Styela plicata

Invasion History

First Non-native North American Tidal Record: 1877
First Non-native West Coast Tidal Record: 1915
First Non-native East/Gulf Coast Tidal Record: 1877

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General Invasion History:

The precise origin of Styela plicata is unknown. The type specimen was described from a ship in the Delaware River, Philadelphia, Pennsylvania in 1823 (Van Name 1912). It was apparently well established south of Cape Hatteras by the late 19th and early 20th centuries before regular collecting began. It is believed to have been introduced to the Mediterranean, probably centuries ago (Monniot, in Food and Agricultural Organization 2000). James T. Carlton considers the species introduced to the Northwest Atlantic, based on the general diversity of this genus in the Pacific (Carlton and Rucklelshaus 1997; Carlton, pers. comm.).  Genetic analysis by de Barros et al. (2009) suggested the Northwest Pacific as the probable native region. However, another study (Pineda et al. 2011) found the highest genetic diversity in the Western Atlantic, although the authors could not definitely identify a native region. The cladistic history of the genus Styela does support a Northwest Pacific ancestry of S. plicata, since it is distinct both from an Eastern Atlantic clade of warm-shallow water Styela, and cold deep-water forms (e.g. S. rustica (Carlton et al., in prep.)). Pineda et al's (2011) genetic analysis is consistent with historical data that suggests that transport of S. plicata began very early, and was very frequent, blurring genetic distinctions among native and introduced populations.

North American Invasion History:

Invasion History on the West Coast:

In the Northeast Pacific, Styela plicata was first collected in San Diego Bay in 1915 and ranges north to Santa Barbara, California (Lambert and Lambert 1998; Nichols et al. 2023). To the south, it has been reported from Baja California, Mexico, being collected in Ensenada in 2000 (Lambert and Lambert 2003) and Bahia San Quintin in 2005 (Rodriguez and Ibarra-Obando 2008). 

Invasion History on the East Coast:

The type specimen for Styela plicata was described from a ship in the Delaware River, Philadelphia, Pennsylvania in 1823 (Van Name 1912; 1945). However, because this specimen was reported from a vessel hull and not as present in the surrounding environment, we do not consider this record to be its date of introduction.  It was also collected in 1880 on Blackfish Bank, off Charleston, South Carolina (USNM 6960, U.S. National Museum of Natural History 2007). The long-established range of S. plicata along the Western Atlantic is from Cape Lookout, North Carolina (USNM 14790, U.S. National Museum of Natural History 2007) to Padre Island, Texas (USNM 14424, U.S. National Museum of Natural History 2007), and south through the Caribbean to Venezuela (Van Name 1945; USNM 14481, U.S. National Museum of Natural History 2007). Recently, however, S. plicata has been collected north of Cape Hatteras, in Lynnhaven Bay, Virginia Beach, at the mouth of Chesapeake Bay (Ruiz et al., unpublished data), in Chincoteague Bay, Virginia (in 1999, O'Beirn et al. 2004, on oyster-culture floats), and in the Mystic River, Long Island Sound (one specimen in 2001, but not yet established in Long Island Sound as of 2011; J.T. Carlton personal communication).  Howver, by 2023, thousands of individuals ranging from juveniles to adults were found attached to many different substrates in the Mystic River Estuary in Mystic, Connecticut. The population is being monitored to determine if it has survived the winter of 2022-2023. (James T. Carlton, perosnal communication 2023)..

Invasion History on the Gulf Coast:

A specimen of Styela plicata was collected from Padre Island, Texas (USNM 14424, U.S. National Museum of Natural History 2007) and Cedar Key, Florida in 1877 (USNM 940, U.S. National Museum of Natural History 2007). This tunicate is established and abundant on the Gulf Coast from Florida through Texas (Van Name 1921; Van Name 1945; Ruiz et al., unpublished data).

Invasion History Elsewhere in the World:

The long-established range of Styela plicata along the Western Atlantic is from Cape Lookout, North Carolina (USNM 14790, U.S. National Museum of Natural History 2007) to Padre Island, Texas (USNM 14424, U.S. National Museum of Natural History 2007), and south through the Caribbean to Venezuela (Van Name 1945; USNM 14481, U.S. National Museum of Natural History 2007). In 1884, S. plicata was reported from St. Thomas, US Virgin Islands/Caribbean Sea (USNM 6916, U.S. National Museum of Natural History 2007), and from Bermuda in 1882 (USNM 2769, U.S. National Museum of Natural History 2007) suggesting that it was established and widespread in warm waters of the Western Atlantic by the late 19th century.

