Skip to main content Skip to main navigation menu Skip to site footer
Type: Articles
Published: 2012-01-06
Page range: 1–35
Abstract views: 283
PDF downloaded: 17

Spionidae (Annelida) from shallow waters around the British Islands: an identification guide for the NMBAQC Scheme with an overview of spionid morphology and biology

A.V. Zhirmunsky Institute of Marine Biology, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, 690059, Russia 2Far Eastern Federal University, 8 Sukhanov Street, Vladivostok 690091, Russia
Annelida morphology ecology reproductive biology polychaetes key to genera

Abstract

This is the third guide in a series aimed to help in the identification of northern European samples through the NationalMarine Biological Analytical Quality Control (NMBAQC) Scheme. A review is presented of general morphology andbiology of one of the most common groups in marine and estuarine communities worldwide, spionid polychaetes(Annelida: Spionidae). An identification key to 20 genera and a list of 80 species likely to occur in waters around Britainand Ireland are provided based on the taxonomic literature, collection records and the author’s examination of various collections. The names are updated but some records need verification.

References

  1. Aguirrezabalaga, F. & Ceberio, A. (2005) Spionidae (Annelida: Polychaeta) from the Capbreton Canyon (Bay of Biscay, NE Atlantic) with description of a new genus and three new species. Marine Biology Research, 1, 267–280.

    Allen, E.J. (1904) The anatomy of Pœcilochætus, Claparède. Quarterly Journal of Microscopical Science, London, 48, 79–151.

    Atkins, S.M., Jones, A.M. & Garwood, P.R. (1987) The ecology and reproductive cycle of a population of Marenzelleria viridis (Annelida: Polychaeta: Spionidae) in the Tay Estuary. Proceedings of the Royal Society of Edinburgh, 92B, 311–322.

    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, 30, 411–425.

    Bellan, G., Dauvin, J.-C. & Laubier, L. (2003) The genus Lindaspio (Annelida: Polychaeta: Spionidae), and a new species from an oil field off Congo, western Africa. Journal of Natural History, 37, 2413–2414.

    Benham, W.B. (1896) Archiannelida, Polychaeta, and Myzostomaria. In: Harmer, S.F. & Shipley, A.E. (Eds.), The Cambridge Natural History. Vol. 2, MacMillan and Co., U.K., pp. 241–344.

    Bick, A. (2005) A new Spionidae (Polychaeta) from North Carolina, and a redescription of Marenzelleria wireni Augener, 1913, from Spitsbergen, with a key for all species of Marenzelleria. Helgoland Marine Research, 59, 265–272.

    Bick, A. & Gosselck, F. (1985) Arbeitsschlüssel zur Bestimmung der Polychaeten der Ostsee. Mitteilungen aus dem zoologischen Museum in Berlin, 61, 171–272.

    Blake, J.A. (1983) Polychaetes of the family Spionidae from South America, Antarctica, and adjacent seas and islands. Biology of the Antarctic Seas XIV. Antarctic Research Series, 39, 205–288.

    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. & Scott, P.H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Volume 6. The Annelida Part 3 - Polychaeta: Orbiniidae to Cossuridae, Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 81–223.

    Blake, J.A. (2006) Spionida. In: Rouse, G. & Pleijel, F. (Eds.), Reproductive Biology and Phylogeny of Annelida. Vol. 4 of Series: Reproductive Biology and Phylogeny, Science Publisher, Enfield, NH, pp. 565–638.

    Blake, J.A. & Arnofsky, P.L. (1999) Reproduction and larval development of the spioniform Polychaeta with application to systematics and phylogeny. Hydrobiologia, 402, 57–106.

    Blake, J.A. & Evans, J.W. (1973) Polydora and related genera as borers in mollusk shells and other calcareous substrates (Polychaeta: Spionidae). Veliger, 15, 235–249.

    Blake, J.A. & Grassle, J.F. (1994) Benthic community structure on the U.S. South Atlantic slope off the Carolinas: Spatial heterogeneity in a current-dominated system. Deep-Sea Research II, 41, 835–874.

    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.

    Bleidorn, C., Vogt, L. & Bartolomaeus, T. (2003a) A contribution to sedentary polychaete phylogeny using 18S rRNA sequence data. Journal of zoological Systematics and evolutionary Research, 41, 186–195.

