Abstract
In this study all available species of the nudibranch family Bornellidae (Bornella Gray, 1850, Pseudobornella Baba, 1932) are re-examined anatomically and their status re-evaluated. Of these, B. hancockana Kelaart, 1859 syn. nov., B. arborescens Pease, 1871 syn. nov., B. caledonica Crosse, 1875a syn. nov., and B. marmorata Collingwood, 1881 syn. nov. are considered to be new synonyms of B. stellifer (Adams and Reeve in Adams, 1848), and B. japonica Baba, 1949 syn. nov. is considered to be a new synonym of B. hermanni Angas, 1864. Two species, B. excepta Bergh, 1884 and B. simplex Eliot, 1904, are at present unrecognizable, B. semperi Crosse 1875b is here considered a nomen nudum, and four new species, B. dotoides sp. nov., B. pele sp. nov., B. valdae sp. nov. and B. johnsonorum sp. nov. are proposed. These are compared with a further four recognized species, B. calcarata Mörch, 1863, B. anguilla Johnson, 1984, B. sarape Bertsch, 1980 and B. irvingi Edmunds and Preece, 1996. The presence of an unpaired oral gland in Bornella is reported for the first time. The reproductive system is diaulic, and the morphology of the penis and the arrangement of the penial spines are considered diagnostic for most species. The monotypic genus Pseudobornella is re-examined for the first time. A comparative table for all recognized species is also provided. The review of the morphological differences within the family provides the basis for a phylogenetic analysis of the group. Bornella is shown to represent a monophyletic clade while the monophyly of the family Bornellidae is not supported.References
Adams, A. (1848) Notes from a journal of research into the natural history of the countries visited during the voyage of H.M.S. Samarang, under the command of Captain Sir Edward Belcher, C.B., F. R. A. S. In: Belcher, E. (Ed.), Narrative of the voyage of H.M.S. Samarang during the years 1843–1846. Vol. 2. Reeve, Benham and Reeve, London. pp. 223–532.
Adams, A. & Reeve, L.A. (1848–1850) Mollusca, pt. 1, pp. 1–24, pls. 1–9 [1848]; pt. 2, pp. 25–44, pls. 10–17 [1850]; pt. 3, pp. 45–87, pls. 18–24 [1850]. In: Adams, A. (Eds.), The zoology of the voyage of H.M.S. Samarang; under the command of Captain Sir Edward Belcher, C.B., F.R.A.S., F.G.S., during the years 1843–1846. London, [Nudibranchia pp. 66–68, pls. 17,19].
Adams, M.J. (2003) Bornella anguilla from Burma. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=9438 (22 March 2003), (accessed 26 October 2008)
Adriance, T. (2006) Bornella anguilla from the Philippines. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=16854 (8 July 2006), (accessed 26 October 2008)
Alder, J. & Hancock, A. (1864) Notice of a collection of nudibranchiate mollusca made in India by Walter Elliot Esq., with descriptions of several new genera and species. Transactions of the Zoological Society of London, 5, 113–147.
Allan, J.K. (1950) Australian shells, with related animals living in the sea, in freshwater and on the land. Georgian House, Melbourne. [Nudibranchia pp. 218–226, pl. 28], 470 pp, 44 pls.
Angas, G,F. (1864) Description d'espèces nouvelles appartenant à plusieurs genres de Mollusques Nudibranches des environs de Port-Jackson (Nouvelle-Galles du Sud), accompagnée de dessins faits d'après nature. Journal de Conchyliologie, 12, 43–70.
Aytur, O. (2003) Bornella anguilla from Burma (Myanmar). [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=9488 (2 April 2003), (accessed 26 October 2008)
Baba, K. (1932) Pseudobornella orientalis, nov. gen. et sp. from Japan. Annotationes Zoologicae Japonenses, 13, 369–376.
Baba, K. (1933) Supplementary note on the nudibranchia collected in the vicinity of the Amakusa Marine Biological Laboratry. Annotationes Zoologicae Japonenses, 14, 273–283.
Baba, K. (1937) Opisthobranchia of Japan (I and II). Journal of the Department of Agriculture, Kyushu Imperial University, 5, 289–344.
Baba, K. (1949) Opisthobranchia of Sagami Bay Collected by: His Majesty The Emperor of Japan. Iwanami Shoten, Tokyo, 194 pp., 50 pls.
