Invasion History

First Non-native North American Tidal Record: 1932
First Non-native West Coast Tidal Record: 1932
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Polydora cornuta was described from Charleston, South Carolina by Bosc in 1802, and later widely found on the East Coast of the United States from the Gulf of St. Lawrence to the Gulf of Mexico (Blake 1971; Blake and Maciolek 1987). It was reported from northern Europe as early as 1859 (Claparede 1861), but its range and invasion status are complicated by confusion with the similar species, P. ciliata, which is usually found boring in shells and calcareous substrates (Wolff 1973; Radashevsky 2005). It is apparently introduced in the Black and Mediterranean Seas (Cinar et al. 2005; Surugiu 2012; Radashevsky and Selifonova 2013), the Southwestern Atlantic (Brazil-Argentina, Orensanz et al. 2002; Radashevsky 2005), the Northeast Pacific (British Columbia to Mexico, Hartman 1941; Light 1977; Carlton 1979; Blake and Ruff 2007), the Northwest Pacific (China-Russia, Sato-Okoshi 1999; Radashvesky 2005), and the Southwest Pacific (Australia-New Zealand, Read 1975; Blake and Kudenov 1978; Walker 2009).

North American Invasion History:

Invasion History on the West Coast:

Polydora cornuta was first collected from Ladysmith, British Columbia (BC) in 1932, and found at the San Juan Islands, Washington (WA), San Francisco Bay, and Mission Bay, California (CA) from 1932 to 1938 (Carlton 1979; Hartman 1941; U.S. National Museum of Natural History 2015). It was subsequently found at many intermediate estuaries, including Grays Harbor, WA (1st record 1953, U.S. National Museum of Natural History 2015); Willapa Bay, WA (1st record 2000, Cohen et al. 2001); the Columbia River estuary, WA – Oregon (OR) (1980, Jones 1983); Coos Bay, OR (Hartman and Reish, 1950; cited by Carlton 1979); Humboldt Bay, CA (Barnhart 1992, cited by Boyd et al. 2002); Elkhorn Slough, CA (1st record 1977, Wasson et al. 2001); and Morro Bay, CA (1st record 1960, Reish and Barnard 1967). Except for the Columbia River, all of these estuaries had received transplants of Eastern Oysters (Crassostrea virginica) (Carlton 1979).

In southern California, although few Eastern Oyster transplants were made, Polydora cornuta has been collected, often in high abundance, both in major harbors for commercial shipping and small bays used by recreational boats. For instance, records are known from Los Angeles-Long Beach Harbor (1st record 1966, Crippen and Reish 1969 cited by Carlton 1979); San Diego Bay (1st record 1994, Levin et al. 1998), Playa del Rey Lagoon, Venice Beach (1st record 1941, Alan Hancock Foundation, cited by Carlton 1979); Alamitos Bay (1st record 1952, Reish and Winter 1954, cited by Carlton 1979); Anaheim Bay (1st record 1972, Reish et al. 1975); and Santa Catalina Island (1st record 1963, Reish et al. 1964, cited by Carlton 1979). Polydora cornuta has also been introduced to the Salton Sea, a land-locked artificial saline lake, and was first collected there in 1961 (Carlton 1979; USNM 39642, U.S. National Museum of Natural History 2015).

It is likely that Polydora cornuta has been transferred from the East to the West Coast many times, with Eastern Oyster transplants, hull fouling, and ballast water, and later moved to additional estuaries by hull fouling, and perhaps by other vectors, such as discarded shellfish, or seaweed used to pack bait.

Invasion History Elsewhere in the World:

In Europe, Polydora cornuta was reported from Scotland in 1859 (Claparede 1861), but was often confused with P. ciliata, although the latter species is predominantly associated with shells and other calcareous substrates (Wolf 1973; Michaelis 1978; Ramberg and Schram 1982; Surugiu 2012). Wolff (1973) notes that it was collected in the Netherlands in 1930, but identified then as P. ciliata. It is known from the Scheldt, Meuse and Rhine estuaries in the Netherlands (Wolff 1973); Langeoog Island and the Jade-Weser Delta, Germany (U.S. National Museum of Natural History 2015; Michaelis 1978); and the inner Oslofjord, Norway (Ramberg and Schram 1982). It is also known from the western Baltic, including the Isefjord, Denmark (Hansen et al. 2010), and the Kiel Bight and Mecklenberg Bay, Germany (Anger 1977; Radashevsky 2005; U.S. National Museum of Natural History 2015). Records on the outer Atlantic coast of Europe are rare and scattered, including the Isle of Skye/Scotland (Claparede 1861), Marennes-Oleron Bay, France (de Montaudouin and Sauriau 2000), and the Óbidos lagoon, Portugal (Carvalho et al. 2011). The frequent confusion between P. ciliata and P. cornuta makes it difficult to determine the invasion status of P. cornuta, at least in northern Europe. Genetic studies would be useful in resolving the origin of these worms.

In the Black Sea, there were apparently no non-boring Polydora collected before 1962, and in the Mediterranean, records of P. cornuta are recent, widely scattered, and located near harbors. These include Valencia, Spain (1st record 1990, Tena et al. 1991); the Venice Lagoon, Italy (1st record 1991, Tagliapietra et al. 1998); Izmir Bay, Turkey (1st record 2003, Cinar et al. 2005); and the Saronikos Gulf, Greece (1st record 2008, Simboura et al. 2008). In the Black Sea proper, P. cornuta was first collected off Odessa, Ukraine in 1962, and initially identified as P. ciliata limicola (Losovskaya and Zolotarev 2003). It is now widespread in the Black Sea, the Sea of Marmara, and the Sea of Azov (Dagli and Ergen 2008; Karhan et al. 2008; Radashevsky and Selifonova 2013).

In the Southwestern Atlantic, Polydora cornuta was first collected in the Mar Chiquita Lagoon, Mar Del Plata, Argentina in 1967 (Orensanz et al. 2002; Radashevsky 2005; U.S. National Museum of Natural History 2015) and later, in Rio de Janeiro Harbor (in 1986) and Paranagua Bay (in 1998), Brazil (Radashevsky et al. 2005; U.S. National Museum of Natural History 2015). In the Northwest Pacific, it was first reported from the east coast of Taiwan, on the Formosa Straits (in 1990, Radashevsky and Hsieh 2000; Radashevsky 2005). Other early records are from Shihwa Lake, South Korea, on the Yellow Sea (in 1994, Lee and Cha, 1997, cited by Radashevsky 2005) and Vladivostok, Russia (in 1994, Radashevsky 2005). Polydora cornuta was collected at several ports on the southeastern and west coasts of Japan (1984-1997, Sato-Okoshi 2000). One puzzling record came from the Sea of Japan, off Russia, at 3334 m depth (in 2010, Alalykina 2013). This polychaete is also known from two ports in New Zealand: Whangarei and Waitemata Harbors (Read 1975; Read and Gordon 1991), and from Victoria (Blake and Kudenov 1978) and Botany Bay, Australia (in 1973, Walker 2009).