    Bleidorn, C., Vogt, L. & Bartolomaeus, T. (2003b) New insights into polychaete phylogeny (Annelida) inferred from 18S rDNA sequences. Molecular Phylogenetics and Evolution, 29, 279–288.

    Böggemann, M. (1998) Polychaeten aus der Deutschen Bucht. Courier Forschungsinstitut Senckenberg, 202, 1–315.

    Böggemann, M. (2009) Polychaetes (Annelida) of the abyssal SE Atlantic. Organisms Diversity & Evolution, 9, 251–428.

    Buchanan, F. (1890) Hekaterobranchus shrubsolii, a new genus and species of the family Spionidae. Quarterly Journal of Microscopical Science, London, 31, 175–200.

    Claparède, E. (1861) Über Polydora cornuta Bosc. Archiv für Anatomie und Physiologie, 3, 542–544.

    Claparède, E. (1868) Les Annélides Chétopodes du Golfe de Naples. Ramboz et Schuchardt, Genève et Bale, 500 pp.

    Claparède, E. (1869) Les Annélides Chétopodes du Golfe de Naples. Seconde partie. Mémoires de la Société de Physique et d'Histoire Naturelle de Genève, 20, 1–225.

    Claparède, E. (1870) Les Annélides Chétopodes du Golfe de Naples. Supplément. Mémoires de la Société de Physique et d'Histoire Naturelle de Genève, 20, 365–542.

    Clark, R.B. (1960) The fauna of the Clyde Sea area. Polychaeta. Scottish Marine Biological Association, Millport, 71 pp.

    Dales, R.P. (1962) The polychaete stomodeum and the inter-relationships of the families of Polychaeta. Proceedings of the Zoological Society of London, 139, 389–428.

    Dales, R.P. (1977) The polychaete stomodeum and phylogeny. In: Reish, D.J. & Fauchald, K. (Eds.), Essays on Polychaetous Annelids in Memory of Dr. Olga Hartman, Allan Hancock Foundation, University of Southern California, Los Angeles, pp. 525–546.

    Dauer, D.M. (1983) Functional morphology and feeding behavior of Scolelepis squamata (Polychaeta: Spionidae). Marine Biology, 77, 279–285.

    Dauer, D.M. (1985) Functional morphology and feeding behavior of Paraprionospio pinnata (Polychaeta: Spionidae). Marine Biology, 85, 143–151.

    Dauer, D.M., Maybury, C.A. & Ewing, R.M. (1981) Feeding behavior and general ecology of several spionid polychaetes from the Chesapeake Bay. Journal of Experimental Marine Biology and Ecology, 54, 21–38.

    Day, J.H. (1934) Development of Scolecolepis fuliginosa (Clpd.). Journal of the Marine Biological Association of the United Kingdom, 19, 633–654.

    Delgado-Blas, V.H. & Díaz-Díaz, Ó. (2010) Description of two new species of Malacoceros and Rhynchospio spionids (Polychaeta: Spionidae) from the Grand Caribbean Region. Revista Chilena de Historia Natural, 83, 249–257.

    Ditlevsen, H. (1929) Polychæta. Zoology of the Faroes, 16, 1–83.

    Dorsett, D.A. (1961) The behavior of Polydora ciliata (Johnst.). Tube-building and burrowing. Journal of the Marine Biological Association of the United Kingdom, 41, 577–590.

    Eisig, H. (1879) Die Seitenorgane und becherförmigen Organe der Capitelliden. Mittheilungen aus der Zoologischen Station zu Neapel, 1, 278–348.

    Eisig, H. (1914) Zur Systematik, Anatomie und Morphologie der Ariciiden nebst Beiträgen zur generellen Systematik. Mittheilungen aus der Zoologischen Station zu Neapel, 21, 153–593.

    Eleftheriou, A. (1970) Notes on the polychaete Pseudopolydora pulchra (Carazzi) from British waters. Cahiers de Biologie Marine, 11, 459–474.

    Eliason, A. (1920) Polychaeta. Biologisch-faunistische Untersuchungen aus dem Öresund. Lunds Universitets Årsskrift, N. F., Avd. 2, 16, 1–103.

