Invasion History

First Non-native North American Tidal Record: 1914
First Non-native West Coast Tidal Record: 1914
First Non-native East/Gulf Coast Tidal Record: 1931

General Invasion History:

Amphibalanus amphitrite was described by Darwin (1854), using specimens collected from Portugal, the Mediterranean, West Africa, the West Indies, the Indo-Pacific, Australia, and New Zealand. However, because A. amphitrite is part of a complex of similar species, and prone to transport on ships' hulls, its native range is difficult to determine (Utinomi 1960; Henry and McLaughlin 1975). In the West Pacific and Indian Ocean, it may be native from Southeastern Africa to Southern China, but it is a recent introduction in Eastern (Panama-California), Northwestern (Korea-Japan-Russia, 1st record Tokyo Bay, 1950), and Southwestern reaches of the Pacific, including New Zealand (Cranfield et al. 1998) and possibly Southern Australia. Amphibalanus amphitrite invaded North American waters in the 20th century (Zullo 1966; Southward 1975; Carlton et al. 2011). It was absent on pre-Columbian oyster shells from the Indian River lagoon examined by Boudreaux et al. (2009). Darwin (1854) observed that it was 'extremely common on ship's bottoms' and it has been introduced over much of its present range, including most of the Atlantic Basin (cryptogenic in the Mediterranean) and the Eastern Pacific, including Hawaii, and the North American Coast, from Panama to San Francisco Bay, California (Zullo et al. 1972; Henry and McLaughlin 1975; Cohen and Carlton 1995; Carlton et al. 2011).

North American Invasion History:

Invasion History on the West Coast:

On the Pacific Coast of North and Central America, Amphibalanus amphitrite is abundant on the coast of Panama and ranges at least as far north as San Francisco Bay, California (Carlton et al. 2011). Its usual habitats are the sheltered waters of harbors. There is an apparent gap in its range between Santa Monica Bay and San Francisco Bay (Carlton 1979; Wasson et al. 2001; Cohen et al. 2002), where it was first collected in 1938. In San Francisco Bay, it is confined to the warmest parts of the estuary (Zullo et al. 1972). A remarkable invasion took place in the 1940s, when A. amphitrite was introduced to the Salton Sea, a salt lake created by a ruptured canal in a desert region of California, near the Colorado River. The barnacles were introduced from San Diego Bay to the Salton Sea with buoys used to mark seaplane landing areas and were very abundant by 1944 (Carlton 1979; Carlton et al. 2011). The adult barnacles have developed a distinctive morphology, and were previously recognized as a subspecies (Henry and McLaughlin 1975). However, rearing experiments and genetic studies indicate that these changes are environmental (Flowerdew 1985; Carlton et al. 2011). The Salton Sea barnacles have lost pigmentation in their cyprid larvae, which is an evolutionary change from their marine ancestors (Raimondi 1992).

Invasion History on the East Coast:

Darwin (1854) reported Amphibalanus amphitrite as occurring in the Caribbean, but the identity of his specimens is uncertain. The earliest verified record from the Northwest Atlantic is a specimen collected from the Dry Tortugas, Florida, in the Gulf of Mexico in 1931 (U.S. National Museum of Natural History collections). However, this could represent an isolated specimen collected from a ship. It is plausible that large-scale introductions of this barnacle to the Northwest Atlantic occurred during and after World War II (Carlton et al. 2011). The Striped Barnacle was common at Beaufort, North Carolina by 1955 (Costlow and Bookhout 1958; Zullo 1966; Henry and McLaughlin 1975) at Biscayne, Florida by 1960 (Moore and Frue 1974), and in Bermuda by 1952 (Henry and McLaughlin 1975; USNM 195758 U.S. National Museum of Natural History 2011). It may have been overlooked at many locations by confusion and lumping with the native A. venustus, then known as 'Balanus amphitrite niveus', and may have been present in Biscayne Bay as early as the 1940s (Moore and Frue 1974). Later collections have shown it to be widely distributed on the East Coast south of Cape Hatteras (Mook 1983; Zullo 1966; Zullo and Lang 1978; Prezant et al. 2002; Crickenberger and Sotka 2009).

