Invasion History

First Non-native North American Tidal Record: 1852
First Non-native West Coast Tidal Record: 1972
First Non-native East/Gulf Coast Tidal Record: 1852

General Invasion History:

Styela canopus (Rough Sea Squirt) is now widely distributed in temperate and tropical coastal waters of the world. It was described from the Red Sea in 1816 and was later reported from both sides of the North Atlantic, Ascension Island in the South Atlantic, the Northwest Pacific, the tropical Indo-Pacific, and temperate waters of Australia. (Kott 1998). In the Atlantic, it was first reported (as Cynthia partita) from Boston Harbor in 1852 (Stimpson 1852) and then from the English Channel (as Styela variabilis) in 1868. It was reported from the Mediterranean (as S. canopoides) in 1877 (Kott 1998). By the late 19th century, it was abundant and widespread on both sides of the Atlantic and was generally regarded as a native species. In the Northeast Pacific, where it is a recent introduction, S. canopus was first reported from San Diego Bay, California, in 1972. It has been collected north to Alamitos Bay but is rare outside San Diego Bay (Lambert and Lambert 1998). James Carlton considers S. canopus to be a native of the Indo-West Pacific (Carlton 2000 personal communication), introduced to the Atlantic (possibly a century or more before its description), a judgment which we have adopted here. Kott (1985) considers S. canopus to be the only species of Styela native to the Indo-West Pacific region.

North American Invasion History:

Invasion History on the West Coast:

On the west coast of North America, Styela canopus was first collected in San Diego, California in 1972. To the south, this tunicate was discovered in Mexico, near Mazatlan, in the Urias Estuary, in 1995 (Salgado-Barragan et al. 2004). In 1996-2000, it was collected in Mission Bay, Oceanside Harbor, Newport Bay, Alamitos Bay, Los Angeles-Long Beach Harbors, and Marina del Rey, off Santa Monica Bay (Lambert and Lambert 1998; Fairey et al. 2002; Lambert and Lambert 2003; Ruiz et al., unpublished data). At most sites, it was present at low to moderate levels of abundance (Lambert and Lambert 2003). In 2015, one specimen of S. canopus was identified from fouling plates at the Oakland Yacht Club, San Francisco Bay, while several nearby immature specimens were probably this species. This is the northernmost record of this species on the West Coast, but the establishment of this tunicate in the Bay is still unconfirmed (Tracy et al. 2017).

Invasion History on the East Coast:

In the northeast Atlantic, S. canopus ranges from the south coast of England and Normandy (Hayward and Ryland 1990; Breton et al. 1996) to the Cape Verde Islands, Sierra Leone and Senegal (Monniot and Monniot 1994) to Egypt and Lebanon (Bitar and Kouli-Bitar 2001). It was reported from the Channel Islands, off France, in 1868, as Cynthia variabilis (Hancock 1868, cited by Kott 1998), and from the Adriatic Sea in 1877 as Styela canopoides (Heller 1877, as cited by Kott 1998). On biogeographic grounds, based on its likely origin in the Indo-Pacific, S. canopus appears to be a very early introduction to European waters (Carlton, personal communication 2000). 
 
In the southwest Atlantic, S. canopus was first collected from an abandoned fishnet in Rio de Janeiro Bay, Brazil (Monniot 1969, cited by da Rocha and Kremer 2005). In Brazil, it appears to be most abundant in port areas (da Rocha and Kremer 2005).

Invasion History on the Gulf Coast:

In the northeast Atlantic, S. canopus ranges from the south coast of England and Normandy (Hayward and Ryland 1990; Breton et al. 1996) to the Cape Verde Islands, Sierra Leone and Senegal (Monniot and Monniot 1994) to Egypt and Lebanon (Bitar and Kouli-Bitar 2001). It was reported from the Channel Islands, off France, in 1868, as Cynthia variabilis (Hancock 1868, cited by Kott 1998), and from the Adriatic Sea in 1877 as Styela canopoides (Heller 1877, as cited by Kott 1998). On biogeographic grounds, based on its likely origin in the Indo-Pacific, S. canopus appears to be a very early introduction to European waters (Carlton, personal communication 2000).