Description

Adults of the 'Polydora cornuta' species complex have up to 90 segments. The prostomium has a lateral incision, forming two lobes and is prolonged back to chaetiger 3 as a caruncle. Most specimens have 4 eyes in a trapezoidal arrangement, on the prostomium, with a small, triangular, occipital tentacle located posteriorly to them. A ciliated groove runs on either side of the caruncle. The prostomium is flanked by a pair of long, prehensile palps, which stretch back to chaetigers 15-35. The palps each have a ciliated groove along their lengths, bordered by a fine black line, but lack conspicuous pigment.

Chaetiger 1 has well-formed parapodial lobes, but has chaetae (fine, capillary) only on the neuropodia. The notopodia lack chaetae, but have long, finger-like lamellae. Chaetigers 2-4 and 6 have bundles of capillary neurochaetae and notochaetae. However, chaetiger 5 is modified and bears a row of 6-8 stout spines with adhering companion chaetae, and with dorsal musculature running back to chaetigers 6 and 7. Chaetigers from 7 on, have long, thin branchiae and up to 15 stout hooded hooks on the neuropodia, while notopodial chaetae decrease in number posteriorly. The pygidium is flared and resembles a suction cup. Adult worms reach a length of 12-32 mm and are light tan in color. Smaller worms have black melanophores laterally from chaetigers 2-7 to 10-22, and yellow chromatophores ventrally from chaetigers 5-7 to 16-21. Larval development is described (as P. ligni) by Blake (1969). Description based on: Blake 1971, Blake and Kudenov 1978, Light 1978, Blake 1983, Blake and Maciolek 1987, Radashevsky and Hsieh 2000, Cinar et al. 2005, Radashevsky 2005, and Radashevsky and Selifonova 2013; Bertasi 2016.

The species was first described in 1805 from Charleston, North Carolina, but was later known more widely under the name Polydora ligni (Webster 1879, described from New Jersey). The name 'Polydora cornuta' was re-instituted and the animal redescribed by Blake and Maciolek (1987). Populations of P. cornuta around the world show small differences in morphology and substantial differences in size which may indicate both genetic and environmental differences (Radashevsky and Hsieh 2000; Cinar et al. 2005). Rice (1991) and Rice et al. (2008) found that populations of P. cornuta from Florida, Maine, and California each showed genetic differences and low frequency of hybridization indicating their status as cryptic species. However worms from North Carolina and California were interfertile (Rice 1991), and California worms were interfertile with worms from New Zealand (Rice et al. 1998). Further genetic comparisons among populations are needed to trace the likely anthropogenic spread of this species.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Annelida
Class:   Polychaeta
Subclass:   Palpata
Order:   Canalipalpata
Suborder:   Spionida
Family:   Spionidae
Genus:   Polydora
Species:   cornuta

Synonyms

Polydora amarincola (Hartman, 1936)
Polydora ciliatum (Agassiz, 1867)
Polydora cornuta (Blake and Maciolek, 1987)
Polydora ligni (Webster, 1809)
Polydora littorea (Verrill, 1881)

Potentially Misidentified Species

Polydora ciliata
Polydora ciliata (Johnston 1833) is primarily a borer in calcareous substrates, native to the Northeast Atlantic. In European waters, P. cornuta has been widely misidentified as P. ciliata (Wolff 1973; Michaelis 1978; Blake and Maciolek 1987; Bertasi 2016). Polydora ciliata and P. websteri are closely related, frequently confused, and can be considered as members of the same species complex of shell-boring Polydora spp. (Sato-Okoshi and Abe 2013; Surugiu 2012).

Polydora limicola
Polydora limicola (Annenkova 1934) was described from the Bering Sea, initially as a subspecies of P. ciliata. It has been reported as an introduction from Vladivostok, Russia (Bagaveeva and Zvyagintsev 2000) and the Black Sea. Specimens from the Black Sea were re-identified as P. cornuta (Radashevsky and Selifonova 2013). Identifications of this polychaete on the West Coast (e.g. Light 1977; Blake and Ruff 2007) probably refer to other species, including P. cornuta (Leslie Harris, personal communication 2014; Vasily Radashevsky, personal communication 2015).

Polydora nuchalis
Polydora nuchalis (Woodwick 1953), native to the Northeast Pacific is morphologically similar to P. cornuta, but differs in the structure of spines and companion chaetae on chaetiger 5. It inhabits estuarine mudflats (Blake and Ruff 2007).

Polydora websteri
Polydora websteri (Hartman 1943) described from Milford, Connecticut, is a reportedly cosmopolitan shell-borer (Blake 1971; Radashevsky 1999; Sato-Okoshi and Abe 2013).

Ecology

General:

'Polydora cornuta' is an estuarine spionid polychaete, which dwells in mud and mucus tubes in sediment, and in fouling communities on hard surfaces (Blake 1971; Blake and Maciolek 1987). 'Polydora cornuta' is a widely distributed complex of cryptic species, probably varying in environmental tolerances and life history (Rice et al. 2008). The sexes are separate (Rice 1991). Eggs are laid in capsules, linked in strings attached to the interior of the tubes. Capsules in Maine estuaries contained up to 132 eggs, but about half of those were unfertilized and used as food by developing larvae (Blake 1969; Gibson et al. 2012). Larvae are released from the capsules at the 3-chaetiger stage, and feed in the plankton (Blake 1969). Larvae settle at about 12-22 chaetigers, and ~1.1-1.3 mm length (Blake 1969; Orth 1971; Levin 1984; Anger et al. 1986; Radashevsky 2005). Development is strongly effected by temperature, with time to settlement shortening with increasing temperatures (60 days at 10C ; 16-28 days at 12C; 12-41 days at 18C; 7-14 days at 20C, 12 days at 28C) (Orth 1979; Levin 1984, Anger et al. 1986; Takata et al. 2011). Food supply can also effect developmen - two weeks of partial starvation led to an approximate doubling of time to settlement, from ~19 days to 42 days at 11C (Qian and Chia 1993). The wide variation in development times with temperature may reflect laboratory food supply, as well as genetic differences in populations from different regions. Settlement is stimulated by the presence of sediment containing microorganisms (Sebesvari et al. 2006). Growth to first reproduction is rapid, 33 days in culture at 18 C. Adults in culture, survived for an average of 13 months (Anger et al. 1986). In a Brazilian population, some males were mature at 2 mm length and females at 5 mm (Radashevsky 2005), but adult lengths of 10-20 mm are more typical (Levin 1984) and adults can reach 32 mm (Radashevsky 2005).

'Polydora cornuta' is known from a wide range of climates, and estuarine-marine habitats from the tropics to cold-temperate waters, although some of this range may result from the occurrence of cryptic species. Its habitats include subtidal and intertidal soft substrates, fouling communities in marinas and harbors, ships' hulls, and oyster beds (Graham and Gay 1945; Wolff 1973; Bagaveeva and Zvyagintsev 2000). This worm's habitats are often exposed to temperature and salinity extremes, as well as hypoxia, exposure and toxic chemicals (Wolff 1973; Levin 1984; Cinar et al. 2005). Polydora cornuta occurs in water prone to sea ice formation (Blake 1969), and has been collected from salinities of 29C (Cory 1967) in Chesapeake Bay, and probably occurs at higher temperatures. It has been collected at a salinity range of 2-75 PSU, and is abundant at salinities ranging from 5 to 75 PSU (Wolff 1973; Simmons 1957). Polydora cornuta is also tolerant of hypoxic conditions and is considered an indicator species for pollution (Anger 1977; Rice et al. 1980; Cinar et al. 2005; Levin et al. 2009).

