Abstract
Morphological and genetic studies of specimens identified as Spiophanes bombyx (Claparède, 1870) revealed that the supposedly cosmopolitan species represents a species complex. Species belonging to this complex are S. bombyx (Claparède, 1870), S. aucklandicus Meißner, 2005, S. uschakowi Zachs, 1933 and the newly described S. norrisi. Morphological differences between these species are slight and concern the start of sabre chaetae, the shape of dorsal ciliated organs, and the start and number of neuropodial hooks in relation to body size. Specimens from the Atlantic Ocean examined in the course of this study also belong to this species complex but could not be assigned to any of the known species. Genetic differences between S. bombyx and S. norrisi in both examined genes (18S, COI) are significant and genetic distances are comparable to distances found between sister species in other spionid taxa. As expected, the nuclear 18S rRNA is generally more conserved than the mitochondrial COI gene. The significance beyond taxonomy of dorsal ciliated organs and of the presence of sabre chaetae in certain body regions is briefly discussed, but conclusions about the phylogenetic relationships and character evolution within Spiophanes must remain tentative.References
Blake, J.A. 2006. Spionida. pp. 566–638, In: Rouse, G. and F. Pleijel, Editors, Reproductive Biology and Phylogeny of Annelida. Volume 4 of Series: Reproductive Biology and Phylogeny, Series Editor, B.G.M. Jamieson. Science Publishers, Enfield, New Hampshire.
Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171–280.
Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In Blake, J. A., Hilbig, B. & P. H. Scott (ed.)]. Taxonomic Atlas of the benthic fauna of the Santa Maria Basin and the western Santa Barbara Channel. Santa Barbara Museum of Natural History, Volume 6, The Annelida Part 3 – Polychaeta: Orbiniidae to Cossuridae, Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 81–223.
Blank, M. & Bastrop, R. (2008) Phylogeny of the mud worm genus Marenzelleria (Polychaeta, Spionidae) inferred from mitochondrial DNA sequences. Zoologica Scripta Doi:10.1111/j.1463–6409.2008.00370.x.
Chamberlin, R. V. (1919) New polychaetous annelids from Laguna Beach, California. Pomona College Journal of Entomology and Zoology, 11, 1–23.
Claparède, E. (1870) Les Annélides Chétopodes du Golfe de Naples. Troisième partie. Mémoires de la Societe de Physique et d'Histoire Naturelle de Geneve, 20 (2), 365–542.
Englisch, U. & Koenemann, S. (2001) Preliminary phylogenetic analysis of selected subterranean amphipod crustaceans, using small subunit rDNA gene sequences. Organisms Diversity and Evolution, 1, 139–145.
Fauvel, P. (1927) Faune de France Vol. 16 Polychètes sedentaires. Addenda aux Errantes, Archiannélides, Myzostomaires. Librairie de la Faculté des Sciences, Paris, Paul Lechevalier, 494 pp.
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using bootstrap. Evolution, 39, 783–791.
Folmer, O., Black, M., Hoen, 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.
Grube, A. E. (1860) Beschreibung neuer oder wenig bekannter Anneliden. Archiv der Naturgeschichte, 26, 71–118.
Halanych, K. M., Bacheller, J. D., Aguinaldo, A. M. A., Liva, S. M., Hillis, D. M.& Lake, J. A. (1995). Evidence from 18S ribosomal DNA that the lophophorates are protostome animals. Science, 267, 1641–1643.
Hall, T.A. 1999. Bioedit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.
Hannerz, L. 1956. Larval development of the polychaete families Spionidae Sars, Disomidae Mesnil and Poecilochaetidae n. fam. in the Gullmar Fjord (Sweden). Zoologiska Bidrag från Uppsala 31, 1–204.
Hartmann-Schröder, G. (1965). Die Polychaeten des Sublitorals. Zur Kenntnis des Sublitorals der chilenischen Küste unter besonderer Berücksichtigung der Polychaeten und Ostracoden (mit Bemerkungen über den Einfluss sauerstoffarmer Strömungen auf die Besiedlung von marinen Sedimenten). Mitteilungen aus dem Hamburger Zoologischen Museum, 62 (Supplement), 59–305.
