Invasion History
First Non-native North American Tidal Record: 1985First Non-native West Coast Tidal Record: 1985
First Non-native East/Gulf Coast Tidal Record:
General Invasion History:
Ciona savignyi appears to be native to Japan and possibly northern Asia. In Japan, it can be found from Mutsu Bay (north end of Honshu) to the Seto Inland Sea (south end of Honshu), both on the Pacific and Sea of Japan coasts (Hoshino and Nishikawa 1980; Nishikawa 1991). In 2004, it was first collected as an introduced species in the most northern reaches of the Sea of Japan, in Peter the Great Bay, Russia (Zvyagintsev et al. 2007). In the northeast Pacific, there are two puzzling records from southeastern Alaska (Inside Passage: Loring, in Behm Canal) and northern British Columbia (Stuart Island, in Queen Charlotte Strait) (AK in 1903, USNM 5633, U.S. National Museum of Natural History 2003; BC in 1937, Lambert 2003). These could represent a cryptic species, a very early introduction, or a relict population. There are also disjunct records from Argentina (Hoshino and Nishikawa 1985) and Spain (Perez et al. 1957) which for the purposes of this database, we consider unverified. However, C. savignyi has recently invaded shallow coastal waters of central and southern California and Puget Sound (Cohen and Carlton 1995; Lambert and Lambert 1998; Lambert 2003; Cohen et al. 2005; Blum et al. 2007).
North American Invasion History:
Invasion History on the West Coast:
The first record of Ciona savignyi in continental US waters was in 1985 in Long Beach Harbor, southern CA (Lambert and Lambert 1998). In 1993, it was collected in central California, San Francisco Bay (Cohen and Carlton 1995). By 1994 the southern California population ranged from San Diego Bay to Santa Barbara Harbor, CA (Lambert and Lambert 1998; Lambert and Lambert 2003). The central California population soon included specimens collected in 1998 Elkhorn Slough surveys (Wasson et al. 2001) and in 2003 Monterey and Moss Landing Harbors samples (deRivera et al. 2005). As of 2005, C. savignyi has been collected in the central and southern sections of San Francisco Bay, but not the more northern San Pablo Bay (Cohen et al. 2005; Ruiz et al. unpublished data). North of San Francisco Bay, C. savignyi has been found in samples collected from Tomales Bay (in 2001, Fairey et al. 2002), Bodega Harbor (deRivera et al. 2005), and Humboldt Bay, CA (in 2001, Fairey et al. 2002; Ruiz et al. unpublished data).
In Washington, C. savignyi has spread throughout Puget Sound. It was first discovered in 1998 in the Des Moines Marina near Seattle and by 1999 it had spread north to the Brownsville and Edmonds Marinas. In 2001, it was found near the Tacoma Yacht Club in Tacoma, WA (Lambert 2003). Subsequently, scuba divers discovered extensive populations in several other parts of Puget Sound, including Hood Canal in the western portion of Puget sound (in 2005, USGS Nonindigenous Aquatic Species Program 2003-2012) and in more northern reaches of the San Juan Islands (USGS Nonindigenous Aquatic Species Program 2003-2012).
Description
Ciona savignyi is a solitary tunicate with a smooth, elongated, cylindrical or vase-shaped body which can reach a length of 150 mm long. It is widest near the permanently attached posterior end and tapers toward the anterior end, compressed laterally. The tunic is transparent, translucent, and white or yellowish-green in color. Much of the tunic is soft, flexible and gelatinous. The muscle bands and organs are often visible beneath the tunic surface. Its siphons are on short, forwardly directed (not divergent) and the oral siphon is larger than the atrial siphon. The oral siphon has 8 lobes, each with a yellow margin containing 8 reddish-orange spots. The atrial siphon has 6 lobes, each with a yellow margin containing 6 reddish-orange spots. There are 5-7 conspicuous longitudinal muscle bands on each side of the body that extend nearly the entire length of the body (Van Name 1945; Kott 1985; Lambert and Lambert 1998).