Its larvae are planktotrophic, feeding on phytoplankton (Anger et al. 1986). Adults of P. cornuta are capable of shifting between deposit-feeding detritus and surface-microalgae, and suspension-feeding (phytoplankton and suspended particles). Their ciliated palps can either stretch over the sediment surface to pick up sediment particles, or wave in the current to pick up phytoplankton (Dauer et al. 2005). Feeding and growth rates of P. cornuta increased in simulated tidal currents (Hentschel and Herrick 2005). Although P. cornuta is generally considered a suspension and deposit-feeder, predation on small bivalves (veligers of several species, juveniles of Gemma gemma) has been documented (Breese and Phibbs 1972; Weinberg 1984). The large populations of P. cornuta in eutrophic waters are a potential food source for fishes and invertebrates (Oviatt et al. 1987). Sublethal predation on P. cornuta, with predators biting off parts of worms, with subsequent regeneration, was common in Long Island Sound (Zajac 1995). However, hypoxic conditions can limit the availability of these worms to fishes and other active predators.

Food:

Phytoplankton; benthic microalgae, detritus

Consumers:

Fishes, shrimps, crabs

Competitors:

Other polychaetes, Macoma spp.

Trophic Status:

Deposit Suspension Feeder

DepSusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatUnstructured BottomNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatSalt-brackish marshNone
General HabitatGrass BedNone
General HabitatVessel HullNone
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEndobenthicNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)-2Based on geographical range
Maximum Temperature (ºC)29Field, Patuxent River, Chesapeake Bay (Cory 1967). This polychaete probably tolerates higher temperatures.
Minimum Salinity (‰)2Field, abundant above 5 PSU (Wolff 1973)
Maximum Salinity (‰)75Field, Laguna Madre, Texas (Simmons 1957)
Minimum Dissolved Oxygen (mg/l)1.4Field, Dauer et al. 1992, cited by Levin et al. 2009
Minimum Reproductive Temperature10Laboratory, Chesapeake Bay (Orth 1971)
Maximum Reproductive Temperature29Field, Patuxent River, Chesapeake Bay (Cory 1967). This polychaete probably tolerates ligher temperatures.
Minimum Reproductive Salinity5Field, Patuxent River, Chesapeake Bay (Cory 1967). This was the approximate lower limit for settlement
Minimum Duration918 C (Anger et al. 1986) Development is probably much shorter at higher temperatures.
Maximum Duration6010 C (Orth 1971)
Minimum Length (mm)2Smallest mature male, Brazil. Females were mature at 5.5 mm (Radashevsky 2005).
Maximum Length (mm)32Radashevsky 2005
Broad Temperature RangeNoneCold temperate-Tropical
Broad Salinity RangeNoneOligohaline-Hyperhaline

General Impacts

Economic Impacts

On the East of North America, Polydora cornuta is widely regarded as a pest in Easter Oyster (Crassostrea gigas) culture. It can develop dense colonies on the outside of the shells of oysters and mussels, smothering them with mud from their tubes. However, it is not capable of boring into shells, and is frequently confused with the specialized shell-borers P. ciliata and P. websteri (Galtsoff 1964; White and Wilson 1996; Sato-Okoshi 2000; Surugiru 2012). The extent to which P. cornuta affects oyster culture elsewhere in the world is unknown. However, P. cornuta was found consuming clam and oyster larvae in a hatchery in Newport, Oregon (Breese and Phibbs 1972).

Ecological Impacts

Polydora cornuta is widely recognized as a 'pollution indicator' species, known to rapidly increase in eutrophic estuaries, and estuaries receiving toxic pollutants, both in its native range and in areas where it has been introduced. This worm's tolerance of environmental extremes of temperature, salinity, and oxygen, as well as its feeding and habitat flexibility have enabled it to colonize harbors where other species have declined due to natural or anthropogenic stresses (Wolff 1973; Anger 1977; Rice and Simon 1980; Oviatt et al. 1987; Tagliapietra et al. 1998). Polydora cornuta occurs both in soft-bottom communities and fouling communities on marinas, docks, shells, ship hulls, etc. (Graham and Gay 1945; Wolff 1973; Bagaveeva and Zvyagintsev 2000), and can alternate between deposit-feeding from the sediment surface, or suspension-feeding on phytoplankton and particles in the water column (Dauer et al. 2005; Hentshel and Herrick 2005). Consequently, P. cornuta is a strong competitor in eutrophic habitats, and can become overwhelmingly dominant in sewage-polluted habitats (Anger 1977; Oviatt et al. 1987).

Competition- Competitive impacts of invading populations of Polydora cornuta are difficult to evaluate, because this polychaete tends to dominate in strongly disturbed and eutrophic environments, in which many polychaetes do poorly. However, its rapid growth, high fecundity, hypoxic tolerance, and trophic flexibility enable it to outnumber most other polychaetes (Wolff 1973; Anger 1977; Oviatt et al. 1987). Evidence of P. cornuta's competitive impacts is based partly on rapid colonization of polluted areas in the Black Sea, Sea of Marmara, Aegean Sea, and the Seto Inland Sea, Japan (Yamada et al. 2001; Losovskaya and Zolotarev 2003; Dagli and Ergen 2008; Karhan et al. 2008; Cinar et al. 2012), but also on eutrophication experiments, where it can reach extraordinary densities (e.g., Oviatt et al. 1987).

Herbivory- Polydora cornuta, reaches high abundances and becomes the dominant suspension-feeder in eutrophic areas, both as adults, with their palps extended in the water column (Dauer et al. 2005; Hentschel and Herrick 2005; Shimeta et al. 2005), and as planktonic larvae (Watras et al. 1985; Hanson 1999; Yamada et al. 2001). However, we have not found estimates of its grazing rates.

