Invasion History

First Non-native North American Tidal Record: 1937
First Non-native West Coast Tidal Record: 1937
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Watersipora subtorquata is an encrusting bryozoan widely distributed around the globe. Its native range is poorly understood because of taxonomic confusion with related species, particularly W. subovoidea, which it has been lumped with in older literature as 'W. cucullata' (Gordon 1989; Gordon and Mawatari 1992), or by treating W. subovoidea as a synonym of W. subtorquata (Seo 1999). This species was described from Rio de Janeiro, Brazil (Gordon 1989). We regard it as cryptogenic in the western Atlantic, where early records (e.g., Osburn 1914, Dry Tortugas; Osburn 1940, Puerto Rico) refer to W. cucullata, and where it has also been recorded as W. subovoidea (Winston 1982). In recent collections, it has been identified in Jamaica (Creary 2003), Puerto Rico (in 2007, Ruiz et al. unpublished data), and in Florida: Biscayne Bay (in 2004, Ruiz et al. unpublished data), Indian River Lagoon (in 2007, Ruiz et al. unpublished data), and Jacksonville (in 2002, Ruiz et al. unpublished data). This bryozoan also occurs in the Southwest Atlantic, on the west coast of South Africa (Florence et al. 2007, cited by Mead et al. 2011b). We also consider it cryptogenic in the Northwest Pacific, where it has been reported from the Sea of Japan and East China Sea coasts of South Korea, the Yellow Sea coast of Korea and China, and the Pacific coast of Japan (Tokyo Bay and southward) south to the Paracel Islands in the South China Sea (Seo 1999; Huang 2001).

In California, molecular surveys identified two clades of Watersipora subtorquata (Clades A and B), and an additional species (Watersipora n. sp.). Clade A is widely distributed globally, including Australia, New Zealand, and South Korea. Clade B is known from China and California (Mackie et al. 2012). In California, Clade A occurred from San Diego to Humboldt Bay, but was most abundant in southern and central-region harbors. Clade B occurred over the same range, but was usually less dominant, and was very spotty in the northern part of the range (Mackie et al. 2012).

Watersipora subtorquata has been introduced to the Northeast Pacific (1st record 1888, Gulf of California, and Cabo San Lucas to Puget Sound, Cohen and Carlton 1995; Cohen 2005; Ruiz et al. unpublished data), much of the coast of Australia (1st record 1950, Sydney Harbor, Winston 1977), New Zealand (1st Record 1983, Gordon and Mawatari 1992), and the Atlantic coast of France (1st Record 1973, d'Hondt 1984, cited by Ryland et al. 2009). This organism has a short planktonic stage (Gordon and Mawatari 1992; Cohen and Carlton 1995) suggesting that ship fouling is its likeliest mode of transport to most locations. However, its appearances in France appear related to culture of the Pacific Oyster (Crassostrea gigas) (Ryland et al. 2009).

North American Invasion History:

Invasion History on the West Coast:

Bryozoans of the Watersipora subtorquata complex have been introduced to the northeast Pacific (1st record 1888, Gulf of California, and Cabo San Lucas to Puget Sound, (Cohen and Carlton 1995; Cohen 2005; Ruiz et al. unpublished data). The invasion history of the Watersipora spp. on the west coast is murky, with conflicting molecular and morphological surveys. In California, molecular surveys identified two clades of Watersipora subtorquata (Clades A and B), and an additional species (Watersipora n. sp.). Clade A is widely distributed globally, including Australia, New Zealand, and South Korea. Clade B is known from China and California (Mackie et al. 2012). In California, Clade A occurred from San Diego to Humboldt Bay, but was most abundant in southern and central-region harbors. Clade B occurred over the same range, but was usually less dominant, and was very spotty in northern part of the range They found a third form, Watersipora n. sp., ranging from southern California to Puget Sound. This species has been named formally or described morphologically (Mackie et al. 2012). The molecularly defined species Clades A and B have not been studied morphologically and are unnamed as present. 
 