In the southwest Atlantic, S. canopus was first collected from an abandoned fishnet in Rio de Janeiro Bay, Brazil (Monniot 1969, cited by da Rocha and Kremer 2005). In Brazil, it appears to be most abundant in port areas (da Rocha and Kremer 2005).

Invasion History in Hawaii:

It was first collected in the Hawaiian Islands on Oahu in 1929 and has also been found in Maui (Coles et al. 2004; Carlton and Eldredge 2009). Styela canopus?has a broad range in the west Pacific and Indian Oceans, from Japan to the Red Sea, South Africa, and Australia, and presumed to be native over most of that range (Nishikawa 1991; Kott 1985; Kott 1998; Monniot and Monniot 2001; Monniot 2002). However, it is introduced on some of the islands of the Central Pacific, including the Hawaiian Islands (Coles et al. 2002; Coles et al. 2004), American Samoa (Coles et al. 2003) and Guam (Lambert 2002; Lambert 2003).

Invasion History Elsewhere in the World:

In the northeast Atlantic, S. canopus ranges from the south coast of England and Normandy (Hayward and Ryland 1990; Breton et al. 1996) to the Cape Verde Islands, Sierra Leone and Senegal (Monniot and Monniot 1994) to Egypt and Lebanon (Bitar and Kouli-Bitar 2001). It was reported from the Channel Islands, off France, in 1868, as Cynthia variabilis (Hancock 1868, cited by Kott 1998), and from the Adriatic Sea in 1877 as Styela canopoides (Heller 1877, as cited by Kott 1998). On biogeographic grounds, based on its likely origin in the Indo-Pacific, S. canopus appears to be a very early introduction to European waters (Carlton, personal communication 2000). 
 
In the southwest Atlantic, S. canopus was first collected from an abandoned fishnet in Rio de Janeiro Bay, Brazil (Monniot 1969, cited by da Rocha and Kremer 2005). In Brazil, it appears to be most abundant in port areas (da Rocha and Kremer 2005).

Description

Styela canopus is a solitary tunicate. Its body shape varies from globular, to elongate, depending on whether it's growing as a single individual or in a dense cluster. When growing singly, it is attached by much of its ventral surface, with the oral siphon slightly below the anterior tip. When crowded, the body is attached by only a small part of the ventral surface, with the oral siphon at the anterior tip, and the atrial siphon slightly below. Both siphons are low and warty. The test is variable in thickness, tough and leathery with rough bumps and wrinkles. Worm tubes, bryozoans and algae frequently grow on the test. The color is usually grayish or yellowish posteriorly, becoming brown, purplish, or red anteriorly, with four dark stripes on the siphons. Internally, the pharynx has four narrow folds on each side, and there are two gonads on the left side and 2-5 on the right. The animal is up to 25-30 mm in length (Van Name 1945; Kott 1985; Lambert and Lambert 1998).

Taxonomy

Taxonomic Tree

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

Synonyms

Cynthia partita (Stimpson, 1852)
Cynthia stellifera (Verrill, 1871)
Halocynthia partita (Verrill, 1879)
Styela aggregata var. americana (Metcalf, 1888)
Styela barbaris (Kott, 1952)
Styela canopoides (Heller, 1877)
Styela marquesana (Michaelsen, 1918)
Styela orbicularis (Sluiter, 1904)
Styela partita (Verrill, 1901)
Styela rectangularis (Kott, 1952)
Styela variabilis (Hancock, 1868)
Tethyum canopus (Hartmeyer, 1915)
Tethyum partitum (Hartmeyer, 1912)
Cynthia canopus (Savigny, 1816)
Styela bicolor (Sluiter, 1887)
Styela pupa (Heller, 1878)
Styela stephensoni (Michaelsen, 1934)
Tethyum orbiculare (Sluiter, 1904)
Tethyum pupa (Heller, 1878)

Potentially Misidentified Species

Styela clava
Native to NW Pacific, widespread in higher latitude harbors

Styela panamensis

Native to Caribbean Panama (De Barros and da Rocha (2021). Records of S.canopus in Boca del Toros, Panama, and in Venezuela, may refer to this newly described native species (de Barros and da Rocha 2021).



Styela plicata
Possibly native to NW Pacific, widespread in warm-temperate and tropical waters.

