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Zootaxa 2995: 64–68 (2011) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2011 · Magnolia Press ISSN 1175-5334 (online edition)

Parazoanthus Haddon & Shackleton, 1891, and Delage & Hérouard, 1901: Conservation of usage by Reversal of Precedence with Bergia Duchassaing & Michelotti, 1860, and Bergiidae Verrill, 1869 (: : Hexacorallia)

MARTYN E. Y. LOW1 & JAMES DAVIS REIMER2,3 1Department of Marine and Environmental Sciences, Graduate School of Engineering and Science, University of the Ryukyus, 1 Sen- baru, Nishihara, Okinawa 903-0213, Japan. E-mail: [email protected] 2Molecular Invertebrate Systematics and Ecology Laboratory, Rising Star Program, Trans-disciplinary Organization for Subtropical Island Studies, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903-0213, Japan; Marine Biodiversity Research Program, Institute of Biogeosciences, Japan Agency for Marine-Earth Science and Technology (JAMSTEC), 2-15 Natsushima, Yokosuka, Kana- gawa 237-0061, Japan. E-mail: [email protected] 3Corresponding author

Abstract

The names Bergia Duchassaing & Michelotti, 1860, and Bergiidae Verrill, 1869, are respectively, senior subjective syn- onyms of Haddon & Shackleton, 1891, and Parazoanthidae Delage & Hérouard, 1901. The junior synonyms Parazoanthus and Parazoanthidae are in current and widespread use. In the interest of nomenclatural stability, we enact Articles 23.9.1 and 23.9.2 to reverse precedence of these names, thereby making Parazoanthus and Parazoanthidae nomi- na protecta, and Bergia and Bergiidae nomina oblita.

Key words: , Zoanthidea, Article 23.9.1, Article 23.9.2, ICZN, nomenclature

The family Parazoanthidae Delage & Hérouard, 1901, and its type Parazoanthus Haddon & Shackleton, 1891, are a group of zoanthids frequently epizoic on sponges or other benthic organisms (see Reimer & Sinniger 2010: 253). The genus Parazoanthus was established by Haddon & Shackleton (1891a: 633) with the type species Palythoa axinellae Schmidt, 1862, by original designation. In addition to the type species, at least nine other species are currently included in the genus (Appeltans et al. 2011): Parazoanthus anguicomus (Norman, 1869), Parazoanthus capensis Carlgren, 1938, Parazoanthus catenu- laris (Duchassaing & Michelotti, 1860), Parazoanthus darwini Reimer & Fujii, 2010, Parazoanthus dichroicus Haddon & Shackleton, 1891b, Parazoanthus elongatus McMurrich, 1904, Parazoanthus parasiticus (Duchassaing & Michelotti, 1860), Parazoanthus puertoricense West, 1979, and (Duchassaing & Michel- otti, 1860) (Sinniger et al. 2010; Swain 2009). The family Parazoanthidae was established by Delage & Hérouard (1901: 665), with only the type genus included. Parazoanthidae now contains five genera (see Sinniger et al. 2010: 58). The family Parazoanthidae and the genus Parazoanthus are still undergoing revision (discussed in Sinniger et al. 2010), and the exact number of species in Parazoanthus is still a matter of debate. The genus Bergia was established by Duchassaing & Michelotti (1860: 54) for two new species, Bergia catenularis and Bergia vialactea. The family Bergiidae was established by Verrill (1869: 494), with the inclusion of only the type genus. Duerden (1903: 496) considered the two species of Bergia to be conspecific, using Bergia catenularis in favour of Bergia vialactea, and transferred Bergia catenularis to the genus Parazoanthus Haddon & Shackleton, 1891. The names Bergia Duchassaing & Michelotti, 1860, and Bergiidae Verrill, 1869, are thus respec- tively, senior subjective synonyms of Parazoanthus Haddon & Shackleton, 1891, and Parazoanthidae Delage & Hérouard, 1901.