Ciona savignyi is very similar in appearance to C. intestinalis, but there are a few notable morphological differences. Ciona savignyi always has white pigmented flecks, or spots, in its body wall while C. intestinalis lacks these. Additionally, C. savignyi has orange pigmentation around the siphon while C. intestinalis has yellow pigmentation (Smith et al. 2010). The number of tentacles around the oral siphon is variable in both species, but generally C. savignyi have fewer (n<50) tentacles than C. intestinalis (Hoshino and Nishikawa 1985). Ciona savignyi does not have an endostylar appendage while C. intestinalis has endostyles. Additionally, the pharyngeo-epicardiac openings in C. savignyi are located close to the oesophageal opening while in C. intestinalis these openings are usually very small and located near its base (Hoshino and Nishikawa 1985). Lastly, the color of the enlarged end of the vas deferens is always white in C. savignyi and red in the West Coast C. intestinalis (Lambert and Lambert 1998).
Taxonomy
Taxonomic Tree
Kingdom: | Animalia | |
Phylum: | Chordata | |
Subphylum: | Tunicata | |
Class: | Ascidiacea | |
Order: | Phlebobranchia | |
Family: | Cionidae | |
Genus: | Ciona | |
Species: | savignyi |
Synonyms
Potentially Misidentified Species
Ritter 1913, mis-identification of specimen from Loring, Alaska (Lambert 2003)
Ciona robusta
None
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 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
SusFedHabitats
General Habitat | Marinas & Docks | None |
General Habitat | Rocky | None |
Salinity Range | Polyhaline | 18-30 PSU |
Salinity Range | Euhaline | 30-40 PSU |
Tidal Range | Subtidal | None |
Vertical Habitat | Epibenthic | None |
Life History
Tolerances and Life History Parameters
Maximum Temperature (ºC) | 26.7 | Field, US East & West Coast marinas (Lord et al. 2015) |
Maximum Length (mm) | 94 | Hoshino and Nishikawa 1985 |
Broad Temperature Range | None | Cold temperate-Warm temperate |
Broad Salinity Range | None | Polyhaline-Euhaline |
General Impacts
Economic Impacts
Shipping and Industry: Ciona savignyi is now an abundant component of the fouling community in southern California and Puget Sound (Lambert and Lambert 2003). It is likely that fouling by C. savignyi affects shipping and harbor uses, but no impacts have been reported.
Ecological Impacts
Competition: Currently, little is known about species competition with C. savignyi despite the fact that it is an abundant component of the fouling community in southern California and Puget Sound (Lambert and Lambert 2003). In southern California, it has been speculated that species competition is possible with C. intestinalis. However, die-offs of both species, due to environmental changes, make it difficult to determine the extent of competition. Lambert and Lambert (2003) state that 'Perhaps C. savignyi would eventually replace C. intestinalis were it not that the frequently changing habitat apparently tends to stabilize their co-occurrence in nearly equal numbers.'
Regional Distribution Map
Bioregion | Region Name | Year | Invasion Status | Population Status |
---|---|---|---|---|
NEP-VI | Pt. Conception to Southern Baja California | 1985 | Non-native | Established |
NEP-V | Northern California to Mid Channel Islands | 1993 | Non-native | Established |
NEP-III | Alaskan panhandle to N. of Puget Sound | 1903 | Crypogenic | Unknown |
NWP-4b | None | 0 | Native | Established |
NWP-3b | None | 1882 | Native | Established |
NWP-4a | None | 0 | Native | Established |
