Skip to main content Skip to main navigation menu Skip to site footer
Type: Articles
Published: 2011-01-28
Page range: 40–62
Abstract views: 198
PDF downloaded: 5

A review of the current status of the Polydora-complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species

School of Biological Sciences, University of Queensland, Brisbane, 4072, QLD, Australia
Annelida Mudworm polydorid marine pest oyster disease estuary aquaculture taxonomy

Abstract

In 1885 Australia’s first recorded marine pest, a mudworm of the Polydora-complex (Spionidae), was identified on Hunter River, New South Wales, oysters (Saccostrea glomerata Gould, 1850). Mudworm is still a serious pest of cultured molluscs in Australia but, although of great concern to many, relatively little progress has been made in resolving the problem. One hundred and thirty years later the identity and life history of this pest remains unclear. The longevity of this problem in Australia is largely due to unaddressed issues of basic taxonomy. This review addresses these issues by presenting the information currently available on the Polydora-complex species in Australia, including taxonomy; history of discovery in the natural environment and in commercial aquaculture facilities; and ecological information. An updated checklist of Polydora-complex species from Australia is also included. It is hoped that by being explicit about the knowledge gaps and problems they can be addressed, ultimately providing a solid taxonomic background for further research into the Polydora-complex species and resolution of the mudworm problem in Australia.

References

  1. Augener, H. (1914) Polychaeta II. Sedentaria. In: W. Michaelsen & R. Hartmeyer (eds), Die Fauna Südwest-Australiens, pp. 1–170.

    ABRS (2010) Australian Faunal Directory. Family Spionidae. Australian Biological Resources Study, Canberra. Available from: http://www.environment.gov.au/biodiversity/abrs/online-resources/fauna/afd/taxa/SPIONIDAE (14 July 2010)

    Australian Museum Business Services Consulting (2002) Port Survey for Introduced Marine Species - Sydney Harbour. Final Report. Australian Museum Business Services for Sydney Ports Corporation. 7 May 2002.

    Bailey-Brock, J.H. (2000) A new record of the polychaete Boccardia proboscidea (Family Spionidae), imported to Hawai'i with oysters. Pacific Science, 54, 27–30.

    Benham, W.B. (1896) Archiannelida, Polychaeta, and Myzostomaria. In: S. F. Harmer & A. E. Shipley (Eds), The Cambridge Natural History. MacMillan & Co. Ltd, London, pp. 241–334.

    Bick, A. (2001) The morphology and ecology of Dipolydora armata (Polychaeta, Spionidae) from the western Mediterranean Sea. Acta Zoologica (Copenhagen), 82, 177–187.

    Blake, J.A. (1969) Reproduction and larval development of Polydora from nothern New England (Polychaeta: Spionidae). Ophelia, 7, 1–63.

    Blake, J.A. (1971) Revision of the genus Polydora from the east coast of North America (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, 75, 1–32.

    Blake, J.A. (1996) Family Spionidae Grube, 1850 In: J. A. Blake, B. Hilbig & P. H. Scott (Eds), The Annelida: Part 3 Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History Santa Barbara, California, pp. 81–224.

    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.P. (1974) Polydora and related genera as borers in mollusk shells and other calcareous substrates. Veliger, 15, 235–250.

    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. & Kudenov, J.D. (1981) Larval development, larval nutrition and growth for two Boccardia species (Polychaeta: Spionidae) from Victoria, Australia. Marine Ecology Progress Series, 6, 175–182, illustr.

    Blake, J.A. & Maciolek, N.J. (1987) A redescription of Polydora cornuta Bosc (Polychaeta: Spionidae) and designation of a neotype. Bulletin of the Biological Society of Washington, 7, 11–15.

    Blake, J.A. & Woodwick, K.H. (1971) A review of the genus Boccardia Carazzi (Polychaeta: Spionidae) with description of two new species. Bulletin of the Southern California Academy of Sciences, 70, 31–42.

    Blake, J.A. & Woodwick, K.H. (1976) A new species of Boccardia (Polychaeta: Spionidae) from two freshwater lakes in southeastern Australia. Records of the Australian Museum, 30, 123–128.