Ecology

General:

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

Solitary ascidians are hermaphroditic, meaning that both eggs and sperm are released to the atrial chamber. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but many species have a partial block to self-fertilization. In internal fertilizers, including S. canopus, eggs are brooded and fertilized within the atrial chamber and then released into the water column upon hatching. Fertilized eggs hatch into a tadpole larva with a muscular tail, notochord, eyespots, and a set of adhesive papillae. The lecithotrophic (non-feeding, yolk-dependent) larva swims briefly before settlement. In S. canopus, the larvae settled after ~2 hours at 25 ?C (Ying et al 2003). Once settled, the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Barnes 1983).

Food:

Phytoplankton, detritus

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
General HabitatMangrovesNone
General HabitatUnstructured BottomNone
General HabitatCoral reefNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone

Life History

Tolerances and Life History Parameters

Maximum Temperature (ºC)30Mazatlan/Mexico/Urias estuary, Gulf of California (Salgado Barragan et al. 2004)
Minimum Salinity (‰)23Mazatlan//Mexico/Urias estuary, Gulf of California (Salgado Barragan et al. 2004)
Maximum Salinity (‰)43Mazatlan/Mexico/Urias estuary, Gulf of California (Salgado Barragan et al. 2004)
Maximum Length (mm)30Van Name 1945; Kott 1985; Lambert and Lambert 1998
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Styela canopus, commonly known as the Rough Sea Squirt, is a widespread member of the fouling community in the world's coastal waters, and has been reported from ships, buoys, piers, and docks (Woods Hole Oceanographic Institution 1952; Lambert and Lambert, 1998). However, its impacts are unknown.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET2 Bay of Fundy to Cape Cod 1852 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 1871 Def Estab
CAR-VII Cape Hatteras to Mid-East Florida 1872 Def Estab
NA-ET4 Bermuda 1901 Def Estab
SP-XXI None 1928 Def Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1879 Def Estab
NWP-3a None 0 Native Estab
AUS-III None 1952 Crypto Estab
AUS-IV None 1952 Crypto Estab
AUS-XIII None 0 Crypto Estab
AUS-XII None 0 Crypto Estab
AUS-XIX None 0 Crypto Estab
SP-IV None 0 Crypto Estab
NWP-4a None 0 Native Estab
EAS-III None 0 Native Estab
EAS-IV None 0 Native Estab
EAS-I None 0 Native Estab
EAS-II None 0 Native Estab
AG-1 None 0 Native Estab
AG-5 None 0 Native Estab
AG-3 None 0 Native Estab
AG-4 None 0 Native Estab
NEP-VI Pt. Conception to Southern Baja California 1972 Def Estab
NEA-II None 1868 Def Estab
MED-VII None 1877 Def Estab
NWP-2 None 0 Native Estab
MED-V None 0 Def Estab
CAR-II None 1984 Def Estab
CAR-III None 2002 Def Estab
CAR-IV None 1921 Def Estab
RS-1 None 1816 Native Estab
RS-2 None 1816 Native Estab
RS-3 None 0 Native Estab
EA-IV None 1918 Crypto Estab
AUS-II None 0 Crypto Estab
AUS-XIV None 0 Crypto Estab
AUS-XI None 0 Crypto Estab
AUS-I None 0 Crypto Estab
MED-III None 0 Def Estab
MED-IV None 0 Def Estab
