Abstract
Metamorphosed, sexually mature Platynereis adults captured swimming nearshore at night off the New Zealand coast had major morphological differences that indicated four similar sympatric species existed, where previously only P. australis had been known. Simultaneous reproductive swarming of all taxa was possible at a locality. The three hitherto cryptic species are described as Platynereis mahanga sp. nov., P. kau sp. nov., and P. karaka sp. nov.. Paragnath arrangement and chaetal morphology are near identical in all four species. Homogomph notopodial falcigers, which are distinctive in shape in some Platynereis, are lacking in adults of this group of species. Pigmentation pattern differences and other minor variations are present in life in atokous (unmetamorphosed) benthic forms, but museum specimen atokes cannot be unambiguously separated into species by morphology. Heteronereid (metamorphosed) natatory forms of the four New Zealand species are distinct in several characters, including morphology of sperm papillae groups in males, and number of pre-natatory anterior segments in both sexes. Differences in size, habitat, peak spawning season, interbreeding, and juvenile development are noted. Morphology and biology are discussed in relation to other Platynereis. Presence of the P. australis group species beyond the continental shelf seas adjoining the three main islands is now undetermined except for P. australis at Chatham Islands, P. mahanga at the subantarctic Auckland and Campbell Islands, and P. karaka at Wanganella Bank; all prior reports of P. australis outside the New Zealand region need re-evaluation from specimens.References
Audouin, J.V. & Milne Edwards, H. (1833) Classification des annélides, et description de celles qui habitent les côtes de la France. Annales des Sciences Naturelles, Paris, 29, 195–269.
Augener, H. (1923) Papers from Dr. Th. Mortensen's Pacific Expedition 1914–1916. No. 14. Polychaeta I. Polychaeten von den Auckland- und Campbell-Inseln. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i København, 75, 1–115.
Augener, H. (1924) Papers from Dr. Th. Mortensen's Pacific Expedition 1914–1916. 18. Polychaeta II. Polychaeten von Neuseeland I: Errantia. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i København, 75, 241-441.
Augener, H. (1926) Papers from Dr. Th. Mortensen's Pacific Expedition 1914–16. 34. Polychaeta III. Polychaeten von Neuseeland. II. Sedentaria. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i København, 81, 157–294.
Augener, H. (1927) Die Polychaeten der Sammlung Thilenius von Neuseeland und Samoa. Mitteilungen aus dem Zoologischen Museum in Berlin, 13, 338-363.
Augener, H. (1932a) Antarktische und antiboreale Polychaeten nebst einer Hirudinee. Scientific results of the Norwegian Antarctic Expeditions 1927–1928 et sqq., 1, 1–86.
Augener, H. (1932b) Zoologische Ergebnisse der Reisen von Dr. Kohl-Larsen nach den Subantarktischen Inseln bei Neuseeland und nach Südgeorgien. 8. Polychaeten. Senckenbergiana, Frankfurt am Main, 14, 95–117.
Avise, J.C. (2000) Phylogeography : the history and formation of species. Harvard University Press, Cambridge, Mass., viii, 447 pp.
Bakken, T. & Wilson, R. (2005) Phylogeny of nereidids (Polychaeta, Nereididae) with paragnaths. Zoologica Scripta, 34, 507-547(41).
Bellan, G. (1964) Contribution a l'étude systématique, bionomique et écologique des Annélides polychètes de la Méditerrannée. Recueil des Travaux de la Station Marine d'Endoume, 49, 1–371.
Benham, W.B. (1909a) Report on the Polychaeta of the sub-Antarctic Islands of New Zealand. In: Chilton, C. (Ed.) The subantarctic Islands of New Zealand. Philosophical institute of Canterbury, Wellington, pp. 236–250.
Benham, W.B. (1909b) Scientific results of the New Zealand Government trawling expedition, 1907. Annelida and Sipunculoidea. Records of the Canterbury Museum, 1, 71–82.
Benham, W.B. (1950) Polychaeta of the Auckland and Campbell Islands. New Zealand Department of Scientific and Industrial Research Cape Expedition Series Bulletin, 10, 3-26.
Clark, R.B. (1961) The origin and formation of the heteronereis. Biological Reviews, 36, 199–236.
Clark, R.B. (1977) Reproduction, speciation and polychaete taxonomy. In: Reish, D.J. & K. Fauchald (Ed.) Essays on Polychaetous Annelids in Memory of Dr. Olga Hartman. Allan Hancock Press, Los Angeles, pp. 477–501.
Daly, J.M. (1973) Behavioural and secretory activity during tube construction by Platynereis dumerilii Aud & M. Edw. [Polychaeta: Nereidae]. Journal of the Marine Biological Association of the United Kingdom, 53, 521–529.
