Abstract
A new species of Ctenodrilus is described based on morphology, anatomy and molecular (COI and 16S) data. Phylogenetic analyses revealed that the newly described Hawaiian species is the sister taxon of the widely distributed C. serratus. Comparisons of K2P distances show distances of at least 10.9 % for 16S and 18.7 % for CO1. Morphology of C. serratus differs from C. pacificus sp. nov. in the shape, number and distribution of multidentate hooks along the body, extent in segments of anterior ventral cilia, and presence of stomach cilia. Ctenodrilus pacificus sp. nov. resembles C. parvulus in terms of the internal anatomy but differs in respect to the shape of multidentate hooks. Asexual reproduction in the new species is described including associated changes in the external morphology and internal anatomy. A revision of the taxonomic characters of Ctenodrilus is also presented in order to provide background information for a better assessment of the widely distributed taxon Ctenodrilus serratus.
References
Bastida-Zavala, J.R. (1994) Hermafroditismo simultáneo y extensión de ámbito de Ctenodrilus serratus (Polychaeta: Ctenodrilidae) en Baja California Sur, Mexico. Revista de Biología Tropical, 42 (3), 737–741.
Ben Eliahu, N.M. (1976) Polychaete cryptofauna from rims of similar intertidal vermetid reefs on the Mediterranean coast of Israel and in the gulf of Elat: Sedentaria. Israel Journal of Zoology, 25, 121–155.
Blake, J.A. (1996) Family Cirratulidae Ryckholdt, 1851. Including a revision of the genera and species from the eastern North Pacific. 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. 6 - The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, pp. 263–384.
Bleidorn, C., Vogt, L. & Bartolomaeus, T. (2003) New insights into polychaete phylogeny (Annelida) inferred from 18S rDNA sequences. Molecular Phylogenetics and Evolution, 29, 279–288.
http://dx.doi.org/10.1016/S1055-7903(03)00107-6Carr, C.M., Hardy, S.M., Brown, T.M, Macdonald, T.A. & Hebert, P.D.N. (2011) A tri-oceanic perspective: DNA barcoding reveals geographic structure and cryptic diversity in Canadian polychaetes. PLoSOne, 6, e22232.
http://dx.doi.org/10.1371/journal.pone.0022232Castresana, J. (2002) Gblocks Server. Castresana Lab, Institut de Biologia Evolutiva, Barcelona.
Claparède, E. (1863) Beobachtungen über Anatomie und Entwicklungsgeschichte der wirbellosen Thiere an der Küste von Normandie angestellt. Leipzig, W. Engelmann, 120 pp., 18pls.
Clark, R.B. (1977) Reproduction, speciation and polychaete taxonomy. In: D.J. Reish & K. Fauchald, (Eds.). Essays on polychaetous annelids in memory of Dr. Olga Hartman. Allan Hancock Foundation, Los Angeles, California, p p. 477–501.
Day, J.H. (1973) New polychaeta from Beaufort, with a key to all species recorded from North Carolina. NOAA Technical Reports, Ser. National Marine Fisheries Service, Circulars, 375, 1–140.
Devaney, D.M. & Bailey-Brock, J.H. (1987) Polychaetes of Enewetak Atoll. Chapter 13. In: The Natural history of Enewetak Atoll, 2, pp 97–103. [DOE/EV/00703-T1-Vol.2 (DE87006111)]
Dean, H.K. & Blake, J.A. (2014) 7.15.5. Ctenodrilidae Kennel, 1882. In: Schmidt-Rhaesa, A. & Walter de Gruyter, A. (Eds.), Handbook of Zoology, A Natural History of the Phyla of the Animal Kingdom. Berlin. [Database]
Delle Chiaje, S. (1828) Memoria sulla storia e notomia degli animali senza vertebre del regno di Napoli. Stamperia della Società Tipografica, Napoli, 3, 232.
Galvagni, E. (1905) Histologie des genus Ctenodrilus Clap. Arbeiten aus den Zoologischen Instituten der Universität Wien und der Zoologischen Station in Triest, 15, 47–80.
Gelder, S.R. & Palmer, R. (1976) The nervous system of the marine polychaete Ctenodrilus serratus and its importance in the taxonomic position of the Ctenodrilidae. Transactions of the American Microscopial Society, 95 (2), 156–164.
http://dx.doi.org/10.2307/3225058George, J.D. & Petersen, M.E. (1991) The validity of the genus Zeppelina Vaillant (Polychaeta: Ctenodrilidae). Ophelia Supplement, 5, 89–100.
