Ciona intestinalis (sea vase)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Biology and Ecology
- Water Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Ciona intestinalis (Linnaeus, 1767)
Preferred Common Name
- sea vase
Other Scientific Names
- Ascidia canina Mueller, 1776
- Ascidia corrugata Mueller, 1776
- Ascidia diaphanea Quoy & Gaimard, 1834
- Ascidia intestinalis Linnaeus, 1767
- Ascidia membranosa Renier, 1807
- Ascidia ocellata Agassiz, 1850
- Ascidia pulchella Alder, 1863
- Ascidia tenella Stimpson, 1852
- Ascidia virens Fabricius, 1779
- Ascidia virescens Pennant, 1812
- Ascidia viridiscens Brugiere, 1792
- Ciona canina Mueller, 1776
- Ciona diaphanea Quoy & Gaimard, 1834
- Ciona ocellata Agassiz, 1850
- Ciona pulchella Alder, 1863
- Ciona robusta Hoshino & Tokioka, 1967
- Ciona roulei Lahille, 1887
- Ciona sociabilis Gunnerus, 1765
- Ciona tenella Stimpson, 1852
- Phallusia intestinalis Linnaeus, 1767
- Tethyum sociabile Gunnerus, 1765
International Common Names
- English: transparent sea squirt; yellow sea squirt
Summary of InvasivenessTop of page
C. intestinalis can grow, reproduce and spread quickly (Ramsay et al., 2009), and can alter benthic communities by either smothering other species (Blum et al., 2007), competing with other suspension-feeders for food (Petersen, 2007; Daigle and Herbinger, 2009) or heavily grazing phytoplankton (Petersen and Riisgård, 1992). The native range of C. intestinalis is currently unresolved, and it is formally considered cryptogenic throughout the northern Atlantic. However, its abundance has increased considerably in the Gulf of St Lawrence and New England, so this species can be considered invasive, and likely exotic, to these areas (Locke, 2009). C. intestinalis invasions often exhibit dramatic boom-bust cycles, but it has established persistent populations in some areas (e.g. in southern California for at least a century (Lambert and Lambert, 1998). C. intestinalis poses a significant economic threat to bivalve aquaculture, and has been documented growing in these areas in Hong Kong, New Zealand, Japan, Canada, Spain, South Africa and Chile (Rocha et al., 2009).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Chordata
- Subphylum: Tunicata
- Class: Ascidiacea
- Order: Enterogona
- Suborder: Aplousobranchia
- Family: Cionidae
- Genus: Ciona
- Species: Ciona intestinalis
Notes on Taxonomy and NomenclatureTop of page
There are about 15 species within the genus Ciona. Ciona intestinalis is a widespread, morphologically variable species that had been frequently misidentified in the past. Further adding confusion, recent molecular studies have identified at least two cryptic C. intestinalis species, formally referred to as type A and type B. Type A is widely distributed in the Mediterranean and the Pacific and type B is widely distributed throughout the northern Atlantic. Although the two types do coexist in part of their range, they are partially reproductively isolated: crosses of type B eggs with type A sperm result in normal fertilization rates, but reciprocal crosses yield very low fertilization rates (Lambert et al., 1990; Suzuki et al., 2005). See Suzuki et al. (2005), Caputi et al. (2007), Iannelli et al. (2007), and Nydam and Harrison (2007) for further information on the C. intestinalis cryptic species complex.
DescriptionTop of page
C. intestinalis is a solitary, translucent ascidian that can have a pale yellow or pale green hue. If individuals are not fouled by algae or invertebrates, ten muscle bands that run the length of the body are visible, and pale orange internal organs are seen through the translucent body. The brachial siphon has eight lobes and the atrial siphon has six lobes. Both siphons may have yellow or orange margins.