Hartmann-Schröder, G. (1971). Annelida, Borstenwürmer, Polychaeta.—Die Tierwelt Deutschlands und der angrenzenden Meeresteile nach ihren Merkmalen und ihrer Lebensweise. Gustav Fischer-Verlag, Jena u.a., Teil 58, 594 pp.
Huelsenbeck, J.P. & Ronquist, F. (2001). MRBAYES: Bayesian inference for phylogenetic trees. Bioinformatics, 17, 745–755.
Imajima, M. (1991). Spionidae (Annnelida, Polychaeta) from Japan VII. The genus Spiophanes. Bulletin of National Science Museum Tokyo Ser. A (Zoology), 17, 115–137.
Jelsing, J. (2002) Ultrastructural investigations on the cephalic and metameric nuchal organs of Spio cf. filicornis (Polychaeta, Spionidae). Zoomorphology, 121, 213–220.
Jelsing, J. (2003) Ultrastructural studies of dorsal ciliated organs in Spionidae (Annelida: Polychaeta). Hydrobiologia, 496, 241–251.
Jobb, G., von Haeseler, A. & Strimmer, K. (2004) Treefinder: a powerful graphical analysis for molecular phylogenetics. BMC Evolutionary Biology, 4, 18.
Light, W. J. (1978) Invertebrates of the San Francisco Bay Estuary System. Family Spionidae (Annelida, Polychaeta). The Boxwood Press, Pacific Grove, California, 211 pp.
Mahon, A.R., Mahon, H.K., Dauer, D.M. & Halanych, K.M. (2009). Discrete genetic boundaries of three Streblospio (Spionidae, Annelida) species and the status of S. shrubsolii. Marine Biology Research, 5, 172–178.
Meißner, K. & Hutchings, P. A. (2003) Spiophanes species (Polychaeta: Spionida) from Eastern Australia—with descriptions of new species, new records and an emended generic diagnosis. Records of the Australian Museum, 55 (2), 117–140.
Meißner, K. (2005) Revision of the genus Spiophanes (Polychaeta, Spionidae); with new synonymies, new records and descriptions of new species. Mitteilungen des Museums für Naturkunde Berlin, Zoologische Reihe, 81 (1), 3–66.
Mesnil, F. (1896) Études de morphologie externe chez les Annélides. Les Spionidiens des côtes de la Manche. Bulletin Scientifique de la France et de la Belgique, 29, 110–287.
Mesnil, F. (1897) Études de morphologie externe chez les Annélides. II. Remarques complémentaires sur les Spionidens. —La famille nouvelle des Disomidiens. — La place des Aonides (sensus Tauber, Levinsen). Bulletin Scientifique de la France et de la Belgique, 30, 83–100.
Mincks, S.L., Dyal, P.L., Paterson, G.L.J., Smith, C.R. & Glover, A.G. (2009) A new species of Aurospio (Polychaeta, Spionidae) from the Antarctic shelf, with analysis of its ecology, reproductive biology and evolutionary history. Marine Ecology, 30, 181–197.
Nylander, J. A. A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University.
Okuda, S. (1937) Spioniform polychaetes from Japan. Journal of the Faculty of Science of the Hokkaido Imperial University, (G), 5(3), 217–254.
Söderström, A. (1920) Studien über die Polychaetenfamilie Spionidae. PhD thesis, Uppsala, 286 pp.
Struck, T.H., Nesnidal, M.P., Purschke, G. & Halanych, K.M. (2008) Detecting possibly saturated positions in 18S and 28S sequences and their influence on phylogenetic reconstruction of Annelida (Lophotrochozoa). Molecular Phylogenetics and Evolution, 48, 628–645.
Swofford, D.L. (2000) PAUP* Phylogenetic analysis using parsimony and other methods, Version 4.0 b10. Sunderland, MA: Sinauer.
Webster, H.E. & Benedict, J.E. (1884) The Annelida Chaetopoda from Provincetown and Wellfleet, Mass. United States Fisheries and Wildlife Service Biological Report, 1881, 699–747.
Zachs I.G. (1933) Polychaeta of the North-Japanese Sea. Explorations of the Seas of the USSR, 19, 125–137.