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Global Diversity of Ascidiacea

By Noa Shenkar and Billie J. Swalla


The class Ascidiacea presents fundamental opportunities for research in the fields of development, evolution, ecology, natural products and more. This review provides a comprehensive overview of the current knowledge regarding the global biodiversity of the class Ascidiacea, focusing in their taxonomy, main regions of biodiversity, and distribution patterns. Based on analysis of the literature and the species registered in the online World Register of Marine Species, we assembled a list of 2815 described species. The highest number of species and families is found in the order Aplousobranchia. Didemnidae and Styelidae families have the highest number of species with more than 500 within each group. Sixty percent of described species are colonial. Species richness is highest in tropical regions, where colonial species predominate. In higher latitudes solitary species gradually contribute more to the total species richness. We emphasize the strong association between species richness and sampling efforts, and discuss the risks of invasive species. Our inventory is certainly incomplete as the ascidian fauna in many areas around the world is relatively poorly known, and many new species continue to be discovered and described each year

Topics: Review
Publisher: Public Library of Science
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Provided by: PubMed Central

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  1. (1816) Histoire naturelle des animaux sans verte `bres. Tome III. Tuniciers. Paris: De ´terville.
  2. (1816) Me ´moires sur les Animoux sans Verte `bres. Part 2.
  3. (1843) E ´le ´ments de zoologie.
  4. (1866) Entwicklungsgeschichte der einfachen Ascidien.
  5. (1881) A treatise of comparative embryology.
  6. (1886) Sur la classification des Tuniciers.
  7. (1898) Note sur la classification des Tuniciers.
  8. A (1850) On the anatomy of the fresh water Bryozoa.
  9. (1955). A collection of ascidians from South Africa.
  10. (2009). A first assessment of invasive marine species on Chinese and Korean Coasts. In:
  11. (2001). A global overview of ascidian introductions and their possible impact on the endemic fauna.
  12. (2001). A macho way to make muscles.
  13. (2010). A recent invader, Perophora japonica, in Europe: Temporal genetic change and spatial distribution of genetic diversity in the English Channel. III IISSC Woods Hole,
  14. (1996). A review and classification of fossil didemnid ascidian spicules.
  15. (1991). A test of criteria for introduced species: the global invasion by the isopod Synidotea laevidorsalis (Miers,
  16. (1994). Additions to the inventory of eastern tropical Atlantic ascidians; arrival of cosmopolitan species.
  17. (2009). Adventures of a sea squirt sleuth: unraveling the identity of Didemnum vexillum, a global ascidian invader.
  18. (2010). Alien species in the Mediterranean Sea by 2010. A contribution to the application of European Union’s Marine Strategy Framework Directive (MSFD). Part I Spatial distribution.
  19. (2004). An overview of the clubbed tunicate (Styela clava) in Prince Edward Island.
  20. (2009). An updated 18S rRNA phylogeny of tunicates based on mixture and secondary structure models.
  21. (1969). Anchor Ice Formation in McMurdo Sound, Antarctica, and Its Biological Effects.
  22. (2009). Antarctic ascidians: an isolated and homogeneous fauna.
  23. (2002). Anti-predatory chemical defenses of ascidians: secondary metabolites or inorganic acids?
  24. (1998). Antifouling defense in a subtidal guild of temperate zone encrusting invertebrates.
  25. (2006). Antimalarial activity from three ascidians: an exploration of different marine invertebrate phyla. Trans Roy Soc Trop Med H
  26. (2009). Antitumor potential of natural products from Mediterranean ascidians.
  27. (1990). Antitumor tetrahydroisoquinoline alkaloids from the colonial ascidian Ecteinascidia turbinata.
  28. (2007). Aquaculture of sea-pineapple, Halocynthia roretzi in Japan.
  29. (2001). Are Antarctic suspension-feeding communities different from those elsewhere in the world?
  30. (2004). Ascidiacea (Tunicata) in Australian waters of the Timor and Arafura Seas. Beagle Rec Mus Art Galleries
  31. (1995). Ascidian fauna of the Aegean sea with a check list of the Eastern Mediterranean and Black Sea species. Annales de l’Institut Oceanographique 71:
  32. (2008). Ascidian recruitment patterns on an artificial reef in Eilat (Red Sea).
  33. (1988). Ascidians collected during the International Indian Ocean Expedition.
  34. (1997). Ascidians collected in Tanzania.
  35. (2005). Ascidians from Bocas del Toro,
  36. (1984). Ascidians from Caribbean shallow water localities.
  37. (1956). Ascidians from Mozambique, East Africa.
  38. (1975). Ascidians from the Indo-West-Pacific region in the Zoologlcal Museum,
  39. (2000). Ascidians from the North-Western Pacific region.
  40. (1999). Ascidians from the NW Pacific region.