SP-XII None 2001 Def Estab
MED-I None 0 Def Estab
SP-XIII None 1955 Crypto Estab
WA-V None 0 Crypto Estab
WA-IV None 0 Crypto Estab
SP-IX None 2002 Def Estab
P020 San Diego Bay 1972 Def Estab
G070 Tampa Bay 2002 Def Estab
G130 Pensacola Bay 1963 Def Estab
M020 Narragansett Bay 1880 Def Estab
M010 Buzzards Bay 1873 Def Estab
M040 Long Island Sound 1871 Def Estab
M130 Chesapeake Bay 2000 Def Estab
S190 Indian River 2005 Def Estab
S180 St. Johns River 2000 Def Estab
G310 Corpus Christi Bay 2002 Def Estab
G260 Galveston Bay 2000 Def Estab
P050 San Pedro Bay 1998 Def Estab
SA-III None 2005 Def Estab
SA-II None 1925 Def Estab
EA-III None 0 Crypto Estab
GAden Gulf of Aden 0 Native Estab
CIO-I None 0 Native Estab
P030 Mission Bay 1996 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 1996 Def Estab
P040 Newport Bay 1997 Def Estab
NEA-IV None 1868 Def Estab
N195 _CDA_N195 (Cape Cod) 1871 Def Estab
NWP-3b None 0 Native Estab
NEA-III None 1868 Def Estab
MED-VI None 0 Def Estab
S030 Bogue Sound 1872 Def Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 1885 Def Unk
S050 Cape Fear River 1885 Def Estab
G080 Suwannee River 1887 Def Estab
N100 Casco Bay 1873 Def Estab
S056 _CDA_S056 (Northeast Cape Fear) 1980 Def Estab
M030 Gardiners Bay 1983 Def Estab
S010 Albemarle Sound 1980 Def Estab
G074 _CDA_G074 (Crystal-Pithlachascotee) 1983 Def Estab
G020 South Ten Thousand Islands 1990 Def Estab
S200 Biscayne Bay 1946 Def Estab
S080 Charleston Harbor 2004 Def Estab
MED-II None 0 Def Estab
G330 Lower Laguna Madre 2003 Def Estab
N170 Massachusetts Bay 1852 Def Estab
N010 Passamaquoddy Bay 2004 Def Estab
P060 Santa Monica Bay 2001 Def Estab
NEP-VIII None 1995 Def Estab
N140 Hampton Harbor 2007 Def Estab
SEP-H None 2008 Def Estab
G030 North Ten Thousand Islands 1982 Def Estab
G045 _CDA_G045 (Big Cypress Swamp) 1982 Def Estab
N180 Cape Cod Bay 1879 Def Estab
WA-VI None 1906 Def Estab
SP-XVI None 1905 Def Estab
G100 Apalachicola Bay 1986 Def Estab
SA-IV None 2009 Def Estab
S206 _CDA_S206 (Vero Beach) 1887 Def Estab
N185 _CDA_N185 (Cape Cod) 1884 Def Estab
CIO-II None 2008 Native Estab
NEP-VII None 2012 Def Estab
WA-I None 1994 Def Estab
NEA-V None 0 Def Estab
PAN_PAC Panama Pacific Coast 2008 Def Estab
PAN_CAR Panama Caribbean Coast 2003 Def Estab
P090 San Francisco Bay 2015 Def Unk
NEP-V Northern California to Mid Channel Islands 2015 Def Unk
SEP-Z None 2016 Def Estab
S045 _CDA_S045 (New) 2018 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
7591 US National Museum of Natural History 2007 1885 1885-01-01 Sable Island, NW Of/ Def 43.9500 -59.9158
7592 Trott 2004 2004 2004-01-01 Cobscook Bay Def 44.9095 -67.0555
7593 US National Museum of Natural History 2007 1873 2007-01-01 Casco Bay Def 43.8023 -70.0564
7594 Stimpson 1852 1852 1852-01-01 Boston Harbor Def 42.3418 -70.9662
7595 Yale Peabody Museum 2009 1879 1879-01-01 Provincetown Harbor Def 42.0418 -70.1661
7596 US National Museum of Natural History 2007 1884 1884-01-01 Nantucket Shoals, South Of Def 39.9417 -69.7222
7597 Verrill and Smith 1873 1873 1873-01-01 Buzzards Bay Def 41.5667 -70.6000