NEP-IV | Puget Sound to Northern California | 2001 | Non-native | Established |
AUS-V | None | 2002 | Non-native | Established |
P050 | San Pedro Bay | 1985 | Non-native | Established |
P130 | Humboldt Bay | 2001 | Non-native | Established |
P020 | San Diego Bay | 1994 | Non-native | Established |
P030 | Mission Bay | 1994 | Non-native | Established |
P023 | _CDA_P023 (San Louis Rey-Escondido) | 1998 | Non-native | Established |
P040 | Newport Bay | 1994 | Non-native | Established |
P060 | Santa Monica Bay | 1995 | Non-native | Established |
P062 | _CDA_P062 (Calleguas) | 1994 | Non-native | Established |
P064 | _CDA_P064 (Ventura) | 1996 | Non-native | Established |
P065 | _CDA_P065 (Santa Barbara Channel) | 1994 | Non-native | Established |
P080 | Monterey Bay | 2003 | Non-native | Established |
P090 | San Francisco Bay | 1993 | Non-native | Established |
P110 | Tomales Bay | 2001 | Non-native | Established |
P290 | Puget Sound | 1998 | Non-native | Established |
P292 | _CDA_P292 (San Juan Islands) | 2008 | Non-native | Established |
P112 | _CDA_P112 (Bodega Bay) | 2004 | Non-native | Established |
NZ-IV | None | 2010 | Non-native | Established |
NWP-3a | None | 0 | Native | Established |
P027 | _CDA_P027 (Aliso-San Onofre) | 2011 | Non-native | Established |
SA-I | None | 1962 | Non-native | Established |
Occurrence Map
OCC_ID | Author | Year | Date | Locality | Status | Latitude | Longitude |
---|---|---|---|---|---|---|---|
767444 | Ruiz et al., 2015 | 2013 | 2013-07-23 | Marina Village, Mission Bay, CA, California, USA | Non-native | 32.7605 | -117.2364 |
767494 | Ruiz et al., 2015 | 2013 | 2013-07-31 | Campland on the Bay, Mission Bay, CA, California, USA | Non-native | 32.7936 | -117.2234 |
767512 | Ruiz et al., 2015 | 2013 | 2013-08-01 | Hyatt Resort Marina, Mission Bay, CA, California, USA | Non-native | 32.7634 | -117.2397 |
767556 | Ruiz et al., 2015 | 2013 | 2013-08-02 | The Dana Marina, Mission Bay, CA, California, USA | Non-native | 32.7671 | -117.2363 |
767681 | Ruiz et al., 2015 | 2013 | 2013-07-17 | Naval Station San Diego, San Diego Bay, CA, California, USA | Non-native | 32.6867 | -117.1333 |
767695 | Ruiz et al., 2015 | 2013 | 2013-07-24 | NAB ACU-1 Docks, San Diego Bay, CA, California, USA | Non-native | 32.6786 | -117.1615 |
767719 | Ruiz et al., 2015 | 2013 | 2013-07-21 | Cabrillo Isle Marina, San Diego Bay, CA, California, USA | Non-native | 32.7272 | -117.1995 |
767732 | Ruiz et al., 2015 | 2013 | 2013-07-22 | Coronado Cays Marina, San Diego Bay, CA, California, USA | Non-native | 32.6257 | -117.1309 |
767747 | Ruiz et al., 2015 | 2013 | 2013-07-18 | NAB Fiddlers Cove, San Diego Bay, CA, California, USA | Non-native | 32.6524 | -117.1486 |
767764 | Ruiz et al., 2015 | 2013 | 2013-07-26 | Pier 32 Marina, San Diego Bay, CA, California, USA | Non-native | 32.6516 | -117.1077 |
767787 | Ruiz et al., 2015 | 2013 | 2013-07-28 | Marriott Marquis and Marina, San Diego Bay, CA, California, USA | Non-native | 32.7059 | -117.1655 |
767804 | Ruiz et al., 2015 | 2011 | 2011-09-15 | Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA | Non-native | 37.9117 | -122.3494 |
767858 | Ruiz et al., 2015 | 2011 | 2011-09-13 | Oyster Point Marina, San Francisco Bay, CA, California, USA | Non-native | 37.6725 | -122.3864 |
767892 | Ruiz et al., 2015 | 2011 | 2012-09-19 | Sausalito Marine Harbor, San Francisco Bay, CA, California, USA | Non-native | 37.8609 | -122.4853 |
767909 | Ruiz et al., 2015 | 2011 | 2011-09-21 | South Beach Harbor, San Francisco Bay, CA, California, USA | Non-native | 37.7797 | -122.3871 |