Regional Impacts

NEP-IVPuget Sound to Northern CaliforniaEconomic ImpactFisheries
IIn a Newport, Oregon shellfish hatchery, in 1970, Polydora cornuta was found to be feeding on larvae of Pacific Oysters (Crassostrea gigas and 'Manila Clams' (Venerupis philippinarum), and causing significant losses (Breese and Phipps 1972).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P023 _CDA_P023 (San Louis Rey-Escondido) 2001 Non-native Established
P062 _CDA_P062 (Calleguas) 2000 Non-native Established
P010 Tijuana Estuary 2000 Non-native Established
P027 _CDA_P027 (Aliso-San Onofre) 2000 Non-native Established
P020 San Diego Bay 1994 Non-native Established
P040 Newport Bay 1994 Non-native Established
P130 Humboldt Bay 1992 Non-native Established
P080 Monterey Bay 1977 Non-native Established
P065 _CDA_P065 (Santa Barbara Channel) 1971 Non-native Established
P095 _CDA_P095 (Tomales-Drakes Bay) 1969 Non-native Established
P061 _CDA_P061 (Los Angeles) 1969 Non-native Established
P058 _CDA_P058 (San Pedro Channel Islands) 1963 Non-native Established
P070 Morro Bay 1960 Non-native Established
P050 San Pedro Bay 1952 Non-native Established
NEP-IV Puget Sound to Northern California 1950 Non-native Established
P060 Santa Monica Bay 1941 Non-native Established
NEP-VI Pt. Conception to Southern Baja California 1938 Non-native Established
P030 Mission Bay 1938 Non-native Established
P090 San Francisco Bay 1933 Non-native Established
P100 Drakes Estero 1933 Non-native Established
NEP-V Northern California to Mid Channel Islands 1933 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
755892 ISS 2000-2002 Survey Data 2000 2000-09-14 Port Hueneme Infaunal 25 Non-native 34.1482 -119.2038
755893 ISS 2000-2002 Survey Data 2000 2000-10-25 Monterey Harbor Epifaunal 01 Non-native 36.6032 -121.8920
755894 ISS 2000-2002 Survey Data 2000 2000-10-25 Monterey Harbor Epifaunal 04 Non-native 36.6039 -121.8896
755895 ISS 2000-2002 Survey Data 2001 2001-08-14 Oceanside Epifaunal 05 Non-native 33.2058 -117.3898
755896 Introduced Species Study 2003 2003-08-26 Ballena Island Non-native 37.7661 -122.2824
755897 Introduced Species Study 2003 2003-08-26 Ballena Island 2 Non-native 37.7665 -122.2851
755898 Introduced Species Study 2003 2003-08-26 BerkeleyHarbor Non-native 37.8666 -122.3176
755899 Introduced Species Study 2003 2003-08-27 Loch Lomond 2 Non-native 37.9717 -122.4811
755900 Introduced Species Study 2005 2005-04-05 Point Dume Non-native 34.0087 -118.7938
755901 Introduced Species Study 2005 2005-06-07 Aquatic Park Non-native 37.8080 -122.4216
755902 Introduced Species Study 2005 2005-06-07 Oakland Inner Harbor - Small marinas Non-native 37.7847 -122.2669
755903 Introduced Species Study 2005 2005-06-08 Crown Beach Non-native 37.7603 -122.2737
755904 Introduced Species Study 2005 2005-06-08 Sea Plane Lagoon Non-native 37.7761 -122.2998
755905 Introduced Species Study 2005 2005-06-09 McNears Beach Non-native 37.9962 -122.4556
755906 Introduced Species Study 2005 2005-06-09 Paradise Area Non-native 37.9062 -122.4768
755907 Introduced Species Study 2005 2005-06-09 Richardson Bay Non-native 37.8588 -122.4798
755908 Introduced Species Study 2005 2005-06-10 Hayward Landing Non-native 37.6447 -122.1543
755909 Introduced Species Study 2005 2005-06-10 Toll Plaza Non-native 37.8266 -122.3166
755910 Introduced Species Study 2005 2005-07-06 Coyote Point Non-native 37.5920 -122.3210
755911 Introduced Species Study 2005 2005-07-07 Tiburon Non-native 37.8883 -122.4445
755912 Introduced Species Study 2005 2005-07-08 Point Richmond Non-native 37.9212 -122.3871
755913 Introduced Species Study 2005 2005-07-08 Richmond Marina Non-native 37.9137 -122.3504
755914 Introduced Species Study 2005 2005-08-19 Ayala Cove Non-native 37.8680 -122.4350
755915 Introduced Species Study 2005 2005-08-25 China Basin Non-native 37.7780 -122.3881
755916 Introduced Species Study 2005 2005-08-25 Ferry Terminal Pier Non-native 37.7945 -122.3917
755917 Introduced Species Study 2005 2005-08-25 Potrero Point Non-native 37.7521 -122.3790
755918 Introduced Species Study 2005 2005-09-07 Dumbarton Bridge Non-native 37.5070 -122.1168
755919 Introduced Species Study 2005 2005-09-07 Redwood Creek - Marina Non-native 37.5021 -122.2130
755920 Introduced Species Study 2005 2005-09-07 Redwood Creek - Shipping Non-native 37.5120 -122.2109
755921 Introduced Species Study 2005 2005-09-08 Cruise Ship Pier Non-native 37.8085 -122.4060
755922 Introduced Species Study 2005 2005-09-08 Pier 39 Non-native 37.8108 -122.4086
755923 Introduced Species Study 2005 2005-09-08 Saint Francis Yacht Harbor Non-native 37.8066 -122.4463
755924 Introduced Species Study 2005 2005-09-08 Treasure Island Non-native 37.8149 -122.3702
755925 Introduced Species Study 2005 2005-09-09 Coyote Point Marina Non-native 37.5905 -122.3177
755926 Introduced Species Study 2005 2005-09-09 San Mateo Bridge Non-native 37.5806 -122.2543
755927 Introduced Species Study 2005 2005-09-09 Sea Plane Harbor Non-native 37.6349 -122.3848
755928 Introduced Species Study 2005 2005-10-04 Berkeley Flats/Berkeley Pier Non-native 37.8600 -122.3256
755929 Introduced Species Study 2005 2005-10-04 Berkeley Marina Non-native 37.8676 -122.3172
755930 Introduced Species Study 2005 2005-10-06 Chevron Pier Non-native 37.9228 -122.4105
755931 Introduced Species Study 2005 2005-10-06 Point Richmond Piers Non-native 37.9085 -122.3913
755932 Introduced Species Study 2005 2005-10-06 Richmond Marina Non-native 37.9137 -122.3504
755933 Introduced Species Study 2005 2005-10-06 Santa Fe Channel - Back Non-native 37.9207 -122.3684
755934 Introduced Species Study 2005 2005-10-07 Benicia Waterfront Non-native 38.0401 -122.1385
755935 Introduced Species Study 2005 2005-10-19 Hercules Wharf Non-native 38.0231 -122.2928
755936 Introduced Species Study 2005 2005-10-19 Mare Island Strait - Navy Non-native 38.1015 -122.2695
755937 Introduced Species Study 2005 2005-10-19 Rodeo Marina Non-native 38.0394 -122.2717
755938 Introduced Species Study 2005 2005-10-20 Loch Lomond Marina Area Non-native 37.9720 -122.4832
755939 Introduced Species Study 2005 2005-10-20 Petaluma River Turning Basin Non-native 38.2344 -122.6354
755940 Introduced Species Study 2005 2005-10-20 Point San Pablo Yacht Harbor Non-native 37.9643 -122.4185
755941 Introduced Species Study 2005 2005-10-20 Port Sonoma/Petaluma R. Non-native 38.1157 -122.5026
755942 Introduced Species Study 2005 2005-10-21 Ayala Cove Non-native 37.8680 -122.4350
755943 Introduced Species Study 2005 2005-10-21 Corinthian Marina Non-native 37.8726 -122.4563
755944 Introduced Species Study 2005 2005-10-21 Paradise Cay Non-native 37.9146 -122.4776
755945 Introduced Species Study 2005 2005-10-21 Richardson Bay Non-native 37.8588 -122.4798
755946 Introduced Species Study 2005 2005-10-21 Romberg Tiburon Center Non-native 37.8906 -122.4458
755947 Introduced Species Study 2005 2005-11-14 Cal Maritime Academy/Vallejo Non-native 38.0661 -122.2299
755948 Introduced Species Study 2005 2005-11-15 China Camp Non-native 38.0025 -122.4617
755949 Introduced Species Study 2006 2006-08-11 Slip In A Harbor Non-native 38.3295 -123.0565
755950 Introduced Species Study 2006 2006-08-22 Fuel Depot Non-native 33.7440 -118.2358
755951 Introduced Species Study 2006 2006-08-25 Northern Corner of Harbor Non-native 33.9830 -118.4465