    Blank, M. & Bastrop, R. (2009) Phylogeny of the mud worm genus Marenzelleria (Polychaeta, Spionidae) inferred from mitochondrial DNA sequences. Zoologica Scripta, 38, 313–321.

    Bondad-Reantaso, M.G., McGladdery, S.E., Ladra, D. & Wang, C. (2007) Pearl oyster health: experiences from the Philippines, China, the Persian Gulf and the Red Sea. FAO Fisheries Technical Paper, 111–121.

    Bower, S.M., Blackbourn, J., Meyer, G.R. & Nishimura, D.J.H. (1992) Diseases of cultured Japanese scallops (Patinopecten yessoensis) in British Columbia, Canada. Aquaculture, 107, 201–210, illustr.

    Carazzi, D. (1895) Revisione del genere Polydora Bosc e cenni su due specie che vivono sulle ostriche. Mittheilungen aus der Zoologischen Station zu Neapel, 11, 4–45,plate42.

    Claparede, E. (1869) Les Annelides Chetopodes du Golfe de Naples. Seconde partie. Annelides sedentaires. Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve, 20, 1–225,pls.217–231.

    Claparede, E. (1870) Les Annelides Chetopodes du Golfe de Naples. Supplement. Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve, 20, 365–542, pls 361–316.

    Colgan, D.J., Hutchings, P.A. & Braune, M. (2006) A multigene framework for polychaete phylogenetic studies. Organisms Diversity & Evolution, 6, 220–235.

    Costa-Pierce, B.A. & Bridger, C.J. (2002) The role of marine aquaculture facilities as habitats and ecosystems. Responsible Marine Aquaculture, 105–144.

    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, illustr.

    Davies, P.J. & Hutchings, P.A. (1983) Initial colonization, erosion and accretion on coral substrate. Experimental results, Lizard Island, Great Barrier Reef. Coral Reefs, 2, 27–35.

    Dinley, J., Hawkins, L., Paterson, G., Ball, A.D., Sinclair, I., Sinnett-Jones, P., et al. (2010) Micro-computed X-ray tomography: a new non-destructive method of assessing sectional, fly-through and 3D imaging of a soft-bodied marine worm. Journal of Microscopy-Oxford, 238, 123–133.

    Dorresteijn, A. & Westheide, W. (1999) Reproductive Strategies and Developmental Patterns in Annelids. In: Hydrobiologia: Developments in Hydrobiology. Kluwer Academic Publishers.

    Dunphy, B.J. & Wells, R.M.G. (2001) Endobiont infestation, shell strength and condition index in wild populations of New Zealand abalone, Haliotis iris. Marine and Freshwater Research, 52, 781–786.

    Ford, E. & Hutchings, P. (2005) An analysis of morphological characters of Owenia useful to distinguish species: description of three new species of Owenia (Oweniidae : Polychaeta) from Australian waters. Marine Ecology-an Evolutionary Perspective, 26, 181–196.

    Gibson, G., Paterson, I.G., Taylor, H. & Woolridge, B. (1999) Molecular and morphological evidence of a single species, Boccardia proboscidea (Polychaeta: Spionidae), with multiple development modes. Marine Biology (Berlin), 134, 743–751.

    Gibson, G.D. & Paterson, I.G. (2003) Morphogenesis during sexual and asexual reproduction in Amphipolydora vestalis (Polychaeta: Spionidae). New Zealand Journal of Marine and Freshwater Research, 37, 741–752.

    Government of South Australia - Marine Parks (2010) The Natural Values of South Australia's Marine Environment. Available from: http://www.environment.sa.gov.au/marineparks/pdfs/natural_wonders.pdf (August 23rd, 2010)

    Grube, A.E. (1855) Beschreibungen neuer oder wenig bekannter Anneliden. Arkiv fur Naturgeschichte, Berlin, 21, 81–136, pls. 133–135.

    Gudmundsson, H. (1985) Life history patterns of polychaete species of the family Spionidae. Journal of the Marine Biological Association of the United Kingdom, 65, 93–112.

