Abstract
Three new Ophryotrocha species are described from sites with high levels of organic carbon flux including a whale-fall at 125 m depth off the Swedish coast and sediment sampled at 104 m depth beneath a fish farm in a Norwegian fjord. Phylogenetic analyses based on the nuclear gene H3 and the mitochondrial genes COI and 16S using MrBayes and Maximum Likelihood analyses show that Ophryotrocha eutrophila sp. nov. is a close relative to Ophryotrocha puerilis, while Ophryotrocha craigsmithi sp. nov. falls together with Palpiphitime lobifera, and Ophryotrocha scutellus sp. nov. occur within the 'hartmanni' clade. The genus Ophryotrocha is in our study monophyletic only if the genera Iphitime and Palpiphitime are included. Two representatives of Ophryotrocha previously described from anthropogenically-enriched sediments are here reported for the first time in very high abundance from a naturally occurring habitat. We suggest that whale falls are important habitats for the evolution of ecosystem services such as the degradation of complex organic compounds.References
Åkesson, B. (1973) Morphology and life history of Ophryotrocha maculata sp. n. (Polychaeta, Dorvilleidae). Zoologica Scripta, 2, 141–144.
Åkesson, B. (1977) Crossbreeding and geographic races: Experiments with the Polychaete genus Ophryotrocha. Mikrofauna des Meeresbodens, 61, 11–18.
Åkesson, B. & Rice, S.A. (1992) Two new Dorvillea (Polychaeta, Dorvilleidae) with obligate asexual reproduction. Zoologica Scripta, 21, 351–362.
Bely, A.E. & Wray, G.A. (2004) Molecular phylogeny of naidid worms (Annelida : Clitellata) based on cytochrome oxidase I. Molecular Phylogenetics and Evolution, 30, 50–63.
Blake, J.A. (1985) Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacific. I. Euphrosinidae, Phyllodocidae, Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae, and Maldanidae. Bulletin of the Biological Society of Washington, 6, 67–101.
Blake, J.A. & Hilbig, B. (1990) Polychaeta from the vicinity of deep-sea hydrothermal vents in the eastern Pacific. II. New species and records from the Juan de Fuca and Explorer Ridge systems. Pacific Science, 44, 219–253.
Claparède, E. & Mecznikow, E. (1869) Beiträge zur Erkenntniss der Entwicklungsgeschichte der Chaetopoden. Zeitschrift für wissenschaftliche Zoologie, 16, 163–205.
Colgan, D.J., Ponder, W.F. & Eggler, P.E. (2000) Gastropod evolutionary rates and phylogenetic relationships assessed using partial 28S rDNA and histone H3 sequences. Zoologica Scripta, 29, 29–63.
Crossland, C. (1924) Polychaeta of tropical East Africa, the Red Sea, and Cap Verde Islands, collected by Cyril Crossland, and of the Maldive Archipelago collected by Professor Stanley Gardiner, M.A., F.R.S. Zoological Society of London, 47, 1–106.
Dahlgren, T.G., Åkesson, B., Schander, C., Halanych, K.M. & Sundberg, P. (2001) Molecular phylogeny of the model annelid Ophryotrocha. Biological Bulletin, 201, 193–203.
Dahlgren, T.G., Glover, A.G., Baco, A. & Smith, C.R. (2004) Fauna of whale falls: systematics and ecology of a new polychaete (Annelida : Chrysopetalidae) from the deep Pacific Ocean. Deep-Sea Research Part I-Oceanographic Research Papers, 51, 1873–1887.
Dahlgren, T.G., Wiklund, H., Källström, B., Lundälv, T., Smith, C.R. & Glover, A.G. (2006) A shallow-water whale-fall experiment in the north Atlantic. Cahiers De Biologie Marine, 47, 385–389.
Drummond, A.J., Ashton, B., Cheung, M., Heled, J., Kearse, M., Moir, R., Stones-Havas, S., Thierer, T. & Wilson, A. (2007) Geneious v3.8, Available from http://www.geneious.com/
Eibye-Jacobsen, D. & Kristensen, R.M. (1994) A New Genus and Species of Dorvilleidae (Annelida, Polychaeta) from Bermuda, with a Phylogenetic Analysis of Dorvilleidae, Iphitimidae and Dinophilidae. Zoologica Scripta, 23, 107–131.
Esmark, L. (1874) Eteonopsis geryonicola. Forhandlinger i Videnskabsselskabet i Christiania, 497–498.