755952 Introduced Species Study 2006 2006-08-25 NW Corner of Harbor Non-native 33.9830 -118.4564
755953 Introduced Species Study 2006 2006-09-14 Back Bay Marina Non-native 33.6194 -117.8933
755954 Introduced Species Study 2006 2006-11-01 The Steelhead/MBARI Bridge Non-native 36.8005 -121.7877
755955 Introduced Species Study 2010 2010-05-31 Dumbarton Bridge Non-native 37.5070 -122.1168
755956 Introduced Species Study 2010 2010-05-31 Railroad Bridge Non-native 37.4602 -121.9750
755957 Introduced Species Study 2010 2010-05-31 Redwood Creek - Marina Non-native 37.5021 -122.2130
755958 Introduced Species Study 2010 2010-05-31 Redwood Creek - Shipping Non-native 37.5120 -122.2109
755959 Introduced Species Study 2010 2010-06-01 Coyote Point Marina Non-native 37.5905 -122.3177
755960 Introduced Species Study 2010 2010-06-01 Sea Plane Harbor Non-native 37.6349 -122.3848
755961 Introduced Species Study 2010 2010-06-02 Ballena Bay Non-native 37.7661 -122.2834
755962 Introduced Species Study 2010 2010-06-02 Coast Guard Island Non-native 37.7812 -122.2457
755963 Introduced Species Study 2010 2010-06-02 Port of Oakland Office Non-native 37.7954 -122.2804
755964 Introduced Species Study 2010 2010-06-03 Berkeley Flats/Berkeley Pier Non-native 37.8600 -122.3256
755965 Introduced Species Study 2010 2010-06-03 Berkeley Marina Non-native 37.8676 -122.3172
755966 Introduced Species Study 2010 2010-06-03 Treasure Island Non-native 37.8149 -122.3702
755967 Introduced Species Study 2010 2010-06-12 China Camp Non-native 38.0025 -122.4617
755968 Introduced Species Study 2010 2010-06-12 McNears Beach Non-native 37.9962 -122.4556
755969 Introduced Species Study 2010 2010-06-13 Coyote Point Non-native 37.5920 -122.3210
755970 Introduced Species Study 2010 2010-06-13 Hayward Landing Non-native 37.6447 -122.1543
755971 Introduced Species Study 2010 2010-06-28 Chevron Pier Non-native 37.9228 -122.4105
755972 Introduced Species Study 2010 2010-06-28 Point Richmond Piers Non-native 37.9085 -122.3913
755973 Introduced Species Study 2010 2010-06-28 Santa Fe Channel - Back Non-native 37.9207 -122.3684
755974 Introduced Species Study 2010 2010-06-28 Santa Fe Channel - Front Non-native 37.9101 -122.3644
755975 Introduced Species Study 2010 2010-06-30 Hercules Wharf Non-native 38.0231 -122.2928
755976 Introduced Species Study 2010 2010-06-30 Mare Island Strait - Marina Non-native 38.1051 -122.2667
755977 Introduced Species Study 2010 2010-06-30 Rodeo Marina Non-native 38.0394 -122.2717
755978 Introduced Species Study 2010 2010-07-01 Loch Lomond Marina Area Non-native 37.9720 -122.4832
755979 Introduced Species Study 2010 2010-07-01 Richardson Bay Non-native 37.8588 -122.4798
755980 Introduced Species Study 2010 2010-07-12 Cruise Ship Pier Non-native 37.8085 -122.4060
755981 Introduced Species Study 2010 2010-07-12 Pier 45 Non-native 37.8111 -122.4196
755982 Introduced Species Study 2010 2010-07-12 Potrero Point Non-native 37.7521 -122.3790
755983 Introduced Species Study 2010 2010-07-12 Saint Francis Yacht Harbor Non-native 37.8066 -122.4463
755984 Introduced Species Study 2010 2010-07-13 Ayala Cove Non-native 37.8680 -122.4350
755985 Introduced Species Study 2010 2010-07-13 Port Sonoma/Petaluma R. Non-native 38.1157 -122.5026
755986 Introduced Species Study 2010 2010-07-14 Paradise Cay Non-native 37.9146 -122.4776
755987 Introduced Species Study 2010 2010-07-14 Point San Pablo Yacht Harbor Non-native 37.9643 -122.4185
755988 Introduced Species Study 2010 2010-07-15 San Pablo Bay Pumphouse Non-native 38.0446 -122.4326
755989 Introduced Species Study 2010 2010-07-29 Mare Island Strait - Navy Non-native 38.1015 -122.2695
755990 Introduced Species Study 2010 2010-07-29 San Mateo Bridge Non-native 37.5806 -122.2543
755991 Introduced Species Study 2011 2011-04-08 Wharf 4 Non-native 34.1500 -119.2100
755992 Introduced Species Study 2011 2011-04-19 Back Bay Marina Non-native 33.6194 -117.8933
755993 Introduced Species Study 2011 2011-04-19 Balboa Coves Non-native 33.6213 -117.9364
755994 Introduced Species Study 2011 2011-04-19 Newport Bay Harbor Entrance Non-native 33.5974 -117.8798
755995 Introduced Species Study 2011 2011-04-19 Pump-A-Head Dock Non-native 33.7026 -118.0542
755996 Introduced Species Study 2011 2011-04-20 Fuel Depot Non-native 33.7440 -118.2358
755997 Introduced Species Study 2011 2011-04-22 Marina del Rey Harbor Entrance Non-native 33.9702 -118.4496
755998 Introduced Species Study 2011 2011-04-22 NW Corner of Harbor Non-native 33.9830 -118.4564
755999 Introduced Species Study 2011 2011-05-05 Middle Harbor Yacht Slip Non-native 33.2106 -117.3960
756000 Introduced Species Study 2011 2011-05-05 Oceanside Commercial Fishing Dock Non-native 33.2057 -117.3897
756001 Introduced Species Study 2011 2011-06-03 NE Corner of Bay Non-native 38.3340 -123.0511
756002 Introduced Species Study 2011 2011-06-29 Eureka Boat Launch Non-native 40.8040 -124.1766
759001 Allan Hancock Foundation Collection record, cited in Carlton 1979 1933 Drakes Estero, Point Reyes National Seashore Non-native 38.0567 -122.9394
759002 Hartman 1936c; Hartman 1969 1933 San Francisco Bay Non-native 37.8494 -122.3681
759003 Hartman 1936c, 1941 1936 Lake Merritt Non-native 37.8025 -122.2578
759004 Hartman 1936c 1936 Oakland Estuary Non-native 37.7866 -122.2654
759005 Hartman 1936c 1936 Berkeley Non-native 37.8897 -122.3293
759006 Allan Hancock Foundation Collection record, cited in Carlton 1979; Hartman 1941 1938 Mission Bay Non-native 32.7791 -117.2288
759007 Allan Hancock Foundation Collection record, cited in Carlton 1979 1941 Playa del Rey Lagoon Non-native 33.9614 -118.4515
759008 Graham and Gay 1945; Light 1977 1941 Fruitvale Avenue Bridge Non-native 37.7689 -122.2296
759009 Light 1977 1951 Wilson Point, San Pablo Bay Non-native 38.0129 -122.3165
759010 Filice 1954; Light 1977 1951 Off Castro Creek, San Pablo Bay Non-native 37.9700 -122.4044
759011 Reish and Winter 1954 1952 1952-07-29 Alamitos Bay, Station 5 (middle of channel at end of 67th Place) Non-native 33.7488 -118.1186
759012 Reish and Winter 1954 1952 1952-07-30 Alamitos Bay, Station 19 (middle of channel at end of Division Street) Non-native 33.7562 -118.1304
759013 Reish and Winter 1954 1952 1952-07-30 Los Cerritos Channel, Station 26 (at confluence of north and south forks) Non-native 33.7649 -118.1123
759014 Reish and Winter 1954 1952 1952-07-30 North Fork, Los Cerritos Channel, Station 27 (about 1,200 yards E of Pacific Coast Highway) Non-native 33.7671 -118.1047
759015 Reish and Winter 1954 1952 1952-07-30 South Fork, Los Cerritos Channel, Station 28 (about 600 feet E of Pacific Coast Highway) Non-native 33.7639 -118.1110
759016 Reish and Winter 1954 1952 1952-07-30 South Fork, Los Cerritos Channel, Station 30 (near terminus, about 700 feet E of Pacific Coast Highway) Non-native 33.7634 -118.1053
759017 Reish and Winter 1954 1952 1952-07-31 Colorado Lagoon, Station 32 (immediately above tidal gates) Non-native 33.7702 -118.1314
759018 Jones 1961; Hopkins 1986 1955 Point Richmond, Station A Non-native 37.9211 -122.3881
759019 Jones 1961; Hopkins 1986 1955 Point Richmond, Station B Non-native 37.9114 -122.3889
759020 Jones 1961; Hopkins 1986 1955 Point Richmond, Station C Non-native 37.9036 -122.3853