Styela plicata was collected from Panama in 1973 (USNM 19744, US National Museum of Natural History 2007). Nearly a century before, in 1883, it was collected from Rio de Janeiro state, Brazil (Trautstedt 1883, cited by da Rocha and Kremer 2005), It ranges south to Uruguay (Orensanz et al. 2002), but in Bahia, northern Brazil, it was known only from 'one individual' (da Rocha and Kremer 2005).

Styela plicata is considered introduced to the Mediterranean: ['This species is not native to the Mediterranean, but was introduced centuries ago. It is present in all warm-temperate and tropical regions, especially in zones of human activity.'] (C. F. Monniot, in Food and Agricultural Organization 2000). It was collected from Naples (Traustedt 1877, cited by Kott 1985) and Trieste, Italy (Heller 1877, cited by Kott 1998). Outside the Mediterranean, it has been collected from Dakar, Senegal (Monniot 1969).

In the Northwest Pacific, S. plicata ranges from Hong Kong (Huang 2001) to Mutsu Bay, at the north end of Honshu, Japan (Oka 1935, cited by Nishikawa 1991). Within the Indian Ocean, it has been collected from Vizhinjam, India, on the Arabian Sea (Abdul and Sivakumar 2007), Somalia (in 1964, USNM 18297, U.S. National Museum of Natural History 2007), and the Gulf of Suez (in 1927, Monniot 2002). These scattered records could represent introductions.

In the Southwest Pacific, S. plicata was first collected at Port Jackson, near Sydney, Australia (Heller 1878, cited by Kott 1985). Its range runs from the mouth of the Calliope River, Queensland to Port Phillip Bay (in 1963, Millar 1966, cited by Keough and Ross 1999), and west to Cockburn Sound and the Perth area (Hartmeyer and Michaelsen 1928, cited by Kott 1985). Within this range, it is strongly associated with harbors and artificial structures (Kott 1985; Keough and Ross 1999). A genetic survey indicates high diversity in Australian populations, with significant genetic structure in more southern latitudes, but no structure in tropical latitudes (David et al. 2010). In New Zealand, it has been reported at several locations on the North Island, including Auckland Harbour and Hauraki Gulf (in 1957, Cranfield et al. 1998), Gulf Harbour Marina (Inglis et al. 2005), and Taranaki (Inglis et al. 2006).

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Description

Styela plicata is a solitary tunicate, variable in shape, but roughly oval. It is fixed to the substrate by the posterior end of its body, usually without roots or stalks. Its tunic is firm and thick, slightly translucent, with deep, irregular, longitudinal furrows, and horizontal creases that form large, irregularly rounded lumps.  Its total body length can reach 90 mm. The oral siphon is terminal, and the atrial siphon is a little behind it – both siphons are short, with square apertures with rounded humps on each side. The color of the tunic is whitish with brown or black stripes radiating from the siphons (Van Name 1945; Kott 1985; Nishikawa 1991; Gretchen Lambert, personal communication 2012).

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Taxonomy

Ecology

General:

Life History- A solitary tunicate is ovoid, elongate or vase-like in shape, with two openings or siphons. Most solitary tunicates attach to substrates by their side or base, but some attach with a conspicuous stalk. They are sessile filter feeders with two siphons, an oral and an atrial siphon. Water is pumped in through the oral siphon, where phytoplankton and detritus is filtered by the gills, and passed on mucus strings to the stomach and intestines. Waste is then expelled in the outgoing atrial water.

Solitary ascidians are hermaphroditic, meaning that both eggs and sperm are released to the atrial chamber. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but many species have a partial block to self-fertilization. Depending on the species, eggs may be externally or internally fertilized. In external fertilizers, eggs and sperm are released through the atrial siphon into the surrounding water column were fertilization takes place. In internal fertilizers, eggs are brooded and fertilized within the atrial chamber and then released into the water column upon hatching. Fertilized eggs hatch into a tadpole larva with a muscular tail, notochord, eyespots, and a set of adhesive papillae. The lecithotrophic (non-feeding, yolk-dependent) larva swims briefly before settlement. Swimming periods are usually less than a day and some larvae settle immediately after release, but the larval period can be longer at lower temperatures. Once settled, the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Barnes 1983).

