Amathia verticillata

Invasion History

First Non-native North American Tidal Record: 1905
First Non-native West Coast Tidal Record: 1905
First Non-native East/Gulf Coast Tidal Record: 2005

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General Invasion History:

Amathia verticillata is an erect, soft-bodied bryozoan, first described from the Mediterranean Sea (Naples, Italy) and now widespread in tropical, subtropical, and warm-temperate waters around the world (Winston 1977; Gordon and Mawatari 1992) We consider it cryptogenic in the tropical-subtropical Atlantic from North Carolina and Bermuda to Santos, Brazil, (Osburn 1914; Marcus 1937; Osburn 1940; Maturo 1958; Winston 1982, but see Farrapeira, 2011) and the coast of west Africa (Senegal and Gana, Cook 1968), Pacific coast of Central America (Soule et al. 1980, cited by Carlton and Cohen 1995), Arabian Sea and Bay of Bengal waters of India (Robertson 1921, cited by Cohen and Carlton 1995); Karande (1968, cited by Winston 1977; Rao and Ganapati 1978), and southern China, in the South China Sea (Huang 2001). 
 
The origin of this bryozoan is unknown, but it is a documented invader in California (1905, Cohen and Carlton 1995), Hawaii (before 1921, Carlton and Eldredge 2007), New Zealand (Gordon and Mawatari 1992), Australia (before 1975, Keough and Ross 1999), Japan (early 1900s, Asakura 1992); South Korea (Je et al. 1988), the Azores (2008, Amat and Tempera 2009), Madeira (Wirtz and Canning-Clode 2009), and the Canary Islands (2010, Canning-Clode et al. 2013; Minchin 2012), and Taiwan (Minchin et al. 2016). Its larval period is brief (Hyman 1959), so that fouling and ballast water (as fragments of adult colonies or attached to bits of flotsam) are likely vectors for its introduction. A recent review documents the probable introduction of this bryozoan to the Mediterranean Sea, where it was first discovered in the early 19th century but is largely restricted to harbors and marinas (Marchini et al. 2015). Galil and Gevli (2014) argued for a Caribbean origin of A. verticillata, because of its strong association with corals, rocks, and seagrasses there, and the presence of a specialized nudibranch predator there (Okenia zoobotryon).

North American Invasion History:

Invasion History on the West Coast:

Robertson (1905, cited by Cohen and Carlton 1995) observed dense growths of Amathia verticillata in San Diego Bay, where it is still abundant in eelgrass beds (Williams 2007). It is found in Mazatlan, Mexico (Alvarez-Leon and Banta 1984), and the Gulf of California (in 1958, Soule 1963). The pattern of its spread along the California coast is unknown, because most of the records in smaller harbors and bays are quite recent. It was found in Mission Bay, La Jolla, and Newport Bay by 1963 (Soule 1963), and Los Angeles Harbor 'in recent years' (Soule et al. 1980, cited by Cohen and Carlton 1995). In 1993, it appeared in Redwood Creek, South San Francisco Bay (Cohen and Carlton 1995), but has not been seen since (James T. Carlton, personal communication 2013). In a 2001 survey, it was found in small harbors in southern California, including Oceanside Harbor, Channel Islands Harbor, and Avalon Harbor, on Catalina Island (Fairey et al. 2002).

Invasion History on the East Coast:

Amathia verticillata was collected in the Dry Tortugas, Florida, as early as 1908 (Osburn 1914), although it was earlier collected in Bermuda in 1876 (Maturo and Schopf, 1968). It has been treated as native, from North Carolina to Venezuela, by most writers (Osburn 1914; Osburn 1940; Weiss 1948; Maturo 1957; Winston 1982; Pederson and Peterson 2002). However, Carlton and Ruckelshaus (1997) and Gossett et al. (2004) list it as an introduction to Florida and the Gulf of Mexico. We consider it cryptogenic here. A colony found in 2005 at the restoration shipyard of Mystic Seaport, Connecticut, on Long Island Sound (James T. Carlton, personal communication), was an introduction by boat fouling. It was collected in the Mystic River again in 2010 (James T. Carlton, personal communication). It was collected in the Mystic River again in 2010 (James T. Carlton, personal communication). Additional occurrences have been seen from 2014 to 2016, from Wachapreague and Kiptopeake, Virginia and from New Bedford, Massachusetts. Establishment of this warm-water bryozoan in Long Island Sound is unlikely until further warming occurs.