  41. (1998). Ascidians from the NW Pacific region. 4. Polyclinidae and Placentelidae.
  42. (2003). Ascidians from the outer slope and bathyal western Pacific.
  43. (1965). Ascidians from the tropical coast of west Africa.
  44. (2001). Ascidians from the tropical western Pacific.
  45. (1946). Ascidians in the vicinity of the Portobello Marine Biological Station, Otago Harbour.
  46. (1982). Ascidians of Hong Kong. In: The Marine Flora and Fauna of Hong Kong and Southern China. Hong Kong: Hong Kong
  47. (1950). Ascidians of New Zealand. Part IV. Ascidians in the vicinity of Christchurch.
  48. (1983). Ascidies littorales de Guadeloupe I. Didemnidae. Bull Mus Natl Hist Nat, Paris 4e ser,
  49. (1974). Ascidies littorales et bathyales recoltees au cours de la campagne Biacores: Phlebobranches et Stolidobranches.
  50. (1985). Ascidies littorals de Guadeloupe. IX: Caracte ´ristiques des populations, e ´cologie, rapports avec la faune mondiale.
  51. (1998). Bacterial suspension feeding by coral reef benthic organisms.
  52. (2004). Biochemical composition, energy content and chemical antifeedant and antifoulant defenses of the colonial Antarctic ascidian Distaplia cylindrica.
  53. (2002). Biodiversity as a barrier to ecological invasion.
  54. (2006). Biological synopsis of the colonial tunicates, Botryllus schlosseri and Botrylloides violaceus. Canadian Manuscript Report of Fisheries and Aquatic Sciences.
  55. (2010). Biology, ecology and trials of potential methods for control of the introduced ascidian Eudistoma elongatum in Northland, New Zealand. III IISSC Woods Hole,
  56. (2009). Bivalve cultures provide habitat for exotic tunicates in southern Brazil.
  57. (2006). Caribbean Sea Squirts: The Goodbody collection. The Mona Institute of Applied Sciences,
  58. (1991). Changes in a Red Sea Coral Community Structure: A LongTerm Case History Study. In:
  59. (2008). Clarke A
  60. (2010). Colonization of Patagonian harbours (SW Atlantic) by an invasive sea squirt (Chordata,
  61. (2010). Comparative phylogeography of two colonial ascidians reveals contrasting invasion histories in North America.
  62. (1977). Competition on marine and hard substrata: the adaptive significance of solitary and colonial strategies.
  63. (1991). Coral Reef Ascidians of New Caledonia.
  64. (2008). Deciphering deuterostome phylogeny: molecular, morphological and palaeontological perspectives.
  65. (2006). Deuterostome phylogeny reveals monophyletic chordates and the new phylum Xenoturbellida.
  66. (2001). Distribution and abundance of ascidian assemblages
  67. (2009). Distribution of Alien and Cryptogenic Ascidians along the Southern Coasts of Indian Peninsula.
  68. (2008). Distribution of the sea squirt Ecteinascidia thurstoni Herdman, 1890 (Ascidiacea: Perophoridae) along Suez Canal and Egyptian Red Sea Coasts.
  69. (2004). Diversity and distribution of ascidians (Tunicata) at Twin Cays,
  70. (1993). Ecological roulette: the global transport and invasion of nonindigenous marine organisms.
  71. (2005). Ecology and natural history ofthe protochordates.
  72. (2008). Ecology and systematics of the ascidian fauna in the Gulf of Eilat (Aqaba).
  73. (2002). ET-743: more than an innovative mechanism of action.
  74. (2000). Evolution of the chordate body plan: new insights from phylogenetic analyses of deuterostome phyla.
  75. (2008). Factors influencing the recruitment and abundance of Didemnum in Narragansett Bay, RhodeIsland.
  76. (2005). First North American record of the ascidian Perophora japonica.
  77. (2010). First occurrence of the invasive tunicate Didemnum vexillum in eelgrass habitat.
  78. (2010). First Pacific record of the north Atlantic ascidian Molgula citrina- bioinvasion or circumpolar distribution?
  79. (2010). First record of the non-native light bulb tunicate Clavelina lepadiformis (Mu ¨ller, 1776) in the northwest Atlantic.
  80. (2009). Fossil didemnid ascidian spicule records in the PlioQuaternary marine clastics of the Antalya Basin (Eastern Mediterranean) and their stratigraphic calibration to new nannofossil data.
  81. (2008). Free-living benthic marine invertebrates in Chile.
  82. (1962). Further descriptions of South African ascidians.
  83. (2009). Genetic conspecificity of the worldwide populations of Didemnum vexillum Kott,
  84. (1998). Guia Ilustrado para Identificac ¸a ˜o das Ascı ´dias do Estado de Sa ˜o
  85. (2005). Guide to Exotic Species of San Francisco Bay. San Francisco Estuary Institute.
