Phallusia nigra

Invasion History

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

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General Invasion History:

Phallusia nigra was described from the Red Sea in 1816, but now has a strongly disjunct distribution, in the Red Sea, Singapore, and the western Atlantic Ocean (Vandepas et al. 2015). A recent molecular survey has re-identified many Indo-Pacific records of P. nigra as P. philippinensis. [The re-identified records include two coasts of India (Abdul et al. 2007; Ali et al. 2009), Guam (Lambert 2002), Hawaii (1930s, Carlton and Eldredge 2009), and the Mediterranean Sea (1952, Peres 1958, cited by Izquierdo-Muñoz 2009; Shenkar and Loya 2009).] However, Phallusia nigra was collected in the late 19th century in the Western Atlantic off Florida (1st Record, 1869) and was present from the Caribbean south to Brazil by 1921 (Van Name 1921). The phylogeography of the genus supports introduced status in the Western Atlantic. The diversity of the genus is greatest in the Indo-Pacific, and P. nigra is the only Western Atlantic species (James T. Carlton, personal communication). This species is genetically homogenous over the range from Florida to Brazil (Nobrega et al. 2004; Vandepas et al. 2015), suggestive of introduced status, although genetic comparisons have not yet been made with Indian Ocean populations.

North American Invasion History:

Invasion History on the East Coast:

Records from the east coast of Florida are more recent than those from the Gulf coast or the Caribbean, but this may reflect patterns of collecting. Weiss (1948) collected Phallusia nigra in 1943-1946 on fouling plates in Biscayne Bay and Mook found it in the Indian River Lagoon in 1976-1980 (Mook 1983). It is common in the Indian River Lagoon, and elsewhere in southern Florida (Bingham 1992; Smithsonian Marine Station at Fort Pierce 2011; Vandepas et al. 2015).

Invasion History on the Gulf Coast:

Phallusia nigra was first collected off the Dry Tortugas, west of the Florida Keys in the Gulf of Mexico in 1869 (USNM 14478, USNM 7261, US National Museum of Natural History 2010). It was found on the west coast of Florida in 1885, off Clearwater (USNM 7261, US National Museum of Natural History 2010), and in Charlotte Harbor in 1938 (USNM 10530, US National Museum of Natural History 2010).

Invasion History in Hawaii:

Carlton and Eldredge (2015) have removed Phallusia nigra from their list of species introduced to the Hawaiian islands.

Invasion History Elsewhere in the World:

As noted above, it was apparently a very early invader in the western Atlantic Ocean, collected in the Gulf of Mexico in 1869, in Bermuda in 1877, Jamaica, the US Virgin Islands, and Curacao in 1884 (US National Museum 2011; Yale Peabody Museum 2011). It was present in Brazil near the mouth of the Amazon River and at Rio de Janeiro before 1921 (Van Name 1921). It has been found in harbors along much of the Brazilian coast from Ceara State (Lotufo and Oliveira Filho 2010) to Sao Paulo State, from 4 to 23 degrees S (da Rocha et al. 1999).

 

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Description

Phallusia nigra is a solitary tunicate. Its tunic is firm, smooth, and rarely encrusted with algae, debris or other invertebrates. The tunic can be dark brown, blue-black or black, but young individuals may be grey in color. Its shape is oval or elongated, and it attaches to the substrate by its base (posterior) or posterior left side. Its oral siphon is usually bent dorsally, bringing it close to the atrial siphon (Bonnet and Rocha 2011). Phallusia nigra occasionally grows up to 100-110 mm in height, and frequently is larger than 60-70 mm (Van Name 1921, Van Name 1945). With the tunic removed, the body is dark or grey in color with a dense network of longitudinal and transverse muscle bands on the right side. Oral tentacles in a range of sizes surround the aperture, ranging in number from 30-100 (Bonnet and Rocha 2011). The digestive tract is large, covering more than half of the left side (Van Name 1945).