Beasley, C.J. (2006) Bornella anguilla from Thailand. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=15697 (8 February 2006), (accessed 26 October 2008)
Bergh, L.S.R. (1874) Malacologische Untersuchungen. In: Semper, C. (Eds.), Reisen im Archipel der Philippinen von Dr.Carl Gottfried Semper. Zweiter Theil. Wissenschaftliche Resultate, 2, pp. 287–314, pls. 36–39.
Bergh, L.S.R. (1884) Report on the Nudibranchiata dredged by H.M.S. Challenger during the years 1873–1876. Challenger Reports, Zoology, 10, 1–154, pls. 1–14.
Bergh, L.S.R. (1890) Malacologische Untersuchungen. In: Semper, C. (Eds.), Reisen im Archipel der Philippinen von Dr.Carl Gottfried Semper. Zweiter Theil. Wissenschaftliche Resultate, 2, pp. 886–889, pls. 88–89.
Bergh, L.S.R. (1902) The Danish Expedition to Siam 1899–1900. 1. Gasteropoda Opisthobranchiata. Det Kongelige Danske Videnskabernes Selskabs Skrifter. 6 Raekke. Naturvidenskabelig og Matematisk Afdeling, 12, 153–218.
Bergh, L.S.R. (1905) Die Opisthobranchiata. Siboga Expeditie Report, 50, 248 pp., pls 1–20.
Bertsch, H. (1980) A new species of Bornella from tropical west-America (Mollusca, Opisthobranchia). Spixiana, 3, 33–42.
Bertsch, H. & Kerstitch. A. (1984) Distribution and radular morphology of various nudibranchs (Gastropoda: Opisthobranchia) from the Gulf of California, Mexico. The Veliger, 26, 264–273.
Bidgrain, P. (2005) Bornella anguilla from Reunion Island. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=13094 (16 February 2005), (accessed 26 October 2008)
Burn, R.F. (1965) A centennial commentary and zoogeographical remarks on Angas' Sydney nudibranchs (molluscs, Gastropoda). Journal de Conchyliologie, 104, 85–93.
Coleman, N. (1989) Nudibranchs of the South Pacific, Vol.1. Springwood, Queensland, Sea Australia Resource Centre, 64 pp.
Coleman, N. (2001) 1001 Nudibranchs. Catalogue of Indo-Pacific Sea Slugs. Neville Coleman´s Underwater Geographic Pty Ltd. Springwood, Queensland, Australia, 144 pp.
Collingwood, C. (1868) Observations on the distribution of some species of nudibranchiate Mollusca in the China Sea. Annals & Magazine of Natural History, 4, 90–94.
Collingwood, C. (1879) New species of Nudibranchs from the Eastern seas. Journal of the Linnean Society of London, Zoology, 14, 737–738.
Collingwood, C. (1881) On some new species of nudibranchiate Mollusca from the eastern seas. Transactions of the Linnean Society of London, Zoology, 2, 123–140, pls. 9–10.
Coze, Y. (2003) Bornella anguilla from Reunion Is. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=10608 (31 July 2003), (accessed 26 October 2008)
Crosse, J.C.H. (1875a) Description de nudibranches inédits, provenant de la Nouvelle-Calédonie, avec le catalogue des espèces actuellement connues. Journal de Conchyliologie, 23, 305–322, Pl. 12.
Crosse, J.C.H. (1875b) Note sur les genres Bornella et Placobranchus, accompagnée du catalogue des espèces actuellement connue. Journal de Conchyliologie, 23, 322–328.
Debelius, H. (1998) Nudibranchs and Sea Snails: Indo Pacific Field Guide. M&G Difusión, S.L. Germany, 321 pp.
Debelius, H. & Kuiter, R.H. (2007) Nudibranchs of the world. IKAN-Unterwasserarchiv, Frankfurt, 360 pp.
Edmunds, M. & Preece, R. (1996) A collection of opisthobranch mollusks from the Pitcairn Islands. Journal of Conchology, 35, 407–420.
Eliot, C. (1904) On some nudibranchs from east Africa and Zanzibar. Part V. Proceedings of the Zoological Society of London, 2, 83–105, pls. 3–4.
Eliot, C. (1905) Nudibranchs from the Indo-Pacific. I., Notes on a collection dredged near Karachi and Maskat. Journal of Conchology, 2, 237–238.
Eliot, C. (1913) Japanese nudibranchs. Journal of the College of Sciences, Imperial University of Tokio, 35, 1–47.