Invasion History in Hawaii:

Robertson (1921) reported having received specimens of A. verticillata from Honolulu Harbor (Robertson 1921, cited by Carlton and Eldredge 2009). It was present in Pearl Harbor by 1921, and in Kaneohe Bay by 1939 (Carlton and Eldredge, 2009), and is also known from Maui (Coles et al. 2004). Recent reports have A. verticillata rapidly becoming abundant at Palmyra Island in the Line Islands, about 2000 km south of Hawaii (Knapp et al. 2011). 

Invasion History Elsewhere in the World:

Amathia verticillata is a recent introduction in the Indian Ocean, the South China Sea, and Japan, appearing in the early 1900s, and first collected in Tokyo Bay in 1963 (Asakura 1992; Jebakumar et al. 2017). This bryozoan was newly recorded on the south coast of South Korea, where it became a pest fouling cultured pearl oysters (Je et al. 1988). However, Otani (2006) lists it as a cryptogenic species in Japan. It was first collected in Auckland Harbor, New Zealand about 1960 and has since been found in several harbors on the North Island (Gordon and Mawatari 1992).  Amathia verticillata was collected in New South Wales before 1975 (Russ and Wake 1975, cited by Keough and Ross 1999) and in South and Western Australia before 1982 (Bock 1982, cited by Keough and Ross 1999; Huisman et al. 2008). 
 
Amathia verticillata was first reported from Europe in Cadiz, Spain, in 1807, and described from Naples in 1822 Marchini et al. (2015) has described this species as 'pseudo-indigenous', probably introduced very early, but still largely confined to harbors and marinas, and preferring artificial substrates (Marchini et al. 2015). Early Mediterranean records, from west to east include Malaga, Spain, 1823, Algiers, Algeria 1848, Livorno, Italy 1848, Naples, Italy 1822 and Alexandria, Egypt 1828 (Marchini et al. 2015). Locally in the Mediterranean, A. verticillata rapidly colonize marinas, and provide habitat for other introduced species (Marchini et al. 2015). A. verticillata is a recent invader in the Azores, where it appeared in marinas in 2008 on three of the islands (Amat and Tempera 2009), and Madeira, where it was first collected in May of 2009 (Wirtz and Canning-Clode 2009). 
 
In the Eastern Pacific, Amathia verticillata was first collected in Los Frailes Bay in the Gulf of California in 1958,), Baja California and at Mazatlán, Sinaloa (Soule 1963). This bryozoan was collected further south in Oaxaca in 2017 and 2018 (Humara-Gil, et al. 2019). It was first reported from the Pacific coast of South America in 2014, in the Salinas Yacht Club, Ecuador (Priscilla Martinez, personal communication, February 2015, cited by McCann et al. 2015).

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Description

Amathia verticillata is an erect, un-calcified bryozoan, with colonies made of irregularly thick transparent, branching or triple-quadruple branching stolons with zooids attached. Colonies can resemble ‘stringy, gelatinous noodles or sauerkraut up to 2 m long’ (Gordon and Mawatari 1992). The zooids are ovoid, about 380 X 580 µm in size, and occur in clusters on either side of the thick transparent stolon. The polypides have eight tentacles and large gizzards. The zooids brood whitish embryos (description from Winston 1982; Gordon and Mawatari 1992; Vieira et al. 2014). Colonies vary greatly in size, 400–700 mm in Brazil (Farrapeira et al. 2010) to 2200 mm (Amat et al. 2011). 