767923 | Ruiz et al., 2015 | 2011 | 2011-09-20 | Jack London Square Marina, San Francisco Bay, CA, California, USA | Non-native | 37.7947 | -122.2822 |
767934 | Ruiz et al., 2015 | 2011 | 2011-09-22 | Ballena Isle Marina, San Francisco Bay, CA, California, USA | Non-native | 37.7676 | -122.2869 |
767953 | Ruiz et al., 2015 | 2011 | 2011-09-12 | Paradise Cay Yacht Harbor, San Francisco Bay, CA, California, USA | Non-native | 37.9156 | -122.4769 |
767969 | Ruiz et al., 2015 | 2011 | 2011-09-12 | Corinthian Yacht Club, San Francisco Bay, CA, California, USA | Non-native | 37.8103 | -122.3228 |
767989 | Ruiz et al., 2015 | 2012 | 2012-08-24 | Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA | Non-native | 37.9134 | -122.3523 |
768008 | Ruiz et al., 2015 | 2012 | 2012-08-23 | Sausalito Marine Harbor, San Francisco Bay, CA, California, USA | Non-native | 37.8609 | -122.4853 |
768023 | Ruiz et al., 2015 | 2012 | 2012-08-28 | San Francisco Marina, San Francisco Bay, CA, California, USA | Non-native | 37.8071 | -122.4341 |
768042 | Ruiz et al., 2015 | 2012 | 2012-08-27 | Port of San Francisco Pier 31, San Francisco Bay, CA, California, USA | Non-native | 37.8078 | -122.4060 |
768065 | Ruiz et al., 2015 | 2012 | 2012-09-11 | Ballena Isle Marina, San Francisco Bay, CA, California, USA | Non-native | 37.7676 | -122.2869 |
768088 | Ruiz et al., 2015 | 2012 | 2012-08-30 | Oyster Point Marina, San Francisco Bay, CA, California, USA | Non-native | 37.6633 | -122.3817 |
768112 | Ruiz et al., 2015 | 2012 | 2012-08-29 | Coyote Point Marina, San Francisco Bay, CA, California, USA | Non-native | 37.5877 | -122.3174 |
768134 | Ruiz et al., 2015 | 2012 | 2012-09-04 | Redwood City Marina, San Francisco Bay, CA, California, USA | Non-native | 37.5023 | -122.2130 |
768178 | Ruiz et al., 2015 | 2012 | 2012-09-05 | Port of Oakland, San Francisco Bay, CA, California, USA | Non-native | 37.7987 | -122.3228 |
768198 | Ruiz et al., 2015 | 2012 | 2012-09-07 | Jack London Square Marina, San Francisco Bay, CA, California, USA | Non-native | 37.7940 | -122.2787 |
768254 | Ruiz et al., 2015 | 2012 | 2012-09-12 | Emeryville, San Francisco Bay, CA, California, USA | Non-native | 37.8396 | -122.3133 |
768280 | Ruiz et al., 2015 | 2013 | 2013-08-15 | Ballena Isle Marina, San Francisco Bay, CA, California, USA | Non-native | 37.7656 | -122.2858 |
768300 | Ruiz et al., 2015 | 2013 | 2013-08-20 | Coyote Point Marina, San Francisco Bay, CA, California, USA | Non-native | 37.5877 | -122.3163 |
768319 | Ruiz et al., 2015 | 2013 | 2013-08-22 | Jack London Square Marina, San Francisco Bay, CA, California, USA | Non-native | 37.7926 | -122.2746 |
768341 | Ruiz et al., 2015 | 2013 | 2013-08-23 | Loch Lomond Marina, San Francisco Bay, CA, California, USA | Non-native | 37.9723 | -122.4829 |
768360 | Ruiz et al., 2015 | 2013 | 2013-08-13 | Oyster Point Marina, San Francisco Bay, CA, California, USA | Non-native | 37.6639 | -122.3821 |
768384 | Ruiz et al., 2015 | 2013 | 2013-08-14 | Redwood City Marina, San Francisco Bay, CA, California, USA | Non-native | 37.5024 | -122.2134 |
768404 | Ruiz et al., 2015 | 2013 | 2013-08-19 | Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA | Non-native | 37.9138 | -122.3522 |
References
Abbott, Donald P.; Lambert, Charles C.; Lambert, Gretchen; Newberry, A. Todd (2007) The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon (4th Edtion), University of California Press, Berkeley, CA. Pp. 949-964Barnes, Robert D. (1983) Invertebrate Zoology, Saunders, Philadelphia. Pp. 883