Fraser, V. (1999a) Bornella from South Africa. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=1469 (7 November 1999), (accessed 26 October 2008)
Fraser, V. (1999b) Head of Bornella from S. Africa. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=1517 (11 November 1999), (accessed 26 October 2008)
Fraser, V. (2000) Bornella anguilla from South Africa. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=2456 (28 May 2000), (accessed 26 October 2008)
Garthwaite, T. (2002) Bornella stellifer from Taiwan's N.E. Coast. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=7039 (28 May 2002), (accessed 26 October 2008)
Gosliner, T.M. (1987a) Nudibranchs of Southern Africa. A guide to Opisthobranch Molluscs of Southern Africa. Sea Challengers, Monterey, California, 136 pp.
Gosliner, T.M. (1987b) Review of the Nudibranch Genus Melibe (Opisthobranchia: Dendronotacea) with descriptions of two new species. The Veliger, 29, 400–414.
Gosliner, T.M., & Bertsch, H. (1985) Records and morphology of Lomanotus stauberi Clark & Goetzfried, 1976, from the Panamic Pacific. Veliger, 27, 397–405.
Gosliner, T.M., Behrens, D. & Willians, G. (1996) Coral reef animals of the Indo-Pacific. Sea Challengers, Monterey, California, 314 pp.
Gosliner, T.M. & Smith, V. (2003) Systematics and phylogenetic analysis of the nudibranchs genus Melibe (Opisthobranchia: Dendronotacea) with descriptions of three new species. Proceedings of the California Academy of Sciences, 54, 302–355.
Gray, M.E. (1850) Figures of molluscous animals, selected from various authors, etched for the use of students, vol. 2, pls. 79–199. Longman, Brown, Green and Longmans, London. [Nudibranchia pp. 14–15, 37–46, 52, 102–110].
Hayashi, S. (2002) Pseudobornella orientalis from the Japan Sea. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=7453 (15 July 2002), (accessed 26 October 2008)
Henke, M. (2006) Bornella anguilla swimming. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=18621 (29 November 2006), (accessed 26 October 2008)
Hermosillo, A. (2002) Bornella sarape from Mexico. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=7179 (15 June 2002), (accessed 26 October 2008)
Hermosillo, A., Behrens, D.W. & Rios Jara, E. (2006) Opisthobranquios de Mexico. Guia de babosas marinas del Pacifico, Golfo de California y las islas oceanicas. CONABIO, Guadalajara, Jalisco, Mexico, 144 pp.
Ianniello, L. (2002) Bornella calcarata from St. Vincent Is. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=7982 (24 September 2002), (accessed 26 October 2008)
Imamoto, J. (2008) Bornella stellifer [In] Umiushi. Available from: http://www.umiushi.info/. (accessed 22 May 2008)
Johnson, S. (1984) A new Indo-West Pacific species of the dendronotacean nudibranch Bornella (Mollusca: Opisthobranchia) with anguilliform swimming behavior. Micronesica, 19, 17–26.
Johnson, S. (2007) Bornella japonica? from the Marshall Islands. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=19286 (25 May 2007), (accessed 26 October 2008)
Kelaart, E.F. (1859) On some additional species of nudibranchiate mollusks from Ceylon. Annals & Magazine of Natural History, 4, 267–270.
Kenny, R. (1970) Queensland faunistic records. Part IX. A second collection of opisthobranch mollusks from Queensland. University Queensland Papers, Department of Zoology, 3, 83–96.
Kinoshita, H. (2002) Pseudobornella orientalis from Izu Peninsula, Japan.. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=6851 (14 May 2002), (accessed 26 October 2008)
Kitagawa, S. (2004) Pseudobornella orientalis from Philippines. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=12463 (19 March 2004), (accessed 26 October 2008)
Koh, D.B. (2006) Sea Slugs of Korea. Published by Pungdeung, Korea, 248 pp.
Krampf, M. (2006) Bornella anguilla from Sulawesi. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=18590 (29 November 2006), (accessed 26 October 2008)
Kurihara, T. (2002) Pseudobornella orientalis from Japan. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=6760 (1 May 2002), (accessed 26 October 2008)
Kurihara, T. (2004) Re: Pseudobornella orientalis from Philippines. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=12521 (25 March 2004), (accessed 26 October 2008)
Maddison, W.P. & Maddison, D.R. (2005) MacClade, Version 4.08. Sunderland, Massachusetts: Sinauer Associates.
MacFarland, F.M. (1923) The morphology of the nudibranch genus Hancockia. Journal of Morphology, 38, 65–104, pls. 1–6.