759021 Jones 1961; Hopkins 1986 1955 Point Richmond, Station D Non-native 37.9058 -122.3850
759022 Reish and Barnard 1967 1960 1960-08-25 Morro Bay Non-native 35.3500 -120.8500
759023 Reish 1964d 1963 1963-08-29 Catalina Harbor, Santa Catalina Island Non-native 33.4303 -118.5075
759024 Crippen and Reish 1969 1967 Los Angeles Harbor, East Basin, Boat Moorage 53 (Station LA 54) Non-native 33.7662 -118.2503
759025 Crippen and Reish 1969 1967 Los Angeles Harbor, in inner reaches of Slip 1 near Pier 160 (Station LA 39) Non-native 33.7628 -118.2662
759026 Crippen and Reish 1969 1967 Fleitz Brothers Yacht Anchorage (Station LA 7) Non-native 33.7203 -118.2758
759027 Crippen and Reish 1969 1967 Norm's Landing, Boat Moorage 1 (Station LA 26) Non-native 33.7345 -118.2777
759028 Light 1969 1967 Bolinas Lagoon Non-native 37.9189 -122.6816
759029 MacGinitie and MacGinitie 1969, cited in Carlton 1979 1969 Mugu Lagoon Non-native 34.1016 -119.0999
759030 Light 1977 1969 Off Standard Oil [Chevron] Refinery, Richmond Non-native 37.9309 -122.4049
759031 Kauwling and Reish 1975 1970 Huntington Harbour, N of Warner Avenue bridge (Station 7) Non-native 33.7123 -118.0607
759032 Kauwling and Reish 1975 1970 Huntington Harbour, slips in SE corner (Station 8) Non-native 33.7128 -118.0541
759033 Kauwling and Reish 1975 1970 Huntington Harbour, slips nearest Harbour Lane and Heil Avenue (Station 9) Non-native 33.7220 -118.0570
759034 Kauwling and Reish 1975 1971 Anaheim Bay, in channel inshore of Pacific Coast Highway bridge (Station 1) Non-native 33.7320 -118.0840
759035 Kauwling and Reish 1975 1971 Huntington Harbour, N of Warner Avenue bridge (Station 7) Non-native 33.7123 -118.0607
759036 Kauwling and Reish 1975 1971 Huntington Harbour, slips nearest Harbour Lane and Heil Avenue (Station 9) Non-native 33.7220 -118.0570
759037 Kauwling and Reish 1975 1971 Huntington Harbour, near slips adjacent ot Bonaire Circle (Station 10) Non-native 33.7277 -118.0595
759038 Reish et al. 1975 1975 Anaheim Bay, in channel inshore of Pacific Coast Highway bridge (Station 1) Non-native 33.7320 -118.0840
759039 J. Nybakken, pers. comm. 1977, in Carlton 1979 1977 Elkhorn Slough General Location Non-native 36.8086 -121.7856
759040 Barnhart et al. 1992 1992 Humboldt Bay Non-native 40.7864 -124.1922
759041 Ranasinghe et al. 2005 1998 Marina del Rey Non-native 33.9759 -118.4482
759042 Ranasinghe et al. 2005 1998 Los Angeles/Long Beach Harbor Non-native 33.7632 -118.2526
759043 Ranasinghe et al. 2005 1998 Newport Bay Non-native 33.6092 -117.9067
759044 Ranasinghe et al. 2005 1998 San Diego Bay Non-native 32.6717 -117.1439
759045 MEC Analytical Systems, Inc. et al. 2002 (Los Angeles/Long Beach Baseline Study of 2000) 2000 Los Angeles/Long Beach Harbor Complex Non-native 33.7632 -118.2526
759046 Boyd et al. 2002 2002 Klopp Lake Non-native 40.8553 -124.0919
759047 Boyd et al. 2002 2002 Southport Landing Non-native 40.6952 -124.2494
759048 Boyd et al. 2002 2002 Upper Elk River Non-native 40.7557 -124.1881
759049 Maloney et al. 2007 2005 2005-06-28 Shelter Island Marina Non-native 32.7180 -117.2255
759050 Maloney et al. 2007 2005 2005-06-29 Bulk Carrier Terminal Non-native 32.6969 -117.1526
759051 Heiman and Micheli 2010; Hobbs et al. 2015 2002 Elkhorn Slough (Upper) Non-native 36.8547 -121.7600
759052 Carpelan 1957 1951 Alviso Salt Evaporation Ponds 1-3 Non-native 37.4424 -122.0701
759053 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1974 1974-06-05 Forbidden Anchorage Zone off Hunter's Point Non-native 37.7375 -122.3417
759054 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1973 1973-09-19 Redwood City Harbor Entrance Channel Non-native 37.5508 -122.1944
759055 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1974 1974-06-06 Redwood City Harbor Entrance Channel Non-native 37.5508 -122.1944
759056 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1973 1973-12-03 Redwood City Harbor Entrance Channel Non-native 37.5508 -122.1944
759057 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1973 1973-09-20 Oakland Inner Harbor Non-native 37.7961 -122.3178
759058 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1973 1973-03-21 Oakland Inner Harbor Non-native 37.7961 -122.3178
759059 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1974 1974-06-05 Oakland Inner Harbor Non-native 37.7961 -122.3178
759060 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1974 1974-03-04 Oakland Inner Harbor Non-native 37.7961 -122.3178
759061 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1973 1973-12-04 Oakland Inner Harbor Non-native 37.7961 -122.3178
759062 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1973 1973-12-03 Main Shipping Channel (E of Foster City; South San Francisco Bay) Non-native 37.5681 -122.2092
759063 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1974 1974-03-05 Main Shipping Channel (E of Foster City; South San Francisco Bay) Non-native 37.5681 -122.2092
759064 Light 1977; California Academy of Sciences, Invertebrate Zoology Collections Database 1973 1973-09-19 Main Shipping Channel (E of Foster City; South San Francisco Bay) Non-native 37.5681 -122.2092
767918 Ruiz et al., 2015 2011 2011-09-20 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7947 -122.2822
768019 Ruiz et al., 2015 2012 2012-08-28 San Francisco Marina, San Francisco Bay, CA, California, USA Non-native 37.8071 -122.4341
768038 Ruiz et al., 2015 2012 2012-08-27 Port of San Francisco Pier 31, San Francisco Bay, CA, California, USA Non-native 37.8078 -122.4060
768080 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Non-native 37.6633 -122.3817
768104 Ruiz et al., 2015 2012 2012-08-29 Coyote Point Marina, San Francisco Bay, CA, California, USA Non-native 37.5877 -122.3174
768152 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9736 -122.4802
768171 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Non-native 37.7987 -122.3228
768210 Ruiz et al., 2015 2012 2012-08-31 Glen Cove Marina, San Francisco Bay, CA, California, USA Non-native 38.0663 -122.2130
768232 Ruiz et al., 2015 2012 2012-09-13 San Leandro Marina, San Francisco Bay, CA, California, USA Non-native 37.6962 -122.1919
768247 Ruiz et al., 2015 2012 2012-09-12 Emeryville, San Francisco Bay, CA, California, USA Non-native 37.8396 -122.3133
768272 Ruiz et al., 2015 2013 2013-08-15 Ballena Isle Marina, San Francisco Bay, CA, California, USA Non-native 37.7656 -122.2858
768335 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9723 -122.4829
768376 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.5024 -122.2134
768539 Ruiz et al., 2021b 2017 2017-07-24 Alamitos Bay Marina, San Pedro Bay, California, USA Non-native 33.7487 -118.1136
770780 Ruiz et al., 2021a 2017 2017-09-14 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.0667 -122.2132
770832 Ruiz et al., 2021a 2017 2017-09-28 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5898 -122.3165
770842 Ruiz et al., 2021a 2017 2017-09-28 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5898 -122.3165
770848 Ruiz et al., 2021a 2017 2017-09-28 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5898 -122.3165
770857 Ruiz et al., 2021a 2017 2017-09-28 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5898 -122.3165