In a global survey of the genus Watersipora, Vieira et al. (2014) mapped three species on the West Coast, the well-defined W. arcuata, W. subatra, and the little-known?W. atrofusca Vieira et al. (2014) identified the well-defined W. arcuata and W. subatra, and W. atrofusca (known from Mexico and possibly southern California, little information available). Watersipora subatra ranges from Mexico to Puget Sound (Vieira et al. 2014; Ruiz et al. unpublished data). The 'true' W. suborquata was not shown on the West Coast of North America on Vieira et al.’s (2014) map. However, it has been found locally in Long Beach and other sites in southern California, but is less widespread (Linda McCann, personal communication).

Invasion History on the East Coast:

Invasion History on the Gulf Coast:

Invasion History in Hawaii:

The earliest record for Watersipora subtorquata in Hawaii are specimens collected in 1966 in Pearl Harbor, and the Ala Wai Marina, near Honolulu, Oahu, in 1966 (Carlton and Eldredge, 2009). This bryozoan was collected on fouling plates in 2007 in Kaneohe Bay, Barbers Point Harbor, and several locations near Honolulu (Ruiz et al., unpublished data).

Invasion History Elsewhere in the World:

Since Watersipora subtorquata is a member of a complex of cryptic species, its native and introduced ranges are uncertain. Possible native regions include the tropical southwest Atlantic (type locality in Brazil), and the Indo-West Pacific (Vieira et al. 2014). It is found in many parts of the of the Mediterranean, from France to Egypt but the dates are unknown due to confusion with the native W. cucullata (Harmelin 2014; Vieira et al. 2014). In the eastern Atlantic, W. subtorquata was found in the Azores in 1888 (Chainho et al. 201), Madeira in 2006 (Canning-Clode et al. 2016), and South Africa (Florence et al. 2007; Mead et al. 2011). So far, specimens of Watersipora on the Atlantic coast of Europe (d'Hondt 1984; Ryland et al. 2009) have been identified or re-identified as W. subatra (Vieira et al. 2014). 

Watersipora subtorquata's occurrence in Europe was obscured by its confusion with W. subovoidea, but it was collected in the Bay of Arcachon, France on the Bay of Biscay in 1973 (d'Hondt 1984, cited by Ryland et al. 2009). It was collected again in the Bay of Arcachon in 2003, and in 1999-2007, found in many sites in Brittany and the Channel Islands (Ryland et al. 2009). Most of the collection sites were located near oyster-culture operations, and this species was most likely introduced to Europe with the Pacific Oyster (Crassostrea gigas) (Ryland et al. 2009). This bryozoan is also introduced on the west coast of South Africa (Florence et al. 2007, cited by Mead et al. 2011b).


Description

Watersipora subtorquata is an encrusting bryozoan, growing in single layers of flat substrates, but becomes multilamellar on rough substrates, and sometimes erect and leaf-like (foliaceous). Zooids are roughly elongate-rectangular, about twice as long as wide, being740–1,500 µm X 290–680 µm in size. The frontal shield is flat to slightly convex, perforated by numerous round pseudopores. Lateral-oral septulae (connecting the zooids) are absent. The orifice is slightly wider than the long, proximal edge, with a V-shaped or rounded sinus and approximately 130 x 260 (mean ~230) µm in size. It occupies less than 10% of the total zooid area with a well-defined proximal sinus demarcated by triangular condyles, ~55 × 115 µm (Ryland et al. 2009). The rim of the orifice has projecting proximo-lateral, and triangular condyles. The operculum has a dark central band and two lucida (transparent spots) adjacent to the condyles (Vieira et al. 2014). There are no oral spines or avicularia, and no ovicells. However, the zooids brood bright orange-red embryos internally. Colonies are orange to brownish-purple or black with dried or dead colonies becoming dark-orange or gray. (Description from Gordon and Mawatari 1992; Seo 1999; Ryland et al. 2009; Vieira et al. 2014). Colonies can become erect and leaf-like, with extensively overlapping calcareous crusts and curled edges. The crusts are often grayish black or dull orange, with bright orange (New Zealand) or purplish red (Korea) growing edges. The operculum is strongly pigmented with a dark, broad, biconcave band proximally, gradually spreading around paired clear areas. The polypides have orange lophophores 740 x 850 µm long, and 24 tentacles (Gordon 1989).  
 
