Invasion History

First Non-native North American Tidal Record: 1899
First Non-native West Coast Tidal Record: 1899
First Non-native East/Gulf Coast Tidal Record: 1975

General Invasion History:

Diplosoma listerianum was first described from the English Channel by Milne-Edwards in 1841. Initially, this species was described, under different names, from many different places around the world, such as Australia (D. rayneri Macdonald 1859) Brazil (D. macdonaldi Herdman 1886), and Japan (D. mitsukurii Oka 1892). These and many other names were synonymized by Kott (1990; 2001). This tunicate is now said to have a cosmopolitan distribution but represents a species complex (Perez-Portela et al. 2013). In most of these regions, D. listerianum was already established when researchers began surveying tunicates and, as a result, many populations have not been recognized as true introductions. One cryptic species (clade A) occurred in most of the sites sampled worldwide, while three other clades had more local distributions (Perez-Portela et al. 2013). We consider this species complex to be cryptogenic (of unknown origin) through most of its range. However, it appears to be a definite introduction in several locations, including the Northeast Pacific (California to British Columbia), the Northeast Atlantic (north of Cape Hatteras, North Carolina), Hawaii, New Zealand, and South Africa. In European waters, near the site of first description, range expansions have been noted in the North Sea (UK, Vance et al. 2008) and in the Netherlands (1st Record 1977, Gittenberger et al. 2007).

North American Invasion History:

Invasion History on the West Coast:

Diplosoma listerianum was collected in San Diego Bay, California in 1917. It was described by Ritter and Forsyth as D. pizoni from a single colony (Van Name 1945). Five additional colonies collected in 1899 were also identified as D. pizoni by Ritter (Eldredge 1966). Diplosoma listerianum apparently expanded its range northward, reaching Monterey Bay by 1939 (Van Name 1945; Eldredge 1966). In 1948, D. listerianum was found in San Francisco Bay, California (Eldredge 1966). In 1960, it was collected on Vancouver Island, British Columbia and by 1966 it was common in Puget Sound, Washington (Lambert 1966). 
 
Since D. listerianum was treated as a native species, we have not been able to find first dates of record for many West Coast water bodies. By 1980, it was considered common from San Diego, California to British Columbia, Canada (Abbott and Newberry 1980). In Southern California, Lambert and Lambert (1998) described it as occurring in 'all harbors, at all times of year' and subsequent surveys have confirmed this (Fairey et al. 2001; de Rivera et al. 2006; Ruiz et al., unpublished data). Further north, it occurs in Humboldt Bay, CA (Fairey et al. 2003; Ruiz et al. unpublished data) and Coos Bay, Oregon (1988, Hewitt 1993, 2001, Ruiz et al., unpublished data). It is widespread throughout Puget Sound (Ruiz et al., unpublished data) and has been collected at as far north as Bamfield, Vancouver Island, British Columbia (Mackie and Singla 1987). 

Invasion History on the East Coast:

Diplosoma listerianum was first collected in 1880 in the Atlantic Ocean off South Carolina, and subsequently found from Cape Fear, NC to Biscayne Bay, FL (U.S. National Museum of Natural History 2002; Van Name 1921; Ruiz et al. unpublished data). We consider it cryptogenic in these southern US waters. The first reported occurrence of D. listerianum north of Cape Hatteras, NC was in Long Island Sound, CT in the 1970s (R. Whitlatch to James T. Carlton, personal communication, 2002). In 1981, it was found at the east end of the Cape Cod Canal, in the Gulf of Maine (R. Whittaker to James T. Carlton, personal communication, 1984). In 1997, it was found in Great Bay, near Portsmouth, New Hampshire (Blezard 1999). In 1998, it was found in Narragansett Bay, Rhode Island and Vineyard Sound, Massachusetts (Whitlatch and Osman 2000). In 1999, it was found near Cape Neddick, Maine (ME) (Harris and Tyrell 2001), and in 2003 it was found in Casco Bay, ME (MIT Sea Grant 2003). It was found in the Gulf of St. Lawrence, in the Magdalene Islands, Quebec in 2008 and in Lunenburg Harbour, Nova Scotia in 2012; however establishment in both locations is uncertain (Willis et al. 2011; Moore et al. 2014).  There are no published records of this tunicate south of Long Island Sound, CT. However, in 2001 and 2002, specimens were found on fouling plates in lower Chesapeake Bay near Cape Charles, VA. Since there were concerns about identifications, specimens were confirmed as D. listerianum by Gretchen Lambert. Colonies in Cape Charles, Virginia (VA) appeared to be abundant (Ruiz et al. unpublished data).