DistributionTop of page
C. intestinalis is now a cosmopolitan species inhabiting sub-arctic, temperate and tropical waters. It is considered to be primarily a cold-water species, but temporary or transient populations have been observed in tropical waters.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Arctic Sea||Present||Native||Denmark Strait, Barents Sea|
|Atlantic - Eastern Central||Present||Cap Verde, east Africa|
|Atlantic - Northeast||Present, Widespread||Native||Sweden, Denmark, Norway, Germany, United Kingdom, Spain|
|Atlantic - Northwest||Present||Native||Bay of Fundy|
|Atlantic - Southeast||Present||Introduced||South Africa|
|Atlantic - Southwest||Present||Introduced||Brazil|
|Indian Ocean - Eastern||Present||Introduced||Papua New Guinea, Western Australia|
|Mediterranean and Black Sea||Present, Widespread||Native|
|Pacific - Eastern Central||Present, Widespread||Introduced||Invasive||California (San Diego, San Francisco)|
|Pacific - Northeast||Present||Introduced||Alaska, British Columbia, Washington|
|Pacific - Northwest||Present||China, Japan, Korea|
|Pacific - Southeast||Present||Introduced||Invasive||Chile|
|Pacific - Southwest||Present||Introduced||New Zealand, Southeast Australia|
|Pacific - Western Central||Present||Introduced||Indonessia|
History of Introduction and SpreadTop of page
C. intestinalis was introduced to non-native ranges of southern California, South Africa and Brazil at least as early as the twentieth century. The main vector of dispersal is widely believed to be ship fouling, including via sea chests (Lambert and Lambert, 2003). Persistent taxonomic confusion and poor early records prevent more detailed understanding of the history of its introduction and spread. Recently, C. intestinalis has become a nuisance species to aquaculture industries in northern Atlantic Canada and in Chile (Uribe and Etchepare, 2002; Carver et al., 2003).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Atlantic, Northwest||2004||Aquaculture (pathway cause); Hitchhiker (pathway cause)||Yes||Locke et al. (2009b)||Brundell River, Prince Edward Island, Canada|
|Pacific, Eastern Central||1917onwards||Yes||Lambert and Lambert (1998); Ritter and Forsyth (1917)||San Diego, CA|
Risk of IntroductionTop of page
The risk of introduction of C. intestinalis is high. It has a broad temperature and salinity, fast growth rate and high fecundity (Dybern, 1965) and can quickly form dense aggregations (Lambert and Lambert, 1998). C. intestinalis can exclude native species (Blum et al., 2007) and reduce the productivity of aquaculture industries (Uribe and Etchepare, 2002; Carver et al., 2003; Ramsay et al., 2009). Total eradication of C. intestinalis would be costly and has never been attempted (Edwards and Leung, 2009), but rapid-response strategies to reduce population sizes and prevent spread have proven successful (Locke et al., 2009b). Anthropogenic dispersal appears to be common, most likely via hull-fouling and transportation of aquaculture and fishing gear (Lambert and Lambert, 1998). Natural long-distance dispersal is rare because adults are sessile and the dispersive larval stage is short (Petersen and Svane, 1995), although long-distance natural dispersal via rafting on eelgrass or kelp is possible (Havenhand and Svane, 1991).
HabitatTop of page
C. intestinalis grows on submerged substrates including rock, eelgrass and kelp, and on anthropogenic substrates such as wood, metal or concrete docks, pilings and aquaculture gear (Dybern, 1963; 1965; Yamaguchi, 1971; McDonald, 2004). One study found that C. intestinalis was more abundant on vertical surfaces (Costelloe et al., 1986), but it is unclear if this is due to preferential larval settlement, preferential growth rates or higher biotic resistance at other orientations. C. intestinalis is most abundant in enclosed or semi-enclosed embayments, but has been found at depths up to 100 m (Dybern, 1965).
Habitat ListTop of page
|Littoral||Coastal areas||Present, no further details|
|Marine||Inshore marine||Present, no further details|
|Marine||Benthic zone||Present, no further details|
Hosts/Species AffectedTop of page
C. intestinalis can quickly form dense aggregations, which can smother and eventually exclude other fouling species and exert heavy grazing pressure on the local phytoplankton and bacterial communities (Peterson and Riisgård, 1992; Lambert and Lambert, 1998; Riisgård et al., 1998; Blum et al., 2007; Petersen, 2007). In Tasmania, C. intestinalis was found to harbour the amoeba, Neoparamoeba pemaquidensis, which is responsible for amoebic gill disease in farmed salmon (Tan et al., 2002).
Biology and EcologyTop of page
C. intestinalis can survive a broad range of temperatures (-1°C to 30°C) (Dybern, 1965; Carver et al., 2003) and salinities (Dybern, 1967). However, survival, growth and filtration rates are reduced at extreme temperatures (Dybern, 1965). Mortality increases at <10°C and filtration rates reduce at >21°C (Petersen et al., 1999) and it appears that this species cannot withstand extended periods with salinity <11 ppt (Dybern, 1967).