MacFarland, F.M. (1966) Studies of opisthobranchiate mollusks of the Pacific coast of North America. Memoirs of the California Academy of Sciences, 6, 1–546, pls. 1–72.
Marcus, Er. (1958) On western Atlantic opisthobranchiate gastropods. American Museum Novitates, 1906, 1–82.
Marcus, Ev. (1976) Opisthobranchia von Santa Marta, Colombia. Studies on the Neotropical Fauna and Environment, 11, 119–150.
Marcus, Ev. (1977) An annotated checklist of the western Atlantic warm water opisthobranchs. The Journal of Molluscan Studies, supplement 4, 1–23.
Marcus, Ev. & Marcus, Er. (1967) Tropical American opisthobranchs. Studies in Tropical Oceanography, 6, 3–137.
Marshall, J.G. & Willan, R.C. (1999) Nudibranchs of Heron Island, Great Barrier Reef. A Survey of the Opisthobranchia (Sea Slugs) of Heron and Wistari Reef. Backhuys Publisher, PO Box 321, 2300 AH Leiden, The Netherlands.
Masayoshi, N. (2002) Bornella stellifer mating. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=7728 (9 October 2002), (accessed 26 October 2008)
Mörch, O. (1863) Contributions à la faune malacologique des Antilles Danoises. Journal de Conchyliologie, 3, 21–43.
Nakano, R. (2001) Bornella japonica? from Hachijo Is. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=5499 (1 November 2001), (accessed 26 October 2008)
Nakano, R. (2004) Opisthobranchs of Japan Islands. Rutles Inc., Tokyo, Japan. 304 pp.
Nishina, M. (2001a) Bornella japonica? from Japan. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=5214 (12 September 2001), (accessed 26 October 2008)
Nishina, M. (2001b) Bornella japonica? from Japan. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=5572 (1 November 2001), (accessed 26 October 2008)
Nijssen-Meyer, J. (1965) Notes on a few opisthobranch Mollusca from Surinam (Guianas). Zoologische Mededelingen uitgegeven door het Rijksmuseum van Natuurlijke Historie te Leiden, 40, 143–150.
Odhner, N.H. (1936) Nudibranchia Dendronotacea—A revision of the system. Mémoires du Musee Royal d'Histoire Naturelle de Belgique, 2, 1057–1128, pl. 1.
Okutani, T. (2000) Marine Mollusks in Japan. Tokay University Press, Tokyo, Japan, 1173 pp.
Orr, J. (1981) Hong Kong Nudibranchs. Urban Council Publisher. Hong Kong, 82 pp.
Pease, W.H. (1871) Descriptions of nudibranchiate Mollusca inhabiting Polynesia. American Journal of Conchology, 6, 299–305, pls. 19–22.
Pola, M. & Scout, C. (In press) Description of the first two tropical Indo-Pacific species of Dendronotus (Gastropoda: Nudibranchia) with the re-description of the poorly known species Dendronotus gracilis Baba, 1949. Zootaxa.
Preece, R.C. (1995) The composition and relationships of the marine molluscan fauna of the Pitcairn Islands. Biological Journal of the Linnean Society, 56, 339–358.
Pruvot-Fol, A., 1930. Diagnose provisoires (incomplètes) des espèces nouvelles et liste provisoire des mollusques nudibranches recueillis par Mme. A. Pruvot-Fol en nouvelle Calédonie (Ile des Pins). Bulletin Muséum National d'Histoire Naturelle Paris, series 2, 2, 229–232.
Risbec, J. (1928) Contribution a l'etude des nudibranches Néo-Calédoniens. Thèse présentée à la faculté des sciences de l'Université de Paris pour obtenir le grade de docteur ès-sciences naturelles. Faune des Colonies Française, 2, 1–328, pls. 1–12.
Risbec, J., (1930) Nouvelle contribution à l'étude des nudibranches Néo-Calédoniens. Annales de l'Institut Océanographique (Monaco) Paris, (N.S.), 7, 263–298, Text-figs 1–69, Pl.1.
Risbec, J. (1932) Notes sur la ponte et le développment de mollusques gastéropodes de Nouvelle-Calédonie. Bulletin de la Société Zoologique de France, 57, 358–374.
Risbec, J. (1953) Mollusques nudibranches de la Nouvelle-Calédonie. Faune de l'Union Française Paris, Libraire Larose, 15, 1–189.