It was collected in 1967 in Virginia Beach (Henry and McLaughlin 1975), but was reported as 'scarce' in lower Chesapeake Bay (Van Engel et al. 1972). It now occurs regularly in the lower Bay on fouling plates (2001-2004, Ruiz et al., unpublished data) and extends south to Biscayne Bay, Florida (Henry and McLaughlin 1975; Ruiz et al., unpublished data). This barnacle may occur sporadically further north, but apparently cannot survive the winter. Zullo (1966) collected four specimens from pilings near Hyannis, Massachusetts, on Cape Cod, and maintained them in a running seawater system. The specimens died when temperatures dropped to 1.5ºC (Zullo 1966). This barnacle has regularly settled in Long Island Sound in small numbers (Carlton et al. 2011), but its winter survival is unknown. The range of this barnacle is expected to move north, with the warming climate (Carlton et al. 2011).

Invasion History on the Gulf Coast:

On the Gulf Coast, as noted above, Amphibalanus amphitrite was collected from the Dry Tortugas in 1931 (this record may have been from a ship), but was not reported from other locations until the 1950s (Carlton et al. 2011). Early locations include Tampa Bay (in 1952), Boca Ciega Bay (in 1955), St. Marks Bay (in 1959), St. Andrews Bay (in 1964) (all in Florida), and Pass Christian, Mississippi Sound (Henry and McLaughlin 1975; Wells 1966; Zullo 1966). Amphibalanus amphitrite is present along the western coast of Texas, from Corpus Christi Bay (in 1971) to Port Mansfield and south into Mexico (in 1965, Vergera and Vercruz) (Henry and McLaughlin 1975; Gittings 1985; Gittings et al. 1986). Gittings et al. (1986) note a gap in distribution from Louisiana to Reddish Bay, Texas, where they suggest A. reticulatus may have excluded A. amphitrite in the more turbid waters of the north-central Gulf coast.

Invasion History in Hawaii:

Amphibalanus amphitrite was first collected in the Hawaiian Islands in Honolulu in 1902 (USNM 32517, Henry and McLaughlin 1975, U.S. National Museum of Natural History 2002). Currently, it is found throughout the main Hawaiian Islands (Carlton and Eldredge 2009), although in some locations, it is now mostly replaced by A. reticulatus and Chthamalus proteus. However, it is still common in low salinity settings (Zabin 2009).

Invasion History Elsewhere in the World:

The status of Amphibalanus amphitrite in the Eastern Atlantic is complicated by its identification in an excavation of a 2000 year-old Carthaginian naval base in Tunis, which leads us to treat this barnacle as cryptogenic in the Mediterranean Sea (Southward 1998), although Carlton et al. (2011) suggest it may have been introduced to Tunis even at this early date. It appears to be a recent invader in other parts of the Atlantic, however, where it is confined to tropical and warm-temperate waters. Specimens were collected in Atlantic France as early as 1914 (Goulletquer et al. 2002), in the Azores in 1920 (Southward 1998), and in the Bay of Biscay in 1934 (Bishop 1950). In Northern Europe, this species is confined to warmer waters, and is abundant in thermal effluents. It was first reported from Le Havre, France in 1928, and has subsequently been found in England, Ireland, the Netherlands, Belgium, and Germany (Bishop 1950; Bishop et al. 1957; Wolff 1999; Kerckhof and Cattrijsse 2001; Kerckhof et al. 2007; Wiegemann 2008). Many of these occurrences are likely to be ephemeral, dependent on effluents, or vulnerable to severe cold weather. There are few records of Amphibalanus amphitrite from the Atlantic Coast of Africa, a likely result of undersampling. It was collected from Madeira in 2005 (Wirtz et al. 2006), but was not abundant. Henry and McLaughlin (1975) reported specimens from Walvis Bay, Namibia, collected in 1969.

In the Caribbean and Southwest Atlantic, the timing of Amphibalanus amphitrite's invasion is uncertain because of taxonomic confusion. Darwin (1854) referred to this species occurring in 'the West Indies', but the earliest dated, verified record which we have is from Curacao in 1957 (Henry and McLaughlin 1975). According to available records, this barnacle is widespread, but scattered, in the western Caribbean. Southward (1975) and Bacon (1976) found it rare and local in Jamaica, largely confined to ships and man-made structures in Bonaire and Trinidad, but locally common on mangroves on Isla Margarita, Venezuela. Amphibalanus amphitrite was common at the eastern entrance to the Panama Canal in 1974 (Spivey 1976). In the Southwest Atlantic, the first definitive record of A. amphitrite was from 1940, in Rio de Janeiro, Brazil (de Oliveira 1941, cited by Carlton et al. 2011). At present, this barnacle ranges from Guimaraes, Maranhao state, Brazil (2⁰S, Young 1994) to Mar del Plata, Argentina (38⁰S, Orensanz et al. 2002), with many occurrences on the Brazilian coast (Young 1994).