C. intestinalis exhibits considerable variation in generation time and spawning phenology across its range, and this variation is believed to be driven by temperature regime (Dybern, 1965). In sub-arctic or deep waters that are cold throughout the year, C. intestinalis can live up to two years and will spawn continuously throughout the year. In waters that exhibit strong seasonal differences in temperature, C. intestinalis lives ~ 1 year and spawns at temperatures >8°C, and in continuously warm waters generation time is short <1 year) and spawning is continuous throughout the year.
C. intestinalis is a suspension-feeder, and uses ciliary action to pump water through its branchial basket, where small particles (0.5-5 µm) including phytoplankton and bacteria are trapped on a mucous sheet (Lambert, 2005, see Petersen, 2007 for a detailed review of feeding anatomy and feeding rates).
C. intestinalis has a wide temperature and salinity tolerance, with adults being more tolerant than larvae. Adults can survive temperatures ranging from -1 to 30°C (Dybern, 1965; Carver et al., 2003) and salinities ranging from 8-40 ppt (Dybern, 1967). However, C. intestinalis does not appear to be able to survive salinities persistently below 11 ppt (Dybern, 1967), and growth and filtration rates are reduced at temperature extremes (Dybern, 1965; Petersen and Riisgård, 1992; Riisgård et al., 1996; Petersen et al., 1999).
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Depth (m b.s.l.)||Optimum||1-100 tolerated|
|Salinity (part per thousand)||8||42||Optimum||34-42 optimal for embryogenesis (Bellas et al., 2003)|
|Velocity (cm/h)||Optimum||Prefers semi-protected embayments with good water flow|
|Water pH (pH)||7||9||Optimum||(Bellas et al., 2003)|
|Water temperature (ºC temperature)||10||20||Optimum||-1-30 tolerated (Dybern, 1965)|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Asterias rubens||Predator||Adult||not specific|
|Aurelia aurita||Predator||Larval||not specific|
|Cancer irroratus||Predator||Adult||not specific|
|Carcinus maenas||Predator||Adult||not specific|
Notes on Natural EnemiesTop of page
Natural enemies include a variety of fish (Yamaguchi, 1971; 1975; Petersen and Svane, 1995), crabs (Carver et al., 2003), seastars (Gulliksen and Skjæveland, 1973; Svane, 1983) and small snails (Mitrella) (Whitlatch and Osman, 2009); Hydrobia and Littorina (Petersen and Svane, 1995) all of which consume adult or newly-settled C. intestinalis. Petersen and Svane (1995) also demonstrated that the jellyfish Aurelia aurita consume substantial amounts of C. intestinalis eggs and larvae.
Means of Movement and DispersalTop of page
Adults are sessile, so natural dispersal occurs primarily by passive drifting of eggs/egg-strings and active swimming by larvae, although dispersal of adults rafting on eelgrass blades and kelp fronds can occur. The major anthropogenic vectors of adult C. intestinalis are widely believed to be boat hulls and aquaculture equipment (Lambert and Lambert, 1998).
Natural Dispersal (Non-Biotic)
C. intestinalis currently has no commercial or aesthetic values, so all introductions are assumed to be accidental.
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
Economic ImpactTop of page
C. intestinalis can reduce the productivity of bivalve aquaculture (Uribe and Etchepare, 2002; Carver et al., 2003; Ramsay et al., 2009) either by fouling aquaculture gear (Yamaguchi, 1975;Kang et al., 1978; Castilla et al., 2005), smothering shellfish or competing with shellfish for food (Daigle and Herbinger, 2009). Edwards and Leung (2009) estimated that fouling by invasive ascidians costs the aquaculture industry on Prince Edward Island approximately Canadian $5 million annually.
Environmental ImpactTop of page
C. intestinalis can significantly alter marine benthic communities by competing with native benthic species for space and food and by consuming phytoplankton and possibly invertebrate larvae. C. intestinalis has been demonstrated to exclude sessile marine invertebrates and reduce native biodiversity (Blum et al., 2007), compete with other suspension-feeding species for food (Daigle and Herbinger, 2009), and its relatively high clearance rate (Petersen and Riisgärd, 1992) indicates that it has the potential to significantly alter the phytoplankton community. Furthermore, solitary ascidians species are known to consume invertebrate larvae (Bingham and Walters, 1989) although this has not yet been demonstrated for C. intestinalis specifically.
Impact on Habitats
C. intestinalis can form dense aggregations that can provide a new substrate for fouling organisms. Ciona intestinalis also has the ability to filter a large amount of water, which could alter phytoplankton communities.
