Invasion HistoryFirst Non-native North American Tidal Record: 2000
First Non-native West Coast Tidal Record: 2000
First Non-native East/Gulf Coast Tidal Record:
General Invasion History:
Polyclinum constellatum was described from Mauritius in 1816. It is widely reported from mangroves, dead corals, rocks, and from man-made structures, such as pilings, floats, buoys, etc. (da Rocha et al. 2010; Carlton and Eldredge 2009). Although its discovery in Mauritius suggests an Indian Ocean origin, native status in the tropical Atlantic cannot be excluded. By the end of the 19th century, it was reported from the Gulf of Mexico (in 1887, Florida), the Caribbean Sea (in 1883, Jamaica), Hawaii (in 1873), and Japan (in 1900) (US National Museum of Natural History 2002; Carlton and Eldredge 2009). We consider it cryptogenic in the western Atlantic, Indian Ocean, and the western Pacific. Polyclinum constellatum is considered introduced in Guam (Lambert 2002; Lambert 2003), Tahiti (Monniot and Monniot 1985), and the Hawaiian Islands (Carlton and Eldredge 2009). It was found at the western entrance to the Panama Canal in 2002-2009 (Ruiz et al., unpublished data; Carman et al. 2010) where Carman et al (2010) considered it cryptogenic. In 2008-2009, it was found in Pacific Mexico, near Mazatlan, at the mouth of the Gulf of California (Tovar-Hernandez et al. 2010). One specimen was identfied on fouling plates in San Diego Bay in 2000 (Ruiz et al., unpublished data).
North American Invasion History:
Invasion History on the West Coast:
In the summer of 2000, a single specimen of Polyclinum constellatum was identified on settlement plates from Navy Pier 14 in San Diego (Ruiz et al., unpublished data). The population status of this species in southern California is not known. In 2008-2009, it was discovered further south, in the Urias estuary, near the port of Mazatlan, Mexico, and in a nearby oyster farm (Tovar-Hernández et al. 2010).
Invasion History on the East Coast:
Van Name (1921; 1945) mentions the occurrence of Polyclinum constellatum on the west coast of Florida, but not on the east coast. However, Weiss (1948) found it on fouling plates in Biscayne Bay in 1944-1946. It was found in Bermuda before 1972 (Monniot 1972). Currently, it is known from waters off Georgia (in 1981, USNM 24319), the Indian River Lagoon (Mook 1983, Ruiz et al. unpublished), and Biscayne Bay (Weiss 1948; Ruiz et al., unpublished data).
Invasion History on the Gulf Coast:
The first collection of Polyclinum constellatum in US waters was in Cedar Key, Florida, in the Gulf of Mexico in 1887 (USNM 6975, US National Museum of Natural History 2010). It was collected in the Dry Tortugas in the 1880s ( USNM 7137, US National Museum of Natural History 2010, exact date unknown); Cedar Key, Florida in 1887; Tampa Bay (Gretchen Lambert 2003, personal communication); and South Padre Island, Texas in 2004 (Lambert et al. 2005). As noted above, we consider this tunicate cryptogenic in the western Atlantic.
Invasion History in Hawaii:
The first Hawaiian record for Polyclinum constellatum is from Oahu in 1873 (Carlton and Eldredge 2009). Subsequently it was reported from Pearl Harbor in 1920 (Coles et al. 1999b, Carlton and Eldredge 2009); Honolulu Harbor in 1997 (Coles et al. 1999a); Ala Wai Harbor in 1998 (Coles et al. 1999a); Kaneohe Bay in 1999 (Coles et al. 2002); and Waikiki in 2001 (Coles et al. 2002). In Hawaii, this tunicate occurs on floats and pilings, but also on dead coral, and under rocks at exposed locations (Carlton and Eldredge 2009).
Invasion History Elsewhere in the World:
Polyclinum constellatum was described from Mauritius in 1816. Although its discovery in Mauritius suggests an Indian Ocean origin, native status in the tropical Atlantic cannot be excluded. Because of this uncertainty, it is considered to be cryptogenic throughout much of its global range. It was reported from Jamaica in 1883 and Japan in 1900 (US National Museum of Natural History 2002; Carlton and Eldredge 2009). Polyclinum constellatum is widely distributed throughout the Caribbean (Van Name 1945) and has also been reported from Tahiti (Monniot and Monniot 1985), New Caledonia (Monniot 1987), Tanzania (Monniot and Monniot 1997) and Brazil (da Rocha and Costa 2005). During 2009, P. constellatum was found on the Atlantic and Pacific sides of the Panama Canal (Carman et al. 2011), where Carman et al (2011) considered it to be cryptogenic.