In the tropical East Pacific, Amphibalanus amphitrite was found in 1946 in the Gulf of California (Henry and McLaughlin, 1975; USNM 173807, US. National Museum of Natural History 2011), in 1960 in Acapulco (Henry and McLaughlin 1975), and 1974, in Balboa, Panama, at the western end of the Panama Canal (Spivey 1976). An isolated occurrence was found in La Punta, Lima, Peru, in 1999, on 'artificial rock' in a harbor (Carlton et al. 2011).

Amphibalanus amphitrite has a wide presumed native/cryptogenic range in the Indo-West Pacific, but is apparently introduced in the Northwest Pacific (northern China, South Korea, Japan, and Pacific Russia), the Southwest Pacific (southern Australia, New Zealand), and central Pacific Islands (Fiji, American Samoa). The first records in Japanese waters were from Kyushu and southeastern Honshu in 1937 (Hiro 1937, cited by Utinomi 1960). It invaded Korean waters in the 1970s (Kim 1992), and by 1975 it occurred seasonally, during warm years, in the Golden Horn Bay, Vladivostok, Russia. Its winter survival here is dependent on thermal effluents (Zvyaginstsev and Korn 2003). In the Southwest Pacific, the boundary between A. amphitrite's native and introduced ranges is unclear. Keough and Ross (1999), Hewitt et al. (2004), and Huisman et al. (2008) consider it introduced to ports of southern Australia, but the date of first invasion is unknown, due in part to confusion with the very similar native species A. variegatus (Keough and Ross (1999). The first record from New Zealand was from Waitemata Harbour, near Auckland in 1960 (Cranfield et al. 1998). In Guam, A. amphitrite appeared confined to harbors, but is abundant as a fossil, and so is considered native (Paulay and Ross 2003). Further east, in Fiji (Foster 1974) and American Samoa (Coles et al. 2003), this barnacle was probably introduced by shipping.


Description

The shell of Amphibalanus amphitrite is usually conical or subcylindrical. The orifice is round or slightly toothed. Its width is usually more than 1/2 its height. The plates have wide longitudinal ribs (radii), narrowing to the tops of the shell plates. The plates are white, with longitudinal lavender or purple stripes. Inside the operculum, on the interior face of the scutum, the adductor ridge is moderately long and usually thick. The tergum has a blunt apex. The length of its spur is about 1/4 of the length of the basal margin, and the spur width is roughly 3/10 of the basal margin (Henry and McLaughlin 1975). The shell ranges up to 30.2 mm basal diameter (Henry and McLaughlin 1975), but adults typically range from 5.5 to 15 mm basal diameter (Shkedy et al. 1995; Shalla et al. 1995). This barnacle is characteristic of sheltered marine habitats, and tolerates some salinity variation. Larval development of A. amphitrite is described and illustrated by Costlow and Bookhout (1958), Lang (1979), and Zvagintsev and Korn (2003).

Amphibalanus amphitrite is a member of the Amphibalanus amphitrite species complex and can be confused with A. improvisus, A. eburneus, A. reticulatus, A. subalbidus, A. variegatus and other closely related species (Henry and McLaughlin 1975). Molecular analysis showed that the subspecies A. amphitrite saltonensis, described by Henry and McLaughlin for specimens from the Salton Sea and a California coast specimen, is identical to A. amphitrite amphitrite, invalidating the status of the subspecies (Flowerdew 1985). Salton Sea and Pacific Coast A. amphitrite show identical morphology when reared together, indicating that morphological differences are environmental, but the cyprid larvae of Salton Sea barnacles lack the green pigmentation of Pacific barnacles. This change in larval morphology has evolved since the Salton Sea population was established in the 1940s (Raimondi 1992).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Maxillopoda
Subclass:   Thecostraca
Infraclass:   Cirripedia
Superorder:   Thoracica
Order:   Sessilia
Suborder:   Balanomorpha
Superfamily:   Balanoidea
Family:   Balanidae
Genus:   Amphibalanus
Species:   amphitrite