Invasion History on the Gulf Coast:

Diplosoma listerianum was first found at Cedar Key, FL in 1885 (U.S. National Museum of Natural History 2002; Van Name 1921). It has been collected from Cape Sable, FL to Laguna Madre, Texas (U.S. National Museum of Natural History 2002; Van Name 1921, Lambert et al. 2005; Ruiz et al. unpublished data). We regard it as cryptogenic in the Atlantic Ocean south of Cape Hatteras, NC.

Invasion History in Hawaii:

Diplosoma listerianum was collected in Honolulu, Oahu as early as 1900. It has also been found on Maui and on remote islands, such as Midway Island and Johnston Atoll (Eldredge 1967, Coles et al. 1999; Coles et al. 2001; Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

Diplosoma listerianum has been reported worldwide from tropical to cold-temperate regions. Among locations where it is considered a recent invader are Guam (1998, Lambert 2002), New Zealand (1946, Cranfield et al. 1998), and South Africa (1st Record 1949, now found from Saldanha Bay to Durban, Monniot et al. 2001; Mead et al. 2011b). In 2018, one specimen was found in a harbor on the southwest coast of Iceland (Ramos-Espla et al. 2020). Genetic studies may result in changes to cryptogenic/introduced status of D. listerianum populations.


Description

Diplosoma listerianum is a colonial ascidian species forming extensive thin, delicate encrusting sheets, rarely more than 2 mm thick and up to 50 mm wide. It can overgrow seagrasses, pilings, crab shells, and other sessile organisms such as corals, gorgonians, and other tunicates. Its tunic is transparent and may have white or yellowish granular bodies suspended in the tunic material. Zooids (individual animals) underneath the tunic may be white, brown, green, or black due to pigment on the abdomen, and sometimes the thoracic epithelium. Other tissues are usually light-colored ranging from white to yellowish or rusty (Van Name 1945; Kott 2001). The colonies have a variable but extensive common cloacal cavity that is subdivided by thin connective tissue in which the zooids are embedded. Each zooid is enclosed by a branch or strip of tunic material (Van Name 1945; Kott 2001). Zooids are small, about 1 mm long. The oral siphon is conspicuous and divided into six deeply divided lobes around the opening. The atrial opening is large and oval, with no atrial tube or languet. This opening has plain edges and exposes most of the branchial sac to the cloacal cavity. There are four rows of stigmata, 10 in the anterior row, and eight in the posterior row. The stomach is rounded and smooth-walled, leading to an intestine of large diameter. Conspicuous stolonic vessels extend from the ventral side of the abdomen where the post-pyloric part of the gut is bent ventrally at right angles to the vertical axis of the zooid. Two oval testis follicles lie against the dorsal side of the gut loop. The vas deferens is hooked around between them, extending anteriorly around the ascending limb of the gut loop. The ovaries and eggs are found behind the intestinal loop (Van Name 1945; Kott 2001). The larvae have a trunk 0.4-6 mm long, with three fixatory papillae (Hayward and Ryland 1990).

A recent study has shown that 'Diplosoma listerianum' consists of at least 4 cryptic species. One of these, clade A, is extremely widespread, occurring in the northeast Atlantic (Plymouth, England; Atlantic and Mediterranean Spain); western Atlantic (Panama); southeast Atlantic-Indian Ocean (South Africa); western Pacific (Japan; Melbourne Australia), and eastern Pacific (Washington, California; Chile) (Perez-Portela et al. 2013). Other clades (B, C, and D) have a more limited distribution, but overlap with clade A. The genetic structure of local populations of clade A is suggestive of anthropogenic dispersal along historic shipping routes but does not indicate a region of origin. The distribution of the clades does not easily correspond with the many historic synonyms (eg., 'D. macdonaldi, D. mitsakurii, etc.). Additional genetic surveys will be needed to determine the identity of populations in many regions of the world.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Chordata
Subphylum:   Tunicata
Class:   Ascidiacea
Order:   Aplousobranchia
Family:   Didemnidae
Genus:   Diplosoma
Species:   listerianum

