information on the biology of species and the ecology of habitats found around the coasts and seas of the British Isles

Flustra foliacea and colonial ascidians on tide-swept exposed circalittoral mixed substrata



UK and Ireland classification

UK and Ireland classification


This variant is typically found on very exposed to moderately exposed, circalittoral mixed substrata subject to moderately strong tidal streams. It most frequently occurs between 10m and 20m water depth. This variant is characterized by a dense hydroid and Flustra foliacea turf, along with other scour-tolerant species, growing on the more stable boulders and cobbles which overlie coarse muddy sand and gravel. Although Nemertesia antennina is the dominant species within the hydroid turf, other species such as Halecium halecinumNemertesia ramosa andHydrallmania falcata may also be present. Other bryozoans found amongst the hydroid and Flustra turf include Cellepora pumicosaBugulina flabellataBugulina turbinata, and a crisiid turf. Encrusting red algae, the polychaete Spirobranchus triqueter and barnacles such as Balanus crenatus may be found on the smaller cobbles and pebbles, which may become mobile during extreme storms. Echinoderms such as Asterias rubens and Ophiothrix fragilis may be present on the boulders, or the coarse sediment in between. On the larger, more stable boulders, isolated sponge communities may develop, with species such as Scypha ciliataDysidea fragilisHemimycale columellaEsperiopsis fucorum andStelligera rigida. In addition, small Alcyonium digitatum, various ascidians (Clavelina lepadiformisBotryllus schlosseri),Pododesmus patelliformis and top shells (Calliostoma zizyphinumGibbula cineraria) may colonise the upper faces and vertical sides of larger boulders. At some shallower sites, the foliose red algae Hypoglossum hypoglossoides may be found on the tops of larger boulders. Within the coarse sediment underlying these boulders and cobbles, anemones such as Cerianthus lloydii and Urticina felina may be recorded. Under-boulder fauna typically consists of terebellid worms, and crabs such as Pisidia longicornis and Cancer pagurus.

Depth range


Additional information


Listed By

Further information sources

Search on:

Sensitivity reviewHow is sensitivity assessed?



Species indicative of sensitivity


Physical Pressures

No sensitivity data available

Chemical Pressures

No sensitivity data available

Biological Pressures

No sensitivity data available

Additional information



  1. Ackers, R.G.A., Moss, D. & Picton, B.E. 1992. Sponges of the British Isles (Sponges: V): a colour guide and working document. Ross-on-Wye: Marine Conservation Society.

  2. Allen, J., Slinn, D., Shummon, T., Hurtnoll, R. & Hawkins, S., 1998. Evidence for eutrophication of the Irish Sea over four decades. Limnology and Oceanography, 43 (8), 1970-1974.

  3. Ambrogi, A.O., 2000. Biotic invasions in a Mediterranean lagoon. Biological Invasions, 2 (2), 165-176.

  4. Aoki, S., Cao, L., Matsui, K., Rachmat, R., Akiyama, S.-i. & Kobayashi, M., 2004. Kendarimide A, a novel peptide reversing P-glycoprotein-mediated multidrug resistance in tumor cells, from a marine sponge of Haliclona sp. Tetrahedron, 60 (33), 7053-7059.

  5. Bakus, G.J., 1968. Sedimentation and benthic invertebrates of Fanning Island, Central Pacific. Marine Geology, 6, 45-51.

  6. Banks, P.D. & Brown, K.M., 2002. Hydrocarbon effects on fouling assemblages: the importance of taxonomic differences, seasonal, and tidal variation. Marine Environmental Research, 53 (3), 311-326.

  7. Bell, J.J. & Barnes, D.K., 2000. The distribution and prevalence of sponges in relation to environmental gradients within a temperate sea lough: inclined cliff surfaces. Diversity and Distributions, 6 (6), 305-323.

  8. Bell, J.J. & Barnes, D.K., 2001. Sponge morphological diversity: a qualitative predictor of species diversity? Aquatic Conservation: Marine and Freshwater Ecosystems, 11 (2), 109-121.

  9. Bell, J.J. & Smith, D., 2004. Ecology of sponge assemblages (Porifera) in the Wakatobi region, south-east Sulawesi, Indonesia: richness and abundance. Journal of the Marine Biological Association of the UK, 84 (3), 581-591.