Synonyms

Balanus amphitrite forma hawaiiensis (Broch, 1922)
Balanus amphitrite ssp. amphitrite (Darwin, 1854)
Balanus amphitrite ssp. franciscanus (Rogers, 1949)
Balanus amphitrite ssp. herzi (Rogers, 1949)
Balanus amphitrite ssp. venustus (Sundra Raj, 1927)
Balanus amphitrite var. aeratus (Oliveira, 1941)
Balanus amphitrite var. cochinensis (Nilsson-Cantell, 1938)
Balanus amphitrite var. communis (Darwin, 1854)
Balanus amphitrite var. denticulata (Broch, 1927)
Balanus amphitrite var. fluminensis (Oliveira, 1941)
Balanus amphitrite ssp. Saltonensis (Rogers, 1949)

Potentially Misidentified Species

Amphibalanus improvisus
Historically treated as a variety of A. amphitrite, (var. pallidus, in part), and also misidentified as B. amphitrite by many workers (Henry and Laughlin 1975).

Amphibalanus reticulatus
Historically treated as a variety of A. amphitrite, (var. variegatus, var. tesselatus, var. cirratus) by Darwin (1854), and also misidentified as A. amphitrite by many workers (Henry and Laughlin 1975).

Amphibalanus subalbidus
Historically treated as a variety of A. amphitrite, (var. pallidus, in part), and also misidentified as A. amphitrite by many workers (Henry and Laughlin 1975).

Amphibalanus venustus
Historically treated as a variety of A. amphitrite, (var. venustus, var. niveus) by Darwin (1854), and also misidentified as A. amphitrite by many workers (Henry and Laughlin 1975).

Ecology

General:

Amphibalanus amphitrite, like many other barnacles, is hermaphroditic, but is capable of cross-fertilization. The fertilized eggs are brooded in the mantle cavity, sometimes for several months, and are released as nauplius larvae with three pairs of appendages (Barnes 1983). This barnacle produced 1,000 to 10,000 eggs per animal, generally increasing with body size (El-Komy and Kajihara 1991; Lee and O'Riordan 2014). The nauplii feed in the plankton and go through five successive molts, spending four to 18 days in the water column before molting into a non-feeding cypris stage, covered with a pair of chitinous shells (Anil et al. 1995). Cyprids swim, investigating suitable surfaces, and then settle, secreting a shell and molting into the first juvenile barnacle stages. Juvenile and adult barnacles are filter feeders, sweeping the water with their long bristled appendages to gather phytoplankton, zooplankton, and detritus.

Amphibalanus amphitrite is typically found in the intertidal and shallow subtidal regions of sheltered marine waters, particularly harbors, and man-made structures, but is rare on open rocky coasts (Henry and McLaughlin 1975). It grows on a wide range of hard surfaces, including docks, ship hulls, logs, mangroves, rocks, oysters, and other shellfish (Utinomi 1960; Henry and McLaughlin 1975; Southward 1975; Gitting 1985). It is sensitive to cold temperatures, and in the northern limits of its range, it is most abundant in the warmest habitats, including thermal effluents (Bishop 1950; Zullo 1966; Raymont 1976). This barnacle prefers marine salinities (30-40 ppt), but tolerates a range from 10-52 ppt (McPherson et al. 1984; Anil et al. 1995; Cohen 2005).

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatMangrovesNone
General HabitatVessel HullNone
General HabitatCoral reefNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Salinity RangeHyperhaline40+ PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)1.5Min. temp. comes from observation of 2 individuals in running seawater, Woods Hole MA (Zullo 1966).
Maximum Temperature (ºC)40Temperature to 50% coma, heated 1 C per minute (Ritz and Foster 1968)
Minimum Salinity (‰)10From lab experiments- Anil et al. 1995
Maximum Salinity (‰)52Field observations, pond, Alviso CA, South San Francisco Bay (Cohen 2005).
Minimum Reproductive Temperature12Im cultures at ambient temperature, Japan (El-Komy and Kajihara 1991)
Minimum Duration6Larval Period, 30 C, laboratory (Anil et al. 1995)
Maximum Duration17At 15 C, 10 PSU ((Anil et al. 1995))
Minimum Length (mm)5.5Minimum adult size (Shalla et al. 1995; Shkedy et al. 1995)
Maximum Length (mm)15Maximum adult size (Shalla et al. 1995; Shkedy et al. 1995)
Maximum Width (mm)30Maximum basal width (Henry and McLauglin 19750
Broad Temperature RangeNoneWarm-temperate-Tropical
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Economic Impacts 