Blum, Julia C.; Chang, Andrew L.; Liljesthröm, Marcela; Schenk, Michelle E. Steinberg, Mia K.; Ruiz, Gregory M. (2007) The non-native solitary ascidian Ciona intestinalis depresses species richness., Journal of Experimental Marine Biology and Ecology 342: 5-14
Braga, Marcus Davis Andrade et al. (2021) Retirement risks: Invasive coral on old oil platform on the Brazilian equatorial continental shelf, Marine Pollution Bulletin 165(112156): Published online
https://doi.org/10.1016/j.marpolbul.2021.112156
California Department of Fish and Wildlife (2014) Introduced Aquatic Species in California Bays and Harbors, 2011 Survey, California Department of Fish and Wildlife, Sacramento CA. Pp. 1-36
Chang, Andrew Louis (2009) <missing title>, University of California at Davis, Davis CA. Pp. <missing location>
Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>
Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>
Cohen, Andrew N.; Chapman, John T. (2005) <missing title>, San Francisco Estuary Institute, San Francisco. Pp. <missing location>
Cohen, Andrew; and 16 authors. (1998) <missing title>, Washington State Department of Natural Resources, Olympia, Washington. Pp. 1-37
Cordell, Jeffery R.; Levy, Claire; Toft, Jason D. (2012) Ecological implications of invasive tunicates associated with artificial structures in Puget Sound, Washington, USA, Biological Invasions 15(6): 1303-1318
de Rivera, Catherine, and 27 authors (2005) Broad-scale non-indigenous species monitoring along the West Coast in National Marine Sanctuaries and National Estuarine Research Reserves report to National Fish and Wildlife Foundation, National Fish and Wildlife Foundation, Washington, D.C.. Pp. <missing location>
Fairey, Russell; Dunn, Roslyn; Sigala, Marco; Oliver, John (2002) Introduced aquatic species in California's coastal waters: Final Report, California Department of Fish and Game, Sacramento. Pp. <missing location>
Green, Stephanie J. and 7 authors (2021) Broad-scale acoustic telemetry reveals long-distance movements and large home ranges for invasive lionfish on Atlantic coral reefs, Marine Ecology Progress Series 673: 117-134
Hoshino, Z.; Nishikawa, T. (1985) Taxonomic Studies of Ciona intestinalis, Publication of the Seto Marine Biological Laboratory 30(1/3): 61-79
Kott, Patricia (1985) The Australian Ascidiacea Part 1, Phlebobranchia and Stolidobranchia., Memoirs of the Queensland Museum 23: 1-440
Lambert, C. C.; Lambert, G. (1998) Non-indigenous ascidians in southern California harbors and marinas., Marine Biology 130: 675-688
Lambert, Charles C; Lambert, Gretchen (2003) Persistence and differential distribution of nonindigenous ascidians in harbors of the Southern California Bight., Marine Ecology Progress Series 259: 145-161
Lambert, Gretchen (2003) New records of ascidians from the NE Pacific: a new species of Tridemnum, range extension and description of Aplidiopsis pannosum (Ritter, 1899), including its larva, and several non-indigenous species., Zoosystema 24(4): 665-675
Lambert, Gretchen (2005) First North American record of the ascidian Perophora japonica., Journal of the Marine Biological Association of the United Kingdom 85(4): 1011-1012
Lee, Taekjun; Shin, Sook (2014) Morphological and molecular identification of an introduced alien sea squirt (Tunicata: Ascidiacea) in Korea, Proceedings of the Biological Society of Washington 127(1): 284-297