NOTE: A recent revision of Watersipora taxonomy   presented a drastic change in the nomenclature and worldwide biogeography of the genus. Vieira et al.'s (2014), map shows W. subtorquata as absent from the West Coast of North America and shows three species on the West Coast: W. subatraW. atrofusca, and W. arcuata, with W. subatra identified only from California and Pacific Mexico. Watersipora subtorquata is widely distributed in the tropical-subtropical Atlantic and Mediterranean, Red Sea, and Indo-West Pacific. Another previously identified species, W. subovoidea, has been abandoned and partially synonymized with an earlier name W. cucullata, a name until recently regarded as obsolete (Vieira et al. 2015). This worldwide revision of the genus is at odds with current molecular studies (Mackie et al. 2006; Mackie et al. 2012). Vieira et al.'s (2014) map includes only a small number of West Coast samples. A larger regional sample will be required to reconcile morphological and molecular taxonomy of Watersipora on the West Coast of North America. Until this is available, we will use the names W. subtorquata and Watersipora n. sp. in NEMESIS. 


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Bryozoa
Class:   Gymnolaemata
Order:   Cheilostomata
Suborder:   Ascophora
Family:   Watersiporidae
Genus:   Watersipora
Species:   subtorquata

Synonyms

Escharina torquata (d'Orbigny, 1542)
Watersipora cucullata (Marcus, 1937)
Watersipora subovoidea (d'Orbigny, 1852)

Potentially Misidentified Species

Watersipora subovoidea
Busk 1854. Now considered synonymous with W. cucullata. Described from the Mediterranean Sea, and largely distributed there (Vieira et al. 2014).

Watersipora arcuata
Banta 1969 Described from San Diego, where it was a recent introduction. Possibly native to tropical-subtropical East Pacific, introduced to Hawaii, Australia, and New Zealand (Banta 1969; Mackie et al. 2006; Mackie et al. 2012).

Watersipora atrofusca
Busk 1856. Reported ony from Mazatlan, Mexico, and Califonria, little information available (Vieira et al. 2014).

Watersipora cucullata
Busk1854, described from the Mediterranean Sea

Watersipora n. sp.
This newly identified, undescribed species is morphologically nearly identical to W. subtorquata but distinct at the molecular level (Mackie 2006; Mackie et al. 2012). So far, it is known only from California and South Korea.

Watersipora nigra
Described by Canu and Bassler 1930. as Pachycleithonia nigra from theGalapagos Islands, and moved into the genus Watersipora, by Vieira et al. (2014). It was not found in recent surveys (Carlton et al. 2019; Vieira et al. 2019).

Watersipora subatra
(d'Orbigny 1852) Described from the Aegean Sea. Re-described and redefined by Ryland et al. (2009). Watersipora subatra occurs in the Western Pacific from Japan to New Zealand as well as the West Coast.

Ecology

General:

Life History- Watersipora subtorquata is an encrusting, calcified 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 (Barnes 1983). Larvae of W. subtorquata are lecithotrophic and have a short planktonic period (less than 1 day, Mackie et al. 2006). Larvae settle on a substrate and metamorphose into the first zooid of a colony, an ancestrula (Barnes 1983). 
 
Ecology- Watersipora subtorquata is known from pilings, rocks, shells, floats, oil platforms, ships' hulls, and fouling plates (Mackie et al. 2006; Page et al. 2006; Cohen and Zabin 2009; Ryland et al. 2009). 

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatVessel HullNone
General HabitatCoral reefNone
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)6.7Field, based on coldest site in geographical range, Port Townsend WA (Zerebecki and Sorte 2011)
Maximum Temperature (ºC)30.6Field, based on warmest site in geographical range, Red Sea (Zerebecki and Sorte 2011)
Minimum Salinity (‰)25Field salinity (California) (Cohen 2005)
Maximum Salinity (‰)40Field salinity (Shark Bay, Western Australia) (Wyatt et al. 2005)
Minimum Duration0Larva- Cohen and Carlton 1995
Maximum Duration1Larva- Cohen and Carlton 1995
Broad Temperature RangeNoneWarm-Temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Watersipora subtorquata is an encrusting bryozoan widely distributed around the globe. Its colonies can be erect and leaf-like, with extensive overlapping calcareous crusts and curled edges, which create secondary habitat for the settlement of other marine invertebrates. Its native range is poorly understood because of taxonomic confusion with related species, particularly W. subatra. However, introduced populations have been recorded on the West coast of the United States, Hawaii, Australia, New Zealand, Europe, and South Africa. This species is known from rocks, oyster shells, pilings, floats, oil platforms, ships' hulls, and fouling plates. It is tolerant of copper and mercury antifouling paints and has outcompeted congeneric species in some areas of its introduced range.  