    Handley, S.J. (2000) Larval development of Boccardia knoxi, a shell-infesting spionid polychaete. New Zealand Journal of Marine and Freshwater Research, 34, 681–687.

    Handley, S.J. & Bergquist, P.R. (1997) Spionid polychaete infestations of intertidal Pacific oysters Crassostrea gigas (Thunberg), Mahurangi harbour, northern New Zealand. Aquaculture, 153, 191–205.

    Hartman, O. (1940) Boccardia proboscidea, a new species of spionid worm from California. Journal of the Washington Academy of Sciences, 30, 382–387.

    Hartman, O. (1943) Description of Polydora websteri. in Loosanoff & Engle Polydora in oysters suspended in the water. Biological Bulletin, 85, 69–78.

    Hartmann-Schröder, G. (1979) Teil 2. Die Polychaeten der tropischen Nordwestküste Australiens (zwischen Derby im Norden und Port Hedland im Süden). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 75–218.

    Hartmann-Schröder, G. (1980) Teil 4. Die Polychaeten der tropischen Nordwestküste Australiens (zwischen Port Samson im Norden und Exmouth im Süden). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 41–110.

    Hartmann-Schröder, G. (1981) Teil 6. Die Polychaeten der tropisch-subtropischen Westküste Australiens (zwischen Exmouth im Norden und Cervantes im Süden). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 19–96.

    Hartmann-Schröder, G. (1982) Teil 8. Die Polychaeten der subtropisch-antiborealen Westküste Australiens (zwischen Cervantes im Norden und Cape Naturaliste im Süden). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 51–118.

    Hartmann-Schröder, G. (1983) Teil 9. Die Polychaeten der antiborealen Südwestküste Australiens (zwischen Dunsborough im Norden und Denmark im Süden). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 123–167.

    Hartmann-Schröder, G. (1984) Teil 10. Die Polychaeten der antiborealen Südküste Australiens (zwischen Albany im Westen und Ceduna im Osten). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 7–62.

    Hartmann-Schröder, G. (1985) Teil 11. Die Polychaeten der antiborealen Südküste Australiens (zwischen Port Lincoln im Westen und Port Augusta im Osten). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 61–99.

    Hartmann-Schröder, G. (1986) Teil 12. Die Polychaeten der antiborealen Südküste Australiens (zwischen Wallaroo im Westen und Port MacDonnell im Osten). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 31–70.

    Hartmann-Schröder, G. (1987) Teil 13. Die Polychaeten der antiborealen Küste von Victoria (Australien) (zwischen Warrnambool im Westen und Port Welshpool im Osten). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 27–66.

    Hartmann-Schröder, G. (1989) Teil 14. Die Polychaeten der antiborealen und subtropisch-tropischen Küste Südost-Australiens zwischen Lakes Entrance (Victoria) im Süden und Maclean (New South Wales) im Norden. In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung der Polychaeten und Ostracoden, pp. 11–63.

    Hartmann-Schröder, G. (1990) Teil 15. Die Polychaeten der subtropisch-tropischen und tropischen Ostküste Australiens zwischen Lake Macquarie (New South Wales) im Süden und Gladstone (Queensland) im Norden. In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksichtigung des Polychaeten und Ostracoden, pp. 41–87.

    Hartmann-Schröder, G. (1991) Teil 16. Die Polychaeten der subtropisch-tropischen bis tropischen Ostküste Australiens zwischen Maclean (New South Wales) und Gladstone (Queensland) sowie von Heron Island (Großes Barriere-Riff). In: G. Hartmann-Schröder & G. Hartmann (Eds), Zur Kenntnis des Eulitorals der australischen Küsten unter besonderer Berücksightigung der Polychaeten und Ostracoden, pp. 17–71.

    Haswell, W.A. (1885) On a destructive parasite of the rock oyster. Proceedings of the Linnean Society of New South Wales, 10, 272–275.