  10. Bell, J.J., Barnes, D. & Shaw, C., 2002. Branching dynamics of two species of arborescent demosponge: the effect of flow regime and bathymetry. Journal of the Marine Biological Association of the UK, 82 (2), 279-294.

  11. Berrill, N.J., 1975. Chordata: Tunicata. In Reproduction of marine Invertebrates, vol. II, (ed. A.C. Geise & J.S. Pearse), pp. 241-282. New York: Academic Press.

  12. Berrill, N.J., 1950. The Tunicata with an account of the British species. London: Ray Society.

  13. Boulcott, P. & Howell, T.R.W., 2011. The impact of scallop dredging on rocky-reef substrata. Fisheries Research (Amsterdam), 110 (3), 415-420.

  14. Bryan, G.W. & Gibbs, P.E., 1991. Impact of low concentrations of tributyltin (TBT) on marine organisms: a review. In: Metal ecotoxicology: concepts and applications (ed. M.C. Newman & A.W. McIntosh), pp. 323-361. Boston: Lewis Publishers Inc.

  15. Budd, G.C. 2008. Alcyonium digitatum Dead man's fingers. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. Available from:

  16. Bullimore, B., 1985. An investigation into the effects of scallop dredging within the Skomer Marine Reserve. Report to the Nature Conservancy Council by the Skomer Marine Reserve Subtidal Monitoring Project, S.M.R.S.M.P. Report, no 3., Nature Conservancy Council.

  17. Castège, I., Milon, E. & Pautrizel, F., 2014. Response of benthic macrofauna to an oil pollution: Lessons from the “Prestige” oil spill on the rocky shore of Guéthary (south of the Bay of Biscay, France). Deep Sea Research Part II: Topical Studies in Oceanography, 106, 192-197.

  18. Cole, S., Codling, I.D., Parr, W., Zabel, T., 1999. Guidelines for managing water quality impacts within UK European marine sites [On-line]. UK Marine SACs Project. [Cited 26/01/16]. Available from:

  19. Connor, D.W., Allen, J.H., Golding, N., Howell, K.L., Lieberknecht, L.M., Northen, K.O. & Reker, J.B., 2004. The Marine Habitat Classification for Britain and Ireland. Version 04.05. Joint Nature Conservation Committee, Peterborough.

  20. Cook, R., Fariñas-Franco, J. M., Gell, F. R., Holt, R. H., Holt, T., Lindenbaum, C.,  Porter, J.S., Seed, R., Skates, L.R., Stringell, T.B. & Sanderson, W.G., 2013. The substantial first impact of bottom fishing on rare biodiversity hotspots: a dilemma for evidence-based conservation. PloS One, 8 (8), e69904.

  21. Crisp, D.J., 1964b. Mortalities in marine life in North Wales during the winter of 1962-63. Journal of Animal Ecology, 33, 190-197.

  22. De Caralt, S., López-Legentil, S., Tarjuelo, I., Uriz, M.J. & Turon, X., 2002. Contrasting biological traits of Clavelina lepadiformis (Ascidiacea) populations from inside and outside harbours in the western Mediterranean. Marine Ecology Progress Series, 244, 125-137.

  23. De Goeij, J.M., Moodley, L., Houtekamer, M., Carballeira, N.M. & Van Duyl, F.C., 2008. Tracing 13C‐enriched dissolved and particulate organic carbon in the bacteria‐containing coral reef sponge Halisarca caerulea: Evidence for DOM‐feeding. Limnology and Oceanography, 53 (4), 1376-1386.

  24. De Kluijver, M. & Leewis, R., 1994. Changes in the sublittoral hard substrate communities in the Oosterschelde estuary (SW Netherlands), caused by changes in the environmental parameters. Hydrobiologia, 282 (1), 265-280.

  25. De Vos, L., Rútzler K., Boury-Esnault, N., Donadey C., Vacelet, J., 1991. Atlas of Sponge Morphology. Atlas de Morphologie des Éponges. Washington, Smithsonian Institution Press.

  26. Diaz, R.J. & Rosenberg, R., 1995. Marine benthic hypoxia: a review of its ecological effects and the behavioural responses of benthic macrofauna. Oceanography and Marine Biology: an Annual Review, 33, 245-303.

  27. Dyrynda, P., Fairall, V., Occhipinti Ambrogi, A. & d'Hondt, J.-L., 2000. The distribution, origins and taxonomy of Tricellaria inopinata d'Hondt and Occhipinti Ambrogi, 1985, an invasive bryozoan new to the Atlantic. Journal of Natural History, 34 (10), 1993-2006.