64 Accepted by M. Daly: 29 Jul. 2011; published: 15 Aug. 2011 The family Parazoanthidae and the genus Parazoanthus are in current and widespread use, with new genera and species of Parazoanthus having been recently described (e.g. Reimer & Fujii 2010; Sinniger et al. 2010). How- ever, the Principle of Priority (Article 23 of the International Code of Zoological Nomenclature, hereafter the Code, ICZN 1999: 24) requires that the oldest available name for the taxon under consideration must be used. Replacing the names Parazoanthus and Parazoanthidae with Bergia and Bergiidae, respectively, is not in the interest of nomenclatural stability. To mediate the Principle of Priority, Article 23.9.1 of the Code (ICZN 1999: 27) allows for a reversal of precedence of a junior synonym when the senior synonym has not been used as a valid name after 1899 (Article 23.9.1.1) and the junior synonym “has been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encom- passing a span of not less than 10 years” (Article 23.9.1.2). Since 1899, the name Bergia has been used in a small number of publications. Duerden (1900: 205) used the name twice; firstly as it was being used by workers at that point in time (without actually considering it valid), and secondly to note that Bergia was undoubtedly synonymous with Paraozoanthus. In Duerden (1903: 495, 496), the two species of Bergia are synonymized (with doubt regarding the family name Bergiidae being raised), and the spe- cies is transferred to Parazoanthus, again clearly showing that Duerden did not consider Bergia to be valid. Bergia was also mentioned in a footnote in Delage & Hérouard (1901:667), but this genus was described as “impossibles a classer” [= “impossible to classify”], and again this is not a clear usage of Bergia as a valid name. Pax (1910: 299), in a list of species of Parazoathus species, which he considered to be valid, listed “P. catenularis (Duch. & Mich.)” as a valid species, and briefly discussed that this species was originally described in the genus Bergia, with no dis- cussion about the latter’s validity. Finally, Volpi & Benvenuti (2003:66) reported on the collection of Duchassaing and Michelotti, and noted that a syntype of Bergia catenularis is extant. However, the same authors specifically stated that they were using the “original names”, with no effort being made to ascertain the valid names for the material (Volpi & Benvenuti 2003: 53). As Ng & Low (2010:37, 38) have argued, “valid usage” of a name must be unambiguous and show clearly that the author both considered it the correct name to be used and adopted the name. Duerden (1900, 1903) and Delage & Hérouard (1901) did not use Bergia and Bergiidae as valid names, and Volpi & Benvenuti (2003) simply reported on a specimen, and made no statement on the validity of Bergia. Thus, the names Bergia Duchassaing & Michelotti, 1860, and Bergiidae Verrill, 1869, have not been used as the valid names for the taxa they denote since 1899 (thus fulfilling Article 23.9.1.1 of the Code). In the Appendix, we list 30 publications, by 69 authors in the past 33 years in which the names Parazoanthus Haddon & Shackleton, 1891, and Parazoanthidae Delage & Hérouard, 1901, have been used as valid names for the taxa they denote (thus fulfilling Article 23.9.1.2 of the Code). As both requirements of Article 23.9.1 are met, and in accordance with Article 23.9.2, the names Parazoanthus Haddon & Shackleton, 1891, and Parazoanthidae Delage & Hérouard, 1901, are considered valid names and take precedence over Bergia Duchassaing & Michelotti, 1860, and Bergiidae Verrill, 1869. Parazoanthus Haddon & Shackleton, 1891, and Parazoanthidae Delage & Hérouard, 1901, now become nomina protecta and Bergia Duch- assaing and Michelotti, 1860, and Bergiidae Verrill, 1869, nomina oblita. If future research demonstrates that Parazoanthus catenularis is not referable to the genus Parazoanthus, the genus name Bergia and the family name Bergiidae remain available as outlined in Article 23.9.2 of the Code (ICZN 1999: 28) that allows “in the case of subjective synonymy, whenever the names are not regarded as syn- onyms the older name may be used as valid”.

Acknowledgments

Carey Ashworth (University of the Ryukyus, Japan) assisted in translating literature. The senior author was sup- ported in part by the Rising Star Program at the University of the Ryukyus.

References

Appeltans, W., Bouchet, P., Boxshall, G.A., Fauchald, K., Gordon, D.P., Hoeksema, B.W., Poore, G.C.B., van Soest, R.W.M.,