Economic Impacts Shipping and Boating- Watersipora spp. have long been known to be tolerant of copper and mercury in antifouling paint (Allen 1953; Ryland 1971; Piola and Johnston 2006). Their hard encrusting colonies are tolerant of moving water, and their colonies also provide non-toxic points of attachment for other organisms, allowing a diverse fouling community to develop (Floerl et al. 2004), which can adversely affect the speed and efficiency of ships. McKenzie et al. (2011) found that colonies from different sites varied in copper tolerance, and that tolerance was heritable in cultures. Colonies that produced large larvae tended to be more copper-tolerant (McKenzie et al. 2011).  

Ecological Impacts Competition- In New Zealand, W. subtorquata (arriving in 1983) quickly replaced W. arcuata at most locations (Gordon and Mawatari 1992). In Port Phillip Bay, Victoria, Australia W. subtorquata (arriving in 1976) also became the dominant Watersipora species (Keough and Ross 1999). In southern California, W. subtorquata was as of 2000–2003 the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008 where W. arcuata was previously predominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates and may be expected to extend its range northward as the climate warms. Watersipora subtorquata was one of a group of seven non-native species in Bodega Harbor, most of which were rare or absent in 1970–1971 but were among the eight most abundant species in 2006. 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).  

Habitat Change- Watersipora subtorquata colonies provide habitat for other organisms. Its colonies often develop elevated leaf-like folds rising above the substrate, creating additional space for colonization by other organisms. In Bodega Harbor, Geller et al. 2008 found where morphologically identical species (either W. subtorquata or Watersipora n. sp.) had more than 10% cover on fouling plates, native diversity was correlated with exotic diversity, as both groups of organisms occupied the expanded area (Sellheim et al. 2010). On the hulls of ships, and other surfaces treated with antifouling paints, W. subtorquata is often the only species able to settle, and its colonies provide surfaces on which more sensitive organisms can settle (Floerl et al. 2004). 