  28. Dyrynda, P.E.J., 1994. Hydrodynamic gradients and bryozoan distributions within an estuarine basin (Poole Harbour, UK). In Proceedings of the 9th International Bryozoology conference, Swansea, 1992. Biology and Palaeobiology of Bryozoans (ed. P.J. Hayward, J.S. Ryland & P.D. Taylor), pp.57-63. Fredensborg: Olsen & Olsen.

  29. Eggleston, D., 1972b. Factors influencing the distribution of sub-littoral ectoprocts off the south of the Isle of Man (Irish Sea). Journal of Natural History, 6, 247-260.

  30. Fariñas-Franco, J.M., Pearce, B., Porter, J., Harries, D., Mair, J.M. & Sanderson, W.G, 2014. Development and validation of indicators of Good Environmental Status for biogenic reefs formed by Modiolus modiolus, Mytilus edulis and Sabellaria spinulosa under the Marine Strategy Framework Directive. Joint Nature Conservation Committee,

  31. Fell, P.E., 1974. Diapause in the gemmules of the marine sponge, Haliclona loosanoffi, with a note on the gemmules of Haliclona oculata. The Biological Bulletin, 147 (2), 333-351.

  32. Fiana-Medioni, A., 1978. A scanning electron microscope study of the branchial sac of benthic filter-feeding invertebrates (ascidians). Acta Zoologica, 59, 1, 1-9.

  33. Fish, J.D. & Fish, S., 1996. A student's guide to the seashore. Cambridge: Cambridge University Press.

  34. Fortunato, H., Schäfer, P. & Blaschek, H., 2013. Growth Rates, Age Determination, and Calcification Levels in Flustra foliacea (L.) (Bryozoa: Cheilostomata): Preliminary Assessment. In Ernst, A., et al. (eds.). Bryozoan Studies 2010, Berlin, Heidelberg: Springer Berlin Heidelberg, pp. 59-74.

  35. Freese, J.L., 2001. Trawl-induced damage to sponges observed from a research submersible. Marine Fisheries Review, 63 (3), 7-13.

  36. Freese, L., Auster, P.J., Heifetz, J. & Wing, B.L., 1999. Effects of trawling on seafloor habitat and associated invertebrate taxa in the Gulf of Alaska. Marine Ecology Progress Series, 182, 119-126.

  37. Gaino, E., Pronzato, R., Corriero, G. & Buffa, P., 1992. Mortality of commercial sponges: incidence in two Mediterranean areas. Italian Journal of Zoology, 59 (1), 79-85.

  38. Galstoff, P., 1942. Wasting disease causing mortality of sponges in the West Indies and Gulf of Mexico.  Proceedings 8th American Scientific Congress, pp. 411-421.

  39. Gerrodette, T. & Flechsig, A., 1979. Sediment-induced reduction in the pumping rate of the tropical sponge Verongia lacunosa. Marine Biology, 55 (2), 103-110.

  40. Gochfeld, D., Easson, C., Freeman, C., Thacker, R. & Olson, J., 2012. Disease and nutrient enrichment as potential stressors on the Caribbean sponge Aplysina cauliformis and its bacterial symbionts. Marine Ecology Progress Series, 456, 101-111.

  41. Gochfeld, D.J., Schlöder, C. & Thacker, R.W., 2007. Sponge community structure and disease prevalence on coral reefs in Bocas del Toro, Panama. Porifera Research: Biodiversity, Innovation, and Sustainability, Série Livros, 28, 335-343.

  42. Griffith, K., Mowat, S., Holt, R.H., Ramsay, K., Bishop, J.D., Lambert, G. & Jenkins, S.R., 2009. First records in Great Britain of the invasive colonial ascidian Didemnum vexillum Kott, 2002. Aquatic Invasions, 4 (4), 581-590.

  43. Gunda, V.G. & Janapala, V.R., 2009. Effects of dissolved oxygen levels on survival and growth in vitro of Haliclona pigmentifera (Demospongiae). Cell and tissue research, 337 (3), 527-535.

  44. Hartikainen, H., Johnes, P., Moncrieff, C. & Okamura, B., 2009. Bryozoan populations reflect nutrient enrichment and productivity gradients in rivers. Freshwater Biology, 54 (11), 2320-2334.