CONSERVATION OF THE NAME PARAZOANTHUS Zootaxa 2995 © 2011 Magnolia Press · 65 Stöhr, S., Walter, T.C. & Costello, M. (eds.) (2011). World Register of Marine Species. Accessed at http://www.marinespe- cies.org on 2011-07-14. Carlgren, O. (1938) South African Actiniaria and Zoantharia. Kungliga Svenska vetenskapsakademiens handlingar, ser. 3, 17 (3), 1–148, figs. 1–83, pls. 1–3. Daly, M., Brugler, M.R., Cartwright, P., Collins, A.G., Dawson, M.N., Fautin, D.G., France, S.C., McFadden, C.S., Opresko, D.M., Rodriguez, E., Romano, S.L. & Stake, J.L. (2007) The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus. Zootaxa, 1668, 127–182. Delage, Y. & Hérouard, E. (1901) Zoanthidés. – Zoanthidae. Pp. 654–667, figs. 907–929, pls. 65, 66. In: Traité de Zoologie concrète. Tome II – 2e Partie. Les Coelentérés. C. Reinwald, Paris, x + [1] + 848 pp., figs. 1–1102, pls. 1–72. Duchassaing, P. de F. & Michelotti, J. (1860) Mémoire sur les Coralliaires des Antilles. Imprimerie Royale, Turin, 112 pp., pls. 1–11. [Note: This is separate of a publication with identical title and text in the Memorie della Reale Accademia delle Sci- enze di Torino, ser. 2, vol. 19, pp. 279–365, that appeared in 1861] Duerden, J.E. (1900) Jamaican Actiniaria. Part II – Stichodactylinae and Zoantheae. Scientific Transactions of the Royal Dublin Society, ser. 2, 7(6), 133–220, pls. 10–15. Duerden, J.E. (1903) West Indian sponge-incrusting [sic] actinians. Bulletin of the American Museum of Natural History, 19 (18), 495–503, pls. 44–47. Haddon, A.C. & Shackleton, A.M. (1891a) The Zoantheae. In: A revision of the British Actiniae. Part II. Reports on the zoo- logical collections made in the Torres Straits by A. C. Haddon, 1888–1889. Scientific Transactions of the Royal Dublin Society, ser. 2, 4(12), 609–672, pls. 58–60. [Published November 1891] Haddon, A.C. & Shackleton, A.M. (1891b) Actiniae: I. Zoantheae. In: Reports on the Zoological Collections Made in the Tor- res Straits by Professor A. C. Haddon, 1888–1889. Scientific Transactions of the Royal Dublin Society, ser. 2, 4(13), 673– 658, pls. 61–64. [Published December 1891] ICZN, International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edi- tion. The International Trust for Zoological Nomenclature and The Natural History Museum, London, xxix + 306 pp. McMurrich, J.P. (1904) The Actiniae of the Plate Collection. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 6(2), 215–306, figs. 1–5, pls. 14–19. Ng, P.K.L. & Low, M.E.Y. (2010) On the generic nomenclature of nine brachyuran names, with four replacement names and two nomina protecta (Crustacea: Decapoda). Zootaxa, 2489, 34-46. Norman, A.M. (1869) Shetland Final Dredging Report—Part II. On the Crustacea, Tunicata, Polyzoa, Echinodermata, Actino- zoa, Hydrozosa, and Porifera. In: Last report on dredging among the Shetland Isles. Report of the thirty-eight meeting of the British Association for the Advancement of Science; held at Norwich in August 1868. John Murray, London, pp. 247– 336. Pax, F. (1910) Aktinien der Aru-Inseln. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 33, 299-304, figs. 1-4. Reimer, J.D. & Fujii, T. (2010) Four new species and one new genus of zoanthids (Cnidaria, Hexacorallia) from the Galápagos Islands. ZooKeys, 42, 1–36, figs. 1–9, tables 1–3. Reimer, J.D. & Sinniger, F. (2010) Unexpected diversity in Canadian Pacific zoanthids (Cnidaria: Anthozoa: Hexacorallia): a molecular examination and description of a new species from the waters of British Columbia. Marine Biodiversity, 40(4): 249–260, figs. 1–4, tables 1–3. Schmidt, O. (1862) Die Spongien des Adriatischen Meeres. Wilhelm Engelmann, Leipzig, viii + 88 pp., pls. 1–7. Sinniger, F., Reimer, J.D. & Pawlowski, J. (2010) The Parazoanthidae (Hexacorallia: Zoantharia) DNA : description of two new genera. Marine Biodiversity, 40(1), 57–70, figs. 1–5, table 1. Swain, T.D. (2009) Phylogeny-based species delimitations and the evolution of host associations in symbiotic zoanthids (Anthozoa, Zoanthidea) of the wider Caribbean region. Zoological Journal of the Linnean Society, 156(2), 223–238, figs. 1–3, table 1. Verrill, A.E. (1867–1871) Notes on the Radiata in the Museum of Yale College, with descriptions of new genera and species. Transactions of the Connecticut Academy, 1(2), 247–611, pl. 4–9. Volpi, C. & Benvenuti, D. (2003) The Duchassaing & Michelotti collection of Caribbean corals: status of the types and location of the specimens. Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale de Milano, 144(1), 51–74, fig. 1. West, D.A. (1979) Symbiotic zoanthids (Anthozoa: Cnidaria) of Puerto Rico. Bulletin of Marine Science, 29(2), 253–271, figs. 1–7.

