Styela canopus

Invasion History

First Non-native North American Tidal Record: 1852
First Non-native West Coast Tidal Record: 1972
First Non-native East/Gulf Coast Tidal Record: 1852

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General Invasion History:

Styela canopus (Rough Sea Squirt) is now widely distributed in temperate and tropical coastal waters of the world. It was described from the Red Sea in 1816 and was later reported from both sides of the North Atlantic, Ascension Island in the South Atlantic, the Northwest Pacific, the tropical Indo-Pacific, and temperate waters of Australia. (Kott 1998). In the Atlantic, it was first reported (as Cynthia partita) from Boston Harbor in 1852 (Stimpson 1852) and then from the English Channel (as Styela variabilis) in 1868. It was reported from the Mediterranean (as S. canopoides) in 1877 (Kott 1998). By the late 19th century, it was abundant and widespread on both sides of the Atlantic and was generally regarded as a native species. In the Northeast Pacific, where it is a recent introduction, S. canopus was first reported from San Diego Bay, California, in 1972. It has been collected north to Alamitos Bay but is rare outside San Diego Bay (Lambert and Lambert 1998). James Carlton considers S. canopus to be a native of the Indo-West Pacific (Carlton 2000 personal communication), introduced to the Atlantic (possibly a century or more before its description), a judgment which we have adopted here. Kott (1985) considers S. canopus to be the only species of Styela native to the Indo-West Pacific region.

North American Invasion History:

Invasion History on the West Coast:

On the west coast of North America, Styela canopus was first collected in San Diego, California in 1972. To the south, this tunicate was discovered in Mexico, near Mazatlan, in the Urias Estuary, in 1995 (Salgado-Barragan et al. 2004). In 1996-2000, it was collected in Mission Bay, Oceanside Harbor, Newport Bay, Alamitos Bay, Los Angeles-Long Beach Harbors, and Marina del Rey, off Santa Monica Bay (Lambert and Lambert 1998; Fairey et al. 2002; Lambert and Lambert 2003; Ruiz et al., unpublished data). At most sites, it was present at low to moderate levels of abundance (Lambert and Lambert 2003). In 2015, one specimen of S. canopus was identified from fouling plates at the Oakland Yacht Club, San Francisco Bay, while several nearby immature specimens were probably this species. This is the northernmost record of this species on the West Coast, but the establishment of this tunicate in the Bay is still unconfirmed (Tracy et al. 2017).

Invasion History on the East Coast:

In the northeast Atlantic, S. canopus ranges from the south coast of England and Normandy (Hayward and Ryland 1990; Breton et al. 1996) to the Cape Verde Islands, Sierra Leone and Senegal (Monniot and Monniot 1994) to Egypt and Lebanon (Bitar and Kouli-Bitar 2001). It was reported from the Channel Islands, off France, in 1868, as Cynthia variabilis (Hancock 1868, cited by Kott 1998), and from the Adriatic Sea in 1877 as Styela canopoides (Heller 1877, as cited by Kott 1998). On biogeographic grounds, based on its likely origin in the Indo-Pacific, S. canopus appears to be a very early introduction to European waters (Carlton, personal communication 2000). 
 
In the southwest Atlantic, S. canopus was first collected from an abandoned fishnet in Rio de Janeiro Bay, Brazil (Monniot 1969, cited by da Rocha and Kremer 2005). In Brazil, it appears to be most abundant in port areas (da Rocha and Kremer 2005).

Invasion History on the Gulf Coast:

In the northeast Atlantic, S. canopus ranges from the south coast of England and Normandy (Hayward and Ryland 1990; Breton et al. 1996) to the Cape Verde Islands, Sierra Leone and Senegal (Monniot and Monniot 1994) to Egypt and Lebanon (Bitar and Kouli-Bitar 2001). It was reported from the Channel Islands, off France, in 1868, as Cynthia variabilis (Hancock 1868, cited by Kott 1998), and from the Adriatic Sea in 1877 as Styela canopoides (Heller 1877, as cited by Kott 1998). On biogeographic grounds, based on its likely origin in the Indo-Pacific, S. canopus appears to be a very early introduction to European waters (Carlton, personal communication 2000).