    Eno, N.C., Clark, R.A. & Sanderson, W.G. (1997) Non-native marine species in British waters: a review and directory. Joint Nature Conservation Committee, Petersborough, 152 pp.

    Fauchald, K. (1977) The polychaete worms. Definitions and keys to the orders, families and genera. Natural History Museum of Los Angeles County, Science Series, 28, 1–188.

    Fauchald, K. & Rouse, G. (1997) Polychaete systematics: Past and present. Zoologica Scripta, 26, 71–138.

    Fauvel, P. (1927) Polychètes sédentaires. Addenda aux Errantes, Archiannélides, Myzostomaires. Faune de France, 16, 1–494.

    Fiege, D., Ramey, P.A. & Ebbe, B. (2010) Diversity and distributional patterns of Polychaeta in the deep South Atlantic. Deep Sea Research Part I: Oceanographic Research Papers, 57, 1329–1344.

    Foster, N.M. (1971) Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Studies on the Fauna of Curaçao and other Caribbean Islands, 36, 1–183.

    Friedrich, H. (1938) Polychaeta. Die Tierwelt der Nord- und Ostsee, 32, 1–201.

    George, J.D. (1974) The marine fauna of Lundy. Polychaeta (marine bristleworms). Lundy Field Society Annual Report, 25, 33–48.

    Gibson, G., MacDonald, K. & Dufton, M. (2010) Morphogenesis and phenotypic divergence in two developmental morphs of Streblospio benedicti (Annelida, Spionidae). Invertebrate Biology, 129, 328–343.

    Gibson, R.J.H. (1886) Report on the Vermes of the L. M. B. C. district. In: Herdman, W.A. (Ed.), The first report upon the fauna of Liverpool Bay and the neighbouring seas, Longmans, Green & Co., London, pp. 144–160.

    Haigler, S.A. (1969) Boring mechanism of Polydora websteri inhabiting Crassostrea virginica. American Zoologist, 9, 821–828.

    Hamond, R. (1966) The Polychaeta of the coast of Norfolk. Cahiers de Biologie Marine, 7, 383–436.

    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.

    Hartman, O. (1960) Systematic account of some marine invertebrate animals from the deep basins off southern California. Allan Hancock Pacific Expeditions, 22, 69–215.

    Hartman, O. (1971) Abyssal polychaetous annelids from the Mozambique Basin off southeast Africa, with a compendium of abyssal polychaetous annelids from world-wide areas. Journal of the Fisheries Research Board of Canada, 28, 1407–1428.

    Hartman, O. & Fauchald, K. (1971) Deep-water benthic polychaetous annelids off New England to Bermuda and other North Atlantic Areas. Part II. Allan Hancock Foundation Monographs, 6, 1–327.

    Hartmann-Schröder, G. (1971) Annelida, Borstenwürmer, Polychaeta. Die Tierwelt Deutschlands und der angrenzenden Meeresteile nach ihren Merkmalen und nach ihrer Lebensweise, 58, 1–594.

    Hartmann-Schröder, G. (1996) Annelida, Borstenwürmer, Polychaeta. Die Tierwelt Deutschlands und der angrenzenden Meeresteile nach ihren Merkmalen und nach ihrer Lebensweise, 58, 1–645.

    Hausen, H. (2005) Chaetae and chaetogenesis in polychaetes (Annelida). Hydrobiologia, 535–536, 37–52.

    Hempel, C. (1957) Über den Röhrenbau und die Nahrungsaufnahme einiger Spioniden (Polychaeta Sedentaria) der deutschen Küsten. Helgoländer Wissenschaftliche Meeresuntersuchungen, 6, 100–135.

    Hempel, C. (1960) Über das Festsitzen der Larven und die Bohrtätigkeit der Jugendstadien von Polydora ciliata (Polychaeta Sedentaria). Helgoländer Wissenschaftliche Meeresuntersuchungen, 7, 80–92.

    Hilbig, B. & Blake, J.A. (2006) Deep-sea polychaete communities in the Northeast Pacific Ocean off the Gulf of the Farallones, California. Bulletin of Marine Science, 78, 243–269.

    Jägersten, G. (1952) Studies on the morphology, larval development and biology of Protodrilus. Zoologiska Bidrag från Uppsala, 29, 426–511.

    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.

