Polyandrocarpa zorritensis

Invasion History

First Galapagos Record: 2015

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General Invasion History:

The colonial tunicate Polyandrocarpa zorritensis was described from Zorritos, Peru in 1931 (Van Name 1945) in the southeastern Pacific. The next published collections were from the southeastern Atlantic, in Brazil, at Santos and Itacurussa (near Cananeia) (Millar 1958). The native region of this tunicate could be either the Southwest Atlantic or Southeast Pacific. However, this tunicate has become widely introduced, as it is found on the East and West coasts of North America, Hawaii, Japan, and the Mediterranean. In the Mediterranean and possibly elsewhere, its growth seems to be favored by eutrophication (Brunetti and Mastrototaro 2004).

Invasion History in the Galapagos:

In 2015 and 2016, Polyandrocarpa zorritensis was collected on fouling plates at several sites on Santa Cruz and Baltra Islands (Lambert 2019).

Invasion history elsewhere in the world:

In 1991, the tunicate was found in southern Japan, at Kitakyushu, Kyushu, on the East China Sea, and at Kochi, on Shikoku (Nishikawa et al. 1993, cited by Iwasaki (2006). Polyandrocarpa zorritensis has also invaded the western Mediterranean Sea. In 1975, it was detected in the harbor of La Spezia, Italy, on the Tyrrhenian Sea (Brunetti 1978). In 2001, it was found in the Gulf of Taranto, near the tip of the Italian Peninsula (2001, Brunetti and Mastrototaro 2004). In Spanish waters, P. zorritensis has been found in the delta of the Ebro River (Turon and Perrera 1998). 

Description

Polyandrocarpa zorritensis is a colonial tunicate, whose colonies consist of a crowded mass or cluster of club-shaped zooids. The zooids are connected at the base of the colony by a tangled mass of root-like stolons. Individual zooids have an oval cross-section (meaning they are slightly compressed from side to side; Van Name 1945) and are free at their anterior end. The anterior end is rounded or truncated, and the oral and atrial siphons have four lobes, with two dark bands per lobe (Brunetti and Mastrototaro, 2004). Each zooid has its own tunic, though tunics of adjacent individuals may adhere to each other. The tunics are thin, tough and can be densely encrusted with sand. Zooids are pale brown to yellowish green in color (Lambert and Lambert 1998) and are up to 20 mm long and 3.8 mm diameter (Van Name 1945).

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Taxonomy

Ecology

General:

Life History- A colonial (or compound) tunicate consists of many zooids, bearing many or all of the organs of a solitary tunicate, but modified to varying degrees for colonial life. Colonial tunicates of the genus Polyandrocarpa have zooids resembling solitary tunicates, rounded, oval, or club-like in shape. Zooids can be crowded together, but do not coalesce. Each zooid has an oral and atrial siphon. Water is pumped into the oral siphon, through finely meshed ciliated gills on the pharynx, where phytoplankton and detritus are filtered, and passed on mucus strings to the stomach and intestines. Excess waste is expelled in the outgoing atrial water (Van Name 1945; Barnes 1983). 
 
Colonial tunicates reproduce both asexually, by budding, and sexually, from fertilized eggs developing into larvae. Colonies vary in size and can range from small clusters of zooids to huge spreading masses. The zooids are hermaphroditic, with eggs and sperm being produced by a single individual. 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 life-history of the species, eggs may be externally or internally fertilized. 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 can 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 (Van Name 1945; Barnes 1983). 

Food:

Phytoplankton, detritus

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Marinas & Docks	None
General Habitat	Vessel Hull	None
General Habitat	Mangroves	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

Minimum Temperature (ºC)	12	Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Maximum Temperature (ºC)	30	Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Minimum Salinity (‰)	22.7	Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Maximum Salinity (‰)	38	Field, Mediterranean Sea (Brunetti, pers. comm., cited by Lambert and Lambert 1998)
Minimum Reproductive Salinity	26	Salinities below 26 PSU halted the swimming of larvae (Vazquez and Young 1998)
Maximum Duration	0.2	Larvae, Laboratory observations (Vazquez and Young 1998)
Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Economic Impacts-

Polyandrocarpa zorritensis was reported to overgrow cultured oysters (Crassostrea gigas) in the Ebro Delta, Spain (Perrera et al. 1990, cited by da Rocha et al. 2009).

Ecological impacts-

Polyandrocarpa zorritensis formed 100% cover in several locations in San Diego and Mission Bays in 1994–2000, suggesting it is capable of outcompeting native and other introduced fouling species (Lambert and Lambert 2003).

