Amphibalanus amphitrite

Invasion History

First Non-native Panama (Pacific) Tidal Record: 1974
First Non-native Panama (Caribbean) Tidal Record: 1957

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Panama Invasion History:

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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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