Shipping- We have not found specific reports of economic impacts for Amphibalanus amphitrite in North American waters. However, A. amphitrite is one of the most abundant fouling barnacles in warmer harbors of the U.S. (Moore and Frue 1959; Carlton 1979), and worldwide (Zevina 1988; Jones 1992; Shkedy et al. 1995). It is a major contributor to fouling of ship and harbor structures. Amphibalanus amphitrite is a frequent test organism for various types of anti-fouling agents and treatments. Hull fouling by barnacles and other organisms has costly impacts for shipping lines and navies, greatly increasing fuel costs, decreasing maneuverability, and fouling internal seawater piping. Barnacles also greatly contribute to fouling of navigational buoys and coastal power station intakes (Haderlie 1984).

Fisheries- Amphibalanus amphitrite is a frequent fouling organism of cultured Pacific Oysters (Crassostrea gigas) in warmer waters (Grizel and Heral 1991; Grizel 1994).

Ecological Impacts 

Competition- Amphibalanus amphitrite, together with A. eburneus and A. improvisus, is a competitor in fouling communities in Beaufort, North Carolina. Amphibalanus spp. however, despite their high recruitment rate, were readily overgrown by other fouling organisms (Sutherland and Karlson 1977). In the Indian River Lagoon, the introduced Amphibalanus amphitrite and the native A. eburneus competed with the Eastern Oyster (Crassostrea virginica) for settlement sites, and also affected survival and growth of oysters by settling on their shells (Boudreaux et al. 2009). Amphibalanus amphitrite is a common organism in fouling communities worldwide. In the harbors of Yokohama and Tokyo, Japan, A. amphitrite is reported to have largely replaced the native A. reticulatus (Zvyagintsev and Korn 2003).