Fage, L. & Legendre, R. (1934) Les annélides polychètes du genre Iphitime. A propos d'une espece nouvelle commensale des pagures, Iphitime paguri n.sp. Bulletin de la Société Zoologique de France, 58, 299–305.
Fauchald, K. (1970) Polychaetous annelids of the families Eunicidae, Lumbrinereidae, Iphitimidae, Arabellidae, Lysaretidae, and Dorvilleidae from western Mexico. Allan Hancock Monographs in Marine Biology, 5, 1–335.
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
Fujikura, K., Fujiwara, Y. & Kawato, M. (2006) A new species of Osedax (Annelida : Siboglinidae) associated with whale carcasses off Kyushu, Japan. Zoological Science, 23, 733–740.
Gaston, G.R. & Benner, D.A. (1981) On Dorvilleidae and Iphitimidae (Annelida: Polychaeta) with a redescription of Eteonopsis geryonicola and a new host record. Proceedings of the Biological Society of Washington, 94, 76–87.
Glover, A.G., Källström, B., Smith, C.R. & Dahlgren, T.G. (2005) World-wide whale worms? A new species of Osedax from the shallow north Atlantic. Proceedings of the Royal Society B-Biological Sciences, 272, 2587–2592.
Graybeal, A. (1998) Is it better to add taxa or characters to a difficult phylogenetic problem? Systematic Biology, 47, 9–17.
Grube, A.E. (1855) Beschreibung neuer oder wenig bekannter Anneliden. Archiv für Naturgeschichte, 21, 81–136.
Heggoy, K.K., Schander, C. & Akesson, B. (2007) The phylogeny of the annelid genus Ophryotrocha (Dorvilleidae). Marine Biology Research, 3, 412–420.
Hilbig, B. (1995) 13. Family Dorvilleidae Chamberlin, 1919. In: Blake, J.A., Hilbig, B. & Scott, P.H. (Eds.) Taxonomic atlas of the benthic fauna of the Santa Maria Basin and western Santa Barbara Channel. Santa Barbara Museum of Natural History, Santa Barbara, pp. 341–364.
Hilbig, B. & Blake, J.A. (1991) Dorvilleidae (Annelida, Polychaeta) from the United-States Atlantic Slope and Rise — Description of 2 New Genera and 14 New Species, with a Generic Revision of Ophryotrocha. Zoologica Scripta, 20, 147–183.
Hoisaeter, T. & Samuelsen, T.J. (2006) Taxonomic and biological notes on a species of Iphitime (Polychaeta, Eunicida) associated with Pagurus prideaux from western Norway. Marine Biology Research, 2, 333–354.
Huth, W. (1933) Ophryotrocha-Studien. Zur Cytologie der Ophryotrochen. Zeithscrift für Zellforschung und Mikroskopische Anatomie, Berlin, 20, 309–381,24figs.
Josefson, A. (1975) Ophryotrocha longidentata sp.n. and Dorvillea erucaeformis (Malmgren) (Polychaeta, Dorvilleidae) from the west coast of Sweden. Zoologica Scripta, 4, 49–54.
Jumars, P.A. (1974) A generic revision of the Dorvilleidae (Polychaeta), with six new species from the deep North Pacific. Zoological Journal of the Linnean Society, 54, 101–135.
Kirkegaard, J.B. (1977) A new species of Iphitime (Polychaeta: Iphitimidae) living under the tail of Hyas (Crustacea: Decapoda) in the Oslo Fjord. In: Fauchald, D.J.R.a.K. (Ed.) Essays on polychaetous annelids in memory of dr. Olga Hartman, Los Angeles, pp. 199–209.
Kiseleva, M.I. (1992) New genus and species of a chrysopetalid polychaete from the Black Sea [in Russian]. Zoologichesky Zhurnal, 72, 128–132.
LaGreca, M. & Bacci, G. (1962) Una nuova specie di Ophryotrocha delle coste tirreniche. Bollettino di Zoologia, 29, 13–23.
Malmgren, A.J. (1865) Nordiska Hafs-Annulater. Öfversigt af K. Vetenskapsakademiens förhandlingar, 22, 181–192.
Marenzeller, E. (1902) Südjapaische Anneliden. III. Aphroditea, Eunicea. Denkschriften der Kaiserlichen Akademie der Wissenschaften (Wien), 563–582.