Day, J.H. (1953) The Polychaete Fauna of South Africa. Part 2. Errant species from Cape shores and estuaries. Annals of the Natal Museum, 12, 397–441.
Day, J.H. (1967) A monograph on the Polychaeta of Southern Africa. Trustees British Museum, London, 878 pp.
Day, J.H. (1971) Polychaeta [Marion and Prince Edward Islands; report on the South African Biological and Geological Expedition]. In: Bakker, E., M.S. van Zinderen, J.M. Winterbottom & R.A. Dyer (Ed.) Marion and Prince Edward Islands. A.A. Balkema, Cape Town, pp. 384–390.
Day, J.H. (1975) On a collection of Polychaeta from intertidal and shallow reefs near Perth, Western Australia. Records of the Western Australian Museum, 3, 167–208.
Denes, A.S., Jekely, G., Steinmetz, P.R.H., Raible, F., Snyman, H., Prud'homme, B., Ferrier, D.E.K., Balavoine, G. & Arendt, D. (2007) Molecular Architecture of Annelid Nerve Cord Supports Common Origin of Nervous System Centralization in Bilateria. Cell, 129, 277-288.
Desbruyères, D. (1977) Bionomie benthique du plateau continental des Iles Kerguelen - Macrofaune 6. Évolution des populations de trois espèces d'Annélides Polychètes en milieu sub-antarctique. CNFRA, 42, 135-172.
Edgar, G.J. (1987) Dispersal of fauna and floral propagules associated with drifting Macrocystis pyrifera plants. Marine Biology (Berlin), 95, 599–610.
Ehlers, E. (1897) Polychaeten. Hamburger Magalhaensische Sammelreise, 1897, 1–148.
Ehlers, E. (1904) Neuseeländische Anneliden. Abhandlungen der Königlichen Gesellschaft der Wissenschaften zu Göttingen. Mathematisch-Physikalische Klasse. Neue Folge, 3, 1–80.
Ehlers, E. (1905) Anneliden der sammlung Schauinsland (Ergebnisse einer Reise nach dem Pacific. Schauinsland 1896/97). Zoologische Jahrbücher. Abteilung für systematik, geographie und biologie der tiere, 22, 281-302.
Ehlers, E. (1907) Neuseeländische Anneliden. II. Abhandlungen der Königlichen Gesellschaft der Wissenschaften zu Göttingen. Mathematisch-Physikalische Klasse. Neue Folge, 5, 1-31.
Ehlers, E. (1913) Die Polychaeten-Sammlungen der Deutschen Südpolar-Expedition 1901–1903. Deutsche Südpolar-Expedition, 13, 397–598.
Estcourt, I.N. (1967) Burrowing polychaete worms from a New Zealand estuary. Transactions Royal Society of New Zealand. Zoology, 9, 65-78.
Fage, L. & Legendre, R. (1927) Pêches planctoniques a la lumière effectuées à Banyuls-sur-Mer et à Concarneau. I. Annélides polychètes. Archives de Zoologie Expérimentale et Générale, Paris, 67, 23-248.
Fauvel, P. (1941) Annélides polychètes de la Mission du Cap Horn (1882–1883). Bulletin de la Musee d'Histoire naturelle, Paris, Serie 2, 18, 272–298.
Fischer, A. (1999) Reproductive and developmental phenomena in annelids: a source of exemplary research problems. Hydrobiologia, 402, 1–20.
Fischer, A. & Dorresteijn, A. (2004) The polychaete Platynereis dumerilii (Annelida): a laboratory animal with spiralian cleavage, lifelong segment proliferation and a mixed benthic/pelagic life cycle. Bioessays, 26, 314–325.
Gravier, C. & Dantan, J.L. (1928) Pêches nocturnes a la lumière dans la Baie d'Alger. Annélides polychètes. Annales de l'Institut Océanographique, Paris, n. s., 5, 1-187.
Hanafiah, Z., Sato, M., Nakashima, H. & Tosuji, H. (2006) Reproductive swarming of sympatric nereidid polychaetes in an estuary of the Omuta-gawa River in Kyushu, Japan, with special reference to simultaneous swarming of two Hediste species. Zoological Science, 23, 205-217.
Hardege, J.D. (1999) Nereidid polychaetes as model organisms for marine chemical ecology. Hydrobiologia, 402, 145–161.
Hartman, O. (1945) The marine annelids of North Carolina. Duke University Marine Station Bulletin, 2, 1–54.
Hartman, O. (1948) The marine annelids erected by Kinberg with notes on some other types in the Swedish State Museum. Arkiv för Zoologi, 42A, 1–137.
Hartman, O. (1953) Non-pelagic polychaeta of the Swedish Antarctic Expedition 1901–1903. Further Zoological Results of the Swedish Antarctic Expedition 1901–1903, 4, 1-83.