    Jelsing, J. & Eibye-Jacobsen, D. (2010) Ultrastructure of the extensively developed nuchal organs of Laonice bahusiensis (Annelida: Canalipalpata: Spionidae). Journal of Morphology, 271, 376–382.

    Johnston, G. (1827) Contributions to the British Fauna. Journal of Zoology, London, 3, 321–336.

    Johnston, G. (1838) Miscellanea Zoologica. III.–The British Ariciadæ. Magazine of Zoology and Botany, Edinburgh, 2, 63–73.

    Johnston, G. (1865) A catalogue of the British non-parasitical worms in the collection of the British Museum. British Museum, London, 365+XX pls. pp.

    Keferstein, W. (1862) Untersuchungen über niedere Seethiere. Zeitschrift für wissenschaftliche Zoologie, 12, 1–147.

    Kirkegaard, J.B. (1996) Havbørsteorme. II. Sedentaria. Danmarks Fauna, 86, 1–451.

    Maciolek, N.J. (1981) Spionidae (Annelida: Polychaeta) from the Galápagos Rift geothermal vents. Proceedings of the Biological Society of Washington, 94, 826–837.

    Maciolek, N.J. (1985) A revision of the genus Prionospio Malmgren, with special emphasis on species from the Atlantic Ocean, and new records of species belonging to the genera Apoprionospio Foster and Paraprionospio Caullery (Polychaeta, Annelida, Spionidae). Zoological Journal of the Linnean Society, London, 84, 325–383.

    Maciolek, N.J. (1987) New species and records of Scolelepis (Polychaeta: Spionidae) from the east coast of North America, with a review of the subgenera. Bulletin of the Biological Society of Washington, 7, 16–40.

    Maciolek, N.J. (1990) A redescription of some species belonging to the genera Spio and Microspio (Polychaeta: Annelida) and descriptions of three new species from the northwestern Atlantic Ocean. Journal of Natural History, 24, 1109–1141.

    Maciolek, N.J. (2000) New species and records of Aonidella, Laonice, and Spiophanes (Polychaeta: Spionidae) from shelf and slope depth of the western north Atlantic. Bulletin of Marine Science, 67, 529–547.

    Mackie, A.S.Y. (1984) On the identity and zoogeography of Prionospio cirrifera Wirén, 1883 and Prionospio multibranchiata Berkeley, 1927 (Polychaeta; Spionidae). In: Hutchings, P.A. (Ed.), Proceedings of the First International Polychaete Conference, Sydney, The Linnean Society of New South Wales, Sydney, pp. 35–47.

    Mackie, A.S.Y. (1996) Taxonomy and phylogeny of spioniform polychaetes (Annelida). Dissertation. Göteborg University, Göteborg,

    Mackie, A.S.Y. & Duff, A.A. (1986) Atherospio disticha gen. et sp. nov. (Polychaeta: Spionidae) from Loch Tuirnaig, west coast of Scotland. Ophelia, 25, 139–146.

    Mackie, A.S.Y. & Erséus, C. (1997) Annelida. In: Howson, C.M. & Picton, B.E. (Eds.), The species directory of the marine fauna and flora of the British Isles and surrounding seas, pp. 102–136.

    Mackie, A.S.Y., Parmiter, C. & Tong, L.K.Y. (1997) Distribution and diversity of Polychaeta in the southern Irish Sea. Bulletin of Marine Science, 60, 467–481.

    Malmgren, A.J. (1867) Annulata polychæta Spetsbergiæ, Grönlandiæ, Islandiæ et Scandinaviæ hactenus cognita. Öfversigt af Konglingar Vetenskaps-Akademiens Förhandlingar, Stockholm, 24, 127–235.

    McIntosh, W.C. (1874) On the Annelida of St. Andrews. Annals and Magazine of Natural History, Series 8, 14, 144–155.

    McIntosh, W.C. (1875) The marine invertebrates and fishes of St. Andrews. Adam and Charles Black, Edinburgh, 186+9 pls. pp.

    McIntosh, W.C. (1909) Notes from the Gatty Marine Laboratory, St. Andrews. No. 31. 1. On a young stage of Gadus luscus with bold transverse bars of pigment. 2. On the British Spionidae. 3. On the Spionidae dredged by H.M.S. 'Porcupine' in 1869 and 1870. Annals and Magazine of Natural History, Series 8, 3, 153–180.