Regional Impacts

AUS-XNoneEconomic ImpactShipping/Boating
Shipping and Boating- Watersipora spp. have long been known to be tolerant of copper and mercury in antifouling paint (Piola and Johnston 2006). Their hard encrusting colonies are tolerant of moving water, and their colonies also provide non-toxic points of attachment for other organisms, allowing a diverse fouling community to develop (Floerl et al. 2007). Experimental studies on W. subtorquata's response to antifouling paint were done in Botany Bay and Port Jackson, near Sydney (Piola and Johnston 2006).
AUS-XIINoneEconomic ImpactShipping/Boating
Watersipora spp. have long been known to be tolerant of copper and mercury in antifouling paint (Allen 1950; Ryland 1971; Piola and Johnston 2006). Their hard encrusting colonies are tolerant of moving water, and their colonies also provide non-toxic points of attachment for other organisms, allowing a diverse fouling community to develop (Floerl et al. 2004). Experimental studies on W. subtorquata's response to antifouling paint were done in Townsville and Cairns, Queensland (Floerl et al. 2007).
NZ-IVNoneEcological ImpactCompetition
In New Zealand, W. subtorquata, arriving in 1983, quickly replaced W. arcuata at most locations (Gordon and Mawatari 1992).
NZ-VINoneEcological ImpactCompetition
In New Zealand, W. subtorquata, arriving in 1983, quickly replaced W. arcuata at most locations (Gordon and Mawatari 1992).
AUS-VIIINoneEcological ImpactCompetition
In Port Phillip bay, W. subtorquata, arriving in 1976, quickly replaced W. arcuata at most locations (Keough and Ross 1999). Heavy recruitment of W. subtorquata can affect subsequent community development. However, impacts varied by site and season (Sams and Keough 2012).
NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactCompetition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms. Experimental clearing and routine cleaning of oil platforms favored increased abundance of Watersipora subatra, recruitng in the cleared area (Viola et al. 2018). Viola et al. (2018) suggest the retention of oil platforms as artificial reefs could favor W. subatra. Removal of the upper portions of the platforms might reduce their potential as a source for coastal populations of the bryozoan.
P023_CDA_P023 (San Louis Rey-Escondido)Ecological ImpactCompetition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms.
P050San Pedro BayEcological ImpactCompetition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms.
P060Santa Monica BayEcological ImpactCompetition
In southern California, W. subtorquata is now (2000-2003) the dominant or only form at several sites (Oceanside Harbor, Alamitos Bay, King Harbor; Geller et al. 2008) where W. arcuata was previously dominant (Banta 1969). Geller et al. (2008) suggest that W. subtorquata may be more competitive than W. arcuata in warm-temperate climates, and may be expected to extend its range northward as the climate warms.
AUS-XIINoneEcological ImpactHabitat Change
On the hulls of ships and other surfaces, treated with antifouling paints, W. subtorquata is often the only species able to settle, and its colonies provide surfaces on which more sensitive organisms can settle. Experimental studies were performed in Cairns and Townsville, Queensland (Floerl et al. 2004).
AUS-XNoneEcological ImpactCompetition