770862 Ruiz et al., 2021a 2017 2017-09-28 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5898 -122.3165
770872 Ruiz et al., 2021a 2017 2017-09-26 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.5869 -122.3237
770889 Ruiz et al., 2021a 2017 2017-09-26 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.5869 -122.3237
770902 Ruiz et al., 2021a 2017 2017-09-26 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.5869 -122.3237
770938 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770946 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770955 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770962 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770971 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770980 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770991 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771004 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771014 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771024 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771046 Ruiz et al., 2021a 2017 2017-09-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9129 -122.3494
771106 Ruiz et al., 2021a 2017 2017-09-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9129 -122.3494
771130 Ruiz et al., 2021a 2017 2017-09-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9129 -122.3494
771139 Ruiz et al., 2021a 2017 2017-09-27 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7837 -122.2640
771149 Ruiz et al., 2021a 2017 2017-09-27 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7837 -122.2640
771242 Ruiz et al., 2021a 2017 2017-09-18 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9721 -122.4818
771246 Ruiz et al., 2021a 2017 2017-09-18 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9721 -122.4818
771252 Ruiz et al., 2021a 2017 2017-09-18 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9721 -122.4818
771258 Ruiz et al., 2021a 2017 2017-09-18 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9721 -122.4818
771272 Ruiz et al., 2021a 2017 2017-09-18 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9721 -122.4818
771277 Ruiz et al., 2021a 2017 2017-09-18 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9721 -122.4818
771327 Ruiz et al., 2021a 2017 2017-09-29 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8612 -122.4851
771346 Ruiz et al., 2021a 2017 2017-09-21 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6977 -122.1912
771356 Ruiz et al., 2021a 2017 2017-09-21 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6977 -122.1912
771370 Ruiz et al., 2021a 2017 2017-09-21 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6977 -122.1912
771375 Ruiz et al., 2021a 2017 2017-09-21 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6977 -122.1912
771401 Ruiz et al., 2021a 2017 2017-09-22 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7658 -122.2850
771416 Ruiz et al., 2021a 2017 2017-09-22 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7658 -122.2850
771421 Ruiz et al., 2021a 2017 2017-09-22 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7658 -122.2850
771429 Ruiz et al., 2021a 2017 2017-09-22 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7658 -122.2850
771448 Ruiz et al., 2021a 2017 2017-09-22 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7658 -122.2850
771474 Ruiz et al., 2021a 2017 2017-09-12 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8063 -122.4403
771488 Ruiz et al., 2021a 2017 2017-09-12 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8063 -122.4403
771529 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771537 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771542 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771549 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771553 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771557 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771561 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771566 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771571 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771579 Ruiz et al., 2021a 2017 2017-09-20 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.5048 -122.2574
771635 Ruiz et al., 2021a 2018 2018-09-12 Benicia Marina, San Francisco Bay, California, USA Non-native 38.0453 -122.1539
771646 Ruiz et al., 2021a 2018 2018-09-12 Benicia Marina, San Francisco Bay, California, USA Non-native 38.0453 -122.1539
771663 Ruiz et al., 2021a 2018 2018-09-12 Benicia Marina, San Francisco Bay, California, USA Non-native 38.0453 -122.1539
771683 Ruiz et al., 2021a 2018 2018-09-12 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.0675 -122.2142
771690 Ruiz et al., 2021a 2018 2018-09-12 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.0675 -122.2142
771704 Ruiz et al., 2021a 2018 2018-09-12 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.0675 -122.2142
771745 Ruiz et al., 2021a 2018 2018-09-19 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.8047 -122.2573
771754 Ruiz et al., 2021a 2018 2018-09-19 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.8047 -122.2573
771762 Ruiz et al., 2021a 2018 2018-09-19 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.8047 -122.2573
771776 Ruiz et al., 2021a 2018 2018-09-19 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.8047 -122.2573
771809 Ruiz et al., 2021a 2018 2018-09-17 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8065 -122.4412
771831 Ruiz et al., 2021a 2018 2018-09-17 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8065 -122.4412
771850 Ruiz et al., 2021a 2018 2018-09-17 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8065 -122.4412
771873 Ruiz et al., 2021a 2018 2018-09-17 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8065 -122.4412
771886 Ruiz et al., 2021a 2018 2018-09-17 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8065 -122.4412
771904 Ruiz et al., 2021a 2018 2018-09-24 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7654 -122.2845
771915 Ruiz et al., 2021a 2018 2018-09-24 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7654 -122.2845
771993 Ruiz et al., 2021a 2018 2018-09-24 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7654 -122.2845
772012 Ruiz et al., 2021a 2018 2018-09-21 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7843 -122.2648
772197 Ruiz et al., 2021a 2018 2018-09-20 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6981 -122.1907
772204 Ruiz et al., 2021a 2018 2018-09-20 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6981 -122.1907
772225 Ruiz et al., 2021a 2018 2018-09-20 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6981 -122.1907
772235 Ruiz et al., 2021a 2018 2018-09-20 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6981 -122.1907
772244 Ruiz et al., 2021a 2018 2018-09-20 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6981 -122.1907
772256 Ruiz et al., 2021a 2018 2018-09-20 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6981 -122.1907
772265 Ruiz et al., 2021a 2018 2018-09-20 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6981 -122.1907
772277 Ruiz et al., 2021a 2018 2018-09-18 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9139 -122.3461
772352 Ruiz et al., 2021a 2018 2018-09-18 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9139 -122.3461