7598 US National Museum of Natural History 2007 1881 1881-01-01 Nyes Neck, North Falmouth Def 41.6376 -70.6536
7599 Verrill and Smith 1873 1873 1873-01-01 Woods Hole Def 41.5168 -70.6828
7600 Verrill and Smith 1873 1873 1873-01-01 Edgartown Def 41.4001 -70.4870
7601 MIT Sea Grant 2003 None 9999-01-01 Tripps Marina, Westport Def 41.5126 -71.0889
7603 US National Museum of Natural History 2002 1880 1880-01-01 Newport Harbor Def 41.4826 -71.3256
7604 MIT Sea Grant 2003 2000 2000-01-01 Warwick Cove Marina Def 41.6840 -71.3912
7605 Ruiz et al., unpublished data None 9999-01-01 Barrington Yacht Club Def 41.7340 -71.2934
7606 Ruiz et al. unpublished data 2003 2003-01-01 Sakonnet River Railroad Bridge Def 41.5673 -71.2239
7607 Verrill 1871, cited by Kott 1998 1871 1871-01-01 New Haven Harbor Def 41.2734 -72.9176
7608 US National Museum of Natural History 2007 1874 1874-01-01 Noank Def 41.3279 -71.9906
7609 MIT Sea Grant 2003 2003 2003-01-01 Brewer Yacht Haven Marine Center, Stamford Def 41.0232 -73.5368
7610 National Museum of Natural History 2007 1983 1983-01-01 Great Peconic Bay Def 40.9407 -72.4831
7611 MIT Sea Grant 2003 2003 2003-01-01 East Creek Marina, Jamesport Def 40.9382 -72.5709
7612 Ruiz et al., unpublished data 2000 9999-01-01 Kiptopeke Def 37.1379 -75.9666
7613 Ruiz et al., unpublished data) 2000 2000-01-01 Silver Beach Def 37.4854 -75.9605
7614 Ruiz et al., unpublished data 2000 2000-01-01 Norfolk Naval Station Def 32.3000 -64.7833
7615 Van Name 1902 1901 1901-01-01 Bermuda Def 32.3000 -64.7833
7616 S National Museum of Natural History 2007 1980 1980-01-01 Off North Carolina Def 35.3417 -75.3600
7617 Coues and Yarrow 1878 1878 1878-01-01 Fort Macon Def 34.6957 -76.6885
7618 US National Museum of Natural History 2007 1885 1885-01-01 Cape Fear, SE Of Def 33.6333 -77.6000
7619 US National Museum of Natural History 2007 1980 1980-01-01 Off North Carolina Def 33.5433 -77.4233
7620 Ruiz et al. unpublished data 2004 2004-01-01 Charleston Harbor Def 32.7638 -79.8973
7621 uiz et al., unpublished data 2000 2000-01-01 Mayport Naval Station Def 30.3796 -81.4286
7622 Ruiz et al., unpublished data 2000 2000-01-01 Browns Creek Bridge, Jacksonville Def 30.4214 -81.5209
7623 da Rocha pers. comm. 2005 2005-01-01 Sebastian Def 27.8164 -80.4706
7624 da Rocha pers. comm. 2005 2005-01-01 Cape Marina, Cape Canaveral/ Def 28.4058 -80.6048
7625 da Rocha pers. comm. 2005 2005-01-01 Pelican Marina, Fort Pierce Def 27.4467 -80.3256
7626 Ruiz et al., unpublished data 2004 2004-01-01 Port of Miami Def 25.7745 -80.1709
7627 Ruiz et al., unpublished data 2004 2004-01-01 Matheson Hammock Marina Def 25.6786 -80.2606
7628 US National Museum of Natural History 2007 1990 1990-01-01 Cape Sable, West Of Def 25.2271 -81.1437
7629 US National Museum of Natural History 2007 1887 1887-01-01 Dry Tortugas Def 24.6652 -82.8554
7630 US National Museum of Natural History 2007 None 9999-01-01 Naples, SW of Def 26.0531 -82.1408
7631 US National Museum of Natural History 2009 1982 1982-01-01 Sanibel Island, SW Of Def 26.4404 -82.1137
7632 Ruiz et al., unpublished data) 2002 2002-01-01 Lands End, Apollo Beach, Tampa Def 27.7717 -82.4062