  86. Hanley SCT (1853) A history of British Mollusca and their shells. 4 vols.
  87. (2008). Heart rate reflects osmotic stress levels in two introduced colonial ascidians Botryllus schlosseri and Botrylloides violaceus.
  88. (1999). Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands.
  89. (2005). Historical introduction, overview, and reproductive biology of the protochordates.
  90. (2011). Introduced and cryptogenic marine and estuarine species of South Africa.
  91. (2004). Introduced and cryptogenic species in Port Phillip
  92. (2009). Introduced aquatic species of the North Sea coasts and adjacent brackish waters. In:
  93. (2005). Introduced ascidians in Paranagua ´ Bay, Parana ´,
  94. (2010). Introduced marine species in South Australia: a review of records and distribution mapping.
  95. (2009). Invasive didemnid tunicate spreading across coral reefs at remote Swains Island, American Sa ¯moa.
  96. (2009). Invasive tunicates on Zeeland and Prince Edward Island mussels, and management practices in The Netherlandss.
  97. (2004). Kowalevsky, comparative evolutionary embryology, and the intellectual lineage of evo-devo.
  98. (1987). Les ascidies de Polynesie francaise. Memoires du Museum national d’Histoire naturelle,
  99. (2002). Linking climate change and biological invasions: Ocean warming facilitates nonindigenous species invasions.
  100. (1996). Long-term changes on coral reefs in booming populations of a competitive colonial ascidian.
  101. Lotufo TMC (2010) New records of introduced ascidians at Ceara State harbors,
  102. (2001). Macho-1 encodes a localized mRNA in ascidian eggs that specifies muscle fate during embryogenesis.
  103. (2009). Marine bioinvasions in the Southern Pacific: status, ecology, economic impacts, conservation and management.
  104. (2010). Maritime Boundaries Geodatabase, version X.
  105. (2010). Mass mortality of cultured ascidians Halocynthia roretzi associated with softening of the tunic and flagellate-like cells.
  106. (2005). Molecular phylogeny of the protochordates: chordate evolution.
  107. (2010). Monitoring ascidians on natural and anthropogenic habitats
  108. (2005). New and little-known species of Didemnidae (Ascidiacea,
  109. (2009). New molecular markers to genetically differentiate populations of Didemnum vexillum - an invasive ascidian species.
  110. (2003). New records of ascidians from the NE Pacific: a new species of Trididemnum, range extension and redescription of Aplidiopsis pannosum (Ritter, 1899) including its larva, and several nonindigenous species.
  111. (2010). Non indigenous ascidians in port and natural environments in a tropical Brazilian bay.
  112. (2009). Non-indigenous ascidians (Chordata: Tunicata) along the Mediterranean coast of Israel.
  113. (1998). Non-indigenous ascidians in southern California harbours and marinas.
  114. (2002). Nonindigenous ascidians in tropical waters.
  115. (2001). Nonindigenous marine invertebrates. In: A Guidebook of Introduced Marine Species
  116. (2006). Observations on non-didemnid ascidians in Australian waters.
  117. (2007). Occurrence and distribution of ascidians in Vizhinjam Bay (south west coast of India).
  118. (2010). Ocean warming increases threat of invasive species in a marine fouling community.
  119. (1984). Osmoregulatory capabilities of three macrosympatric stolidobranch ascidians, Styela clava
  120. (1962). Pe ´re `s JM
  121. (2003). Persistence and differential distribution of nonindigenous ascidians
  122. (2008). Phylogeography of the widespread marine invader Microcosmus squamiger (Ascidiacea) reveals high genetic diversity of introduced populations and non-independent colonisations.
  123. (1987). Phylum Urochordata, Class Ascidiacea. In:
  124. (2008). Population dynamics of a coral reef ascidian in a deteriorating environment.
  125. (1995). Potential antifouling mechanisms using toxic chemicals in some British ascidians.
  126. (2008). Predicting the potential distribution of the vase tunicate Ciona intestinalis in Canadian waters: informing a risk assessment.
  127. (1990). Protochordate biomineralization.
  128. (1993). Ramos-Espla A
  129. (2005). Rapid assessment survey for exotic organisms in southern California bays and harbors, and abundance in port and non-port areas.
  130. (2009). Recent nonindigenous ascidians in the Mediterranean Sea.
  131. (1997). Records of ascidians from Bahrain, Arabian Gulf with three new species.
  132. (1997). Records of ascidians from Bahrain, Arabian Gulf, with three new species.
  133. (1973). Redescription de six ascidies du golfe d’Elat re ´colte ´es par
  134. (2007). Reproduction and early life history of an invasive ascidian Styela clava Herdman in Prince Edward Island,
  135. (2010). Response to trabectedin treatment in a highly pretreated patient with an advanced meningeal hemangiopericytoma.