Synonyms

Diplosoma okai (Tokioka, 1949)
Diplosoma atropunctatum (Van Name, 1902)
Diplosoma carnosum (Drasche, 1884)
Diplosoma chamaelean (Drasche, 1884)
Diplosoma crystallinum (Drasche, 1884)
Diplosoma gelatinosum (Milne-Edwards, 1841)
Diplosoma lacteum (Van Name, 1902)
Diplosoma listeri (Lahille, 1890)
Diplosoma macdonaldi (Herdmann, 1886)
Diplosoma mitsakurii (Oka, 1892)
Diplosoma pizoni (Ritter and Forsyth, 1917)
Diplosoma rayneri (Macdonald, 1859)
Leptoclinum mitsakurii (Tokioka, 1953)
Leptoclinum gelatinosum (Alder and Hancock, 1912)
Leptoclinum listerianum (Milne-Edwards, 1841)
Leptoclinum macdonaldi (Hartmeyer, 1909)
Leptoclinum macrolobium (Tokioka, 1949)
Leptoclinum punctatum (Forbes, 1828)
Leptoclinum punctatum-listeri (Lahille, 1890)
Pseudodidemnum crystallinum (Giard, 1872)
Pseudodidemnum listerianum (Della Valle, 1877)
Pseudodidemnum zosterarium (Jourdain, 1885)
Didemnum gelatinosum (Milne-Edwards, 1841)
Leptoclinum rayneri (Kott, 1962)

Potentially Misidentified Species

Diplosoma spongiforme
Synonymized by Giard (1872, as Astellium spongifrome) and Lahille (1890), probable misidentifications

Ecology

General:

Life History- A colonial tunicate consists of many zooids, bearing most or all the organs of a solitary tunicate, but modified to varying degrees for colonial life. Colonial tunicates of the family Didemnidae have small zooids, completely embedded in an encrusting and thin tunic. Each zooid has an oral siphon and an atrial aperture which opens to a shared local chamber. Water is pumped into the oral siphon, through finely meshed ciliated gills on the pharynx, where phytoplankton and detritus are 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 that develop into larvae. Buds can form from the body wall of the zooids. Colonies vary in size ranging from small clusters of zooids to huge spreading masses. The zooids are hermaphroditic, which means both eggs and sperm are released into the atrial chamber. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but some species have a partial block to self-fertilization. Fertilized eggs are brooded within the tunic until they hatch into lecithotrophic (non-feeding, yolk-dependent) tadpole larvae. The larva has a muscular tail and a notochord, eyespots, and a set of adhesive papillae. The larvae are expelled upon hatching and swim briefly before settlement. Swimming periods are usually less than a day, but some larvae settle immediately after release or swim for longer periods if the water temperature is low. On settlement the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Van Name 1945; Barnes 1983). Populations of D. listerianum in Millport, Scotland had 1-2 generations per year, while those in Plymouth, England had up to 3 generations per year, and could produce larvae in 2-4 weeks after settlement (Millar 1954). 

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatGrass BedNone
General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatMangrovesNone
General HabitatVessel HullNone
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

Minimum Temperature (ºC)2.2Field, based on coldest site in geographical range, Boston MA (Zerebecki and Sorte 2011
Maximum Temperature (ºC)30Field, based on warmest site in geographical range, Miami FL (Zerebecki and Sorte 2011) Temperature tolerances vary with acclimation and geographical location. For D. listerianum, from Lynn Harbor MA, acclimated at 17 C, the median lethal 24 h temperature (LT50) was 29.41C, but signficantly lower (27.9) for this species from Bodega Bay CA (Sorte et al. 2013).
Minimum Salinity (‰)18Typical salinity, Black Sea. In Great Bay NH, D. listerianum occurred at a station where salinity dropped to 24 ppt in spring, but not further upstream (Blezard 1999). A population from Holyhead Harbour, Wales (Irish Sea) showed dramatic mortality when transferred from 34 to 27 and 20 PSU (Gröner et al. 2011).
Maximum Salinity (‰)40Field salinity (Shark Bay, Australia, Wyatt et al. 2005)
Minimum Reproductive Temperature15Field, larval release, Mediterranean (Brunetti et al. 1988)
Minimum Duration0Brunetti et al. 1988 (80% settlement within 24 hours in the light)
Maximum Duration3Brunetti et al. 1988 (only 40% settlement with 3 days in the dark)
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Economic Impacts