In the southwest Atlantic, S. canopus was first collected from an abandoned fishnet in Rio de Janeiro Bay, Brazil (Monniot 1969, cited by da Rocha and Kremer 2005). In Brazil, it appears to be most abundant in port areas (da Rocha and Kremer 2005).

Invasion History in Hawaii:

It was first collected in the Hawaiian Islands on Oahu in 1929 and has also been found in Maui (Coles et al. 2004; Carlton and Eldredge 2009). Styela canopus?has a broad range in the west Pacific and Indian Oceans, from Japan to the Red Sea, South Africa, and Australia, and presumed to be native over most of that range (Nishikawa 1991; Kott 1985; Kott 1998; Monniot and Monniot 2001; Monniot 2002). However, it is introduced on some of the islands of the Central Pacific, including the Hawaiian Islands (Coles et al. 2002; Coles et al. 2004), American Samoa (Coles et al. 2003) and Guam (Lambert 2002; Lambert 2003).

Invasion History Elsewhere in the World:

In the northeast Atlantic, S. canopus ranges from the south coast of England and Normandy (Hayward and Ryland 1990; Breton et al. 1996) to the Cape Verde Islands, Sierra Leone and Senegal (Monniot and Monniot 1994) to Egypt and Lebanon (Bitar and Kouli-Bitar 2001). It was reported from the Channel Islands, off France, in 1868, as Cynthia variabilis (Hancock 1868, cited by Kott 1998), and from the Adriatic Sea in 1877 as Styela canopoides (Heller 1877, as cited by Kott 1998). On biogeographic grounds, based on its likely origin in the Indo-Pacific, S. canopus appears to be a very early introduction to European waters (Carlton, personal communication 2000). 
 
In the southwest Atlantic, S. canopus was first collected from an abandoned fishnet in Rio de Janeiro Bay, Brazil (Monniot 1969, cited by da Rocha and Kremer 2005). In Brazil, it appears to be most abundant in port areas (da Rocha and Kremer 2005).

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Description

Styela canopus is a solitary tunicate. Its body shape varies from globular, to elongate, depending on whether it's growing as a single individual or in a dense cluster. When growing singly, it is attached by much of its ventral surface, with the oral siphon slightly below the anterior tip. When crowded, the body is attached by only a small part of the ventral surface, with the oral siphon at the anterior tip, and the atrial siphon slightly below. Both siphons are low and warty. The test is variable in thickness, tough and leathery with rough bumps and wrinkles. Worm tubes, bryozoans and algae frequently grow on the test. The color is usually grayish or yellowish posteriorly, becoming brown, purplish, or red anteriorly, with four dark stripes on the siphons. Internally, the pharynx has four narrow folds on each side, and there are two gonads on the left side and 2-5 on the right. The animal is up to 25-30 mm in length (Van Name 1945; Kott 1985; Lambert and Lambert 1998).

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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. In internal fertilizers, including S. canopus, 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. In S. canopus, the larvae settled after ~2 hours at 25 ?C (Ying et al 2003). Once settled, the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Barnes 1983).

Food:

Phytoplankton, detritus

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
General Habitat	Mangroves	None
General Habitat	Unstructured Bottom	None
General Habitat	Coral reef	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	Mazatlan/Mexico/Urias estuary, Gulf of California (Salgado Barragan et al. 2004)
Minimum Salinity (‰)	23	Mazatlan//Mexico/Urias estuary, Gulf of California (Salgado Barragan et al. 2004)
Maximum Salinity (‰)	43	Mazatlan/Mexico/Urias estuary, Gulf of California (Salgado Barragan et al. 2004)
Maximum Length (mm)	30	Van Name 1945; Kott 1985; Lambert and Lambert 1998
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Styela canopus, commonly known as the Rough Sea Squirt, is a widespread member of the fouling community in the world's coastal waters, and has been reported from ships, buoys, piers, and docks (Woods Hole Oceanographic Institution 1952; Lambert and Lambert, 1998). However, its impacts are unknown.

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