Exposure to copper anti-fouling paint resulted in enhanced recruitment of W. subtorquata, despite post-settlement mortality. Surviving colonies had shorter ancestrulae and were smaller (MacKenzie et al. 2012). The ability to tolerate and even be favored by copper pollution gives W. subtorquata a competitive advantage in polluted habitats (MacKenzie et al. 2011; MacKenzie et al. 2012a)
P065_CDA_P065 (Santa Barbara Channel)Ecological ImpactCompetition
Experimental clearing and routine cleaning of oil platforms favored increased abundance of Watersipora subatra, recruitng in the cleared area (Viola et al. 2018). Viola et al. (2018) suggest the retention of oil platforms as artificial reefs could favor W. subatra. Removal of the upper portions of the platforms might reduce their potential as a source for coastal populations of the bryozoan.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEP-VI Pt. Conception to Southern Baja California 1937 Def Estab
AUS-VIII None 1976 Def Estab
AUS-VII None 1977 Def Estab
AUS-V None 0 Def Estab
AUS-IV None 2001 Def Estab
AUS-XII None 2000 Def Estab
AUS-X None 1950 Def Estab
NZ-IV None 1983 Def Estab
SA-II None 1862 Crypto Estab
NWP-4a None 1941 Crypto Estab
NWP-3a None 0 Crypto Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 0 Crypto Estab
CAR-VII Cape Hatteras to Mid-East Florida 2002 Crypto Estab
CAR-IV None 1940 Crypto Estab
RS-3 None 1924 Crypto Estab
NWP-3b None 0 Crypto Estab
SP-IX None 2002 Def Estab
AUS-III None 2002 Def Estab
NEP-VII None 1888 Def Estab
P020 San Diego Bay 1997 Def Estab
P030 Mission Bay 1989 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 2000 Def Estab
P027 _CDA_P027 (Aliso-San Onofre) 2001 Def Estab
P040 Newport Bay 2001 Def Estab
P058 _CDA_P058 (San Pedro Channel Islands) 2001 Def Estab
P050 San Pedro Bay 1964 Def Estab
P060 Santa Monica Bay 1972 Def Estab
P062 _CDA_P062 (Calleguas) 2001 Def Estab
P064 _CDA_P064 (Ventura) 1963 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 2001 Def Estab
NZ-VI None 1982 Def Estab
AUS-IX None 2001 Def Estab
AUS-II None 2001 Def Estab
AUS-I None 1999 Def Estab
NWP-2 None 0 Crypto Estab
EAS-I None 0 Crypto Estab
S180 St. Johns River 0 Crypto Estab
MED-VI None 1996 Def Estab
S200 Biscayne Bay 2004 Crypto Estab
CAR-II None 0 Crypto Estab
S190 Indian River 0 Crypto Estab
NA-ET4 Bermuda 0 Crypto Estab
WA-IV None 1937 Def Estab
MED-V None 0 Def Estab
AUS-XIII None 1972 Def Estab
SP-XXI None 1966 Def Estab
SP-XIII None 2010 Def Estab
SA-III None 1842 Crypto Estab
WA-I None 2006 Def Estab
NEP-VIII None 1937 Def Estab
MED-II None 1971 Def Estab
NEA-VI None 1888 Def Estab
MED-VII None 0 Def Estab
MED-IV None 2015 Def Estab
CIO-II None 2015 Def Estab
P095 _CDA_P095 (Tomales-Drakes Bay) 2015 Def Estab
NEP-V Northern California to Mid Channel Islands 2015 Def Estab
SEP-Z None 1987 Def Estab
WA-VI None 2008 Def Estab
MED-III None 0 Def Estab
P030 Mission Bay 2000 Def Estab
P023 _CDA_P023 (San Louis Rey-Escondido) 2000 Def Estab
P050 San Pedro Bay 2000 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
7745 Ruiz et al., unpublished data 2002 2002-01-01 Pt. Defiance, Tacoma Def 47.3179 -122.5462
7746 Ruiz et al., unpublished data 2002 2002-01-01 Bremerton Def 47.5673 -122.6326
7750 Cohen and Carlton 1995 1994 1994-01-01 Tomales Bay Def 38.1696 -122.9100
7751 Cohen 2005 2005 2005-01-01 Richmond Def 37.9110 -122.3511
7752 Cohen 2005 2005 2005-01-01 Tiburon Def 37.8774 -122.4450
7753 Cohen 2005 2005 2005-01-01 San Leandro Def 37.7249 -122.1561
7754 Cohen 2005 2005 2005-01-01 San Mateo Bridge Def 37.5797 -122.2555
7755 Cohen et al. 2005 2004 2004-05-24 Coast Guard Island, Alameda Def 37.7812 -122.2457