  45. Hayward, P.J. & McKinney, F.K., 2002. Northern Adriatic Bryozoa from the vicinity of Rovinj, Croatia. Bulletin of the American Museum of Natural History, 1-139.

  46. Hayward, P.J. & Ryland, J.S. 1998. Cheilostomatous Bryozoa. Part 1. Aeteoidea - Cribrilinoidea. Shrewsbury: Field Studies Council. [Synopses of the British Fauna, no. 10. (2nd edition)]

  47. Hayward, P.J. & Ryland, J.S. (ed.) 1995a. The marine fauna of the British Isles and north-west Europe. Volume 2. Molluscs to Chordates. Oxford Science Publications. Oxford: Clarendon Press.

  48. Herreid, C.F., 1980. Hypoxia in invertebrates. Comparative Biochemistry and Physiology Part A: Physiology, 67 (3), 311-320.

  49. Hincks, T., 1880. A history of British marine Polyzoa, vol. I & II. London: John van Voorst.

  50. Hiscock, H., 1985b. Aspects of the ecology of rocky sublittoral areas. In The Ecology of Rocky Coasts: essays presented to J.R. Lewis, D.Sc. (ed. P.G. Moore & R. Seed), pp. 290-328. London: Hodder & Stoughton Ltd.

  51. Hiscock, K. & Hoare, R., 1975. The ecology of sublittoral communities at Abereiddy Quarry, Pembrokeshire. Journal of the Marine Biological Association of the United Kingdom, 55 (4), 833-864.

  52. Hiscock, K., 1981. Marine life on the wreck of the M.V. "Robert". Report of the Lundy Field Society, 32, 40-44.

  53. Hiscock, K., 1983. Water movement. In Sublittoral ecology. The ecology of shallow sublittoral benthos (ed. R. Earll & D.G. Erwin), pp. 58-96. Oxford: Clarendon Press.

  54. Hoare, R. & Hiscock, K., 1974. An ecological survey of the rocky coast adjacent to the effluent of a bromine extraction plant. Estuarine and Coastal Marine Science, 2 (4), 329-348.

  55. Holme, N.A. & Wilson, J.B., 1985. Faunas associated with longitudinal furrows and sand ribbons in a tide-swept area in the English Channel. Journal of the Marine Biological Association of the United Kingdom, 65, 1051-1072.

  56. Holt, T.J., Jones, D.R., Hawkins, S.J. & Hartnoll, R.G., 1995. The sensitivity of marine communities to man induced change - a scoping report. Countryside Council for Wales, Bangor, Contract Science Report, no. 65.

  57. Jennings, S. & Kaiser, M.J., 1998. The effects of fishing on marine ecosystems. Advances in Marine Biology, 34, 201-352.

  58. Jones, J., Bunker, F., Newman, P., Burton, M., Lock, K., 2012. Sponge Diversity of Skomer Marine Nature Reserve. CCW Regional Report,  CCW/WW/12/3.

  59. Kayser, H., 1990. Bioaccumulation and transfer of cadmium in marine diatoms, Bryozoa, and Kamptozoa. In Oceanic processes in marine pollution, vol. 6. Physical and chemical processes: transport and transformation (ed. D.J. Baumgartner & I.W. Duedall), pp. 99-106. Florida: R.E. Krieger Publishing Co.

  60. Knight-Jones, E.W. & Nelson-Smith, A., 1977. Sublittoral transects in the Menai Straits and Milford Haven. In Biology of benthic organisms (ed. B.F. Keegan, P. O Ceidigh & P.J.S. Broaden), pp. 379-390. Oxford: Pergamon Press.

  61. Koopmans, M. & Wijffels, R.H., 2008. Seasonal Growth Rate of the Sponge Haliclona oculata (Demospongiae: Haplosclerida). Marine Biotechnology, 10 (5), 502-510.

  62. Lengyel, N.L., Collie, J.S. & Valentine, P.C., 2009. The invasive colonial ascidian Didemnum vexillum on Georges Bank - Ecological effects and genetic identification. Aquatic Invasions, 4(1), 143-152.

  63. Lysek, N., Rachor, E. & Lindel, T., 2002. Isolation and structure elucidation of Deformylflustrabromine from the North Sea bryozoan Flustra foliacea. Zeitschrift für Naturforschung, C: Biosciences, 57, 1056-1061.