    Hayes, K., Sliwa, C., Migus, S., McEnnulty, F. & Dunstan, P. (2005) National priority pests: Part II. Ranking of Australian marine pests. An independent report undertaken for the Department of Environment and Heritage by CSIRO Marine Research. Feb 2005

    Hewitt, C.L. & Martin, R.B. (1996) Port surveys for introduced marine species—background considerations and sampling protocols. In: CRIMP Technical Report No. 4. Division of Fisheries, CSIRO, Hobart.

    Hewitt, C.L. & Martin, R.B. (2001) Revised protocols for baseline port surveys for introduced marine species—design considerations, sampling protocols and taxonomic sufficiency. In: CRIMP Technical Report No. 22. CSIRO Marine Research, Hobart.

    Hilliard, R. (2004) Best practice for the management of introduced marine pests. A review. Prepared for Global Invasive Species Programme by URS Australia Pty. Ltd. Published by the GISP Secretariat in 2004.

    Hutchings, P. & Glasby, C.J. (2004) Port suveys for introduced marine species-and fate of the material collected. Marine Pollution Bulletin, 48, 1009–1011.

    Hutchings, P.A. & Karageorgopoulos, P. (2003) Designation of a neotype of Marphysa sanguinea (Montagu, 1813) and a description of a new species of Marphysa from eastern Australia. Hydrobiologia, 496.

    Hutchings, P.A. & Murray, A. (1984) Taxonomy of polychaetes from the Hawkesbury River and the southern estuaries of New South Wales, Australia. Records of the Australian Museum Supplement, 3, 1–118.

    Hutchings, P.A. & Rainer, S.F. (1979) The polychaete fauna of Careel Bay, Pittwater, New South Wales, Australia. Journal of Natural History, 13, 745–796.

    Hutchings, P.A. & Recher, H.F. (1974) The fauna of Careel Bay with comments on the ecology of mangrove and sea-grass communities. Australian Zoologist, 18, 99–128.

    Hutchings, P.A. & Turvey, S.P. (1984) The Spionidae of South Australia (Annelida: Polychaeta). Transactions of the Royal Society of South Australia, 108, 1–20.

    Johnston, G. (1838) Miscellanea Zoologica Ariciadae. Magazine of Zoology and Botany, Edinburgh, 2, 63–73, plates62–63.

    Kaluza, P., Kölzsch, A., Gastner, M.T. & Blasius, B. (2010) The complex network of global cargo ship movements. Journal of the Royal Society Interface. Retrieved 04-06-2010 from http://arxiv.org/abs/1001.2172.

    Kent, R.M.L. (1981) The effect of Polydora ciliata on the shell strength of Mytilus edulis. Journal Du Conseil Conseil International Pour L'Exploration De La Mer, 39, 252–255, illustr.

    Kojima, H. & Imajima, M. (1982) Burrowing polychaetes in the shells of the abalone Haliotis diversicolor aquatilis chiefly on the species of Polydora. Bulletin of the Japanese Society of Scientific Fisheries, 48, 31–35,illustr.

    Kostromkina, T.B., Manchenko, G.P. & Radashevsky, V.I. (1991) Chromosome number in the polychaete species Boccardiella hamata and Polydora glycymerica (Spionidae). Biologiya Morya, Vladivostok, 1991, 101–103,illustr.

    Labura, Z. & Hrs-Brenko, M. (1990) Infestation of European flat oyster (Ostrea edulis) by polychaete (Polydora hoplura) in the northern Adriatic Sea. Acta Adriatica, 31, 173–181,illustr.

    Levin, L.A., Ekau, W., Gooday, A.J., Jorissen, F., Middelburg, J.J., Naqvi, S.W.A., et al. (2009) Effects of natural and human-induced hypoxia on coastal benthos. Biogeosciences, 6, 2063–2098.

    Lleonart, M., Handlinger, J. & Powell, M. (2003a) Spionid mudworm infestation of farmed abalone (Haliotis spp.). Aquaculture, 221, 85–96.

    Lleonart, M., Handlinger, J. & Powell, M. (2003b) Treatment of spionid mud worm (Boccardia knoxi Rainer) infestation of cultured abalone. Aquaculture, 217.