In 1998, Polyclinum constellatum was collected on man-made substrates in Apra Harbor, Guam (USNM 25073, US National Museum of Natural History 2010; Lambert 2002; Lambert 2003) and we consider it introduced here. In 2016, it was collected from docks in two harbors on Santa Cruz Island in the Galapagos, its first record there (Lambert 2019).
Polyclinum constellatum is a colonial encrusting tunicate. It can be grey, purplish brown or green in color, with white or beige systems of zooids visible on the surface. The attachment area of the colony can be small, so that only a small part of the base is directly attached to the substratum (Van Name, 1945). Colonies are firm and cartilaginous, often without attached debris, but sometimes colonized by hydroids or other epifauna (da Rocha and Costa 2005). Colonies range in size from 25-65 mm long and 5-20 mm thick. The zooids of P. constellatum are about 5-7 mm long when straightened. Zooids are arranged in circular systems, of about 20 zooids each, which border a circular common cloaca (or atrial siphon). The oral siphons are tubular and fringed by six long and triangular lobes (da Rocha and Costa 2005). The reproductive organs are located in the post-abdomen, with the testes forward to the ovaries (Van Name 1945; da Rocha and Costa 2005).
Polyclinum brasiliense (Michaelsen, 1923)
Potentially Misidentified Species
A colonial (or compound) tunicate consists of many zooids, bearing most or all of the organs of a solitary tunicate, but modified to varying degrees for colonial life. Colonial tunicates of the family Polyclinidae have zooids organized around cloacal systems. Each zooid has a thorax, an abdomen and a posterior abdomen. The thorax has an oral siphon and an atrial aperature with an anterior lip, which open to the surface. Below the thorax, the abdomen contains the stomach and intestines, while the posterior abdomen contains the ovaries, testis, and heart. Water is pumped into the oral siphon, through finely meshed ciliated gills along the pharynx, where it is filtered, and passed on mucus strings to the stomach and intestines. Excess waste is then expelled in the outgoing atrial water (Van Name 1945; Barnes 1983).
Colonial tunicates reproduce both asexually by budding and sexually from fertilized eggs that develop into larvae. Buds can form from the body wall of the zooids. Colonies vary in size ranging from small clusters of zooids to huge spreading masses. The zooids are hermaphroditic, which means both eggs and sperm are released into the atrial chamber. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but some species have a partial block to self-fertilization. Eggs are brooded in the atrial chamber until they hatch into lecithotrophic (non-feeding, yolk-dependent) tadpole larvae. The larva has a muscular tail and a notochord, eyespots, and a set of adhesive papillae. The larvae are expelled upon hatching and swim briefly before settlement. Swimming periods are usually less than a day, but some larvae settle immediately after release or swim for longer periods if the water temperature is low. On settlement the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Van Name 1945; Barnes 1983).
|General Habitat||Marinas & Docks||None|
|General Habitat||Coral reef||None|
|Salinity Range||Polyhaline||18-30 PSU|
|Salinity Range||Euhaline||30-40 PSU|
Tolerances and Life History Parameters
|Minimum Temperature (ºC)||17||Field, near southern range limit, Santa Catarina, Brazil (da Rocha et al. 2009)|
|Maximum Temperature (ºC)||29||Field, near southern range limit, Santa Catarina, Brazil (da Rocha et al. 2009)|
|Minimum Salinity (‰)||24||Field, Ilha Grande Bay, Brazil (Marins et al. 2010)|
|Maximum Salinity (‰)||40||Gulf of California, Mazatlan, Mexico (Tovar-Hernandez et al. 2010)|
|Broad Temperature Range||None||Warm temperate-Tropical|
|Broad Salinity Range||None||Polyhaline-Euhaline|
Economic Impacts – Polyclinum constellatum has been reported as fouling on cultured mussels (Perna perna) in Hong Kong (da Rocha et al. 2009).