  64. Marin, A., Lopez, M., Esteban, M., Meseguer, J., Munoz, J. & Fontana, A., 1998. Anatomical and ultrastructural studies of chemical defence in the sponge Dysidea fragilis. Marine Biology, 131 (4), 639-645.

  65. McKinney, F.K. & Jaklin, A., 2000. Spatial niche partitioning in the Cellaria meadow epibiont association, northern Adriatic Sea. Cahiers de biologie marine, 41 (1), 1-17.

  66. McKinney, F.K., 1986. Evolution of erect marine bryozoan faunas: repeated success of unilaminate species The American Naturalist, 128, 795-809.

  67. Menon, N.R., 1975. Observations on growth of Flustra foliacea (Bryozoa) from Helgoland waters. Helgolander Wissenschaftliche Meeresuntersuchungen, 27, 263-267.

  68. Millar, R., 1971. The biology of ascidians. Advances in marine biology, 9, 1-100.

  69. Millar, R.H., 1970. British Ascidians London: Academic Press.[Synopses of the British Fauna, no. 1.]

  70. Mita, K., Kawai, N., Rueckert, S. & Sasakura, Y., 2012. Large-scale infection of the ascidian Ciona intestinalis by the gregarine Lankesteria ascidiae in an inland culture system. Diseases of aquatic organisms, 101 (3), 185-195.

  71. Mustapha K., Zarrouk, S., Souissi, A. & El Abed, A., 2003. Diversité des Démosponges Tunisiennes. Bulletin Institut national des Sciences et Technologies de la mer de Salammbô, 30, 55-78.

  72. Naranjo, S.A., Carballo, J.L., & Garcia-Gomez, J.C., 1996. Effects of environmental stress on ascidian populations in Algeciras Bay (southern Spain). Possible marine bioindicators? Marine Ecology Progress Series, 144 (1), 119-131.

  73. NBN, 2015. National Biodiversity Network 2015(20/05/2015).

  74. O'Dea, A. & Okamura, B., 2000. Life history and environmental inference through retrospective morphometric analysis of bryozoans: a preliminary study. Journal of the Marine Biological Association of the United Kingdom, 80, 1127-1128.

  75. Okamura, B., 1984. The effects of ambient flow velocity, colony size and upstream colonies on the feeding success of Bryozoa, Bugula stolonifera Ryland, an arborescent species. Journal of the Experimental Marine Biology and Ecology, 83, 179-193.

  76. Ooishi, S., 2010. Enterocola hessei Chatton & Harant (Copepoda: Cyclopoida: Ascidicolidae) living in the compound ascidian Clavelina lepadiformis (Müller). Proceedings of the Biological Society of Washington, 123 (2), 137-148.

  77. Pajic, I., Kljajic, Z., Dogovic, N., Sladic, D., Juranic, Z. & Gasic, M.J., 2002. A novel lectin from the sponge Haliclona cratera: isolation, characterization and biological activity. Comparative Biochemistry and Physiology Part C: Toxicology & Pharmacology, 132 (2), 213-221.

  78. Peters, L., König, G.M., Wright, A.D., Pukall, R., Stackebrandt, E., Eberl, L. & Riedel, K., 2003. Secondary metabolites of Flustra foliacea and their influence on bacteria. Applied and Environmental Microbiology, 69, 3469-3475.

  79. Picton, B.E. & Morrow, C.C. (2004b). Polyclinum aurantium.Encyclopedia of Marine Life of Britain and Ireland. Accessed on 2016-06-15

  80. Picton, B.E. & Morrow, C.C. (2004c). Clavelina lepadiformis (O F Müller, 1776). Encyclopedia of Marine Life of Britain and Ireland. Accessed on 2016-06-15

  81. Picton, B.E. & Morrow, C.C., 2004. Nemertesia ramosa Lamouroux, 1816., 2004-09-14

  82. Porter, J., 2012. Seasearch Guide to Bryozoans and Hydroids of Britain and Ireland.   Ross-on-Wye: Marine Conservation Society.

  83. Preston J. & Burton, M., 2015. Marine microbial assemblages associated with diseased Porifera in Skomer Marine Nature Reserve (SMNR), Wales. Aquatic Biodiversity and Ecosystems, 30th August – 4th September,  Liverpool.,  pp. p110.

  84. Pukall, R., Kramer, I., Rohde, M. & Stackebrandt, E., 2001. Microbial diversity of cultivatable bacteria associated with the North Sea bryozoan Flustra foliacea. Systematic and applied microbiology, 24 (4), 623-633.