  136. (1996). Responses of compound ascidian larvae to haloclines.
  137. (2007). Sanamyan E ˆE, Kashenko SD
  138. (1997). Section 6B: Ascidians (Urochordata). Reef and Shore Fauna of Hawaii. Bishop Museum Special Publ.
  139. (1967). Settlement of sessile animals on eternite slabs in two polls near Bergen.
  140. (1992). Some biogeographical remarks about the ascidian littoral fauna of the Straits of Gibraltar (Iberian sector). Bull de l’Inst Oce ´an Monaco 9:
  141. (2007). Some comments on ascidians of New Caledonia. In:
  142. (1964). South African ascidians collected by Th. Mortensen, with some additional material. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, bd.,
  143. (2007). Spatial and temporal patterns of recruitment of the tunicate Ciona intestinalis on a mussel farm in Nova Scotia,
  144. (2010). Species
  145. (1999). Species diversity and invasion resistance in a marine ecosystem.
  146. (2007). Spread of Microcosmus squamiger (Ascidiacea: Pyuridae) in the Mediterranean Sea and adjacent waters.
  147. (2002). Stolidobranch ascidians from the tropical western Indian Ocean.
  148. (2009). Styela clava (Tunicata, Ascidiacea) - a new threat to the Mediterranean shellfish industry?
  149. (2010). Sua ´rez-Morales E, Ya ´n ˜ez-Rivera B
  150. (2010). Survey of harbours in Newfoundland for indigenous and non-indigenous ascidians and an analysis of their cytochrome c oxidase I gene sequences.
  151. (1996). Taxonomy, abundance, and vertical distribution of the ascidians from Tagus Cove, Isabela Island. Charles Darwin Research Station Annual
  152. (1990). The ascidians of the Japan Sea.
  153. (1981). The ascidians of the reef flats of Fiji.
  154. (2001). The Australian Ascidiacea. Part 4: Aplousobranchia (3),
  155. (2010). The biodiversity of the Mediterranean Sea: estimates, patterns, and threats.
  156. (1971). The biology of ascidians.
  157. (2007). The colonial ascidian Didemnum sp. A: Current distribution, basic biology and potential threat to marine communities of the northeast and west coasts of North America.
  158. (1989). The ecology and behavior of ascidian larvae.
  159. (2003). The encyclopedia of life.
  160. (2003). The first tunicate from the Early Cambrian of South China.
  161. (2008). The marine invertebrate species associated with the biofouling of derelict fishing gear in the Papahanoumokuakea–Marine National Monument. A focus on marine nonnative species transport. KaneoheHI: Hawaii Institute for Marine Biology.
  162. (2007). The nature and extent of organisms in vessel sea-chests: A protected mechanism for marine bioinvasions.
  163. (2006). The non-indigenous ascidian Distaplia bermudensis in the Mediterranean: comparison with the native species Distaplia magnilarva and Distaplia lucillae sp nov.
  164. (2007). The nonindigenous ascidian Molgula ficus in California.
  165. (1945). The North and South American ascidians. Bull Am Mus Nat Hist 84: 1–476.
  166. (2008). The solitary ascidian Herdmania momus: native (Red Sea) vs. non-indigenous (Mediterranean) populations.
  167. (2004). The south temperate and Antarctic ascidian Corella eumyota reported in two harbours in northwestern France.
  168. (1950). The Tunicata with an Account of the British Species. London: The Ray Soc.,
  169. (1994). Toxicity and palatability of some British ascidians.
  170. (2010). Trabectedin therapy for sarcomas.
  171. (2009). Tunicata (Urochordata) of the Gulf of Mexico.
  172. (2003). Tunicate feeding filters.
  173. (2006). Tunicates and not cephalochordates are the closest living relatives of vertebrates.
  174. (2007). Washington state 2006 survey for invasive tunicates with records from previous surveys. Updated
  175. (2010). What is natural? The scale and consequences of marine bioinvasions in the North Atlantic Ocean. Van Denderen BV, Groningen, The Netherlandss.
  176. (2003). What is the view from the boat? Atlantic Canadian Tunicate Workshop Proceedings,
  177. (1988). Where are all the fossil sea squirts?
  178. (2010). Widespread occurrence of the southern hemisphere ascidian Corella eumyota Traustedt, 1882 on the Atlantic coast of Iberia.
  179. (2010). World Ascidiacea Database.
  180. (2007). Zoogeography of the Antarctic ascidian fauna in relation to the sub-Antarctic and South America.
  181. (2004). Zoogeography of the southern African ascidian fauna.
  182. (2008). Zoogeography of the southern New Zealand, Tasmanian and southern African ascidian fauna.

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