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References

Brunetti, Riccardo (1978) A colonial ascidian new to the Mediterranean., Vie et Milieu 28-29(4): 647-652

Brunetti, Riccardo; Mastrototaro, Francesco (2004) The non-indigenous stolidobranch ascidian Polyandrocarpa zorritensis in the Mediterranean: description, larval morphology and pattern of vascular budding, Zootaxa 528: 1-8

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

Carman, Mary, and 8 authors (2011) Ascidians at the Pacific and Atlantic entrances to the Panama Canal, Aquatic Invasions 6(4): 371-380

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. (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

Dalby, James E. Jr.; Young, Craig M. (1992) Role of early post-settlement mortality in setting the upper depth limit of ascidians in Florida epifaunal communities, Marine Ecology Progress Series 80: 221-228,

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>

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

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>

Iwasaki, Keiji (2006) Assessment and Control of Biological Invasion Risks., Shoukadoh Book Sellers,and IUCN, Kyoto and Gland, Switzerland. Pp. 104-112

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

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 (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

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

Lopez-Legentil, Susanna; Legentil, Miquel L.; Erwin, Patrick M.; Turon, Xavier (2015) Harbor networks as introduction gateways: contrasting distribution patterns of native and introduced ascidians, Biological Invasions 17: 1623-1638
DOI 10.1007/s10530-014-0821-z

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

Low-Pfeng, Antonio; Recagno, Edward M. Peters (2012) <missing title>, Geomare, A. C., INESEMARNAT, Mexico. Pp. 236

Lutaenko, Konstantin A.; Furota,Toshio; Nakayama, Satoko; Shin, Kyoungsoon; Xu, Jing (2013) <missing title>, Northwest Pacific Action Plan- Data and Information Network Regional Activity Center, Beijing, China. Pp. <missing location>

Mastrototaro, F.; D’Onghia, G.; Tursi, A. (2008) Spatial and seasonal distribution of ascidians in a semi-enclosed basin of the Mediterranean Sea., Journal of the Marine Biological Association of the United Kingdom 88(5): 1053-1061

Millar, R. H. (1958) Some ascidians from Brazil., Annals and Magazine of Natural History 13(1): 497-514

Monniot, Françoise (2018) Ascidians collected during the Madibenthos expedition in Martinique: 2. Stolidobranchia, Styelidae, Zootaxa 2: 291-318

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

Otani, Michio (2006) Assessment and Control of Biological Invasion Risks., Shoukadoh Book Sellers and IUCN, Kyoto, Japan, and Gland, Switzerland. Pp. <missing location>

Pestana, Lueji Barros; Dias, Gustavo Muniz; Marquesa, Antonio Carlos (2017) A century of introductions by coastal sessile marine invertebrates in Angola, South East Atlantic Ocean, Marine Pollution Bulletin 125: 426-a432

Quintanilla, Elena; Thomas Wilke; Ramırez-Portilla, Catalina; Sarmiento, Adriana; Sanchez, Juan A. () , None <missing volume>: <missing location>

Ramalhosa Patrício; Gestoso, Ignacio Rocha Rosana M.; Lambert, Gretchen; Canning-Clode, João (2021) Ascidian biodiversity in the shallow waters of the Madeira Archipelago: Fouling studies on artificial substrates and new records, Regional Studies in Marine Science 43(1901672): Published online

Rodriguez, Laura F.; Ibarra-Obando, Silvia E. (2008) Cover and colonization of commercial oyster (Crassostrea gigas) shells by fouling organisms in San Quintin Bay, Mexico, Journal of Shellfish Research 27(2): 337-343

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.

Stabili, Loredana; Licciano, Margherita; Longo, Caterina; Lezzi, Marco; Giangrande, Adriana (2015) The Mediterranean non-indigenous ascidian Polyandrocarpa zorritensis: Microbiological accumulation capability and environmental implications, Marine Pollution Bulletin 101: 146-152

Streit, Olivia T; Lambert, Gretchen; Erwin, Patrick M.; Lopez-Legentil, Susanna (2021) Diversity and abundance of native and non-native ascidians in Puerto Rican harbors and marinas, Marine Pollution Bulletin 167(112262): Published online

Tobias-Santos, Vitoria; Andreoni-Pham, Rita; El Gharbi, Dany; Lebel, Marie; Tiozzo, Stefano; Alie´, Alexandre (2024) Salinity-mediated limitation of asexual reproduction in the colonial ascidian Polyandrocarpa zorritensis, Frontiers in Ecology and Evolution 12(1332780.): Published online
DOI 10.3389/fevo.2024.1332780

Tracy, Brianna M.; Reyns, Nathalie B. (2014) Spatial and temporal patterns of native and invasive ascidian assemblages in a Southern California embayment, Aquatic Invasions 9: In press

Van Name, Willard G. (1945) The North and South American ascidians, Bulletin of the American Museum of Natural History 84: 1-462

Vazquez, Elsa; Young, Craig M. (1996) Responses of compound ascidian larvae to haloclines., Marine Ecology Progress Series 133: 179-190

Vazquez, Elsa; Young, Craig M. (1998) Ontogenetic changes in phototaxis during larval life of the Ascidian Polyandrocarpa zorritensis, Journal of Marine Biology and Ecology 231: 267-277

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