Rudman, W.B. (1984) The date and authorship of Bornella and Ceratosoma (Nudibranchia) and other molluscs collected during the voyage of H.M.S. Samarang 1843–1846. Malacological Review, 17, 103–104.
Rudman, W.B. (1998) Suborder Dendronotina. In: Beesley, P.L., Ross, G.J.B. & Wells, A. (Eds.), Mollusca: The Southern Synthesis. Fauna of Australia, Vol 5. CSIRO Publishing: Melbourne. Part B, pp. 1001–1007.
Rudman, W.B. (2000) Dendronotus sp. 1 [In] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/factsheet.cfm?base=dendron (22 March 2000), (accessed 26 October 2008)
Schrödl, M. (1999) Hancockia schoeferti, spec. nov., a new dendronotoidean nudibranch species from central Chile. Spixiana, 22, 247–254.
Smith, E.A. (1884) Report on the zoological collections made in the Indo-Pacific Ocean during the voyage of H.M.S. 'Alert' 1881-2. The Collections from Melanesia. Mollusca. Trustees British Museum, London. [Nudibranchia pp. 90–92, 506]
Smith, V. & Gosliner, T.M. (2003) A new species of Tritonia from Okinawa (Mollusca: Nudibranchia), and its Association with a Gorgonian Octocoral. Proceedings of the California Academy of Sciences, 54, 255–278.
Smith, V. & Gosliner, T.M. (2005) Two new species of Marionia (Mollusca: Nudibranchia) from the Indo-Pacific Region. The Veliger, 48, 260–275.
Suzuki, K. (2000) Opistobranchs of Izu Peninsula. TSB-Britannica, Tokyo, 178 pp.
Swofford, D.L. (2001) PAUP*. Phylogenetic analysis using parsimony (*and other methods)Ó. Version 4. Sinauer Associates, Sunderland, Massachussetts.
Taylor, D. (2001) Bornella anguilla from North West Australia. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from: http://www.seaslugforum.net/find.cfm?id=4331 (23 May 2001), (accessed 26 October 2008)
Thompson, T.E. (1961) The structure and mode of functioning of the reproductive organs of Tritonia hombergi (Gastropoda Opisthobranchia). Quarterly Journal of Microscopical Science, 102, 1–14.
Thompson, T.E. (1980) Jamaican opisthobranch molluscs II. Journal of Molluscan Studies, 46, 74–99.
Tucker, K. (2001) Bornella anguilla from Fiji. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from http://www.seaslugforum.net/find.cfm?id=4824 (18 July 2001), ( accessed 26 October 2008)
Valdés, Á., Hamann, J., Behrens, D. & Dupont, A. (2006) Caribbean Sea Slugs. A field guide to the opisthobranch mollusks from the tropical northwestern Atlantic. Sea Challengers Natural History Books, Etc. Gig Harbor, Washington, USA, 289 pp.
Vayssière, A. (1912) Recherches zoologiques et anatomiques sur les Opisthobranches de la Mer Rouge et du Golfe d'Aden. Deuxieme Partie: opisthobranches (suite et fin) Marseniades, Oncidiides. Annales de la Faculte des Sciences de Marseille, 20 (Suppl.), 5–157, pls. 1–11.
Warren, L. (2002) Bornella sp. 1 from South Africa. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from http://www.seaslugforum.net/find.cfm?id=6076 (23 February 2002), (accessed 26 October 2008)
Wiens, J.J. (2004) The Role of Morphological Data in Phylogeny Reconstruction. Systematic Biology, 53, 653–661.
Wilk, L. (2005) Bornella calcarata from Utila, Honduras. [Message in] Sea Slug Forum. Australian Museum, Sydney. Available from http://www.seaslugforum.net/find.cfm?id=15366 (1 December 2005), (accessed 26 October 2008)
Willan, R.C. (1988) The taxonomy of two host-specific, cryptic dendronotoid nudibranch species from Australia, including a new species description. Zoological Journal of the Linnean Society, 94, 39–63.
Yonow, N. (1990) Red Sea Opisthobranchia 4: The orders Cephalaspidea, Anaspidea, Notaspidea and Nudibranchia: Dendronotacea and Aeolidacea. Fauna of Arabia, 18, 87–131.
Yonow, N. & Hayward, P.J. (1991) Opisthobranches de I'Ile Maurice, avec la description de deux especes nouvelles (Mollusca:Opisthobranchia). Revue francaise d’aquariologie, 18, 1–30.