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Marinas & Docks	None
General Habitat	Oyster Reef	None
General Habitat	Vessel Hull	None
General Habitat	Canals	None
General Habitat	Mangroves	None
General Habitat	Rocky	None
General Habitat	Salt-brackish marsh	None
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

Maximum Temperature (ºC)	30.2	Field, US East & West Coast marinas (Lord et al. 2015)
Minimum Salinity (‰)	17.5	Experimental- This was tested experimentally with animals at Santa Barbara CA. Only '75% seawater' (26 ppt), '50% seawater' (18ppt), and '110% seawater' (38.5 ppt) were used (Sims 1984).
Maximum Salinity (‰)	40	Field salinity (Shark Bay, Western Australia) (Wyatt et al. 2005)
Minimum Reproductive Temperature	18	Experimental- Lowest temperature tested for embryonic development and metamorphosis (Thiyagarajan and Qian 2003).
Maximum Reproductive Temperature	30	Experimental- Highest temperature tested for embryonic development and metamorphosis. Only 30% of larvae successfully completed development (Thiyagarajan and Qian 2003).
Minimum Reproductive Salinity	30	Experimental- Development was observed at 22, 26, 30, and 34 ppt. Embryonic development was unsuccessful at 22 and 26 ppt (Thiyagarajan and Qian 2003).
Maximum Reproductive Salinity	34	Experimental- Highest salinity tested (Thiyagarajan and Qian 2003).
Minimum Duration	0	Larvae can attach immediately after hatching (Thiyagarajan and Qian 2003).
Maximum Duration	2	Larvae prevented from settling (Thiyagarajan and Qian 2003).
Maximum Length (mm)	90	Van Name 1945
Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Economic Impacts

Fisheries: Styela plicata is known to foul cultured bivalves, interfering with their growth in Brazil, Hong Kong, Japan, and Spain (da Rocha et al. 2009).

Ecological Impacts

Competition: Styela plicata has been a dominant fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It dominates fouling communities from Ensenada, Mexico to Santa Barbara (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes from southern California harbors (Lambert and Lambert 1998). Styela plicata was one of the dominant fouling organisms on fouling plates at Beaufort, North Carolina. It invaded plates initially dominated by other species and created 'monopolies,' for up to four months (Sutherland and Karlson 1977). Settled juveniles of Styela plicata inhibited settlement of the native Microcosmus squamiger (from Brisbane, Australia), in a laboratory experiment (Ruis et al. 2009). The mechanism was not clear, but could involve competition for food or allelopathy (inhibitory chemicals).

Fisheries: Styela plicata is known to foul cultured bivalves, interfering with their growth in Brazil, Hong Kong, Japan, and Spain (da Rocha et al. 2009). On the positive side, S. plicata is extensively cultured on long lines in Korea and Japan (Lambert et al. 2016).

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Regional Impacts

NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Competition
Styela plicata has been the dominant marina fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It dominates fouling communities from Ensenada, Mexico to Santa Barbara (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P020	San Diego Bay		Ecological Impact	Competition
Styela plicata has been the dominant marina fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It produced large monospecific patches in several locations during serveral years in San Diego Bay (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P030	Mission Bay		Ecological Impact	Competition
Styela plicata has been the dominant marina fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It produced large monospecific patches in several locations during serval years in San Diego Bay (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P023	_CDA_P023 (San Louis Rey-Escondido)		Ecological Impact	Competition
Styela plicata has been the dominant marina fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It produced large monospecific patches in Oceanside Harbor (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P027	_CDA_P027 (Aliso-San Onofre)		Ecological Impact	Competition
It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P040	Newport Bay		Ecological Impact	Competition
It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P050	San Pedro Bay		Ecological Impact	Competition
It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P060	Santa Monica Bay		Ecological Impact	Competition
It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P062	_CDA_P062 (Calleguas)		Ecological Impact	Competition
It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P064	_CDA_P064 (Ventura)		Ecological Impact	Competition
It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
P065	_CDA_P065 (Santa Barbara Channel)		Ecological Impact	Competition
It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).
CA	California		Ecological Impact	Competition
Styela plicata has been the dominant marina fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It produced large monospecific patches in several locations during serveral years in San Diego Bay (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., Styela plicata has been the dominant marina fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It produced large monospecific patches in several locations during serval years in San Diego Bay (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., Styela plicata has been the dominant marina fouling organism in southern California harbors since 1960 (Lambert and Lambert 1998). It produced large monospecific patches in Oceanside Harbor (Lambert and Lambert 2003). It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998)., It and other introduced ascidians have replaced the native species Pyura haustor and Ascidia ceratodes in southern California harbors (Lambert and Lambert 1998).

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