|Polyclinum constellatum was found fouling mussels (Perna perna) in an aquaculture operation in Santa Catarina, Brazil (da Rocha et al. 2009).|
|Fouling cultured shellfish in Hong Kong (Huang 2003, cited by da Rocha et al. 2009).|
Regional Distribution Map
|Bioregion||Region Name||Year||Invasion Status||Population Status|
|CAR-I||Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida||1887||Crypto||Estab|
|S206||_CDA_S206 (Vero Beach)||1890||Crypto||Estab|
|CAR-VII||Cape Hatteras to Mid-East Florida||1981||Crypto||Unk|
|G330||Lower Laguna Madre||2004||Crypto||Estab|
|NEP-VI||Pt. Conception to Southern Baja California||2000||Def||Unk|
|P020||San Diego Bay||2000||Def||Unk|
|PAN_PAC||Panama Pacific Coast||2008||Crypto||Estab|
|PAN_CAR||Panama Caribbean Coast||1884||Crypto||Estab|
ReferencesAbdul, Jaffar Ali H.; Sivakumar, V. (2007) Occurrence and distribution of ascidians in Vizhinjam Bay (south west coast of India)., Journal of Experimental Marine Biology and Ecology 342: 189-190
Bastida-Zavala, Rolando; de León-González, Jesús Ángel; Carballo Cenizo, José Luis; Moreno-Dávila, Betzabé (2014) [Aquatic Invasive Species in Mexico], Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, <missing place>. Pp. 317-336
Caraballo-Pérez, Virginia; Díaz, Oscar (2011) [Tunicates (Tunicata: Ascidiacea) of the Gulf of Cariaco, Venezuela], Boletino del Instituto Oceanographico de Venezuela 50(2): 233-244
Carlton, James T.; Eldredge, Lucius (2009) Marine bioinvasions of Hawaii: The introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian archipelago., Bishop Museum Bulletin in Cultural and Environmental Studies 4: 1-202
Carman, Mary R. and 7 authors 2010 Ascidians at the Pacific and Atlantic entrances to the Panama Canal. Abstract- International Invasive Sea Squirt Conference-III - Woods Hole, April 26-28 http://www.whoi.edu/page.do?pid=33341&tid=282&cid=72322
Carman, Mary, and 8 authors (2011) Ascidians at the Pacific and Atlantic entrances to the Panama Canal, Aquatic Invasions 6(4): 371-380
Çinar, Melih Ertan and 7 authors (2021) Current status (as of end of 2020) of marine alien species in Turkey, PLOS ONE 16: Published online
Coles S. L., DeFelice R. C., Eldredge, L. G. (1999a) Nonindigenous marine species introductions in the harbors of the south and west shores of Oahu, Hawaii., Bishop Museum Technical Report 15: 1-212
Coles S. L., DeFelice R. C., Eldredge, L. G. (2002b) Nonindigenous marine species at Waikîkî and Hawai`i kai, Oahu, Hawai`i, Bishop Museum Technical Report 25: 1-255
Coles, S. L.; DeFelice, R. C. : Eldredge, L. G. (2002a) Nonindigenous marine species in Kaneohe Bay, Oahu, Hawai`i, Bishop Museum Technical Report 24: 1-364
Coles, S. L.; DeFelice, R. C.; Eldredge, L. G.; Carlton, J. T. (1999b) Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands., Marine Biology 135(1): 147-158
da Rocha, Rosa Morales and 13 authors (2010) Inventory of ascidians (Tunicata, Ascidiacea) from the National Park La Restinga, Isla Margarita, Venezuela, Biota Neotropica 10: published online
da Rocha, Rosana M.; Costa, Luciana (2005) Ascidians (Urochordata: Ascidiacea) from Arriaal do Cabo, Rio de Janeiro, Brazil., Iheringia Series Zoologie 95(1): 57-64
da Rocha, Rosana M.; Kremer, Laura P.; Baptista, Mariah S.; Metri, Rafael (2009) Bivalve cultures provide habitat for exotic tunicates in southern Brazil., Aquatic Invasions 4(1): 195-205
da Rocha, Rosana M.; Kremer, Laura P. (2005) Introduced ascidians in Paranagua Bay, Parana, southern Brazil., Revista Brasileira da Zoologia 22(4): 1170-1184
Darrigran, Gustavo and 29 authora (2023) Species movements within biogeographic regions: exploring the distribution of transplanted mollusc species in South America, Biological Invasions 25: 673-691
DeFelice, Ralph C.; Coles, Steve L.; Muir, David, Eldredge, L. G. (1998) <missing title>, Hawaii Biological Survey, Bishop Museum, <missing place>. Pp. 1-30
Glenner, Henrik; Lützen, Jørgen; , Pacheco-Riaño, Laura Camila; Noever, Christoph (2021) Expansion of the barnacle Austrominius modestus (Darwin, 1854) (Cirripedia, Thoracica, Balanidae) into Scandinavian waters based on collection data and niche distribution modeling, Aquatic Invasions 16: 675-689: <missing location>