7633 Ruiz et al., unpublished data) 2002 2002-01-01 Port Manatee Def 27.6336 -82.5615
7634 Ruiz et al., unpublished data 2002 9999-01-01 St. Petersburg Municipal Marina Def 27.7609 -82.6298
7635 US National Museum of Natural History 2007 1983 1983-01-01 Crystal Bay Def 28.9233 -82.7137
7636 US National Museum of Natural History 2007 1887 1887-01-01 Cedar Key, Def 29.1486 -83.0185
7637 Dalby and Young 1992 1986 1986-01-01 Carrabelle Def 29.8533 -84.6643
7638 Cooley 1978 1963 1963-01-01 Santa Rosa Sound Def 30.3724 -86.9719
7639 Ruiz et al., unpublished data None 2002-01-01 Port of Pensacola Def 30.4044 -87.2114
7640 Ruiz et al., unpublished data 2002 2002-01-01 Port of Texas City Def 29.3669 -94.8166
7641 Ruiz et al., unpublished data None 9999-01-01 sland Moorings Marina, Port Aransas Def 27.8075 -97.0867
7642 Ruiz et al., unpublished data 2002 2002-01-01 Corpus Christi Naval Air Station Def 27.6941 -97.2840
7644 Lambert et al. 2005 2003 2003-01-01 Sea Ranch Marina, Port Isabel Def 26.0759 -97.1647
7645 US National Museum of Natural History 2007 1914 1914-01-01 Los Arroyos Def 22.3533 -84.3825
7646 Goodbody 2003 2003 2003-01-01 Kingston Harbor Def 17.9539 -76.8037
7647 US National Museum of Natural History 2007 1984 1984-01-01 Pelican Cays Def 16.6500 -88.2000
7648 Goodbody 2004) None 9999-01-01 Twin Cays Def 16.8167 -88.1000
7649 Van Name 1921 1921 1921-01-01 Guanica Harbor, Caribbean Sea Def 17.8605 -66.9163
7650 da Rocha et al. 2005 2005 2005-01-01 St. Thomas Def 18.3533 -64.9365
7652 Monniot and Monniot 1985 1985 1985-01-01 Guadeloupe Def 16.2500 -61.5833
7653 da Rocha et al. 2005 2005 2005-01-01 Martinique Def 14.6667 -61.0000
7654 Van Name 1945 1945 1945-01-01 Curacao Def 12.1833 -69.0000
7655 da Rocha et al. 2005 2003 2003-01-01 Bocas del Toro Def 9.3333 -82.2500
7656 da Rocha et al. 2005 2005 2005-01-01 Bonaire Def 12.1833 -68.2500
7657 da Rocha et al. 2005 2005 2005-01-01 Isla Margarita Def 10.9869 -63.9356
7658 Carman et al. 2011 2009 2009-01-01 Colon Def 9.3333 -79.9000
7659 Ruiz et al. unpublished data 2008 2008-01-01 Panama City Def 8.9833 -79.5167
7660 Tovar-Hernandez et al. 2012 2012 2012-01-01 Puerto Ángel, Def 15.6667 -96.4906
7661 Tovar-Hernandez et al. 2012 2012 2012-01-01 Puerto Vallarta, Def 20.6667 -105.2667
7662 Tovar-Hernandez et al. 2012 1995 1995-01-01 Mazatlan Def 23.2200 -106.4200
7663 Tovar-Hernandez et al. 2012 2012 2012-01-01 Isla Talchichitle, Navolato Def 24.9833 -108.0667
7664 Lambert and Lambert 1998 1972 1972-01-01 South San Diego Bay Def 32.6098 -117.1300
7666 Lambert and Lambert 2003 1997 1997-01-01 Bahia Point, San Diego Def 32.7756 -117.2467
7667 Lambert and Lambert 2003 1996 1996-01-01 Oceanside Harbor Def 33.2078 -117.3950
7669 Lambert and Lambert 2003 1998 1998-01-01 Watchorn Marina, Los Angeles Def 33.7200 -118.2770
7670 Lambert and Lambert 2003 2000 2000-01-01 Long Beach Marina Def 33.7528 -118.1112
7672 Carlton and Eldredge 2009 1928 1928-01-01 Pearl Harbor Def 21.3550 -157.9722
7673 Coles et al. 2002 2002 2002-01-01 Kuapa Pond, Kaneohe Bay Def 21.2897 -157.7047
7674 Coles et al. 2004 2003 2003-01-01 Kahului Harbor Def 20.8817 -156.4675