    McIntosh, W.C. (1915) A monograph of the British Marine Annelids. Vol. 3. Part 1. - Text. Polychaeta. Opheliidae to Ammocharidae. Ray Society, London, 368 pp.

    McIntosh, W.C. (1922) Notes from the Gatty Marine Laboratory, St. Andrews. No. XLIV. 1. On new and rare Polychaeta, Gephyrea, etc., from various regions. 2. Recent additions to the British marine Polychaeta (continued). Annals and Magazine of Natural History, Series 9, 9, 1–30.

    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, 3–66.

    Meißner, K., Bick, A. & Bastrop, R. (2011) On the identity of Spio filicornis (O.F. Müller, 1776) — with the designation of a neotype, and the description of two new species from the North East Atlantic Ocean based on morphological and genetic studies. Zootaxa, 2815, 1–27.

    Meißner, K. & Hutchings, P.A. (2003) Spiophanes species (Polychaeta: Spionidae) from Eastern Australia - with description of new species, new records and an emended generic diagnosis. Records of the Australian Museum, 55, 117–140.

    Mesnil, F. (1896) Études de morphologie externe chez les Annélides. I. Les Spionidiens des côtes de la Manche. Bulletin scientifique de la France et de la Belgique, 29, 110–287.

    Moore, H.B. (1937) Marine fauna of the Isle of Man. University Press of Liverpool, Liverpool, 293 pp.

    Nelson-Smith, A., Knight-Jones, P. & Knight-Jones, E.W. (1990) Annelida. In: Hayward, P.J. & Ryland, J.S. (Eds.), The Marine Fauna of the British Isles and North-West Europe. Vol. 1. Introduction and Protozoans to Arthropods, Clarendon Press, Oxford, pp. 201–306.

    Orrhage, L. (1964) Anatomische und morphologische Studien über die polychaetenfamilien Spionidae, Disomidae und Poecilochaetidae. Zoologiska Bidrag från Uppsala, 36, 335–405.

    Örsted, A.S. (1843) Annulatorum danicorum conspectus. Fasc. I. Maricolæ. Sumtibus Librariæ Wahlianæ, Hafniæ, 52 pp.

    Pardo, E.V. & Amaral, A.C.Z. (2004) Feeding behavior of Scolelepis sp. (Polychaeta: Spionidae). Brazilian Journal of Oceanography, 52, 75–79.

    Pettibone, M.H. (1962) New species of polychaete worms (Spionidae: Spiophanes) from the east and west coast of North America. Proceedings of the Biological Society of Washington, 75, 77–88.

    Pettibone, M.H. (1963) Revision of some genera of polychaete worms of the family Spionidae, including the description of a new species of Scolelepis. Proceedings of the Biological Society of Washington, 76, 89–104.

    Purschke, G. & Hausen, H. (2007) Lateral organs in sedentary polychaetes (Annelida) - Ultrastructure and phylogenetic significance of an insufficiently known sense organ. Acta Zoologica (Stockholm), 88, 23–39.

    Radashevsky, V.I. (1993) Revision of the genus Polydora and related genera from the North West Pacific (Polychaeta: Spionidae). Publications of the Seto Marine Biological Laboratory, 36, 1–60.

    Radashevsky, V.I. (2007) Morphology and biology of a new Rhynchospio species (Polychaeta: Spionidae) from the South China Sea, Vietnam, with the review of Rhynchospio taxa. Journal of Natural History, 41, 985–997.

    Radashevsky, V.I. & Fauchald, K. (2000) Chaetal arrangement and homology in spionids (Polychaeta: Spionidae). Bulletin of Marine Science, 67, 13–23.

    Radashevsky, V.I. & Lana, P.C. (2009) Laonice (Annelida: Spionidae) from South and Central America. Zoosymposia, 2, 265–295.

    Radashevsky, V.I. & Olivares, C. (2005) Polydora uncinata (Polychaeta: Spionidae) in Chile: an accidental transportation across the Pacific. Biological Invasions, 7, 489–496.

    Rainer, S. (1973) Polydora and related genera (Polychaeta: Spionidae) from Otago waters. Journal of the Royal Society of New Zealand, 3, 545–564.