Grube, A.E. (1859) Annulata Örstediana. Enumeratio Annulatorum, quae in itinere Indiam Occidentalem et Americam Centralem annis 1845-1848 suscepto legit cl. A.S. Oersted, adjectis speciebus nonnullis a cl. H. Kroyero in itinere ad Americam meridionalem collectis. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Köbenhavn, 3, 105–120.
Halt, M., Rouse G.W., Petersen, M.E. & Pleijel, F. (2006) Cirratuliformia. In: Rouse, G.W. & Pleijel, F. (Eds.), Reproductive biology and phylogeny of Annelida. Science Publishers, Enfield, NH, pp. 497–520.
Hartman, O. (1961) Polychaetous annelids from California. Allan Hancock Pacific Expeditions, 25, 1–226.
Hobson, K.D. & Banse, K. (1981) Sedentariate and archiannelid polychaetes of British Columbia and Washington. Canadian Bulletin of Fisheries and Aquatic Sciences, 209, 1–144.
Johnson, H.P. (1901) The Polychaeta of the Puget Sound region. Proceedings of the Boston Society for Natural History, 29 (18), 381–437.
Katoh, K. & Standley, D.M. (2013) MAFFT Multiple Sequence Alignment Software Version 7: Improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.
http://dx.doi.org/10.1093/molbev/mst010Kennel, J. (1882) Über Ctenodrilus pardalis Clap. Ein Beitrag zur Kenntnis der Anatomie und Knospung der Anneliden. Mit 1 Taf. Arbeiten aus dem zoolog-zootom. Institut in Würzburg, V, 273–429.
Kück, P. & Meusemann, K. (2010) FASconCAT. Zool. Forschungsmuseum A. Koenig, Bonn.
Lankester, E.R. (1867) Contribution to the knowledge of the lower annelids. Transactions of the Linnean Society, Vol. XXVI, S. 631–646, Taf. 48–49.
Magalhães, W.F., Bailey-Brock, J.H. & Davenport, J.S. (2011) On the genus Raphidrilus Monticelli, 1910 (Polychaeta: Ctenodrilidae) with description of two new species. Zootaxa, 2804, 1–14.
Malmgren, A.J. (1867) Annulata Polychaeta Spetsbergiae, Groenlandiae, Islandiae et Scandinaviae hactenus cognita. Ex Officina Frenckelliana, Helsingfors, 127 pp.
Mesnil, F. & Caullery, M. (1867) Sur la position systématique du genre Ctenodrilus Clap.; ses affinités avec les Cirratuliens. Comptes Rendus de l’Académie des Sciences Paris, CXXV, 542–544.
Mesnil, F. & Caullery, M. (1898) Les formes epitoques et l'évolution des Cirratuliens. In: Annales de l'Université de Lyon, XXXIX, Chapitre IV genre Ctenodrilus, pp. 1–200.
Montagu, G. (1808) Description of several marine animals found on the south coast of Devonshire. Transactions of the Linnean Society of London, 9, 81–114.
http://dx.doi.org/10.1111/j.1096-3642.1818.tb00327.xMonticelli, F.S. (1892) Notizia preliminare intorno ad alcuni inquilini degli Holothurioidea del golfo di Napoli. Monitore Zoologico Italiano, III, 12, 246–256.
Monticelli, F.S. (1893) Sullo Ctenodrilus serratus O. Schmidt. Bollettino della Società dei naturalisti in Napoli, 7, 39–44.
Monticelli, F.S. (1910) Raphidrilus nemasoma Montic. Nuovo Ctendorilide del Golfo di Napoli. Revisione de' Ctenodrilidi. Archivio zoologico: pubblicato sotto gli auspicii della Unione Zoologica Italiana, 4, 401–436.
Müller, O.F. (1774) Vermium terrestrium et fluviatilium, seu animalium infusoriorum, helminthicorum, et testaecorum, non marinorum, succincta historia. Vol. 2. Heineck & Faber, Havniæ & Lipsiæ, 214 pp.
Müller O.F. (1776) Zoologiae Danicae Prodromus seu Animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. Typiis Hallageriis, Hafniae, 274 pp.
Örsted, A.S. (1843) Annulatorum danicorum conspectus. Volume Fasc. 1 Maricolae (Quæstio ab universitate Hafniensi ad solvendum proposita et proemio ornata). Librariae Wahlianae, Hafniae (Copenhagen), 52 pp.
Osborn, K.J., Haddock, S.H.D., Pleijel, F., Madin, L.P. & Rouse, G.W. (2009) Deep-sea, swimming worms with luminescent "bombs". Science, 325 (5943), 964.
http://dx.doi.org/10.1126/science.1172488Palumbi, S.R. (1996) Nucleic acids II: The polymerase chain reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular Systematics. Sinauer Ass., Sunderland, Ma, pp. 205–247.