    Loosanoff, V.L. & Engle, J.B. (1943) Polydora in oysters suspended in, the water. Biological Bulletin, 85, 69–78.

    Lunz, G.R., Jr. (1941) Polydora, a pest in South Carolina oysters. Journal of the Elisha Mitchell Scientific Society, 57, 273–283.

    Manchenko, G.P. (2001) Unusual isozyme patterns of glucose-6-phosphate isomerase in Polydora brevipalpa (Polychaeta: Spionidae). Biochemical Genetics, 39, 285–288.

    Manchenko, G.P. & Radashevsky, V.I. (1987) Allozymic variation in polychaetes of the genus Polydora from the Sea of Japan. Biologiya Morya, 5, 52–58.

    Manchenko, G.P. & Radashevsky, V.I. (1993) Genetic differences between two sibling species of the Polydora ciliata complex (Polychaeta: Spionidae). Biochemical Systematics and Ecology, 21, 543–548.

    Manchenko, G.P. & Radashevsky, V.I. (1994) Genetic differences between two allopatric sibling species of the genus Polydora (Polychaeta: Spionidae) from the west Pacific. Biochemical Systematics and Ecology, 22, 767–773.

    Manchenko, G.P. & Radashevsky, V.I. (2002) Genetic differences between two sibling sympatric Dipolydora species (Polychaeta : Spionidae) from the Sea of Japan, and a new species description. Journal of the Marine Biological Association of the United Kingdom, 82, 193–199.

    McDiarmid, H., Day, R. & Wilson, R. (2004) The ecology of polychaetes that infest abalone shells in Victoria, Australia. Journal of Shellfish Research, 23, 1179–1188.

    McIntosh, W.C. (1868) On the boring of certain annelids. Annals and Magazine of Natural History, 2, 276–295.

    Miller, D.C. & Jumars, P.A. (1986) Pellet accumulation, sediment supply, and crowding as determinants of surface deposit-feeding rate in Pseudopolydora kempi japonica Imajima & Hartman (Polychaeta: Spionidae). Journal of Experimental Marine Biology and Ecology, 99, 1–17,illustr.

    Moreno, R.A., Neill, P.E. & Rozbaczylo, N. (2006) Native and non-indigenous boring polychaetes in Chile: a threat to native and commercial mollusc species. Revista Chilena De Historia Natural, 79, 263–278.

    Mustaquim, J. (1986) Morphological variation in Polydora ciliata complex (Polychaeta: Annelida). Zoological Journal of the Linnean Society, 86, 75–88.

    Mustaquim, J. (1988) Electrophoretic variation of isozymes in Polydora ciliata complex (Polychaeta: Spionidae). Comparative Biochemistry and Physiology B-Biochemistry & Molecular Biology, 91, 197–205, illustr.

    National Introduced Marine Pest Information System (2002) NIMPIS. An information system for marine introductions in Australia. CSIRO Australia. Available from: http://www.marine.csiro.au/crimp/nimpis/Default.htm (12th January, 2008).

    Nell, J.A. (2001) The history of oyster farming in New South Wales. Marine Fisheries Review, 63, 14–25.

    Nell, J.A. (2007a) Diseases of Sydney rock oysters. Primefact 289, NSW Department of Primary Industries, Port Stephens Fisheries Centre, Nelson Bay, NSW, 4pp.

    Nell, J.A. (2007b) Controlling mudworm in oysters. Primefact 290. NSW Department of Primary Industries, Port Stephens Fisheries Centre, Nelson Bay, NSW, 4pp.

    Nell, J.A. & Smith, G.S. (1988) Management, production and disease interactions in oyster culture. Fish Diseases, Refresher Course for Veterinarians 23–27 May. Proceedings Volume 106. In, pp. 127–134.

    Ogburn, D.M., White, I. & McPhee, D.P. (2007) The disappearance of oyster reefs from eastern Australian estuaries - impact of colonial settlement or mudworm invasion? Coastal Management, 35, 271–287.

    Okuda, S. (1937) Spioniform polychaetes from Japan. Journal of the Faculty of Science of the Hokkaido Imperial University, series 6, 5, 217–254.