Hartman, O. (1954) Australian Nereidae including descriptions of three new species and one genus, together with summaries of previous records and keys to species. Transactions of the Royal Society of South Australia, 77, 1–41.
Hartman, O. (1959) Catalogue of the polychaetous annelids of the world. Parts 1 & 2. Occasional Papers of the Allan Hancock Foundation, 23, 1–628.
Hartman, O. (1965) Catalogue of the polychaetous annelids of the world. Supplement 1960–1965 and index. Occasional Papers of the Allan Hancock Foundation, 23, 1–197.
Hartmann-Schröder, G. (1962) Zur Kenntnis der Nereiden Chiles (Polychaeta errantia), mit Beschreibung epitoker Stadien einiger Arten und der Jugendentwicklung von Perinereis vallata (Grube). Zoologischer Anzeiger, 168, 389–441.
Hauenschild, C. (1951) Nachweis der sogenannten atoken Geschlechtsform des Polychaeten Platynereis dumerilii Aud. et M.-Edw. als eigene Art aufgrund von Zuchtversuchen. Zoologische Jahrbücher. Abteilung für Allgemeine Zoologie und Physiologie der Tiere, 63, 107–128.
Hauenschild, C. & Fischer, A. (1969) Platynereis dumerilii. Mikroskopische Anatomie, Fortpflanzung und Entwicklung. Grosses Zoologisches Praktikum, 10b, 1–55.
Helmuth, B., Veit, R.R. & Holberton, R. (1994) Long-distance dispersal of a Sub-Antarctic brooding bivalve (Gaimardia trapesina) by kelp-rafting. Marine Biology, 120, 421–426.
Hemplemann, F. (1911) Zur Naturgeschichte von Nereis dumerilii Aud. et Edw. Zoologica (Stuttgart), 25 (Heft 62), 1–135.
Holly, M. (1935) Polychaeta from Hawaii. Bulletin of the Bernice P. Bishop Museum, 129, 1–33.
Hutchings, P.A. & Reid, A. (1991) The Nereididae, Polychaeta, from Australia: Leonnates, Platynereis and Solomononereis. Records of the Australian Museum, 43, 47–62.
Huvet, A., Fabioux, C., McCombie, H., Lapègue, S. & Boudry, P. (2004) Natural hybridization between genetically differentiated populations of Crassostrea gigas and C. angulata highlighted by sequence variation in flanking regions of a microsatellite locus. Marine Ecology Progress Series, 272, 141–152.
Imajima, M. (1972) Review of the annelid worms of the family Nereidae of Japan, with descriptions of five new species or subspecies. Bulletin of the National Science Museum, Tokyo, 15, 37–153.
Just, E.E. (1914) Breeding habits of the heteronereis form of Platynereis megalops at Woods Hole, Mass. Biological Bulletin, 28, 201–212.
Just, E.E. (1915) An experimental analysis of fertilization of Platynereis megalops. Biological Bulletin, 28, 93–114.
Just, E.E. (1929) Breeding habits of Nereis dumerilii at Naples. Biological Bulletin, 57, 307–310.
Kinberg, J.G.H. (1865) Annulata nova. Öfversigt af Konglia Vetenskaps- Akadamiens Förhandlingar, 22, 167–179.
Knox, G.A. (1951) The polychaetous annelids of Banks Peninsula. Part I. Nereidae. Records of the Canterbury Museum, 5, 213–229.
Knox, G.A. (1960) Biological results of the Chatham Islands 1954 Expedition. Part 3. Polychaeta Errantia. New Zealand Oceanographic Institute Memoir, 6, 1–140.
Knox, G.A. & Cameron, D.B. (1970) Polychaeta from the Snares Islands, New Zealand. Transactions of the Royal Society of New Zealand, 12, 73-85.
Knox, G.A. & Cameron, D.B. (1971) Port Phillip Survey 2. Polychaeta. Memoirs of the National Museum of Victoria, 32, 21–41.
Korringa, P. (1947) Relations between the moon and periodicity in the breeding of marine animals. Ecological Monographs, 17, 347–381.
Kott, P. (1951) Nereidae and Eunicidae of south Western Australia; also notes on the ecology of Western Australian limestone reefs. Journal of the Royal Society of Western Australia, 35, 85–130.
de León González, J.A., Solis-Weiss, V. & Valadez-Rocha, V. (2001) Two new species of Platynereis (Polychaeta: Nereididae) from eastern Mexican shores. Proceedings of the Biological Society of Washington, 114, 389–395.
McIntosh, W.C. (1876) Descriptions of some new species of Annelida from Kerguelen Island. Annals and Magazine of Natural History, (series 4) 17, 318–323.
Monro, C.C.A. (1926) Polychaeta of the 'Alert' Expedition. Families Hesionidae and Nereidae. Zoological Journal of the Linnean Society, 36, 311–323.