Granthom-Costa, Luciana Vieira; Werner Ferreira, Carlos Gustavo; Dias, Gustavo Muniz (2016) Biodiversity of ascidians in a heterogeneous bay from southeastern Brazil, Management of Biological Invasions 7: 5-12
Huang, Xuguang, Bingyu;; Guo, Donghu; Zhong;, Yanping; Li, Shunxing; Liu, Xin;; Laws, Edward A.; Huang, Bangqin (2021) Blackfordia virginica blooms shift the trophic structure to smaller size plankton in subtropical shallow waters, Marine Pollution Bulletin 182(111990): Published online
Lambert, Gretchen (2002) Nonindigenous ascidians in tropical waters., Pacific Science 56(3): 191-298
Lambert, Gretchen (2003) Marine biodiversity of Guam: the Ascidiacea., Micronesica 35-36: 584-593
Lambert, Gretchen (2019) Fouling ascidians (Chordata: Ascidiacea) of the Galápagos: Santa Cruz and Baltra Islands, Aquatic Invasions 14: 132-149
Lambert, Gretchen; Faulkes, Zen; Lambert, Charles C.; Scofield, Virginia L. (2005) Ascidians of South Padre Island, Texas, with a key to species., Texas Journal of Science 57(3): 251-262
Low-Pfeng, Antonio; Recagno, Edward M. Peters (2012) <missing title>, Geomare, A. C., INESEMARNAT, Mexico. Pp. 236
Marins, Flavia O.; Novaes, Roberto L. M.; Rocha, Rosana M.; Junquiera, Andrea O. R. (2010) Non indigenous ascidians in port and natural environments in a tropical Brazilian bay, Zoologia 27(2): 213-222
Monniot, C.; Monniot, F. (1985) [Littoral ascidians of Guadeloupe Island: IX. Characteristics of populations, ecology, relationships with the world fauna] (French), Tethys 11(3-4): 203-213
Monniot, C.; Monniot, F. (1997) Records of ascidians from Bahrain, Arabian Gulf with three new species., Journal of Natural History 31: 1623-1643
Monniot, Claude (1972) [Stolidobranch Ascidians of Bermuda[ (French), Bulletin du Museum National d'Histoire Naturelle. 4e Serie. Section A. Zoologie, Biologie et Ecologie Animales 43: 617-643
Monniot, Claude; Monniot, Francoise (1987) [The Ascidians of French Polynesia] (French), Memoires da Museum d'Histoire Naturelle 136: 106-119
Monniot, Claude; Monniot, Francoise; Griffiths, Charles; Schleyer, Michael (2001) South African Ascidians., Annals of the South African Museum 108(1): 1-141
Monniot, Claude; Monniot, Francoise; Laboutte, Pierre (1985) [Ascidians of the port of Papeete (French Polynesia); Relation to the environment and to intercontinental transport by navigation] (French), Bulletin du Museum National d'Histoire Naturelle. 4e Serie. Section A. Zoologie, Biologie et Ecologie Animales 7(3): 481-495
Monniot, Francoise; Monniot, Claude (1997) Ascidians collected in Tanzania, Journal of East African Natural History 86: 1-35
Montesanto, F.; Chimient, G.; Gissi, C,; Mastrototaro, F, (2022) Polyclinum constellatum (Tunicata, Ascidiacea), an emerging non-indigenous species of the Mediterranean Sea: integrated taxonomy and the importance of reliable DNA barcode data, Mediterranean Marine Science 23(1): 69-83
Mook, David (1983) Indian River fouling organisms, a review, Florida Scientist 26(3/4): 162-167
Stewart, Joan G. (1991) Seaweeds and seagrasses of San Diego County, California Sea Grant College, <missing place>. Pp. <missing location>
Tokioka, Takasi (1967) Pacific Tunicata of the United States National Museum, United States National Museum Bulletin 251: 1-247
Tovar-Hernández, M. A.; Villalobos-Guerrero, T. F.; Yáñez-Rivera, B., Aguilar-Camacho, J. M.; Ramírez-Santana, I. D. (2012) [Guide to exotic aquatic invertebrates in Sinaloa] , Geomare, A. C., USFWS, INE-SEMARNAT, Mazatlán, México. Pp. 41
Tovar-Hernández,María Ana; Suárez-Morales, Eduardo; Yáñez-Rivera, Beatriz (2010) The parasitic copepod Haplostomides hawaiiensis (Cyclopoida) from the invasive ascidian Polyclinum constellatum in the southern Gulf of California, Bullerin of Marine Science 86(3): 637-648
U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database. <missing description>
Van Name, Willard G. (1921) Ascidians of the West Indian region and southeastern United States., Bulletin of the American Museum of Natural History 44: 283-494
Van Name, Willard G. (1945) The North and South American ascidians, Bulletin of the American Museum of Natural History 84: 1-462
Weiss, Charles M. (1948) The seasonal occurrence of sedentary marine organisms in Biscayne Bay, Florida., Ecology 29(2): 153-172