Fisheries: Diplosoma listerianum has been reported to foul cultured shellfish in the United Kingdom (Isle of Man), Croatia, Japan, and Hong Kong (Ross et al. 2004, Igic 1972, Arakawa 1990, Huang 2003, cited by da Rocha et al. 2009). Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impacts on mussel growth might be small, but this hypothesis still needs to be tested.

Shipping and Industry: In many parts of the world, the colonial tunicate Diplosoma listerianum is an abundant fouling organism (Lambert 2002), known from docks, buoys, floats, and ship hulls (Woods hole Oceanographic Institution 1951).

Ecological Impacts

Competition: In experiments in Long Island Sound, Diplosoma listerianum significantly reduced recruitment of native Spirorbis spp, Bugula spp, and Balanus spp, mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Diplosoma listerianum also significantly reduced recruitment of Botryllus schlosseri and non-indigenous Botrylloides violaceus, mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Disturbance enhanced the spread of D. listerianum (Altman and Whitlatch 2007). Diplosoma listerianum was a strong competitor in utilizing empty space on fouling plates (Stachowicz et al. 2002). In Eel Pond, Woods Hole, Massachusetts, Diplosoma listerianum outgrew Botrylloides violaceus and other organisms on fouling plates, covering ~72% of the plates at the end of the experiment. In cooler water, north of Cape Cod, in Lynn Harbor, Massachusetts Bay, D. listerianum achieved a stand-off with B. violaceus, becoming co-dominant, covering ~37% of the plates at the peak of its growth (Agius 2007). While working in the English Channel, Schmidt and Warner (1986) found that D. listerianum outcompeted Trididemnum tenerum, Botryllus schlosseri and Botrylloides leachii in 39% of trials on fouling plates and reached stand-offs in 61%.  Vance et al. (2008) observed that D. listerianum rapidly overgrew other fouling organisms on plates in newly colonized regions on the North Sea coast of England. Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010). 