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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, 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	Mangroves	None
General Habitat	Coral reef	None
General Habitat	Vessel Hull	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	19	Field data, da Rocha et al. 1999, São Sebastião, Brazil, 23°53'S, near southern range limit
Maximum Temperature (ºC)	29	Field data, da Rocha et al. 1999, São Sebastião, Brazil, 23°53'S
Minimum Salinity (‰)	24	Field (Brazil, Marins et al. 2010)
Maximum Salinity (‰)	40	Field, typical Red Sea salinity
Maximum Duration	1	Larval duration, Goodbody 1963
Broad Temperature Range	None	Subtropical-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Throughout most of its range, Phallusia nigra is a common and conspicuous fouling organism, known from natural surfaces, such as rocks, mangroves and coral reefs, and man-made structures, such as pilings, buoys, and floats (Van Name 1921; Van Name 1945; Lambert 2002; Woods Hole Ocenographic Institution 1952). Eldredge and Smith (2002) suggest that it may be a competitor with native sessile organisms in Hawaii. However, specific economic and ecological impacts have not been studied or reported.

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References

Ali, H. Abdul Jaffar; Sivakumar, V.; Tamilselvi, M. (2009) Distribution of alien and cryptogenic ascidians along the southern coasts of Indian peninsula, World Journal of Fish and Marine Sciences 1(4): 305-312

Barnes, Robert D. (1983) Invertebrate Zoology, Saunders, Philadelphia. Pp. 883

Berrill, N. J. (1932) Ascidians of the Bermudas, Biological Bulletin 62(1): 77-88

Bingham, Brian L. (1992) Life histories in an epifaunal community: Coupling of adult and larval processes., Ecology 73(6): 2244-2259

Bitar, Ghazi; Kouli-Bitar, Souha (2001) Nouvelles donnees sur la faune et la flore benthiques de la cote libanaise. Migration Lessepsienne, Thalassia Salentina 25: 71-74

Bonnet, Nadia Y. K.; Rocha, Rosana M. (2011) The family Ascidiidae (Herdman) (Tunicata:Ascidiacea) in Bocas del Toro, Panama. Description of six new species., Zootaxa 2864: 1-33
https://pubmed.ncbi.nlm.nih.gov/26167590/

Bonnet, Nadia Yukiji Koto; da Rocha, Rosana Moreira (2011) The Ascidiidae (Ascidiacea: Tunicata) of Coastal Brazil, Zoological Studies 50(6): 809-825

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

Çinar, M. E.; Noglu, M. Bilece; Özturk, B.; Katagan, T. ; Aysel, V. (2005) Alien species on the coasts of Turkey, Mediterranean Marine Science 6/2: 119-146

Çinar, Melih Ertan; Bilecenoglu, Murat; Öztürk, Bilal; Can, Alp (2006) New records of alien species on the Levantine coast of Turkey, Aquatic Invasions 1(2): 84-90

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.; Reath, P. R.; Longenecker, K.; Bolick, Holly; Eldredge, L. G. (2004) <missing title>, Hawai‘i Community Foundation and the U. S. Fish and Wildlife Service, Honolulu. Pp. 1-187

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.; Faria, Suzana B.; Moreno, Tatiane R. (2005) Ascidians from Bocas del Toro, Panama, Caribbean Journal of Science 41(3): 600-612

da Rocha, Rosana Moreira; Lotufo, Tito Monteiro da Cruz; Rodrigues, Sérgio de Almeida (1999) The biology of Phallusia nigra Savigny, 1816 (Tunicata: Ascidiacea) in southern Brazil: Spatial distribution and reproductive cycle, Bulletin of Marine Science 64(1): 77-87

Dias, G. M.; Rocha, R. M.; Lotufo, T. M. C.; Kremer, L. P. (2013) Fifty years of ascidian biodiversity research in Sao Sebastiao, Brazil, Journal of the Marine Biological Association of the United Kingdom 93(1): 273-282

Eldredge, L. G.; Smith, C. M. (2001) Introduced marine species of Hawaii, Bishop Museum Technical Report 21: 1-60

Freestone, Amy L.; Ruiz, Gregory M.; Torchin, Mark E. (2013) Stronger biotic resistance in tropics relative to temperate zone: Effects of predation on marine invasion dynamics, Ecology 94(6): 1370-1377

Goodbody, Ivan (1962) The biology of Ascidia nigra (Savigny). I. Survival and mortality in an adult population, Biological Bulletin 122: 40-51

Goodbody, Ivan (1963) The biology of Ascidia nigra (Savigny). II. The development and survival of young ascidians, Biological Bulletin 124(1): 31-44