7756 Cohen et al. 2005 2004 2004-05-24 Presidio Yacht Club, San Francisco Def 37.8326 -122.4741
7757 Cohen et al. 2005 2004 2004-05-23 Brisbane Lagoon Def 37.6862 -122.3906
7758 Cohen and Carlton 1995 1994 1994-01-01 Moss Landing Harbor Def 36.8002 -121.7872
7759 de Rivera et al. 2005 2003 2003-01-01 Kirby Park, Elkhorn Slough Def 36.8410 -121.7463
7760 Cohen and Carlton 1995 1994 1994-01-02 Monterey Harbor Def 36.6116 -121.8955
7761 de Rivera et al. 2005; Mackie et al. 2006 2003 2003-01-01 Santa Cruz Harbor Def 36.9658 -122.0016
7764 Cohen 2005 1965 1963-01-01 Rincon Beach Def 34.3356 -119.4071
7768 de Rivera et al. 2005 2003 2003-01-01 Ventura West Marina Def 34.2464 -119.2611
7769 Page et al. 2006 2002 2002-01-01 Oil Platform 'Grace', off Ventura/ Def 34.1667 -119.4833
7770 Page et al. 2006, Goddard and Love 2010 2002 2002-01-01 Oil Platform 'Gilda', off Ventura Def 34.1667 -119.4167
7773 Cohen et al. 2002 2000 2000-01-01 Marina del Rey Def 33.9722 -118.4522
7776 Cohen et al. 2002 2000 2000-01-01 Cabrillo Boat Ramp, Los Angeles Def 33.7131 -118.2683
7777 Cohen 2005 1972 1972-01-01 Long Beach Marina Def 33.7528 -118.1112
7778 Cohen 2005 1964 1964-01-01 Huntington Beach Def 33.7508 -118.0242
7782 Cohen et al. 2002 2000 2000-01-01 Snug Harbor Marina, Carlsbad Def 33.1478 -117.3322
7783 Cohen 2005 1989 1989-01-01 Mission Bay Def 32.7792 -117.2342
7785 Cohen et al. 2002 2000 2000-01-01 Chula Vista Boat Ramp, San Diego Def 32.6211 -117.1031
7786 de Rivera et al. 2005 2003 2003-01-01 Sunroad Marina, San Diego Bay Def 32.7263 -117.1922
7787 Rodriguez and Ibarra-Obando 2008 2008 2008-01-01 Bahia San Quintin Def 30.5603 -115.9425
7788 Cohen 2005 1937 1937-01-01 Cabo San Lucas Def 22.8897 -109.9156
7790 Soule 1966, cited by Carlton and Eldredge 2009 1966 9999-01-01 Pearl Harbor Def 21.3550 -157.9722
7791 Ruiz et al., unpublished data 2007 2007-01-01 Kane' ohe Bay Yacht Club Def 21.4628 -157.8103
7792 Coles et al. 2003 2002 2002-01-01 Pago Pago Harbor Def -14.2814 -170.6742
7793 Ryland et al. 2009 None 9999-01-01 Bermuda Crypto 32.3000 -64.7833
7794 Ruiz et al., unpublished data 2002 2002-01-01 None Crypto 30.3841 -81.6336
7795 Winston 1982; Winston 2009 1974 1974-01-01 Fort Pierce Crypto 27.4703 -80.2934
7796 Ruiz et al., unpublished data 2004 2004-01-01 Miami Crypto 25.7745 -80.1709
7797 Creary 2003; Creary and Webber 2009 None 9999-01-01 Kingston Harbor Crypto 17.9539 -76.8037
7798 Osburn 1940; Ruiz et al. 2007, unpublished data 1940 1940-01-01 Puerto Rico Crypto 18.4500 -64.0667
7799 Vieira et al. 2008 1842 1842-01-01 Espritio Santo state Crypto -20.0000 -40.0000
7800 d'Orbigny 1842, cited by Gordon 1989 1842 1842-01-01 Rio de Janeiro Crypto -22.9083 -43.1964
7801 d'Orbigny 1842, cited by Vieira et al. 2008 1842 1842-01-01 Sao Paulo Crypto -23.9672 -46.2926
7802 Florence et al. 2007, cited by Mead et al. 2011b 1937 9999-01-01 west coast, South Africa Def -33.9253 18.4239
7803 Canning-Clode et al. in prep. 2007 2007-01-01 Madeira Def 32.6511 -16.9097
7804 Ryland 2009 2003 2003-01-01 oyster farm between Arcachon and Gujan-Mestras Def 44.6364 -1.0667
7805 Ryland et al. 2009 2007 2007-05-01 St Peter Port harbour, Guernsey, Def 49.5500 -2.5500
7806 Ryland et al. 2009 2003 2003-03-01 Port-Navale/ Def 47.5500 -2.9100
7807 Ryland et al. 2009 2002 2002-02-03 Chausey, 28 km northeast of St Malo Def 48.8700 -1.8200
7808 Ryland et al. 2009, 1999 1999-01-01 St-Jacut-de-la-Mer, Def 48.6200 -2.2000
7812 Seo 1999 None 9999-01-01 Chumunjin Crypto 37.8895 128.8177
7813 Seo 1999 None 9999-01-01 Wolsong Crypto 35.7111 129.4743