Regional Impacts

NA-ET3Cape Cod to Cape HatterasEcological ImpactCompetition
In experiments in Long Island Sound, Diplosoma listerianum significantly reduced recruitment of native Spirorbis spp., Bugula spp., and Balanus spp., mostly through overgrowth of newly settled individuals (Osman and Whitlach 1995). Diplosoma listerianum also significantly reduced recruitment of non-indigenous Botryllus schlosseri and Botrylloides violaceus, mostly through overgrowth of newly settled individuals (Osman and Whitlach 1995). Disturbance enhanced the spread of D. listerianum (Altman and Whitlach 2007). Diplosoma listerianum was a strong competitor in filling up empty space on fouling plates (Stachowicz et al. 2002). In experiments in Eel Pond, Woods Hole, Diplosoma listerianum outgrew Botrylloides violaceus and other organisms on fouling plates, covering ~72% of the plates at the peak of its growth (Agius 2007). In years with abnormally warm winters, D. listerianum creates a community which it dominates, having lower biodiversity than native-dominated communities, especially in homogenously developed areas with extensive hard substrate (Munguia et al. 2011).
M040Long Island SoundEcological ImpactCompetition
In experiments in Long Island Sound, Diplosoma listerianum significantly reduced recruitment of native Spirorbis spp., Bugula spp., and Balanus spp., mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Diplosoma listerianum also significantly reduced recruitment of non-indigenous Botryllus schlosseri and Botrylloides violaceus, mostly through overgrowth of newly settled individuals (Osman and Whitlatch 1995). Diplosoma listerianum was a strong competitior in filling up empty space on foulng plates (Stachowicz et al. 2002). In years with abnormally warm winters, D. listerianum creates a community which it dominates, having lower biodiversity than native-dominated communities (Munguia et al. 2011).
NEA-IINoneEcological ImpactCompetition
Diplosoma listerianum has been reported to foul cultured shellfish in the United Kingdom (Isle of Man) (da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested.  Vance et al. (2008) observed that D. listerianum rapidly overgrew other fouling organisms on plates in newly colonized regions on the North sea coast of England. Schmidt and Warner (1986) found that D. listerianum outcompeted Trididemnurn tenerurnBotryllus schlosseri and Botrylloides leachii in 39% of trials on fouling plates and reached stand-offs in 61%.
NEA-IINoneEconomic ImpactFisheries
Diplosoma listerianum has been reported to foul cultured shellfish in the United Kingdom (Isle of Man) (da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested. 
MED-VIINoneEconomic ImpactFisheries
Diplosoma listerianum has been reported to foul cultured shellfish in Croatia (cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested. 
NWP-3bNoneEconomic ImpactFisheries
Diplosoma listerianum has been reported to foul cultured shellfish in Japan (Arakawa 1990, cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested. 
NWP-3bNoneEcological ImpactCompetition
Diplosoma listerianum has been reported to foul cultured shellfish in Japan (Arakawa 1990, cited by da Rocha et al. 2009). Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested. 
NWP-2NoneEcological ImpactCompetition
Diplosoma listerianum has been reported to foul cultured shellfish in Hong Kong (Huang 2003, cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested. 
NWP-2NoneEconomic ImpactFisheries
Diplosoma listerianum has been reported to foul cultured shellfish in Hong Kong (Huang 2003, cited by da Rocha et al. 2009).  Da Rocha et al. (2009) suggest that because of the thinness of D. listerianum colonies, its impact on mussel growth might be small, but this hypothesis still needs to be tested. 
M010Buzzards BayEcological ImpactCompetition
In experiments in Eel Pond, Woods Hole, Diplosoma listerianum outgrew Botrylloides violaceus and other organisms on fouling plates, covering ~72% of the plates at the peak of its growth (Agius 2007).