    Oliva, M.E. & Sanchez, M. (2005) Metazoan parasites and commensals of the northern Chilean scallop Argopecten purpuratus (Lamarck, 1819) as tools for stock identification. Fisheries Research, 71, 71–77.

    Örsted, A.S. (1843) Annulatorum danicorum conspectus. Maricolae Volume Fasc. 1 Librariae Wahlianae. Hafniae. 1–52.

    Petch, D.A. (1991) Larval development in Boccardia proboscidea (Polychaeta: Spionidae) from southeastern Australia. In: M. E. Petersen & J. B. Kirkegaard (Eds), Ophelia, Suppl. 5. Systematics, Biology and Morphology of World Polychaeta; 2nd International Polychaete Conference, Copenhagen, Denmark, August 1823, 1986. Ophelia Publications, Helsingor, Denmark. Illus. Maps, p. 698.

    Petch, D.A. (1995) Morphological variation in the spionid polychaete Boccardia proboscidea. Proceedings of the Royal Society of Victoria, 107, 25–30.

    Poore, G.C.B. & Kudenov, J.D. (1978a) Benthos around an outfall of the Werribee sewage-treatment farm, Port Phillip Bay, Victoria. Australian Journal of Marine and Freshwater Research, 29, 157–167.

    Poore, G.C.B. & Kudenov, J.D. (1978b) Benthos of the Port of Melbourne: the Yarra River and Hobsons Bay, Victoria. Australian Journal of Marine and Freshwater research, 29, 141–155.

    Poore, G.C.B., Rainer, S.F., Spies, R.B. & Ward, E. (1975) The zoobenthos programs in Port Phillip Bay, 1969–1973. Fisheries and Wildlife Technical Paper, 7, 1–78.

    Quatrefages, M.A.d. (1865) Histoire naturelle des annelés marins et d'eau douce. Annélides et Géphyriens (Vol. 1). Paris: Librairie Encyclopédique de Roret.

    Radashevsky, V.I. (1996) Morphology, ecology and asexual reproduction of a new Polydorella species (Polychaeta: Spionidae) from the South China Sea. Bulletin of Marine Science, 58, 684–693.

    Radashevsky, V.I. (1999) Description of the proposed lectotype for Polydora websteri Hartman in Loosanoff & Engle, 1943 (Polychaeta: Spionidae). Ophelia, 51, 107–113.

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

    Radashevsky, V.I. & Hsieh, H.-L. (2000) Pseudopolydora (Polychaeta: Spionidae) species from Taiwan. Zoological Studies, 39, 218–235.

    Radashevsky, V.I., Lana, P. & Rosebel, C.N. (2006) Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa, 1353, 1–37.

    Radashevsky, V.I. & Nogueira, J.M. (2003) Life history, morphology and distribution of Dipolydora armata (Polychaeta: Spionidae). Journal of the Marine Biological Association of the United Kingdom, 83, 375–384.

    Radashevsky, V.I. & Pankova, V.V. (2006) The morphology of two sibling sympatric Polydora species (Polychaeta: Spionidae) from the Sea of Japan. Journal of the Marine Biological Association of the United Kingdom, 86, 245–252.

    Radashevsky, V.I. & Petersen, M.E. (2005) On the morphology and distribution of Dipolydora giardi and status of D. trilobata (Polychaeta: Spionidae). Zootaxa, 25–36.

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

    Read, G.B. (2010) Comparison and history of Polydora websteri and P. haswelli (Polychaeta:Spionidae) as mud-blister worms in New Zealand shellfish. In: New Zealand Journal of Marine and Freshwater Research. First published on: 04 June 2010 (iFirst), pp. 1–18.

    Rice, S.A. (1978) Spermatophores and sperm transfer in spionid polychaetes. Transactions of the American Microscopical Society, 97, 160–170, illustr.

    Rice, S.A. (1980) Ultrastructure of the male nephridium and its role in spermatophore formation in spionid polychaetes (Annelida). Zoomorphologie, 95, 181–194.