66 · Zootaxa 2995 © 2011 Magnolia Press LOW & REIMER Appendix

A list of 30 publications by 69 different authors published in the last 33 years using Parazoanthus Haddon & Shackleton, 1891, and Parazoanthidae Delage & Hérouard, 1901, as valid names to fulfil the requirements of Article 23.9.1.2 of the Code (ICZN 1999: 28). Acosta, A., Casas, M., Vargas, C.A. & Camacho, J.E. (2005) Lista de Zoantharia (Cnidaria: Anthozoa) del Caribe y de Colom- bia. Biota Colombiana, 6(2), 147–162. [pp. 147, 148, 150, 154, 159–161] Bibiloni, M.A. & Cornet, C. (1982) Estudio faunístico del litoral de Blanes: III. Sistemática de cnidarios, briozoos y equinoder- mos. Miscel·lània Zoològica, 6, 19–25. [p. 21] Burnett, W.J., Benzie, J.A.H., Beardmore, J.A. & Ryland, J.S. (1997) Zoanthids (Anthozoa, Hexacorallia) from the Great Bar- rier Reef and Torres Strait, Australia: systematics, evolution and a key to species. Coral Reefs, 16(1), 55–68. [p. 57, 58, 61–64, 66] Cachet, N., Genta-Jouve, G., Regalado, E.L., Mokrini, R., Amade, P., Culioli, G. & Thomas, O.P. (2009) Parazoanthines A–E, hydantoin alkaloids from the Mediterranean sea . Journal of Natural Products, 72(9), 1612–1615. [pp. 1612–1614] Daly, M., Fautin, D.G. & Cappola, V.A. (2003) Systematics of the Hexacorallia (Cnidaria: Anthozoa). Zoological Journal of the Linnean Society, 139(3), 419–437. [pp. 424, 425, 427, 428] Fautin, D.G. & Daly, M. (2009) Actinaria, Corallimorpharia, and Zoanthidea (Cnidaria) of the Gulf of Mexico. In: Felder, D.L. & Camp, D.K. (eds.), Gulf of Mexico. Origins, Waters and Biota. Volume 1, Biodiversity. Texas A & M University Press, College Station, Texas, pp. 349–357. [p. 356] Filho, J.M.B., Medeiros, K.C.P., Diniz, M. de F.F.M., Batista, L.M., Athayde-Filho, P.F., Silva, M.S., da-Cunha, E.V.L., Silva Almeida, J.R.G. & Quintans-Júnior, L.J. (2006) Natural products [sic] inhibitors of the enzyme acetylcholinesterase. Revista Brasileira de Farmacognosia, 16(2), 258–285. [p. 274] Fujii, T. & Reimer, J.D. (2011) Phylogeny of the highly divergent zoanthid family Microzoanthidae (Anthozoa, Hexacorallia) from the Pacific. Zoologica Scripta, 40(4), 418–431. [pp. 420, 427, 429] Herrera-Moreno, A. & Betancourt, L. (2002) Especies de anémonas (Coelenterata: Actiniaria: Corallimorpharia, Zoanthidea y Ceriantharia) conocidas para la Hispaniola. Revista Ciencia y Sociedad, 27(3), 439–453. [p. 4] Kružič, P. (2007) Anthozoan fauna of the Telaščica Nature Park (Adriatic Sea, Croatia). Natura Croatica, 16(4), 233–266. [pp. 235, 240, 263, 266] Manuel, R L. (1979) Some new records of Anthozoa from British waters. Journal of Marine Biological Association of the United Kingdom, 59(2), 393–397. [pp. 395, 296] Moreno, A.H. & Fernández, L.B. (2002) Anémonas (Anthozoa: Actinaria, Corallimorpharia, Ceriantharia y Zoanthidea) cono- cidas para la Hispaniola. Ciencia y Sociedad, 27(3), 439–452. [pp. 444, 445] Philipp, N.A. & Fautin, D.G. (2009) Three new species of shallow water, yellow zoanthids (Hexacorallia: Zoanthidea: Epizoan- thidae) from southern California, USA, and southern Australia. Zootaxa, 2058, 53–61. [pp. 54, 57, 58, 60] Previati, M., Palma, M., Bavestrello, G., Falugi, C. & Cerrano, C. (2010) Reproductive biology of Parazoanthus axinellae (Schmidt, 1862) and savaglia (Bertoloni, 1819) (Cnidaria, Zoantharia) from the NW Mediterranean coast. Marine Ecology, 31(4), 555–565. [pp. 555–563] Reimer, J. D. & Fujii, T. (2010) Four new species and one new genus of zoanthids (Cnidaria, Hexacorallia) from the Galápagos Islands. ZooKeys, 42, 1–36. [pp. 1, 3, 4, 6, 9, 10, 11, 14, 15, 18, 19, 21–25, 27, 29–32] Reimer, J.D., Hirose, M., Irei, Y., Obuchi, M. & Sinniger, F. (2011) The sands of time: rediscovery of the genus Neozoanthus (Cnidaria: Hexacorallia) and evolutionary aspects of sand incrustation in brachynemic zoanthids. Marine Biology, 158(5), 983–993. [pp. 985, 986, 988, 989] Reimer, J.D., Nakachi, S., Hirose, M., Hirose, E. & Hashiguchi, S. (2010) Using hydrofluoric acid for morphological investiga- tions of zoanthids (Cnidaria: Anthozoa): a critical assessment of methodology and necessity. Marine Biotechnology, 12(5), 605–617. [pp. 606–615] Reimer, J.D., Nonaka, M., Sinniger, F. & Iwase, F. (2008) Morphological and molecular characterization of a new genus and new species of parazoanthid (Anthozoa: Hexacorallia: Zoantharia) associated with Japanese red coral (Paracorallium japonicum) in southern Japan. Coral Reefs, 27(4), 935–949. [pp. 935–939, 942–947] Reimer, J.D., Ono, S., Sinniger, F. & Tsukahara, J. (2008) Distribution of zooxanthellate zoanthid species (Zoantharia: Antho- zoa: Hexacorallia) in southern Japan limited by cold temperatures. Galaxea, Journal of Coral Reef Studies, 10(2), 57–67. [p. 58] Reimer, J.D. & Sinniger, F. (2010) Unexpected diversity in Canadian Pacific zoanthids (Cnidaria: Anthozoa: Hexacorallia): a molecular examination and description of a new species from the waters of British Columbia. Marine Biodiversity, 40(4), 249–260. [pp. 251–253, 256–259] Reimer, J.D., Sinniger, F. & Hickman, C.P., Jr. (2008) Zoanthid diversity (Anthozoa: Hexacorallia) in the Galapagos Islands: a molecular examination. Coral Reefs, 27(3), 641–654. [pp. 641–653] Searle, P.A. & Molinski, T.F. (1995) 4-dehydroecdysterone, a new ecdysteroid from the zoanthid Parazoanthus sp. Journal of Natural Products, 58(2), 264–268. [pp. 264] Sinniger, F. & Häussermann, V. (2009) Zoanthids (Cnidaria: Hexacorallia: Zoantharia) from shallow waters of the southern Chilean fjord region, with descriptions of a new genus and two new species. Organisms Diversity & Evolution, 9(1), 23–