7675 Monniot and Monniot 1985 1905 905-01-01 Outer reef, Hao Def -18.0753 -140.9453
7676 Monniot and Monniot 1985 1984 1984-01-01 Port of Papeete Def -17.5350 -149.5697
7677 Coles et al. 2003 2002 2002-01-01 Pago Pago Harbor Def -14.2814 -170.6742
7678 Monniot and Monniot 2001 1995 1995-01-01 Koror, Goby Lake Crypto 7.3153 134.5019
7679 Lambert 2002; Lambert 2003 2002 2002-01-01 Apra Harbor Def 13.4497 144.6500
7680 Kott 1985 None 9999-01-01 New Caledonia Crypto -21.2500 165.3000
7681 Kott 1985 None 9999-01-01 Heron Island Crypto -23.4420 151.9140
7682 Kott 1985 None 9999-01-01 Lizard Island Crypto -14.6689 145.4567
7683 Kott 1998 None 9999-01-01 Torres Strait Crypto -10.0000 142.0000
7684 Kott 1985 None 9999-01-01 Broome Crypto -17.9619 122.2361
7685 Kott 1953, cited by Kott 1998 None 9999-01-01 Fremantle Crypto -32.0569 115.7439
7686 Monniot and Monniot 2001 None 9999-01-01 Davao, Mindinao Def 7.1042 125.6632
7687 US National Museum of Natural History 2007 1909 9999-01-01 Luzon Island Native 17.0000 122.5000
7688 Sluiter 1904, cited by Kott 1985 1904 1904-01-01 Java Sea Native -7.3233 116.8250
7689 Millar1975, cited by Kott 1985 None 9999-01-01 Gulf of Thailand Native 13.0000 101.0000
7690 Kott and Goodbody 1982, cited by Kott 1985 None 9999-01-01 Hong Kong Native 22.2783 114.1589
7691 Huang 2001 None 9999-01-01 Luoyuan Bay Native 26.4169 119.7160
7692 Nishikawa 1991 None 9999-01-01 Ariake Sea Native 32.9069 130.3722
7693 Rho and Lee 1991 None 9999-01-01 Komundo Native 34.0264 127.3125
7694 Rho et al. 2000 None 9999-01-01 Chindo Islands Native 34.4831 126.2619
7695 Nishikawa 1992 None 9999-01-01 Tottori Native 35.5000 134.2333
7696 Huang 2001 None 9999-01-01 Lianyungang, Native 34.6000 119.1667
7697 Huang 2001) None 9999-01-01 Quindao Native 36.0667 120.3833
7698 Huang 2001 None 9999-01-01 Yantai Native 37.4000 121.2667
7699 Huang 2001 None 9999-01-01 Penglai Native 37.8167 120.7333
7700 Rho and Lee 1991 None 9999-01-01 Anmado Native 35.5142 126.0186
7701 Rho and Lee 1991 None 9999-01-01 Toksan Native 36.7000 126.6680
7702 Nishikawa 1991 None 9999-01-01 Kagoshima Bay Native 31.4697 130.6356
7703 Nishikawa 1991) None 9999-01-01 Seto Inland Sea Native 34.1667 133.3333
7704 Tokioka 1953, cited by Nishikawa 1991 None 9999-01-01 Osaka Bay Native 34.5000 135.3000
7705 Tokioka 1953, cited by Nishikawa 1991) None 9999-01-01 Ise Bay Native 34.7500 136.7500
7706 Tokioka 1953, cited by Nishikawa 1991 None 9999-01-01 Sagami Bay Native 35.1167 139.3833
7707 Oka 1934, cited by Nishikawa 1991 None 9999-01-01 Tokyo Bay Native 35.4167 139.7833
7708 Tamilselvi, et al. 2011 None 9999-01-01 Tuticorin Native 8.8100 78.1400
7709 Ali et al. 2009) None 9999-01-01 Azhikkal Native 11.9000 75.3500
7710 Abdul and Sivakumar 2007 None 9999-01-01 Vizhinjam Native 8.3792 76.9914
7711 Millar1975, cited by Kott 1985 None 9999-01-01 Persian Gulf Native 26.0000 52.0000
7712 Savigny 1816, cited by Kott 1998 1816 9999-01-01 Red Sea Native 20.0000 38.0000
7713 Monniot et al. 2002 None 9999-01-01 Ibo Island Crypto -12.3500 40.6333
7714 Michaelsen 1918, cited by Kott 1998 1918 1918-01-01 Laurenco Marques (Maputo) Crypto -25.6333 32.5833