Habitat Change- In Tampa Bay, Amphibalanus amphitrite affected the composition of the fouling community, mainly by creating additional structure for the recruitment and colonization of motile species. Removing barnacle shells inhibited recruitment, while adding barnacle shells increased recruitment (Bros 1987).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P130 Humboldt Bay 2015 Non-native Unknown
NEP-IV Puget Sound to Northern California 2015 Non-native Unknown
P064 _CDA_P064 (Ventura) 2004 Non-native Established
P010 Tijuana Estuary 2003 Non-native Established
P030 Mission Bay 1977 Non-native Established
P060 Santa Monica Bay 1957 Non-native Established
P040 Newport Bay 1956 Non-native Established
NEP-V Northern California to Mid Channel Islands 1939 Non-native Established
P090 San Francisco Bay 1939 Non-native Established
P020 San Diego Bay 1927 Non-native Established
P022 _CDA_P022 (San Diego) 1919 Non-native Established
NEP-VI Pt. Conception to Southern Baja California 1914 Non-native Established
P050 San Pedro Bay 1914 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
697096 Introduced Species Study 2005 2005-08-25 Ferry Terminal Pier Non-native 37.7945 -122.3917
697423 Introduced Species Study 2011 2011-04-19 Newport Bay Harbor Entrance Non-native 33.5974 -117.8798
697468 Cohen et al. 2002 (So Cal Exotics RAS) 2000 2000-08-30 Pilot's Dock at Pier F Non-native 33.7472 -118.2156
697807 Cohen et al. 2002 (So Cal Exotics RAS) 2000 2000-08-29 Marina del Rey Non-native 33.9722 -118.4522
697817 Cohen et al. 2002 (So Cal Exotics RAS) 2000 2000-08-24 Watchorn Basin Non-native 33.7203 -118.2764
698008 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-24 Fruitvale Bridge, San Francisco Bay Non-native 37.7690 -122.2296
699100 Cohen et al. 2002 (So Cal Exotics RAS) 2000 2000-08-26 Chula Vista Boat Ramp Non-native 32.6211 -117.1031
699320 Introduced Species Study 2010 2010-07-15 San Pablo Bay Pumphouse Non-native 38.0446 -122.4326
699680 Cohen et al. 2002 (So Cal Exotics RAS) 2000 2000-08-30 Long Beach Downtown Marina Non-native 33.7581 -118.1894
700230 Introduced Species Study 2006 2006-10-10 The Tuna Club Non-native 33.3461 -118.3268
700624 Carlton 1979 1940 San Pedro Non-native 33.7385 -118.2760
700921 Newman 1967, cited in Carlton 1979 1967 South San Francisco Bay Non-native 37.5457 -122.1645
701016 Newman 1967, cited in Carlton 1979 1967 North San Francisco Bay Non-native 37.9586 -122.4646
701048 Cohen et al. 2002 (So Cal Exotics RAS) 2000 2000-08-31 Colorado Lagoon Non-native 33.7711 -118.1347
701212 Rodgers 1939, 1949, cited in Zullo et al. 1972 1938 San Francisco Bay Non-native 37.8494 -122.3681
701964 Introduced Species Study 2006 2006-08-25 Overnight Guest Dock Non-native 33.9761 -118.4461
702488 Zullo et al. 1972; Carlton 1979 1927 1927-12-09 San Diego Bay Non-native 32.6717 -117.1439
702774 Newman 1967, cited in Carlton 1979 1967 Central San Francisco Bay Non-native 37.8595 -122.3884
702876 Cohen et al. 2002 (So Cal Exotics RAS) 2000 2000-08-26 Fiddler's Cove Non-native 32.6519 -117.1494
702998 Introduced Species Study 2005 2005-08-25 China Basin Non-native 37.7780 -122.3881
703424 Zullo et al. 1972; Carlton 1979 1921 La Jolla Non-native 32.8479 -117.2776
703458 Introduced Species Study 2011 2011-04-19 Back Bay Marina Non-native 33.6194 -117.8933
703946 Henry and McLaughlin 1975; United States National Museum Collections Database 2018 1914 1914-09-02 Wilmington [Los Angeles] Harbor Non-native 33.7356 -118.2758
704064 Newman 1967, cited in Carlton 1979 1967 San Pablo Bay Non-native 38.0600 -122.3900
712510 de Rivera et al. 2005 2003 Tijuana Estuary, Tidal Linkage Non-native 32.5742 -117.1270
712514 de Rivera et al. 2005 2003 Sunroad Marina, San Diego Non-native 32.7263 -117.1922
712517 de Rivera et al. 2005 2003 Dana Marina, San Diego Non-native 32.7675 -117.2365
712523 Carlton 1979 1977 Colorado Lagoon Non-native 33.7711 -118.1347
712525 Henry and McLaughlin 1975; Carlton 1979 1961 1961-09-27 San Pedro Non-native 33.7333 -118.2000
712536 Rodgers 1939, 1949, cited in Zullo et al. 1972 1939 Aquatic Park, San Francisco Non-native 37.8087 -122.4242
757898 Carlton 1979 1929 Mare Island Naval Shipyard Non-native 38.1015 -122.2695
757899 Carlton 1979 1939 Wilmington [Los Angeles] Harbor Non-native 33.7356 -118.2758
757900 Graham and Gay 1945 1941 Fruitvale Bridge Non-native 37.7689 -122.2296
757901 Hartman 1941, cited in Carlton 1979 1941 Lake Merritt Non-native 37.8025 -122.2578
757902 WHOI 1952 1943 San Diego Non-native 32.6717 -117.1439
757903 Marchette 1953, cited in Carlton 1979 1952 Lake Merritt Non-native 37.8025 -122.2578
757904 Henry and McLaughlin 1975 1919 1919-04-12 La Jolla Non-native 32.8479 -117.2776
757905 Henry and McLaughlin 1975 1952 1952-05-23 Berkeley Aquatic Park Non-native 37.8567 -122.2992
757906 Henry and McLaughlin 1975 1955 1955-12-31 Cerritos Channel Non-native 33.7665 -118.2375
757907 Henry and McLaughlin 1975 1956 1956-11-03 Huntington Beach Non-native 33.6530 -118.0060