Lord, Joshua P.; Calini, Jeremy M.; Whitlatch, Robert B. (2015) Influence of seawater temperature and shipping on the spread and establishment of marine fouling species, Marine Biology 162: 2481-2492
Monniot, Francoise (2018) Ascidians collected during the Madibenthos expedition in Martinique: 1—Phlebobranchia, Zootaxa 4387(3): 451-472
https://doi.org/10.11646/zootaxa.4387.3.3
Nichols, Claire L.; Lambert, Gretchen; Nydam, Marie L. (2023) Continued persistence of non-native ascidians in Southern California harbors and marinas, Aquatic Invasions 18(1): 1-22
https://doi.org/10.3391/ ai.2023.18.1.101962
Nishikawa, T. (1991) The ascidians of the Japan Sea. II., Publications of the Seto Marine Biological Laboratory 35: 25-170
Nishikawa, Teruaki (1992) The Ascidians of the Japan Sea III., Publications of the Seto Marine Biological Laboratory 35(6): 303-334
Pool, Thomas K.; Luis, Sean; Olden, Julian D. (2013) Assessing lethal dissolved oxygen tolerance for invasive tunicate Ciona savignyi in Puget Sound, Northwest Science 87(2): 106-113
Preisler, Rikke K.; Wasson, Kerstin; Wolff, Wim J.; Tyrrell, Megan C. (2009) Biological Invasions in Marine Ecosystems., Springer-Verlag, Berlin Heidelberg. Pp. 587-617
Rho, Boon Jo; Lee, Ji-Eun (1991) A systematic study of the Ascidians in Korea, Korean Journal of Systematic Zoology 7(2): 195-220
Ruiz, Gregory M.; Geller, Jonathan (2018) Spatial and temporal analysis of marine invasions in California, Part II: Humboldt Bay, Marina del Re, Port Hueneme, and San Francisco Bay, Smithsonian Environmental Research Center & Moss Landing Laboratories, Edgewater MD, Moss Landing CA. Pp. <missing location>
Ruiz, Gregory; Geller, Jonathan (2021) Spatial and temporal analysis of marine invasions: supplemental studies to evaluate detection through quantitative and molecular methodologies, Marine Invasive Species Program, California Department of Fish and Wildlife, Sacramento CA. Pp. 153 ppl.
Silva, Nathan; Smith, William C. (2008) Inverse correlation of population similarity and introduction date for invasive ascidians, PLOSOne 3(6): available online
Simkanin, Christina; Fofonoff, Paul W.; Larson, Kriste; Lambert, Gretchen; Dijkstra, Jennifer A.; Ruiz, Gregory M. (2016) Spatial and temporal dynamics of ascidian invasions in the continental United States and Alaska, Marine Biology 163: Published online
Smith, Kirsty F.; Cahill, Patrick L.; Fidler, Andrew E. (2010) First record of the solitary ascidian Ciona savignyi Herdman, 1882 in the Southern Hemisphere, Aquatic Invasions 5(4): 363-368
Smith, Kirsty F.; Thia, Joshua; Gemmill, Chrissen E. C.; Cary, S. Craig; Fidler, Andrew E. (2012) Barcoding of the cytochrome oxidase I (COI) indicates a recent introduction of Ciona savignyi into New Zealand and provides a rapid method for Ciona species discrimination, Aquatic Invasions 7: in press
U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database. http://collections.nmnh.si.edu/search/iz/
USGS Nonindigenous Aquatic Species Program 2003-2024 Nonindigenous Aquatic Species Database. https://nas.er.usgs.gov/
Van Name, Willard G. (1945) The North and South American ascidians, Bulletin of the American Museum of Natural History 84: 1-462
Zvyaginstev, A. Yu.; Radashevsky, V. I.; Ivin, V. V.; Kashin, I. A.; Gorodkov, A. N. (2011) Nonindigenous species in the far-eastern seas of Russia, Russian Journal of Biological Invasions 2(1): 164-182
Zvyagintsev, A. Yu.; Sanamyan, K. E.; Kashenko, S.D. (2007) On the introduction of the ascidian Ciona savignyi Herdman, 1882 into Peter the Great bay, Sea of Japan., Russian Journal of Marine Biology 33(2): 133-136