772381 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772392 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772405 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772412 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772430 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772452 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772459 Ruiz et al., 2021a 2018 2018-09-28 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4818
772466 Ruiz et al., 2021a 2018 2018-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8609 -122.4855
772472 Ruiz et al., 2021a 2018 2018-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8609 -122.4855
772508 Ruiz et al., 2021a 2018 2018-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8609 -122.4855
772517 Ruiz et al., 2021a 2018 2018-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8609 -122.4855
772531 Ruiz et al., 2021a 2018 2018-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8609 -122.4855
772649 Ruiz et al., 2021a 2018 2018-09-27 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6634 -122.3823
772669 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772679 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772732 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
774941 Ruiz et al., 2022 2014 2014-09-08 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8078 -122.4354
774942 Ruiz et al., 2022 2014 2014-09-08 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8078 -122.4354
774943 Ruiz et al., 2022 2014 2014-09-09 San Leandro Marina, San Francisco Bay, California, USA Non-native 37.6978 -122.1928
774944 Ruiz et al., 2022 2014 2014-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8612 -122.4851
774945 Ruiz et al., 2022 2014 2014-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8612 -122.4851
774946 Ruiz et al., 2022 2014 2014-09-19 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8612 -122.4851
774947 Ruiz et al., 2022 2014 2014-09-16 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5895 -122.3160
774948 Ruiz et al., 2022 2014 2014-09-17 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6639 -122.3758
774949 Ruiz et al., 2022 2014 2014-09-17 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6639 -122.3758
774950 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
774951 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
774952 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
774953 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
774954 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
774955 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
774956 Ruiz et al., 2022 2014 2014-09-04 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.0674 -122.2131
774957 Ruiz et al., 2022 2014 2014-09-04 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.0674 -122.2131
774958 Ruiz et al., 2022 2014 2014-09-15 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4829
774959 Ruiz et al., 2022 2014 2014-09-15 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9723 -122.4829
774960 Ruiz et al., 2022 2015 2015-09-24 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7839 -122.2641
774961 Ruiz et al., 2022 2015 2015-09-15 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8066 -122.4331
774962 Ruiz et al., 2022 2015 2015-09-15 San Francisco Marina, San Francisco Bay, California, USA Non-native 37.8066 -122.4331
774963 Ruiz et al., 2022 2015 2015-09-21 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5022 -122.2129
774964 Ruiz et al., 2022 2015 2015-09-21 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5022 -122.2129
774965 Ruiz et al., 2022 2015 2015-09-17 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7679 -122.2863
774966 Ruiz et al., 2022 2015 2015-09-10 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.4096 -122.1247
774967 Ruiz et al., 2022 2015 2015-09-10 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.4096 -122.1247
774968 Ruiz et al., 2022 2015 2015-09-10 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.4096 -122.1247
774969 Ruiz et al., 2022 2015 2015-09-10 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.4096 -122.1247
774970 Ruiz et al., 2022 2015 2015-09-10 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.4096 -122.1247
774971 Ruiz et al., 2022 2015 2015-09-10 Glen Cove Marina, San Francisco Bay, California, USA Non-native 38.4096 -122.1247
774972 Ruiz et al., 2022 2015 2015-09-23 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6630 -122.3798
774973 Ruiz et al., 2022 2015 2015-09-23 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6630 -122.3798
774974 Ruiz et al., 2022 2015 2015-09-23 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6630 -122.3798
774975 Ruiz et al., 2022 2015 2015-09-23 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6630 -122.3798
774976 Ruiz et al., 2022 2015 2015-09-22 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5887 -122.3164
774977 Ruiz et al., 2022 2015 2015-09-22 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5887 -122.3164
774978 Ruiz et al., 2022 2015 2015-09-18 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9731 -122.4827
774979 Ruiz et al., 2022 2015 2015-09-08 Benicia Marina, San Francisco Bay, California, USA Non-native 38.0458 -122.1553
774980 Ruiz et al., 2022 2015 2015-09-08 Benicia Marina, San Francisco Bay, California, USA Non-native 38.0458 -122.1553
774981 Ruiz et al., 2022 2015 2015-09-08 Benicia Marina, San Francisco Bay, California, USA Non-native 38.0458 -122.1553
774982 Ruiz et al., 2022 2016 2016-09-08 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9727 -122.4816
774983 Ruiz et al., 2022 2016 2016-09-08 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9727 -122.4816
774984 Ruiz et al., 2022 2016 2016-09-08 Loch Lomond Marina, San Francisco Bay, California, USA Non-native 37.9727 -122.4816
774985 Ruiz et al., 2022 2016 2016-09-23 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8606 -122.4853
774986 Ruiz et al., 2022 2016 2016-09-23 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8606 -122.4853
774987 Ruiz et al., 2022 2016 2016-09-23 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8606 -122.4853
774988 Ruiz et al., 2022 2016 2016-09-23 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8606 -122.4853
774989 Ruiz et al., 2022 2016 2016-09-23 Sausalito Marine Harbor, San Francisco Bay, California, USA Non-native 37.8606 -122.4853
774990 Ruiz et al., 2022 2016 2016-09-21 Oyster Point Marina, San Francisco Bay, California, USA Non-native 37.6633 -122.3759
774991 Ruiz et al., 2022 2016 2016-09-22 Coyote Point Marina, San Francisco Bay, California, USA Non-native 37.5898 -122.3165
774992 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
774993 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
774994 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
774995 Ruiz et al., 2022 2016 2016-09-13 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7679 -122.2863
774996 Ruiz et al., 2022 2016 2016-09-27 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7831 -122.2630
774997 Ruiz et al., 2022 2016 2016-09-27 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7831 -122.2630

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