757908 Henry and McLaughlin 1975 1956 1956-11-03 Newport Bay Non-native 33.6092 -117.9067
757909 Henry and McLaughlin 1975 1957 1957-01-03 Venice Non-native 33.9767 -118.4709
757910 Henry and McLaughlin 1975 1975 Newport Bay Non-native 33.6092 -117.9067
757911 W.E. Newman pers. comm. 1977, in Carlton 1979 1977 Mission Bay Non-native 32.7791 -117.2288
757912 Reish et al. 1975 1975 Anaheim Bay Non-native 33.7333 -118.0894
757913 de Rivera et al. 2005 2003 Seaforth Marina, San Diego Non-native 32.7630 -117.2373
757914 de Rivera et al. 2005 2003 Model Marsh Non-native 32.5597 -117.1066
757915 de Rivera et al. 2005 2003 Oneonta Slough Non-native 32.5678 -117.1315
757916 de Rivera et al. 2005 2003 Chula Vista Marina, San Diego Non-native 32.6244 -117.1038
757917 de Rivera et al. 2005 2004 Ventura West Harbor Non-native 34.2464 -119.2611
757918 Rodgers 1949, cited in Carlton 1979 1939 Bayview Park, San Francisco Non-native 37.7087 -122.3881
757919 Carlton 1979 1960 Lake Merritt Non-native 37.8025 -122.2578
757920 Carlton 1979 1970 Lake Merritt Non-native 37.8025 -122.2578
767842 Ruiz et al., 2015 2011 2011-09-16 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9724 -122.4796
767867 Ruiz et al., 2015 2011 2011-09-13 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.8046 -122.3985
767916 Ruiz et al., 2015 2011 2011-09-20 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7947 -122.2822
768312 Ruiz et al., 2015 2013 2013-08-22 Jack London Square Marina, San Francisco Bay, CA, California, USA Non-native 37.7926 -122.2746
768329 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Non-native 37.9723 -122.4829
768371 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Non-native 37.5024 -122.2134
768508 Ruiz et al., 2021b 2017 2017-07-24 Alamitos Bay Marina, San Pedro Bay, California, USA Non-native 33.7487 -118.1136
768517 Ruiz et al., 2021b 2017 2017-07-24 Alamitos Bay Marina, San Pedro Bay, California, USA Non-native 33.7487 -118.1136
768570 Ruiz et al., 2021b 2017 2017-07-24 Alamitos Bay Marina, San Pedro Bay, California, USA Non-native 33.7487 -118.1136
768581 Ruiz et al., 2021b 2017 2017-07-24 Alamitos Bay Marina, San Pedro Bay, California, USA Non-native 33.7487 -118.1136
768623 Ruiz et al., 2021b 2017 2017-07-25 Cabrillo Marina, San Pedro Bay, California, USA Non-native 33.7198 -118.2773
768789 Ruiz et al., 2021b 2017 2017-07-30 Leeward Island Marina, San Pedro Bay, California, USA Non-native 33.7767 -118.2428
768841 Ruiz et al., 2021b 2017 2017-08-02 Shoreline Public Marina, San Pedro Bay, California, USA Non-native 33.7598 -118.1877
768868 Ruiz et al., 2021b 2017 2017-08-02 Shoreline Public Marina, San Pedro Bay, California, USA Non-native 33.7598 -118.1877
768877 Ruiz et al., 2021b 2017 2017-08-02 Shoreline Public Marina, San Pedro Bay, California, USA Non-native 33.7598 -118.1877
768902 Ruiz et al., 2021b 2017 2017-08-02 Shoreline Public Marina, San Pedro Bay, California, USA Non-native 33.7598 -118.1877
769017 Ruiz et al., 2021b 2017 2017-07-27 Peter's Landing Marina, San Pedro Bay, California, USA Non-native 33.7256 -118.0758
769028 Ruiz et al., 2021b 2017 2017-07-27 Peter's Landing Marina, San Pedro Bay, California, USA Non-native 33.7256 -118.0758
769091 Ruiz et al., 2021b 2017 2017-07-27 Peter's Landing Marina, San Pedro Bay, California, USA Non-native 33.7256 -118.0758
769246 Ruiz et al., 2021b 2017 2017-07-28 Port of LB Pier F, San Pedro Bay, California, USA Non-native 33.7473 -118.2131
769382 Ruiz et al., 2021b 2018 2018-08-06 Shoreline Public Marina, San Pedro Bay, California, USA Non-native 33.7586 -118.1856
769407 Ruiz et al., 2021b 2018 2018-08-10 Alamitos Bay Marina, San Pedro Bay, California, USA Non-native 33.7494 -118.1154
769776 Ruiz et al., 2021b 2018 2018-08-13 Cabrillo Marina, San Pedro Bay, California, USA Non-native 33.7219 -118.2779
769796 Ruiz et al., 2021b 2018 2018-08-13 Cabrillo Marina, San Pedro Bay, California, USA Non-native 33.7219 -118.2779
769814 Ruiz et al., 2021b 2018 2018-08-13 Cabrillo Marina, San Pedro Bay, California, USA Non-native 33.7219 -118.2779
769830 Ruiz et al., 2021b 2018 2018-08-13 Cabrillo Marina, San Pedro Bay, California, USA Non-native 33.7219 -118.2779
769991 Ruiz et al., 2021b 2018 2018-08-08 Port of LB Matson, San Pedro Bay, California, USA Non-native 33.7729 -118.2178
770447 Ruiz et al., 2021b 2017 2017-08-06 Harbor Patrol Public Dock, Newport Bay, California, USA Non-native 33.6023 -117.8835
770936 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770942 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770951 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770959 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770967 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770976 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
770987 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771000 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771011 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771020 Ruiz et al., 2021a 2017 2017-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2124