7715 Monniot et al. 2001; Mead et al. 2011b None 9999-01-01 Port Elizabeth Crypto -33.9581 25.6000
7716 Monniot et al. 2001 None 9999-01-01 Kommetjie Crypto -34.1403 18.3292
7717 da Rocha and Bonnet 2009 2009 2009-01-01 Arquipélago de Alcatrazes Def -24.1000 -45.7000
7718 da Rocha et al. 2005 1998 1998-09-28 Arvoredro Def -24.2856 -48.3736
7719 da Rocha et al. 2009 1999 1999-01-01 Santa Catarina State Def -26.7694 -48.7844
7720 Marins et al. 2010 None 9999-01-01 Angra dos Reis, Def -23.0067 -44.3181
7721 US National Museum of Natural History 2007 1925 1925-01-01 Rio de Janeiro Def -22.9083 -43.1964
7722 Rocha et al. 2012 1999 1999-07-08 Salvador Def -12.8000 -38.6333
7723 da Rocha and Kremer 2005 2005 2005-01-01 Pernambuco Def -8.0706 -35.2658
7724 da Rocha and Kremer 2005 2005 9999-01-01 Rio Grande de Norte Def -5.0000 -36.0000
7725 Oliveira Filho and Lotufo 2010 2009 2009-01-01 Pecem and Mucuripe Harbors Def -3.7183 -38.5428
7726 Rennie and Wiseman, 1906, cited by Van Name 1921 1906 1906-01-01 Cape Verde Islands Def 15.1111 -23.6167
7727 Monniot and Monniot 1994 1990 1990-01-01 Senegal Def 14.6928 -17.4467
7728 Canning-Clode et al., in prep 2006 2006-01-01 Madeira Def 32.6511 -16.9097
7729 Naranjo et al. 1996 1996 1996-01-01 Algeciras Bay Def 36.1275 -5.4539
7730 Clausade 1969 1969 1969-01-01 Mediterranean Sea Def 43.2964 5.3700
7731 Peres 1957 1957 1957-01-01 Balearic Islands Def 39.5000 3.0000
7732 Chimenz 1985 1985 1985-01-01 Ischia Def 40.7313 13.8957
7733 Ghannudi et al. 1978 1978 1978-01-01 Tripoli/ Def 32.9022 12.1858
7734 Heller 1877, cited by Kott 1998 1877 1877-01-01 Lesina Def 41.7667 15.4333
7735 Tursi et al. 1979 1979 1979-01-01 Bari Def 41.1253 16.8667
7736 Manoudis et al. 2005 2005 2005-01-01 Cape Fanari/ Def 40.9500 25.1333
7737 Antoniadou and Chintiroglou 2007 2007 2007-01-01 Chalkidiki Peninsula, Thessalonica Def 40.3333 23.5000
7738 Koukouras et al. 1995 1995 1995-01-01 Port Said Def 31.2600 32.2900
7739 Bitar and Kouli-Bitar 2001 2001 2001-01-01 Lebanon Def 33.8869 35.5131
7740 Hancock 1868, cited by Kott 1998 1868 1868-01-01 Guernsey Island Def 49.4500 -2.5500
7741 Alder and Hancock 1868, cited by Kott 1985 1868 1868-01-01 English Channel Def 50.8000 -1.0000
7742 Appeltans et al. 2012 1975 1975-01-01 N of Crab Rock, Sherkin Island Def 51.4570 -9.4420
7743 Breton et al. 1996 1996 1996-01-01 Le Havre Def 42.4900 0.1000
7744 Appletans et al. 2012 1983 1983-01-01 Carreg-yr-Afr (Bardsey, Lleyn Peninsula) Def 52.9272 4.6793
8104 Turon 1987 None 9999-01-01 Tossa de Mar Def 41.7206 2.9322
767685 Ruiz et al., 2015 2013 2013-07-17 Naval Station San Diego, San Diego Bay, CA, California, USA Def 32.6867 -117.1333
767735 Ruiz et al., 2015 2013 2013-07-22 Coronado Cays Marina, San Diego Bay, CA, California, USA Def 32.6257 -117.1309
767752 Ruiz et al., 2015 2013 2013-07-18 NAB Fiddlers Cove, San Diego Bay, CA, California, USA Def 32.6524 -117.1486
767777 Ruiz et al., 2015 2013 2013-07-20 Chula Vista Marina, San Diego Bay, CA, California, USA Def 32.6252 -117.1036
767792 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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