Monro, C.C.A. (1930) Polychaete worms. Discovery Reports, 2, 1–222.
Monro, C.C.A. (1936) On the heteronereid of Platynereis pulchella Gravier, a polychaete belonging to the family Nereidae. A preliminary note. Annals and Magazine of Natural History, 18, 380–384.
Monro, C.C.A. (1939) Polychaeta. British and New Zealand Antarctic Research Expedition 1929–1931. Reports, Series B (Zoology and Botany), 4, 89–156.
Ojeda, F.P. & Santelices, B. (1984) Invertebrate communities in holdfasts of the kelp Macrocystis pyrifera from southern Chile. Marine Ecology Progress Series, 16, 65–73.
Pettibone, M.H. (1963) Marine polychaete worms of the New England region. 1. Families Aphroditidae through Trochochaetidae. Bulletin United States National Museum, 227, 1–356.
Pratt, E.M. (1901) A collection of Polychaeta from the Falkland Islands. Memoirs and Proceedings of the Manchester Literary and Philosophical Society, 45, 1–18.
Ramsay, L.N.G. (1914) Polychaeta of the family Nereidae, collected by the Scottish National Antarctic Expedition (1902–1904). Transactions of the Royal Society of Edinburgh, 50, 41–48.
Rullier, F. (1964) Résultats scientifiques des campagnes de la Calypso: Iles du Cap Vert. 5. Annélides Polychètes. Annales de l'Institute Océanographique, Paris, 41, 113–218.
Rullier, F. (1972) Annélides polychètes de Nouvelle-Calédonie recueillies par Y. Plessis et B. Salvat. Expédition Française sur les Récifs Coralliens de la Nouvelle-Calédonie, 6, 1–167.
Santos, G.S.G., Pleijel, F., Lana, P. & Rouse, G.W. (2005) Phylogenetic relationships within Nereididae (Annelida : Phyllodocida). Invertebrate Systematics, 19, 557-576.
Schmarda, L.K. (1861) Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erdr [sic] 1853 bis 1857 von Ludwig K. Schmarda. Erster Band. Turbellarien, Rotatorien und Anneliden. Zweite Hälfte. Wilhelm Engelmann, Leipzig, 164 pp.
Schneider, S., Fischer, A. & Dorresteijn, A.W.C. (1992) A morphometric comparison of dissimilar early development in sibling species of Platynereis (Annelida, Polychaeta). Roux's Archives of Developmental Biology, 201, 243-256.
Schulz, G., Ulbrich, K.P., Hauenschild, C. & Pfannenstiel, H.D. (1989) The atokous-epitokous border is determined before the onset of heteronereid development in Platynereis dumerilii (Annelida, Polychaeta). Roux's Archives of Developmental Biology, 198, 29–33.
Smith, R.I. (1958) On reproductive pattern as a specific characteristic among nereid polychaetes. Systematic Zoology, 7, 60–73.
Smith, S.D.A. (2002) Kelp rafts in the Southern Ocean. Global Ecology and Biogeography, 11, 67–69.
Tessmar-Raible, K. & Arendt, D. (2003) Emerging systems: Between vertebrates and arthropods, the Lophotrochozoa. Current Opinion in Genetics & Development, 13, 331-340.
Tessmar-Raible, K., Steinmetz, P.R.H., Snyman, H., Hassel, M. & Arendt, D. (2005) Fluorescent two-color whole mount in situ hybridization in Platynereis dumerilii (Polychaeta, Annelida), an emerging marine molecular model for evolution and development. Biotechniques, 39, 460-462.
Tosuji, H., Miyamoto, J., Hayata, Y. & Sato, M. (2004) Karyotyping of female and male Hediste japonica (Polychaeta, Annelida) in comparison with those of two closely related species, H. diadroma and H. atoka. Zoological Science, 21, 147–152.
Tosuji, H. & Sato, M. (2006) Salinity favorable for early development and gamete compatibility in two sympatric estuarine species of the genus Hediste (Polychaeta : Nereididae) in the Ariake Sea, Japan. Marine Biology, 148, 529-539.
Verrill, A.E. (1873) Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical characters of the region. In: Baird, S.F. (Ed.) Report of the United States Commissioner of Fish and Fisheries 1871/1872. Government Printing Office, Washington, pp. 295-778.
Verrill, A.E. (1876) Contributions to the Natural History of Kerguelen Island. Annelids and echinoderms. Bulletin United States National Museum, 3, 64–67.
Watson, G.J., Bentley, M.G., Gaudron, S.M. & Hardege, J.D. (2003) The role of chemical signals in the spawning induction of polychaete worms and other marine invertebrates. Journal of Experimental Marine Biology and Ecology, 294, 169–187.