NA-ET2Bay of Fundy to Cape CodEcological ImpactCompetition
In experiments in Lynn Harbor, Massachusetts Bay, Diplosoma listerianum achieved a stand-off with Botrylloides violaceus, becoming co-dominant, covering ~37% of the plates at the peak of its growth (Agius 2007).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010).
P112_CDA_P112 (Bodega Bay)Ecological ImpactCompetition
Diplosoma listerianum was one of several invasive fouling species which showed increased growth (% coverage) at temperatures 3.5 and 4.5°C above the ambient temperature in Bodega Harbor (13.5°C), while the native Distaplia occidentalis showed reduced survival (Sorte et al. 2010).
N170Massachusetts BayEcological ImpactCompetition
In experiments in Lynn Harbor, Massachusetts Bay, Diplosoma listerianum achieved a stand-off with Botrylloides violaceus, becoming co-dominant, covering ~37% of the plates at the peak of its growth (Agius 2007).
N135_CDA_N135 (Piscataqua-Salmon Falls)Ecological ImpactFood/Prey
Increasing abundance of the introduced colonial tunicates Didemnum vexillum and Diplosoma listerianum has resulted in population growth of the native Bloodstar starfish Henricia sanguinolenta (Dijkstra et al. 2012).
NA-ET2Bay of Fundy to Cape CodEcological ImpactFood/Prey
Increasing abundance of the introduced colonial tunicates Didemnum vexillum and Diplosoma listerianum has resulted in population growth of the native Bloodstar starfish Henricia sanguinolenta (Dijkstra et al. 2012).
N135_CDA_N135 (Piscataqua-Salmon Falls)Ecological ImpactTrophic Cascade
Increased abundance of the Bloodstar starfish (Henricia sanguinolenta), supported by growing populations of Didemnum vexillum and Diplosoma listerianum, has resulted in increased predation and near-disapearance of the cryptogenic sponge Halichondria panicea (Dijkstra et al. 2012).
NA-ET2Bay of Fundy to Cape CodEcological ImpactTrophic Cascade
Increased abundance of the Bloodstar starfish (Henricia sanguinolenta), supported by growing populations of Didemnum vexillum and Diplosoma listerianum, has resulted in increased predation and near-disappearance of the cryptogenic sponge Halichondria panicea (Dijkstra et al. 2012).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEA-II None 1834 Crypto Estab
MED-II None 1872 Crypto Estab
MED-I None 0 Crypto Estab
MED-III None 0 Crypto Estab
MED-IV None 0 Crypto Estab
MED-VII None 1883 Crypto Estab
NA-ET3 Cape Cod to Cape Hatteras 1975 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 1981 Def Estab
CAR-VII Cape Hatteras to Mid-East Florida 1880 Crypto Estab
NA-ET4 Bermuda 1902 Crypto Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1885 Crypto Estab
CAR-IV None 1915 Crypto Estab
SP-XXI None 1900 Def Estab
CAR-II None 1884 Crypto Estab
SA-III None 1886 Crypto Estab
SA-II None 1886 Crypto Estab
NEA-III None 0 Crypto Estab
NEA-IV None 0 Crypto Estab
NEP-VI Pt. Conception to Southern Baja California 1899 Def Estab
NEP-V Northern California to Mid Channel Islands 1939 Def Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1960 Def Estab
NEP-IV Puget Sound to Northern California 1988 Def Estab
NWP-4a None 0 Crypto Estab
WA-V None 0 Def Estab
WA-I None 2005 Crypto Estab
AUS-II None 0 Crypto Estab
AUS-III None 0 Crypto Estab
AUS-V None 0 Crypto Estab
AUS-VII None 0 Crypto Estab
AUS-VIII None 1966 Crypto Estab
AUS-X None 1859 Crypto Estab
AUS-XII None 0 Crypto Estab
AUS-XIII None 0 Crypto Estab
SP-VII None 1981 Crypto Estab
NZ-IV None 1944 Def Estab
NZ-VI None 1946 Def Estab
EAS-II None 1898 Crypto Estab
EAS-VII None 1898 Crypto Estab
EAS-VIII None 0 Crypto Estab
EAS-III None 1908 Crypto Estab
NWP-3b None 1893 Crypto Estab
AG-3 None 0 Crypto Estab
AG-4 None 0 Crypto Estab
NWP-3a None 0 Crypto Estab
SP-XII None 1998 Def Estab
CAR-III None 1921 Crypto Estab
EA-III None 1964 Crypto Estab
RS-2 None 1965 Crypto Estab
SEP-B None 1959 Crypto Estab
CAR-VI None 1970 Crypto Estab
NEA-V None 0 Crypto Estab
CIO-I None 1996 Crypto Estab
WA-IV None 1949 Def Estab
SEP-C None 2002 Crypto Estab
S190 Indian River 1977 Crypto Estab
G260 Galveston Bay 2002 Crypto Estab
