebrate Zoology, Saunders, Philadelphia. Pp. 883

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2021 Invasive Algae Response (Aulerpa spp.). Poster, CDFW Invasive Algae Caulerpa Informational Flyer ... - CA.govhttps://nrmsecure.dfg.ca.gov › FileHandler

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