Recent molecular work has confirmed that Amathia verticillata is a single global species (Nascimento et al. 2021). A recent genetic analysis of the family Vesiculariidae supports a synonymy of the genera Amathia, Bowerbankia, and Zoobotryon (Waeschenbach et al. 2015). 

 

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Taxonomy

Ecology

General:

Life History- Amathia verticillata is an erect, uncalcified bryozoan, composed of many individual zooids. The zooids feed by extending the ciliated tentacles of the lophophore as a funnel, creating a current, and driving food particles into their mouths. The food is guided along the tentacles and through the pharynx by the cilia. Larger food particles can be moved or captured by flicking or contracting the tentacles. The zooids are hermaphroditic, and produce large yolky eggs, which hatch into lecithotrophic larvae, which are planktonic for short periods (less than 1 day). Larvae settle on a substrate and metamorphose into the first zooid of a colony, an ancestrula (Barnes 1983). 
 
Ecology- This bryozoan is found on wood, rock, vegetation (including eelgrass and mangroves), shells, and man-made structures, including vessel hulls (Winston 1982; Gordon and Mawatari 1992; Minchin 2012). In an investigation of chemical compounds affecting feeding on A. verticillata, a Brazilian population had alkaloids that deterred feeding, while Florida populations had variable resistance to feeding. Dos Santos et al. (2017) suggest that the success of this bryozoan is due to factors other than toxicity. 

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Grass Bed	None
General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Canals	None
General Habitat	Mangroves	None
General Habitat	Vessel Hull	None
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Salinity (‰)	15	Nair et al. 1992
Maximum Salinity (‰)	40	Field salinity (Shark Bay, Western Australia) (Wyatt et al. 2005)
Minimum Length (mm)	100	Colony length, Azores (Amat et al. 2009, on marina floats)
Maximum Height (mm)	2,200	Colony length, Azores (Amat et al. 2009, on marina floats)
Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Economic Impacts

Shipping/Boating- Amathia verticillata is known from ship fouling (Woods Hole Oceanographic Institution 1952) and is reported to cause fouling problems to boats and marinas in southern California (Johnson et al. 2006).

Fisheries- Amathia verticillata causes major problems by clogging shrimp-fishing gear in Galveston Bay, Texas (Gossett et al. 2004). It was reported as a pest species fouling cultured pearl oysters, Pinctada fuca, on the south coast of Korea (Je et al. 1988).

Ecological Impacts

Habitat Change- Amathia verticillata grows epiphytically on the leaves of eelgrass (Zostera marina), killing plants in San Diego Bay, which creates gaps in the seagrass canopy, leaving bare patches of sediment available for colonization by other algal species (Williams 2007).

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Regional Impacts

NEP-VI	Pt. Conception to Southern Baja California		Economic Impact	Shipping/Boating
Amathia verticillata is reported to cause fouling problems to boats and marinas in southern California (Johnson et al. 2006).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Competition
Amathia verticillata grows epiphytically on the leaves of eelgrass (Zostera marina) in San Diego Bay, killing them (Williams 2007).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Habitat Change
Amathia verticillata kills eelgrass plants in San Diego Bay, creating gaps in the seagrass canopy and bare patches of sediment, which are then colonized by adult plants (Williams 2007).
P020	San Diego Bay		Ecological Impact	Competition
Amathia verticillata grows epiphytically on the leaves of eelgrass (Zostera marina) in San Diego Bay, killing them (Williams 2007).
P020	San Diego Bay		Ecological Impact	Habitat Change
Amathia verticillata kills eelgrass plants in San Diego Bay, creating gaps in the seagrass canopy and bare patches of sediment, which are then colonized by adult plants (Williams 2007).
CA	California		Ecological Impact	Competition
Amathia verticillata grows epiphytically on the leaves of eelgrass (Zostera marina) in San Diego Bay, killing them (Williams 2007).
CA	California		Ecological Impact	Habitat Change
Amathia verticillata kills eelgrass plants in San Diego Bay, creating gaps in the seagrass canopy and bare patches of sediment, which are then colonized by adult plants (Williams 2007).

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