P020 San Diego Bay 1899 Def Estab
N130 Great Bay 1996 Def Estab
G070 Tampa Bay 2001 Crypto Estab
M010 Buzzards Bay 2000 Def Estab
M040 Long Island Sound 1989 Def Estab
P170 Coos Bay 1988 Def Estab
M130 Chesapeake Bay 2001 Def Estab
M020 Narragansett Bay 1998 Def Estab
G130 Pensacola Bay 2002 Crypto Estab
G310 Corpus Christi Bay 1977 Crypto Estab
P090 San Francisco Bay 1948 Def Estab
S200 Biscayne Bay 2004 Crypto Estab
S080 Charleston Harbor 1880 Crypto Estab
N100 Casco Bay 2003 Def Estab
SEP-H None 2005 Crypto Estab
SP-XIII None 1954 Def Estab
G020 South Ten Thousand Islands 1983 Crypto Estab
G050 Charlotte Harbor 1885 Crypto Estab
G074 _CDA_G074 (Crystal-Pithlachascotee) 1885 Crypto Estab
G108 _CDA_G108 (St. Andrew-St. Joseph Bays) 0 Crypto Estab
G080 Suwannee River 1885 Crypto Estab
G330 Lower Laguna Madre 2004 Crypto Estab
P030 Mission Bay 1994 Def Estab
N170 Massachusetts Bay 2000 Def Estab
N180 Cape Cod Bay 1981 Def Estab
N185 _CDA_N185 (Cape Cod) 2003 Def Estab
N195 _CDA_N195 (Cape Cod) 2003 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 1994 Def Estab
P040 Newport Bay 1936 Def Estab
P027 _CDA_P027 (Aliso-San Onofre) 1994 Def Estab
P290 Puget Sound 1966 Def Estab
S050 Cape Fear River 1886 Crypto Estab
P130 Humboldt Bay 2001 Def Estab
P050 San Pedro Bay 1960 Def Estab
P058 _CDA_P058 (San Pedro Channel Islands) 1978 Def Estab
P060 Santa Monica Bay 1994 Def Estab
P062 _CDA_P062 (Calleguas) 1994 Def Estab
P064 _CDA_P064 (Ventura) 1994 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 1977 Def Estab
P286 _CDA_P286 (Crescent-Hoko) 2003 Def Estab
P110 Tomales Bay 2001 Def Estab
P112 _CDA_P112 (Bodega Bay) 1969 Def Estab
P080 Monterey Bay 1939 Def Estab
S090 Stono/North Edisto Rivers 0 Crypto Estab
CIO-II None 0 Crypto Estab
AR-V None 2001 Crypto Estab
SEP-Z None 1999 Def Estab
N135 _CDA_N135 (Piscataqua-Salmon Falls) 1993 Def Estab
NWP-2 None 0 Crypto Estab
SP-XVI None 1962 Def Estab
P076 _CDA_P076 (Carmel) 1947 Def Estab
RS-3 None 1962 Crypto Estab
SA-IV None 2009 Crypto Estab
MED-V None 1976 Crypto Estab
NA-S3 None 2008 Def Estab
N165 _CDA_N165 (Charles) 2006 Def Estab
MED-IX None 0 Crypto Estab
M023 _CDA_M023 (Narragansett) 2010 Def Estab
NEA-VI None 1971 Def Estab
P292 _CDA_P292 (San Juan Islands) 2013 Def Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 2012 Def Unk
N120 Wells Bay 2013 Def Estab
SA-I None 2004 Def Estab
M050 Great South Bay 2013 Def Estab
NEP-VIII None 2007 Crypto Estab
PAN_PAC Panama Pacific Coast 2005 Crypto Estab
PAN_CAR Panama Caribbean Coast 1921 Crypto Estab
P070 Morro Bay 2001 Def Unk
RS-1 None 0 Crypto Estab
B-II None 0 Native Estab
B-I None 0 Native Estab
AUS-IV None 1947 Crypto Estab
AUS-VI None 0 Crypto Estab
WA-VI None 2012 Crypto Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 2013 Def Estab
AR-IV None 2018 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
767325 Ruiz et al., 2015 2012 2012-08-13 Coast Guard, Bodega Bay, California, USA Def 38.3126 -123.0512
767331 Ruiz et al., 2015 2012 2012-08-14 Spud Point South, Bodega Bay, California, USA Def 38.3281 -123.0574
767338 Ruiz et al., 2015 2012 2012-08-14 Spud Point North, Bodega Bay, California, USA Def 38.3301 -123.0572
767356 Ruiz et al., 2015 2012 2012-08-21 Porto Bodega, Bodega Bay, California, USA Def 38.3333 -123.0525
767370 Ruiz et al., 2015 2012 2012-08-22 Tomales-Marshall, Bodega Bay, California, USA Def 38.1514 -122.8888
767381 Ruiz et al., 2015 2012 2012-08-21 Tomales-Nick's Cove, Bodega Bay, California, USA Def 38.1980 -122.9222
767401 Ruiz et al., 2015 2012 2012-08-16 Tomales-SNPS, Bodega Bay, California, USA Def 38.1359 -122.8719
767426 Ruiz et al., 2015 2013 2013-07-19 SeaWorld Marina, Mission Bay, CA, California, USA Def 32.7676 -117.2314
767445 Ruiz et al., 2015 2013 2013-07-23 Marina Village, Mission Bay, CA, California, USA Def 32.7605 -117.2364
767462 Ruiz et al., 2015 2013 2013-07-29 Mission Bay Yacht Club, Mission Bay, CA, California, USA Def 32.7778 -117.2485
767481 Ruiz et al., 2015 2013 2013-08-04 Bahia Resort Marina, Mission Bay, CA, California, USA Def 32.7731 -117.2478