    Rice, S.A. & Rice, K.A. (2009) Variable modes of larval development in the Polydora cornuta complex (Polychaeta: Spionidae) are directly related to stored sperm availability. Zoosymposia, 2, 397–414.

    Roughley, T.C. (1922) Oyster culture on the George's River, New South Wales. Technical Education Series, Technical Museum, Sydney, 25, 1–69.

    Roughley, T.C. (1925) The story of the oyster. Australian Natural History, Sydney, 2, 163–168.

    Rouse, G.W. & Pleijel, F. (2003) Problems in polychaete systematics. Hydrobiologia, 496, 175–189.

    Rousset, V., Rouse, G.W., Siddall, M.E., Tillier, A. & Pleijel, F. (2004) The phylogenetic position of Siboglinidae (Annelida) inferred from 18S rRNA, 28S rRNA and morphological data. Cladistics, 20, 518–533.

    Saenger, P. (1988) Gladstone Environmental Survey. Final report to Queensland Electricity Commission

    Sato-Okoshi, W. (1998) Three new species of polydorids (Polychaeta: Spionidae) from Japan. Species Diversity, 3, 277–288.

    Sato-Okoshi, W. & Nomura, T. (1990) Infestation of the Japanese scallop Patinopecten yessoensis by the boring polychaetes Polydora on the coast of Hokkaido and Tohoku District. Bulletin of the Japanese Society of Scientific Fisheries, 56, 1593–1598,illustr.

    Sato-Okoshi, W., Okoshi, K. & Shaw, J. (2008) Polydorid species (Polychaeta: Spionidae) in south-western Australian waters with special reference to Polydora uncinata and Boccardia knoxi. Journal of the Marine Biological Association of the United Kingdom, 88, 491–501.

    Sato-Okoshi, W. & Takatsuka, M. (2001) Polydora and related genera (Polychaeta, Spionidae) around Puerto Montt and Chiloe Island (Chile), with description of a new species of Dipolydora. Bulletin of Marine Science, 68, 485–503.

    Schmarda, L.K. (1861) Neue wirbellose thiere beobachtet und gesammelt auf einer reise un die erdr 1853 bis 1857. Vol. I. Turbellarian, Rotatorien und Anneliden. Leipzig: Wilhelm Engelmann.

    Schulze, S.R., Rice, S.A., Simon, J.L. & Karl, S.A. (2000) Evolution of poecilogony and the biogeography of North American populations of the polychaete Streblospio. Evolution, 54, 1247–1259.

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

    Silina, A.V. (2006) Tumor-like formations on the shells of Japanese scallops Patinopecten yessoensis (Jay). Marine Biology, 148, 833–840.

    Simon, C.A. (2009) Pseudopolydora species associated with mollusc shells on the south coast of South Africa, with the description of Ps. dayii, sp nov. Journal of the Marine Biological Association of the United Kingdom, 89, 681–687.

    Simon, C.A. & Booth, A.J. (2007) Population structure and growth of polydorid polychaetes that infest cultured abalone Haliotis midae. African Journal of Marine Science, 29, 499–509.

    Simon, C.A., Ludford, A. & Wynne, S. (2006) Spionid polychaetes infesting cultured abalone Haliotis midae in South Africa. African Journal of Marine Science, 28, 167–171.

    Simon, C.A., Thornhill, D.J., Oyarzun, F. & Halanych, K.M. (2009) Genetic similarity between Boccardia proboscidea from Western North America and cultured abalone, Haliotis midae, in South Africa. Aquaculture, 294, 18–24.

    Simon, C.A., Worsfold, T.M., Lange, L. & Sterley, J. (2010) The genus Boccardia (Polychaeta: Spionidae) associated with mollusc shells on the south coast of South Africa. Journal of the Marine Biological Association of the United Kingdom, 90, 585–598.

    Skeel, M. (1977) Further investigations on mudworms in oysters. Australian Fisheries, 36, 22–23.

    Skeel, M. (1978) Water clean up helps keep down pests. Australian Fisheries, 37, 12–13.

    Skeel, M.E. (1979) Shell boring worms (Spionidae: Polychaeta) infecting cultivated bivalve molluscs in Australia. In: Tenth Annual Meeting World Mariculture Society, pp. 529–533.

