Amphibalanus amphitrite

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

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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.

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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).

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Taxonomy

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

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Mangroves	None
General Habitat	Vessel Hull	None
General Habitat	Coral reef	None
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Salinity Range	Hyperhaline	40+ PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	1.5	Min. temp. comes from observation of 2 individuals in running seawater, Woods Hole MA (Zullo 1966).
Maximum Temperature (ºC)	40	Temperature to 50% coma, heated 1 C per minute (Ritz and Foster 1968)
Minimum Salinity (‰)	10	From lab experiments- Anil et al. 1995
Maximum Salinity (‰)	52	Field observations, pond, Alviso CA, South San Francisco Bay (Cohen 2005).
Minimum Reproductive Temperature	12	Im cultures at ambient temperature, Japan (El-Komy and Kajihara 1991)
Minimum Duration	6	Larval Period, 30 C, laboratory (Anil et al. 1995)
Maximum Duration	17	At 15 C, 10 PSU ((Anil et al. 1995))
Minimum Length (mm)	5.5	Minimum adult size (Shalla et al. 1995; Shkedy et al. 1995)
Maximum Length (mm)	15	Maximum adult size (Shalla et al. 1995; Shkedy et al. 1995)
Maximum Width (mm)	30	Maximum basal width (Henry and McLauglin 19750
Broad Temperature Range	None	Warm-temperate-Tropical
Broad Salinity Range	None	Mesohaline-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).

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Regional Impacts

CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Economic Impact	Shipping/Boating
Amphibalanus amphitrite is probably an important hull-fouling organism over most of its range. At a commercial paint-testing station in Biscayne Bay, Miami, it comprised 12% of barnacles maturing on fouling panels (Moore and Frue 1959).
CAR-VII	Cape Hatteras to Mid-East Florida		Ecological Impact	Competition
Amphibalanus amphitrite, together with A. eburneus and A. improvisus, is a competitor in fouling communities in Beaufort, North Carolina. However, despite their high recruitment rate, Amphibalanus spp. were readily overgrown by other fouling organisms, except when present in very high densities (Sutherland and Karlson 1977).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Ecological Impact	Habitat Change
In Tampa Bay, A. 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).
G070	Tampa Bay		Ecological Impact	Competition
In Tampa Bay, A. amphitrite strongly 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).
NEA-IV	None		Economic Impact	Fisheries
Balanus amphitrite is a frequent fouling organism of cultured Pacific Oysters (Crassostrea gigas) in warmer waters (Grizel and Heral 1991; Grizel 1994).
NEA-V	None		Economic Impact	Fisheries
Amphibalanus amphitrite is a frequent fouling organism of cultured Pacific Oysters (Crassostrea gigas) in warmer waters (Grizel and Heral 1991; Grizel 1994).
NWP-3b	None		Ecological Impact	Competition
Amphibalanus amphitrite is a common organism in fouling communities worldwide. In the harbors of Yokohama and Tokyo, A. amphitrite is reported to have largely replaced the native A. reticulatus (Zvyagintsev 2003)
S190	Indian River		Ecological Impact	Competition
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. However, impacts did not differ between the native and non-native barnacle species (Boudreaux et al. 2009).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Ecological Impact	Competition
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).
S030	Bogue Sound		Ecological Impact	Competition
Amphibalanus amphitrite, together with A. eburneus and A. improvisus, is a competitor in fouling communities in Beaufort, North Carolina. However, despite their high recruitment rate, Amphibalanus spp. were readily overgrown by other fouling organisms (Sutherland and Karlson 1977).
S200	Biscayne Bay		Economic Impact	Shipping/Boating
At a commercial paint-testing station in Biscayne Bay, Miami, Amphibalanus amphitrite comprised 12% of barnacles maturing on fouling panels (Moore and Frue 1959).
NWP-2	None		Ecological Impact	Competition
Barnacles, dominated by A. amphitrite overgrew newly recruited corals on settling plates during winter periods of cool weather (14-16⁰C). Competition by barnacles may be one factor limiting corals at the northern limit of their range (Chui and Ang 2010).
NWP-3b	None		Economic Impact	Industry
Amphibalanus amphitrite is reported to foul 'sluice systems' in Japan (Chavanich et al. 2010).
NC	North Carolina		Ecological Impact	Competition
Amphibalanus amphitrite, together with A. eburneus and A. improvisus, is a competitor in fouling communities in Beaufort, North Carolina. However, despite their high recruitment rate, Amphibalanus spp. were readily overgrown by other fouling organisms (Sutherland and Karlson 1977).
FL	Florida		Ecological Impact	Competition
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. However, impacts did not differ between the native and non-native barnacle species (Boudreaux et al. 2009)., In Tampa Bay, A. amphitrite strongly 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).
FL	Florida		Economic Impact	Shipping/Boating
At a commercial paint-testing station in Biscayne Bay, Miami, Amphibalanus amphitrite comprised 12% of barnacles maturing on fouling panels (Moore and Frue 1959).

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