    Ramberg, J.P. & Schram, T.A. (1983) A systematic review of the Oslofjord species of Polydora Bosc and Pseudopolydora Czerniavsky, with some new biological and ecological data (Polychaeta: Spionidae). Sarsia, 68, 233–247.

    Ranade, M.R. & Colman, J.S. (1957) Notes on the west pond at Port Erin. Report of the Port Erin Marine Biological Station, 69, 34–39.

    Rasmussen, E. (1973) Systematics and ecology of the Isefjord marine fauna (Denmark). Ophelia, 11, 1–495.

    Read, G.B. (1975) Systematics and biology of polydorid species (Polychaeta: Spionidae) from Wellington Harbour. Journal of the Royal Society of New Zealand, 5, 395–419.

    Rouse, G.W. & Fauchald, K. (1997) Cladistics and polychaetes. Zoologica Scripta, 26, 139–204.

    Rouse, G.W. & Pleijel, F. (2001) Polychaetes. Oxford University Press, New York, 354 pp.

    Rousset, V., Pleijel, F., Rouse, G.W., Erséus, C. & Siddall, M.E. (2007) A molecular phylogeny of annelids. Cladistics, 23, 41–63.

    Rullier, F. (1951) Étude morphologique, histologique et physiologique de l'organe nucal chez les Annélides Polychètes Sédentaires. Annales de l'Institut océanographique, Paris, 25, 207–341.

    Sæmundsson, B. (1918) Bidrag til Kundskaben om Islands polychæte Børsteorme. (Annulata polychæta Islandiæ). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening I Kjøbenhavn, 69, 165–241.

    Saint-Joseph, A.d. (1894) Les Annélides Polychètes des côtes de Dinard, pt. 3. Annales des Sciences Naturelles, Zoologie, VII Série, Paris, 17, 1–395.

    Sars, M. (1873) Bidrag til Kundskaben om Christianiafjordens Fauna. III. Johan Dahl, Christiania, 81 pp.

    Sato-Okoshi, W. & Okoshi, K. (2000) Structural characteristics of self-excavated burrows by boring polydorid species (Polychaeta, Spionidae). Bulletin of Marine Science, 67, 235–248.

    Sigvaldadóttir, E. (1998) Cladistic analysis and classification of Prionospio and related genera (Polychaeta, Spionidae). Zoologica Scripta, 27, 175–187.

    Sigvaldadóttir, E. (2002) Polychaetes of the genera Prionospio and Aurospio (Spionidae, Polychaeta) from Icelandic waters. Sarsia, 87, 207–215.

    Sigvaldadóttir, E., Mackie, A.S.Y. & Pleijel, F. (1997) Generic interrelationships within the Spionidae (Annelida: Polychaeta). Zoological Journal of the Linnean Society, London, 119, 473–500.

    Sikorski, A.V. (1994a) Malacoceros (Polychaeta, Spionidae) in the Arctic Ocean. Zoologichesky Zhurnal, 73, 21–35. (In Russian with English Summary).

    Sikorski, A.V. (1994b) New arctic species of Scolelepis (Polychaeta, Spionidae). Mémoires du Muséum National d'Histoire Naturelle, Paris, 162, 279–286.

    Sikorski, A.V. (2001) Spionidae Grube, 1850. In: Zhirkov, I.A. (Ed.), Polychaeta of the Arctic Ocean, Yanus-K, Moscow, pp. 271–332.

    Sikorski, A.V. (2003) Laonice (Polychaeta, Spionidae) in the Arctic and the North Atlantic. Sarsia, 88, 316–345.

    Sikorski, A.V. (2011) Review of Laonice (Spionidae, Annelida) with remarks on several species and a description of a new species from South Africa. Italian Journal of Zoology, 78, 201-214.

    Sikorski, A.V. & Bick, A. (2004) Revision of Marenzelleria Mesnil, 1896 (Spionidae, Polychaeta). Sarsia, 89, 253–275.

    Söderström, A. (1920) Studien über die Polychätenfamilie Spionidae. Inaugural-Dissertation. Almquist & Wicksells, Uppsala, 286 pp.