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Highly adaptable to different environments
- Pioneering in disturbed areas
- Tolerant of shade
- Has high reproductive potential
- Ecosystem change/ habitat alteration
- Host damage
- Infrastructure damage
- Modification of natural benthic communities
- Monoculture formation
- Negatively impacts livelihoods
- Negatively impacts aquaculture/fisheries
- Reduced native biodiversity
- Threat to/ loss of native species
- Competition - monopolizing resources
- Competition - smothering
- Pest and disease transmission
- Rapid growth
UsesTop of page
C. intestinalis is used a model organism (Corbo et al., 2001).
Uses ListTop of page
- Laboratory use
- Research model
Similarities to Other Species/ConditionsTop of page
Recent molecular studies have identified at least two cryptogenic species within C. intestinalis: type A which is distributed in the Mediterranean and Pacific and type B which is distributed in the north Atlantic (Suzuki et al., 2005; Caputi et al., 2007; Iannelli et al., 2007; Nydam and Harrison, 2007). There are no obvious morphological differences between these two types, although Caputi et al. (2007) suggest that the two forms can be distinguished by looking at spermiduct pigmentation. With one exception, they found that papillae at the end of the vas deferens (which is located inside the atrial siphon just below the anus) of C. intestinalis type A had bright orange pigmentation whereas the remainder of the duct remained uncoloured. In C. intestinalis type B, bright orange pigmentation is confined to the duct only. Caputi et al. (2007) also report molecular characters that can be used to identify the two types.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
A C. intestinalis prevention strategy should focus on reducing anthropogenic transport of this species. The main vectors are boat-hulls and aquaculture gear, which should be cleared before transportation from infested areas. On Prince Edward Island (PEI), the Introduction and Transfers committee (I&T) composed of federal and provincial resources managers, academic and governmental scientists, and mussel aquaculture industry representatives enforce aquaculture containment and equipment cleaning laws to prevent the spread of invasive solitary ascidians within the region. These laws were very successful at limiting the spread of another solitary ascidian, Styela clava, but only modestly successful in limiting the spread of C. intestinalis (Locke et al., 2009b).
Locke et al. (2009b) present a superb review of rapid response efforts against four different ascidian species, including C. intestinalis, in Prince Edward Islands, Canada.
Cultural control and sanitary measures
Exposure to 5% acetic acid is 100% effective against C. intestinalis after 1 minute of exposure, and was found to be a more effective than hydrated lime, saturated brine, or hypochlorite solution treatments (Carver et al., 2003; Locke et al., 2009a).
In the Prince Edward Islands (PEI), Canada, the regulation of aquaculture transport and harvests has been modestly successful at containing the spread of C. intestinalis. Aquaculture transport, harvest and equipment cleaning regulations were mandated by the PEI Introductions and Transfer Committee under section 55 of Canada’s federal Fishing General Regulations in 2001 and 2002 (Locke et al., 2009). These regulations were actually developed to prevent the spread of an earlier invasive solitary tunicate, Styela clava, and were already in place when C. intestinalis invaded in 2004. Since 2004, C. intestinalis has spread to other rivers and bays in PEI, but it is believed that these regulations have helped slow the invasion.
Gaps in Knowledge/Research NeedsTop of page
In contrast to many other invasive marine species, the biology and ecology of C. intestinalis is well known (Berrill, 1947; Dybern, 1965; Dybern, 1967; Lambert and Brandt, 1967; Gulliksen, 1972; Petersen and Svane, Blum et al., 2007; Daigle et al., 2009).
ReferencesTop of page
Bellas J; Vázquez E; Beiras R, 2001. Toxicity of Hg, Cu, Cd, and Cr on early developmental stages of Ciona intestinalis (Chordata, Ascidiacea) with potential application in marine water quality assessment. Water Research, 35:2905-2912.
Blum JC; Chang AL; Liljeshröm M; Schenk ME; Steinberg MK; Ruiz GM, 2007. The non-native Ciona intestinalis (L.) depresses species richness. Journal of Experimental Marine Biology and Ecology, 342:5-14.
Carman MR; Hoagland KE; Green-Beach E; Grunden DW, 2009. Tunicate faunas of two North Atlantic-New England islands: Martha's Vineyard Massachusetts and Black Island, Rhode Island. Aquatic Invasions, 4:65-70.
Castilla JC; Uribe M; Bahamonde N; Clarke M; Desqueyroux-Faúndez R; Kong I; Moyano H; Rozbaczylo N; Santelices B; Valdovinos C; Zavala P, 2005. Down under the southeastern Pacific: marine non-indigenous species in Chile. Biological Invasions, 7:213-232.