    Söderström, A. (1920) Studien über die Polychätenfamilie Spionidae. Uppsala University: Almquist and Wicksells.

    Southern, R. (1921) Polychaeta of the Chilka Lake and also of fresh and brackish waters in other parts of India. Memoirs of the Indian Museum Calcutta, 5, 563–659.

    Stephenson, W., Cook, S.D. & Newlands, S.J. (1978) The macrobenthos of the Middle Banks area of Moreton Bay. Memoirs of the Queensland Museum, 18, 185–212.

    Stephenson, W., Raphael, Y.I. & Cook, S.D. (1976) The macrobenthos of Bramble Bay, Moreton Bay, Queensland. Memoirs of the Queensland Museum, 17, 425–447.

    Stephenson, W., Williams, W.T. & Cook, S.D. (1974) The benthic fauna of soft bottoms, southern Moreton Bay. Memoirs of the Queensland Museum, 17, 73–123.

    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, 57.

    Templeton, R. (1836) A catalogue of the species of annulose animals and of rayed ones found in Ireland. Magazine of Natural History and Journal of Zoology, Botany, Mineralogy, Geology and Meteorology. London, 9, 233–243.

    Tkachuk, L.P. (1986) Shell borers of the Black Sea edible mussel. Rybnoe Khozyaistvo, 1986, 34–35.

    Turner, H.J. & Hawks, J.E. (1959) Infestation of Pecten irradians by Polydora. Nautilus, 72, 109–111.

    Tzetlin, A.B. & Britayev, T.A. (1985) A new species of the Spionidae (Polychaeta) with asexual reproduction associated with sponges. Zoologica Scripta, 14, 177–181.

    Verrill, A.E. (1879) Notice of recent additions to the marine invertebrata of the northeastern coast of America, with descriptions of new genera and species and critical remarks on others. Part I. Annelida, Gephyraea, Nemertina, Nematoda, Polyzoa, Tunicata, Mollusca, Anthozoa, Echinodermata, Porifera. Proceedings of the United States National Museum, 2, 165–205.

    Verrill, A.E. (1880) Notice of recent additions to the marine invertebrata of the northeastern coast of America, with descriptions of new genera and species and critical remarks on others. Part 2. Mollusca, with notes on Annelida, Echinodermata, etc., collected by the United States Fish Commission. Proceedings of the United States National Museum, 3, 356–405.

    Verrill, A.E. (1882) Notice on recent additions to the marine Invertebrata of the northeastern coast of America, with descriptions of new genera and species and critical remarks on others. Smithsonian Miscellaneous Collections, 22, 356–405.

    Walker, L.M. (2009) Polydora and Dipolydora (Polychaeta: Spionidae) of estuaries and bays of subtropical eastern Australia: A review and morphometric investigation of their taxonomy and distribution. Unpublished MSc thesis, School of Environmental Science and Management, Southern Cross University, Lismore, pp. 180.

    Webster, H.E. (1880) The Annelida Chaetopoda of New Jersey. Annual Report of the New York State Museum of Natural History, 32, 101–128.

    Webster, H. E. (1879) Fam. Spionidae. In: On the Annelida Chaetopoda of the Virginian coast. In Advance of Volume 1X of the Transactions Albany Institute New York, 49–53, pls viii–ix.

    Whitelegge, T. (1890) Report on the worm disease affecting the oysters on the coast of New South Wales. Records of the Australian Museum, 1, 41–54, Plates III – VI.

    Williams, J.D. (2004) Reproduction and morphology of Polydorella (Polychaeta: Spionidae), including the description of a new species from the Philippines. Journal of Natural History, 38, 1339–1358.

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

    Wilson, R.S., Hutchings, P.A. & Glasby, C.J. (2003) Polychaetes: An Interactive Identification Guide. CSIRO Publishing, Melbourne.

    Worsaae, K. (2001) The systematic significance of palp morphology in the Polydora-complex (Polychaeta : Spionidae). Zoologischer Anzeiger, 240, 47–59.