    Soto, E.H., Paterson, G.L.J., Billett, D.S.M., Hawkins, L.E., Galéron, J. & Sibuet, M. (2010) Temporal variability in polychaete assemblages of the abyssal NE Atlantic Ocean. Deep Sea Research Part II: Topical Studies in Oceanography, 57, 1396–1405.

    Southern, R. (1910) The marine worms (Annelida) of Dublin Bay and the adjoining district. Proceedings of the Royal Irish Academy, 28B, 215–264.

    Southern, R. (1914) Archiannelida and Polychaeta. Proceedings of the Royal Irish Academy, 31, 1–160.

    Southward, E.C. (1956) On some Polychaeta of the Isle of Man. Annals and Magazine of Natural History, Series 12, 9, 257–270.

    Storch, V. & Schlötzer-Schrehardt, U. (1988) Sensory structures. In: Westheide, W. & Hermans, C.O. (Eds.), The Ultrastructure of Polychaeta. Microfauna Marina 4, Gustav Fischer Verlag, Stuttgart-New York, pp. 121–133.

    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.

    Struck, T.H., Paul, C., Hill, N., Hartmann, S., Hösel, C., Kube, M., Lieb, B., Meyer, A., Tiedemann, R., Purschke, G. & Bleidorn, C. (2011) Phylogenomic analyses unravel annelid evolution. Nature, 471, 95–98.

    Struck, T.H., Schult, N., Kusen, T., Hickman, E., Bleidorn, C., McHugh, D. & Halanych, K.M. (2007) Annelid phylogeny and the status of Sipuncula and Echiura. BMC Evolutionary Biology, 7. Available from: http://www.biomedcentral.com/1471-2148/7/57 (accessed 19 February 2011).

    Taghon, G.L., Nowell, A.R.M. & Jumars, P.A. (1980) Induction of suspension feeding in spionid polychaetes by high particulate fluxes. Science, New York , 210, 562–564.

    Tauber, P. (1879) Annulata Danica. En kritisk Revision af de i Danmark fundne Annulata, Chaetognatha, Gephyrea, Balanoglossi, Discophorea, Oligochaeta, Gymnocopa og Polychaeta. Copenhagen, 143 pp.

    Templeton, R. (1836) A catalogue of the species of annulose animals, and of rayed ones, found in Ireland, as selected from the papers of the late J. Templeton, Esq., of Cranmore, with localities, descriptions, and illustrations. Magazine of Natural History, London, 9, 233–243.

    Warner, G.F. (1997) Occurrence of epifauna on the periwinkle, Littorina littorea (L.), and interactions with the polychaete Polydora ciliata (Johnston). Hydrobiologia, 355, 41–47.

    Wesenberg-Lund, E. (1951) Polychaeta. Zoology of Iceland, 2, 1–182.

    Wilson, D.P. (1928) The larvae of Polydora ciliata Johnston and Polydora hoplura Claparède. Journal of the Marine Biological Association of the United Kingdom, 15, 567–603.

    Wilson, D.P. & Tebble, N. (1957) Annelida. In: Bruce, J.R., Colman, J.S. & Jones, N.S. (Eds.), Plymouth Marine Fauna, Latimer, Trend & Co., Plymouth, pp. 109–151.

    Würzberg, L., Peters, J., Schüller, M. & Brandt, A. (2011) Diet insights of deep-sea polychaetes derived from fatty acid analyses. Deep Sea Research Part II: Topical Studies in Oceanography, 58, 153–162.

    Yokoyama, H. (2007) A revision of the genus Paraprionospio Caullery (Polychaeta: Spionidae). Zoological Journal of the Linnean Society, 151, 253–284.

    Zottoli, R.A. & Carriker, M.R. (1974a) Burrow morphology, tube formation, and microarchitecture of shell dissolution by the spionid polychaete Polydora websteri. Marine Biology, 27, 307–316.

    Zottoli, R.A. & Carriker, M.R. (1974b) External release of protease by stationary burrow-dwelling polychaetes. Journal of Marine Research, 32, 331–342.

    Zrzavý, J., Ríha, P., Piálek, L. & Janouskovec, J. (2009) Phylogeny of Annelida (Lophotrochozoa): total-evidence analysis of morphology and six genes. BMC Evolutionary Biology, 9. Available from: http://www.biomedcentral.com/1471-2148/9/189 (accessed 19 February 2011).