7814 Seo 1999 None 9999-01-01 Kyokpo Crypto 35.8000 126.2000
7815 Rucker 1969, cited by Winston 1977 None 9999-01-01 Maizuru Crypto 35.4667 135.3833
7816 Huang 2001 None 9999-01-01 Eastern Liaonging Peninsula Crypto 40.0000 124.0000
7817 Huang 2001 None 9999-01-01 Hebei Province Crypto 39.0000 118.0000
7818 Huang 2001 None 9999-01-01 Shangdong Peninsula Crypto 37.4000 121.2667
7819 Long and Rucker 1969 None 9999-01-01 Yokosuka Crypto 35.2500 139.6667
7820 Raveendran and Harada 2002 None 9999-01-01 Kanayama Bay Crypto 33.6819 135.3372
7821 Seo 1999 None 9999-01-01 Nodo Crypto 35.9875 129.1236
7822 Seo 1999 None 9999-01-01 Hoenggando Crypto 34.2406 126.6090
7823 Seo 1999 None 9999-01-01 Sogwepo, Chejedo Island Crypto 33.2528 126.5611
7827 Long and Rucker 1969 None 9999-01-01 Sasebo Crypto 33.1833 129.6700
7828 Huang 2001 None 9999-01-01 Jiangsu Province Crypto 34.0000 121.0000
7829 Huang 2001 None 9999-01-01 Zhejiang Province Crypto 28.0000 122.0000
7830 Huang 2001 None 9999-01-01 Fujian Province Crypto 25.0000 120.0000
7831 Huang 2001 None 9999-01-01 Guangdong Province Crypto 22.0000 114.0000
7832 Huang 2001 None 9999-01-01 Guangxi Province Crypto 21.0000 111.0000
7833 Huang 2001 None 9999-01-01 Hainan Island Crypto 18.5000 109.0000
7834 Huang 2001 None 9999-01-01 Xisha (=Paracel) Islands Crypto 16.5000 112.0000
7835 Huang 2001 None 9999-01-01 Nansha (Spratly) Islands Crypto 10.0000 114.0000
7836 P. Colin, personal communication None 9999-01-01 Palau Crypto 7.3500 134.4667
7837 Wyatt et al. 2005 1999 1999-01-01 Darwin Def -12.4500 130.8333
7838 Mackie et al. 2006 2001 2001-01-01 Dampier Def -20.6631 116.7081
7839 Wyatt et al. 2005 2002 2002-01-01 Carnarvon Def -24.8839 113.6611
7840 Wyatt et al. 2005 2002 2002-01-01 Denham Def -25.9269 113.5339
7841 Wyatt et al. 2005 2002 2002-01-01 Monkey Mia Def -25.7947 113.7172
7842 Mackie et al. 2006 2001 2001-01-01 Fremantle Def -32.0569 115.7439
7843 Hewitt et al., unpublished None 9999-01-01 Esperance Def -33.8778 121.8919
7844 Brock 1983; Keough and Ross 1999 1977 1977-01-01 Adelaide Def -34.9289 138.6011
7845 Keough and Ross 1999 1999 1999-01-01 Port Lincoln Def -34.7322 135.8586
7846 Mackie et al. 2006 2001 2001-01-01 Port Noarlunga Def -35.1500 138.4667
7847 Keough and Ross 1999 1982 1982-01-01 Hobsons Bay, Port Phillip Bay Def -37.8547 144.9358
7848 Keough and Ross 1999 1999 1999-01-01 Queenscliff Def -38.2667 144.6500
7849 Mackie et al. 2006 2001 2001-01-01 Mersey Yacht Club, Devonport Def -41.1806 146.3464
7850 Mackie et al. 2007 2001 2001-01-01 Coningham Def -43.0770 147.2750
7851 Allen and Wood 1950, cited by Winston 1977 1950 1950-01-01 Sydney Harbour Def -33.8583 151.2333
7852 Mackie et al. 2006 2001 2001-01-01 Wollongong Def -34.4331 150.8831
7853 Mackie et al. 2006 2001 2001-01-01 Merimbula Def -36.9000 149.9000
7854 Floerl et al. 2004 2000 200-01-01 Townsville Harbour Def -19.2564 146.8183
7855 Ryland et al. 2009 1972 1972-01-01 Low Isles Def -16.3861 145.5589
7856 Gordon and Mawatari 1992 1983 1983-01-01 Wellington Def -41.2889 174.7772
7857 Cranfield et al. 1998 1998 1998-01-01 Whangarei Harbour Def -35.7167 174.3000
7861 Cranfield et al. 1998 1998 1998-01-01 Auckland Def -36.8404 174.7397
7862 Gordon and Mawatari 1992 1985 1985-01-01 Nelson Def -41.2833 173.2833
7863 Gordon and Mawatari 1992 1982 1982-01-01 Victoria Wharf, Dunedin Def -45.8667 170.5000
7864 Gordon and Mawatari 1992 1983 1983-01-01 Port Chalmers Def -45.8178 170.6188

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