Goodbody, Ivan (1965) The biology of Ascidia nigra (Savigny). III. The annual pattern of colonization, Biological Bulletin 129(1): 128-133

Goodbody, Ivan (2004) Diversity and distribution of ascidians (Tunicata) at Twin Cays, Belize, Atoll Research Bulletin 524: 1-22

Ignacio, Barbara L.; Julio, Luciana M.; Junqueira, Andrea O. R; Ferreira-Silva, Maria A. G. (2010) Bioinvasion in a Brazilian Bay: filling gaps in the knowledge of southwestern Atlantic biota, PLOS ONE 5(9): <missing location>

Izquierdo-Muñoz, Andrés; Díaz-Valdés, Marta; Ramos-Esplá, Alfonso A. (2009) Recent non-indigenous ascidians in the Mediterranean Sea., Memoirs of the Museum of Victoria 4(1): 59-64

Kondilatos, Gerasimos; Corsini-Foka, Maria; Pancucci-Papadopoulou, Maria-Antonietta (2010) Occurrence of the first non-indigenous ascidian Phallusia nigra Savigny, 1816 (Tunicata: Ascidiacea) in Greek waters, Aquatic Invasions 5(2): 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

Lindeman, Kenyon C.; McCarthy, Daniel A.; Holloway-Adkins, Karen; Snyder, David B. (2009) <missing title>, CSA International, Inc., Stuart FL. Pp. <missing location>

Locke, Andrea (2009) A screening procedure for potential tunicate invaders of Atlantic Canada., Aquatic Invasions 4(1): 71-79

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; 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

Mook, David (1983) Indian River fouling organisms, a review, Florida Scientist 26(3/4): 162-167

Nobrega, Rejane; Sole-Cava, Antonio M.; Russo, Claudia A. M. (2004) High genetic homogeneity of an intertidal marine invertebrate along 8000 km of the Atlantic coast of the Americas., Journal of Marine Biology and Ecology 303: 173-181

Oliveira Filho, R. R.; Lotufo, T. M. C.; 2010 New records of introduced ascidians at Ceara State harbors, Northern Brazil. <missing URL>



Rocha, Rosana M.; Bonnet, Nadia Y. K.; Baptista, Mariah S.; Beltramin, Fabiele S. (2012) Introduced and native phlebobranch and stolidobranch solitary ascidians (Tunicata: Ascidiacea) around Salvador, Bahia, Brazil, Zoologia 29(1): 39-53

Shenkar, N.; Loya, Y. (2008) The solitary ascidian Herdmania momus: native (Red Sea) versus non-indigenous (Mediterranean) populations., Biological Invasions Published online: <missing location>

Shenkar, N.; Loya, Y. (2008) Aqaba-Eilat- the improbable gulf. Envrionment, biodiversity, and perservation., Magnes Press, Jerusalem. Pp. 197-237

Smithsonian Marine Station at Fort Pierce 2011 Field Guide to the Indian River Lagoon. <missing URL>



Tamilselvi, M. ; Sivakumar, V.; Ali, H. Abdul Jaffar; Thilaga, R. D. (2011) Distribution of alien tunicates (ascidians) in Tuticorin coast, India, World Journal of Zoology 6(2): 164-172

Thessalou-Legaki, M. and 38 authors (2012) New Mediterranean biodiversity records (December 2012), Mediterranean Marine Science 13(2): 312-327

U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database. http://collections.nmnh.si.edu/search/iz/



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

Vandepas, Lauren E.; Oliveira, Livia M.; Lee, Serina S. C.; Hirose, Euichi; Rocha, Rosana M.; Swalla, Billie J. (2015) Biogeography of Phallusia nigra: Is it really black and white?, Biological Bulletin 228: 52-64

Weiss, Charles M. (1948) The seasonal occurrence of sedentary marine organisms in Biscayne Bay, Florida., Ecology 29(2): 153-172

Woods Hole Oceanographic Institution (1952) Marine fouling and its prevention., United States Naval Institute., Washington, D.C.. Pp. 118-164

Yale Peabody Museum of Natural History 2008-2016 YPM Invertebrate Zoology - Online Catalog. <missing URL>

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