McIntosh, W.C. (1869) On the structure of the British nemerteans, and some new British annelids.
Transactions of the Royal Society of Edinburgh, 25, 305–433.
McIntosh, W.C. (1885) Report on the Annelida Polychaeta collected by H.M.S. Challenger during the years 1873–76. Report of the Scientific Results of the Voyage of H.M.S. Challenger 1873–76, 12, 1–554.
Miura, T. (1997) Two new species of the Genus Ophryotrocha (Polychaeta, Iphitimiidae) from Kagoshima Bay. Bulletin of Marine Science, 60, 300–305.
Müller, O.F. (1776) Zoologicae Danicae Prodromus, seu Animalium Daniae et Norvegiae indigenarum characteres, nomina et synonyma imprimis popularium. Hallageriis, Havniae [Copenhagen], 282 pp.
Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University
Orensanz, J.M. (1990) The eunicemorph polychaete annelids from Antarctic and Subantarctic Seas. Antarctic Research Series, 52, 1–183.
Oug, E. (1978) New and lesser known Dorvilleidae (Annelida, Polychaeta) from Scandinavian and Northeast American waters. Sarsia, 63, 285–303.
Palumbi, S.R. (1996) Nucleic acid II: the polymerase chain reaction. In: Hillis, D.M., Moritz, G. & Mable, B.K. (Eds.) Molecular Systematics. Sinauer Associates, Sunderland, Mass, pp. 205–247.
Parfitt, E. (1866) Description of a Nereis new to science. Zoologist, Serie 2, volume 1, 113–114.
Paxton, H. (2009) A new species of Palpiphitime (Annelida: Dorvilleidae) from western Canada. Proceedings of the Biological Society of Washington, 122, 26–31.
Paxton, H. & Åkesson, B. (2007) Redescription of Ophryotrocha puerilis and O. labronica (Annelida, Dorvilleidae). Marine Biology Research, 3, 3–19.
Paxton, H. & Morineaux, M. (2009) Three species of Dorvilleidae (Annelida: Polychaeta) associated with Atlantic deep-sea reducing habitats, with the description of Ophryotrocha fabriae, new species. Proceedings of the Biological Society of Washington, 122, 14–25.
Pfannenstiel, H.-D. (1972) Eine neue Ophryotrocha-Art (Polychaeta, Eunicidae) aus Japan. Helgoländer wiss. Meresunters., 23, 117–124.
Pfannenstiel, H.-D., Grothe, C. & Kegel, B. (1982) Studies on Ophryotrocha geryonicola (Polychaeta: Dorvilleidae). Helgoländer wissenschaftliche Meeresuntersuchungen, 35, 119–125.
Pleijel, F. & Eide, R. (1996) The phylogeny of Ophryotrocha (Dorvilleidae: Eunicida: Polychaeta). Journal of Natural History, 30, 647–659.
Pleijel, F., Rouse, G.W., Ruta, C., Wiklund, H. & Nygren, A. (2008) Vrijenhoekia balaenophila, a new hesionid polychaete from a whale fall off California. Zoological Journal of the Linnean Society, 152, 625–634.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
Rouse, G.W., Goffredi, S.K. & Vrijenhoek, R.C. (2004) Osedax: Bone-eating marine worms with dwarf males. Science, 305, 668–671.
Rouse, G.W., Worsaae, K., Johnson, S.B., Jones, W.J. & Vrijenhoek, R.C. (2008) Acquisition of dwarf male "Harems" by recently settled females of Osedax roseus n. sp. (Siboglinidae; Annelida). Biological Bulletin, 214, 67–82.
Struck, T.H., Purschke, G. & Halanych, K.M. (2006) Phylogeny of Eunicida (Annelida) and exploring data congruence using a partition addition bootstrap alteration (PABA) approach. Systematic Biology, 55, 1–20.
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Version 4. Sinauer Associates, Sunderland, MA
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882.
Wiklund, H., Glover, A.G., Johannessen, P.J. & Dahlgren, T.G. (2009) Cryptic speciation at organic-rich marine habitats: a new bacteriovore annelid from whale-fall and fish farms in the North-East Atlantic. Zoological Journal of the Linnean Society, 4, 774–785.
Wolf, P.S. (1986) Four new genera of Dorvilleidae (Annelida: Polychaeta) from the Gulf of Mexico. Proceedings of the Biological Society of Washington, 99, 616–626.