Petersen, M.E. & George, J.D. (1991) A new species of Raricirrus from northern Europe, with notes on its biology and a discussion of the affinities of the genus (Polychaeta: Ctenodrilidae). Ophelia, Supplement 5 (Systematics, Biology and Morpholology of World Polychaeta), 185–208.
Quatrefages, A. (1866) Histoire naturelle des Annelés marins et d'eau douce. Annélides et Géphyriens. Vol. 1. Librarie Encyclopédique de Roret, Paris, 588 pp.
Reish, D.J. (1980) The effect of different pollutants on ecologically important polychaete worms. United States Environmental Protection Agency, Cincinnati, OH, EPA 600/3-80-053.
Reish, D.J. & Barnard, J.L. (1967) The benthic polychaeta and amphipoda of Morro Bay, California. Proceedings of the United States National Museum, 120 (3565), 1–26.
http://dx.doi.org/10.5479/si.00963801.120-3565.1Riser, N.W. 1984. General observations on the intertidal interstitial fauna of New Zealand. Tane, 30, 239–249.
Rosa, D. (1889) Il Ctenodrilus pardalis Clap, a Rapallo. Bollettino dei Musei di Zoologia ed Anatomia Comparata, 4:69, 1–4 pp. [Torino]
Rouse, G.W. & Pleijel, F. (2001) Polychaetes. Oxford University Press, Oxford, 354 pp.
Scharff, R. (1887) On Ctenodrilus parvulus n.sp. Quarterly Journal of Microscopical Science, 27, 591–604.
Schmidt, O. (1857) Zur Kenntnis der Turbellaria, Rhabdocoela und einiger anderer Wuermer des Mittelmeeres. Sitzungsberichte der Kaiserliche Akademie der Wissenschaften, Wien, Mathematisch-Naturwissenschaftliche Klasse, 23 (2), 347–366, pls. 1–5.
Sokolow, I. (1911a) Über eine neue Ctenodrilusart und ihre Vermehrung. Zeitschrift für wissenschaftliche Zoologie, Leipzig, 97 (3), 546–603.
Sokolow, I. (1911b) Über eine neue Ctenodrilusart. Nachtrag zu einer früheren Arbeit. Zoologischer Anzeiger, Leipzig 38 (9/10), 222–226.
Stamatakis A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
http://dx.doi.org/10.1093/bioinformatics/btl446Sudzuki, M. & Sekiguchi, K. (1972) Some remarks on five aberrant annelids from the culture water of Japanese horse-shoe crabs. The Science Reports of the Tokvo Kvoiku Daigaku Section B, 15, 39–56.
Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
http://dx.doi.org/10.1093/molbev/mst197Vaillant, L. (1890) Histoire naturelle des Annelés marins et d'eau douce. Lombriciniens, Hirudiniens, Bdellomorphes, Térétulariens et Planariens. Paris, Libraire Encyclopédique de Roret, 3, 1–454.
Vejdovský, F. (1884) System und Morphologie der Oligochaeten. Franz Řivnáč, Prag, 166 pp., 16 leaves of plates.
Wakeman, K.C. & Leander, B.S. (2013) Molecular phylogeny of marine gregarine parasites (Apicomplexa) from tube-forming polychaetes (Sabellariidae, Cirratulidae, and Serpulidae), including descriptions of two new species of Selenidium. The Journal of Eukaryotic Microbiology, 60, 514–525.
http://dx.doi.org/10.1111/jeu.12059Weidhase, M., Bleidorn, C. & Simon, C.A. (2016) On the taxonomy and phylogeny of Ctenodrilus (Annelida: Cirratulidae) with a first report from South Africa. Marine Biodiversity, 1–10. [published online]
http://dx.doi.org/10.1007/s12526-015-0355-3Westheide, W. (1981) Interstitielle Fauna von Galapagos. XXVI. Questidae, Cirratulidae, Acrocirridae, Ctenodrilidae (Polychaeta). Mikrofauna des Meeresbodens, 82, 1–24.
Westheide, W., Hass-Cordes, E., Krabusch, M. & Müller, M.C.M. (2003) Ctenodrilus serratus (Polychaeta: Ctenodrilidae) is a truly amphi-Atlantic meiofauna species – evidence from molecular data. Marine Biology, 142, 637–642.
Wilfert, M. (1973) Ein Beitrag zur Morphologie, Biologie und systematischen Stellung des Polychaeten Ctenodrilus serratus. Helgoländer wiss Meeresunters, 25, 332–346.
http://dx.doi.org/10.1007/BF01611202Zeppelin, G.M. (1883) Über den Bau und die Teilungsvorgänge in Ctenodrilus monostylus n. sp. Zeitschrift für wiss. Zoologie, 39, 615–652.