CONSERVATION OF THE NAME PARAZOANTHUS Zootaxa 2995 © 2011 Magnolia Press · 67 36. [pp. 23–35] Sinniger, F., Montoya-Burgos, J.I., Chevaldonné, P. & Pawlowski, J. (2005) Phylogeny of the order Zoantharia (Anthozoa, Hexacorallia) based on the mitochondrial ribosomal genes. Marine Biology, 147(5), 1121–1128. [pp. 1121–1125, 1127, 1128] Sinniger, F., Reimer, J.D. & Pawlowski, J. (2010) The Parazoanthidae (Hexacorallia: Zoantharia) DNA taxonomy: description of two new genera. Marine Biodiversity, 40(1), 57–70. [pp. 57, 58, 59, 60, 61, 63–69] Swain, T.D. (2009a) antumbrosus, a new species of zoanthid (Cnidaria: Anthozoa: Zoanthidea) symbiotic with Hydrozoa from the Caribbean, with a key to hydroid and sponge-symbiotic zoanthid species. Zootaxa, 2051, 41–48. [pp. 41, 42, 44–47] Swain, T.D. (2009b) Phylogeny-based species delimitations and the evolution of host associations in symbiotic zoanthids (Anthozoa, Zoanthidea) of the wider Caribbean region. Zoological Journal of the Linnean Society, 156(2), 223–238. [pp. 224–226, 230–232, 234, 235] Varela, C., Ortiz, M. & Lalana, R. (2003) Primer registro de Parazoanthidae y Parazoanthus parasiticus (Cnidaria: Anthozoa). Cocuyo, 13, 5. [p. 5] West, D.A. (1979) Symbiotic zoanthids (Anthozoa: Cnidaria) of Puerto Rico. Bulletin of Marine Science, 29(2), 253–271. [pp. 253, 254, 256, 258, 260, 262, 264–267] Won, J.H., Rho, B.J. & Song, J.I. (2001) A phylogenetic study of the Anthozoa (phylum Cnidaria) based on morphological and molecular characters. Coral Reefs, 20(1), 39–50. [pp. 40, 42–47]

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