Dehal P; Satou Y; Campbell RK; Chapman J; Degnan B; Tomaso ADe; Davidson B; Gregorio ADi; Gelpke M; Goodstein DM; Harafuji N; Hastings KEM; Ho I; Hotta K; Huang W; Kawashima T; Lemaire P; Martinez D; Meinertzhagen IA; Necula S; Nonaka M; Putnam N; Rash S; Saiga H; Satake M; Terry A; Yamada L; Wang H-G; Awazu S; Azumi K; Boore J; Branno M; Chin-bow S; DeSantis R; Doyle S; Francino P; Keys DN; Haga S; Hayashi H; Hino K; Imai KS; Inaba K; Kano S; Kobayashi K; Kobayashi M; Lee B-I; Makabe KW; Manohar C; Matassi G; Medina M; Mochizuki Y; Mount S; Morishita T; Miura S; Nakayama A; Nishizaka S; Nomoto H; Ohta F; Oishi K; Rigoutsos I; Sano M; Sasaki A; Sasakura Y; Shoguchi E; Shin-I T; Spagnuolo A; Stainier D; Suzuki MM; Tassy O; Takatori N; Tokuoka M; Yagi K; Yoshizaki F; Wada S; Zhang C; Hyatt PD; Larimer F; Detter C; Doggett N; Glavina T; Hawkins T; Richardson P; Lucas S; Kohara Y; Levine M; Satoh N; Rokhsar DS, 2002. The draft genome of Ciona intestinalis: Insights into chordate and vert
Hewitt CL; Campbell ML; Thresher RE; Martin RB; Boyd S; Cohen BF; Currie DR; Gomom MF; Keough MJ; Lewis JA; Lockett MM; Mays N; MacArthur MA; O'Hara TD; Poore GCB; Ross DJ; Story MJ; Watson JE; Wilson RS, 2004. Introduced and cryptogenic species in Port Phillip Bay, Victoria, Australia. Marine Biology, 144:183-202.
Jordan H, 1908. [English title not available]. (Uber reflezarme tiere. Ein Beitrag zur vergleihende physiologie des zentralen nervensystems, vornehmlich auf grund von versuche an Ciona intestinalis und Oktopoden) Zeitschrift für Allgemeine Microbiologie, 7:86-135.
Kang PA; Bae PA; Pyen CK, 1978. Studies on the suspended culture of oyster, Crassostrea gigas in Korean coastal waters. On the fouling organisms associated with culturing oysters at the oyster culture farms in Chungmu. Bulletin of Fisheries Research and Development Agency, 20:121-127.
Kott P, 1997. The Tunicates. Adelaide, South Australia: Government Printer, pp. In: Marine Invertebrates of Southern Australia, Part I [ed. by Shepherd SA, Thomas IM] Adelaide, South Australia, : Government Printer, 1092-1255.
Lesser MP; Shumway SE; Cucci T; Smith J, 1992. Impact of fouling organisms on mussel rope culture: interspecific competition for food among suspension-feeding invertebrates. Journal of Experimental Marine Biology and Ecology, 165:91-102.
Lo Bianci S, 1909. [English title not available]. (Notizie biologiche riguardanti specialmente il periodic di maturita sessuale degli animali del golfo di Napoli) Mittheilungen aus der Zoologischen Staatsinst zu Neapel, 19:513-761.
Locke A; Doe KG; Fairchild WL; Jackman PM; Reese EJ, 2009. Preliminary evaluation of effects of invasive tunicate management with acetic aid and calcium hydroxide on non-target marine organisms in Prince Edward Island, Canada. Aquatic Invasions, 4:237-247.
Ooishi S; O'Reilly MG, 2004. Redescription of Haplostoma eruca (Copepoda: Cyclopoida: Ascidicolidae) living in the intestine of Ciona intestinalis from Clyde Estuary, Scotland. Journal of Crustacean Biology, 24:9-16.
Pastore M, 2001. Copepods associated with Phallusia mamillata and Ciona intestinalis (Tunicata) in the area of Taranto (Ionian Sea, southern Italy). Journal of the Marine Biological Associations of the United Kingdom, 81:427-432.
Petersen JK; Mayer S; Knudson MA, 1999. Beat frequency of cilia in the branchial basket of the ascidian Ciona intestinalis in relation to temperature and algal cell concentration. Marine Biology, 133:185-192.
Ramsay A; Davidson J; Bourque D; Stryhn H, 2009. Recruitment patterns and population development of the invasive ascidian Ciona intestinalis in Prince Edward Island, Canada. Aquatic Invasions, 4:169-176.