771096 Ruiz et al., 2021a 2017 2017-09-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9129 -122.3494
771127 Ruiz et al., 2021a 2017 2017-09-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, California, USA Non-native 37.9129 -122.3494
772660 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772668 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772678 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772696 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772706 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772720 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772740 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
772751 Ruiz et al., 2021a 2018 2018-09-25 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5023 -122.2114
775502 Ruiz et al., 2022 2015 2015-08-11 Coast Guard, Humboldt Bay, California, USA Non-native 40.7669 -124.2174
775503 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775504 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775505 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775506 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775507 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775508 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775509 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775510 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775511 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775512 Ruiz et al., 2022 2014 2014-09-11 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5024 -122.2134
775513 Ruiz et al., 2022 2014 2014-09-12 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7662 -122.2656
775514 Ruiz et al., 2022 2014 2014-09-12 Ballena Isle Marina, San Francisco Bay, California, USA Non-native 37.7662 -122.2656
775515 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775516 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775517 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775518 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775519 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775520 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775521 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775522 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775523 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775524 Ruiz et al., 2022 2014 2014-09-03 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0187 -121.8221
775525 Ruiz et al., 2022 2014 2014-09-18 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7835 -122.2630
775526 Ruiz et al., 2022 2014 2014-09-18 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7835 -122.2630
775527 Ruiz et al., 2022 2014 2014-09-18 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7835 -122.2630
775528 Ruiz et al., 2022 2014 2014-09-18 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7835 -122.2630
775529 Ruiz et al., 2022 2014 2014-09-18 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7835 -122.2630
775530 Ruiz et al., 2022 2014 2014-09-18 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7835 -122.2630
775531 Ruiz et al., 2022 2014 2014-09-18 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7835 -122.2630
775532 Ruiz et al., 2022 2015 2015-09-21 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5022 -122.2129
775533 Ruiz et al., 2022 2015 2015-09-21 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5022 -122.2129
775534 Ruiz et al., 2022 2015 2015-09-21 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5022 -122.2129
775535 Ruiz et al., 2022 2015 2015-09-21 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5022 -122.2129
775536 Ruiz et al., 2022 2015 2015-09-09 Pittsburg Marina, San Francisco Bay, California, USA Non-native 38.0354 -121.8822
775537 Ruiz et al., 2022 2015 2015-09-09 Pittsburg Marina, San Francisco Bay, California, USA Non-native 38.0354 -121.8822
775538 Ruiz et al., 2022 2015 2015-09-09 Pittsburg Marina, San Francisco Bay, California, USA Non-native 38.0354 -121.8822
775539 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775540 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775541 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775542 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775543 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775544 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775545 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775546 Ruiz et al., 2022 2015 2015-09-09 Antioch Marina, San Francisco Bay, California, USA Non-native 38.0204 -121.8212
775547 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
775548 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
775549 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
775550 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
775551 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138
775552 Ruiz et al., 2022 2016 2016-09-26 Redwood City Marina, San Francisco Bay, California, USA Non-native 37.5027 -122.2138

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