767495 Ruiz et al., 2015 2013 2013-07-31 Campland on the Bay, Mission Bay, CA, California, USA Def 32.7936 -117.2234
767513 Ruiz et al., 2015 2013 2013-08-01 Hyatt Resort Marina, Mission Bay, CA, California, USA Def 32.7634 -117.2397
767528 Ruiz et al., 2015 2013 2013-08-03 Mission Bay Sport Center, Mission Bay, CA, California, USA Def 32.7857 -117.2495
767541 Ruiz et al., 2015 2013 2013-07-30 Hilton Resort Docks, Mission Bay, CA, California, USA Def 32.7791 -117.2128
767557 Ruiz et al., 2015 2013 2013-08-02 The Dana Marina, Mission Bay, CA, California, USA Def 32.7671 -117.2363
767568 Ruiz et al., 2015 2013 2013-08-05 Paradise Point Resort, Mission Bay, CA, California, USA Def 32.7730 -117.2406
767582 Ruiz et al., 2015 2013 2013-08-30 201 Main, Morro Bay, CA, California, USA Def 35.3564 -120.8474
767593 Ruiz et al., 2015 2013 2013-08-27 City Harbor, Morro Bay, CA, California, USA Def 35.3709 -120.8582
767615 Ruiz et al., 2015 2013 2013-08-29 Moorings, Morro Bay, CA, California, USA Def 35.3619 -120.8548
767646 Ruiz et al., 2015 2013 2013-09-03 State Park Marina, Morro Bay, CA, California, USA Def 35.3459 -120.8423
767670 Ruiz et al., 2015 2013 2013-07-16 Naval Base Point Loma, San Diego Bay, CA, California, USA Def 32.6886 -117.2343
767683 Ruiz et al., 2015 2013 2013-07-17 Naval Station San Diego, San Diego Bay, CA, California, USA Def 32.6867 -117.1333
767696 Ruiz et al., 2015 2013 2013-07-24 NAB ACU-1 Docks, San Diego Bay, CA, California, USA Def 32.6786 -117.1615
767709 Ruiz et al., 2015 2013 2013-07-25 Navy Ammo Dock, Pier Bravo, San Diego Bay, CA, California, USA Def 32.6939 -117.2276
767720 Ruiz et al., 2015 2013 2013-07-21 Cabrillo Isle Marina, San Diego Bay, CA, California, USA Def 32.7272 -117.1995
767733 Ruiz et al., 2015 2013 2013-07-22 Coronado Cays Marina, San Diego Bay, CA, California, USA Def 32.6257 -117.1309
767748 Ruiz et al., 2015 2013 2013-07-18 NAB Fiddlers Cove, San Diego Bay, CA, California, USA Def 32.6524 -117.1486
767765 Ruiz et al., 2015 2013 2013-07-26 Pier 32 Marina, San Diego Bay, CA, California, USA Def 32.6516 -117.1077
767774 Ruiz et al., 2015 2013 2013-07-20 Chula Vista Marina, San Diego Bay, CA, California, USA Def 32.6252 -117.1036
767788 Ruiz et al., 2015 2013 2013-07-28 Marriott Marquis and Marina, San Diego Bay, CA, California, USA Def 32.7059 -117.1655
767991 Ruiz et al., 2015 2012 2012-08-24 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9134 -122.3523
768067 Ruiz et al., 2015 2012 2012-09-11 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
768090 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6633 -122.3817
768114 Ruiz et al., 2015 2012 2012-08-29 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3174
768180 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Def 37.7987 -122.3228
768200 Ruiz et al., 2015 2012 2012-09-07 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7940 -122.2787
768255 Ruiz et al., 2015 2012 2012-09-12 Emeryville, San Francisco Bay, CA, California, USA Def 37.8396 -122.3133
768281 Ruiz et al., 2015 2013 2013-08-15 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7656 -122.2858
768302 Ruiz et al., 2015 2013 2013-08-20 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3163
768320 Ruiz et al., 2015 2013 2013-08-22 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7926 -122.2746
768342 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9723 -122.4829
768362 Ruiz et al., 2015 2013 2013-08-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6639 -122.3821
768385 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.5024 -122.2134
768406 Ruiz et al., 2015 2013 2013-08-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9138 -122.3522
768423 Ruiz et al., 2015 2013 2013-08-12 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8078 -122.4354
768436 Ruiz et al., 2015 2013 2013-08-21 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6980 -122.1908
768454 Ruiz et al., 2015 2013 2013-08-16 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8611 -122.4851

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