Runnström S, 1936. [English title not available]. (Die Anpassung der Fortpflanzung und Entwicklung mariner Tiere an die Temperaturverhältnisse verschiedener Verbreitungsgebiete) Bergens Museum Arbok 3, 1-36.
Sordino P; Heisenberg CP; Cirino P; Toscano P; Giuliano A; Marino R; Pinto MR; De-Santis R, 2000. A mutational approach to the study of development of the protochordate Ciona intestinalis (Tunicate, Chordata). Sarsia, 85:173-176.
Therriault TW; Herborg L-M, 2008. Predicting the potential distribution of the vase tunicate Ciona intestinalis in Canadian waters: informing a risk assessment. ICES Journal of Marine Science, 65:788-794.
Uribe E; Etchepare I, 2002. Effects of biofouling by Ciona intestinalis on suspended aquaculture of Agropecten purpuratus in Bahia Inglesa, Chile. Bulletin of the Aquaculture Association of Canada, 102:93-95.
Yamaguchi M, 1975. Growth and reproductive cycles of the marine fouling ascidians Ciona intestinalis, Styela plicata, Botrylloides violaceus and Leptoclinum mitsukurii at Aburatsubo-Moroiso inlet (Central Japan). Marine Biology, 29:253-259.
Blum J C, Chang A L, Liljeshröm M, Schenk M E, Steinberg M K, Ruiz G M, 2007. The non-native Ciona intestinalis (L.) depresses species richness. Journal of Experimental Marine Biology and Ecology. 5-14.
CABI, Undated. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Carman M R, Hoagland K E, Green-Beach E, Grunden D W, 2009. Tunicate faunas of two North Atlantic - New England islands: Martha's Vineyard Massachusetts and Black Island, Rhode Island. Aquatic Invasions. 65-70.
Castilla J C, Uribe M, Bahamonde N, Clarke M, Desqueyroux-Faúndez R, Kong I, Moyano H, Rozbaczylo N, Santelices B, Valdovinos C, Zavala P, 2005. Down under the southeastern Pacific: marine non-indigenous species in Chile. Biological Invasions. 213-232.
Jordan H, 1908. About reflex-poor animals. A contribution to comparative physiology of the central nervous system, primarily on the basis of experiments on Ciona intestinalis and octopuses. (Über reflezarme tiere. Ein Beitrag zur vergleichende physiologie des zentralen nervensystems, vornehmlich auf grund von versuche an Ciona intestinalis und Oktopoden.). Zeitschrift für Allgemeine Microbiologie. 86-135.
Kang P A, Bae P A, Pyen C K, 1978. Studies on the suspended culture of oyster, Crassostrea gigas in Korean coastal waters. On the fouling organisms associated with culturing oysters at the oyster culture farms in Chungmu. Bulletin of Fisheries Research and Development Agency. 121-127.
Linnaeus C, 1758. [English title not available]. (Systema Naturae per Regna Tria Naturae, Secundum classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. ed. X.). In: Systema Naturae per Regna Tria Naturae, Secundum classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. ed. X. Holmiae,
Lo Bianci S, 1909. Biological information regarding the frequency of sexual maturity of animals in the Gulf of Naples. (Notizie biologiche riguardanti specialmente il periodic di maturita sessuale degli animali del golfo di Napoli.). Mittheilungen aus der Zoologischen Staatsinst zu Neapel. 513-761.
Pérès J M, 1951. A new contribution to the study of the ascidians of the coast of West Africa. (Nouvelle contribution à l'éude des ascidies de la côte occidentale d'Afrique.). In: Bulletin de la Institute France Afrique Noir.
Uribe E, Etchepare I, 2002. Effects of biofouling by Ciona intestinalis on suspended aquaculture of Agropecten purpuratus in Bahia Inglesa, Chile. Bulletin of the Aquaculture Association of Canada. 93-95.
OrganizationsTop of page
Canada: Department of Fisheries and Aquaculture, Department of Fisheries, Aqualculture and Rural Development Fifth Floor, Jones Building, 11 Kent St, PO Box 2000, Charlottetown, PEI
Ontario: Department of Fisheries and Ocean - Canada, Fisheries and Oceans Canada Communications Branch 200 Kent Street, 13th Floor, Station 13E228, Ottawa, http://www.dfo-mpo.gc.ca
ContributorsTop of page
19/06/09 Original text by:
Erin Grey, University of Chicago, USA
Distribution MapsTop of page
Select a dataset
CABI Summary Records
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