  85. Rees, H.L., Waldock, R., Matthiessen, P. & Pendle, M.A., 2001. Improvements in the epifauna of the Crouch estuary (United Kingdom) following a decline in TBT concentrations. Marine Pollution Bulletin, 42, 137-144.

  86. Reinhardt, J.F., Stefaniak, L.M., Hudson, D.M., Mangiafico, J., Gladych, R. & Whitlatch, R.B., 2010. First record of the non-native light bulb tunicate Clavelina lepadiformis (Müller, 1776) in the northwest Atlantic. Aquatic Invasions, 5 (2), 185-190.

  87. Riisgård, H.U., Bondo Christensen, P., Olesen, N.J., Petersen, J.K, Moller, M.M. & Anderson, P., 1993. Biological structure in a shallow cove (Kertinge Nor, Denmark) - control by benthic nutrient fluxes and suspension-feeding ascidians and jellyfish. Ophelia, 41, 329-344.

  88. Riley, K. 2008. Clavelina lepadiformis Light bulb sea squirt. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. Available from:

  89. Roberts, D., Cummins, S., Davis, A. & Chapman, M., 2006. Structure and dynamics of sponge-dominated assemblages on exposed and sheltered temperate reefs. Marine Ecology Progress Series, 321, 19-30.

  90. Rose, C.S. & Risk, M.J., 1985. Increase in Cliona delitrix infestation of Montastrea cavernosa heads on an organically polluted portion of the Grand Cayman fringing reef. Marine Ecology, 6 (4), 345-363.

  91. Rosenberg, R., Hellman, B. & Johansson, B., 1991. Hypoxic tolerance of marine benthic fauna. Marine Ecology Progress Series, 79, 127-131.

  92. Ryland, J.S. & De Putron, S., 1998. An appraisal of the effects of the Sea Empress oil spillage on sensitive invertebrate communities. Countryside Council for Wales Sea Empress Contract Report, no. 285, 97pp.

  93. Ryland, J.S., 1970. Bryozoans. London: Hutchinson University Library.

  94. Ryland, J.S., 1976. Physiology and ecology of marine bryozoans. Advances in Marine Biology, 14, 285-443.

  95. Ryland, J.S., Holt, R., Loxton, J., Spencer Jones, M. & Porter, J.S., 2014. First occurrence of the non-native bryozoan Schizoporella japonica Ortmann (1890) in Western Europe. Zootaxa, 3780 (3), 481-502.

  96. Sagasti, A., Schaffner, L.C. & Duffy, J.E., 2000. Epifaunal communities thrive in an estuary with hypoxic episodes. Estuaries, 23, 474-487.

  97. Santalova, E.A., Makarieva, T.N., Gorshkova, I.A., Dmitrenok, A.S., Krasokhin, V.B. & Stonik, V.A., 2004. Sterols from six marine sponges. Biochemical Systematics and Ecology, 32 (2), 153-167.

  98. Schönberg, C.H.L., 2015. Happy relationships between marine sponges and sediments–a review and some observations from Australia. Journal of the Marine Biological Association of the United Kingdom, 1-22.

  99. Silén, L., 1981. Colony structure in Flustra foliacea (Linnaeus) (Bryozoa, Cheilostomata). Acta Zoologica (Stockholm.), 62, 219-232.

  100. Sipkema, D., Van Wielink, R., Van Lammeren, A., Tramper, J., Osinga, R. & Wijffels, R., 2003. Primmorphs from seven marine sponges: formation and structure. Journal of Biotechnology, 100 (2), 127-139.

  101. Soule, D.F. & Soule, J.D., 1979. Bryozoa (Ectoprocta). In Pollution ecology of estuarine invertebrates (ed. C.W. Hart & S.L.H. Fuller), pp. 35-76.

  102. Stanley, J.A., Wilkens, S.L. & Jeffs, A.G., 2014. Fouling in your own nest: vessel noise increases biofouling. Biofouling, 30 (7), 837-844.

  103. Stebbing, A.R.D., 1971a. Growth of Flustra foliacea (Bryozoa). Marine Biology, 9, 267-273.

  104. Stebbing, A.R.D., 1971b. The epizoic fauna of Flustra foliacea [Bryozoa]. Journal of the Marine Biological Association of the United Kingdom, 51, 283-300.

  105. Stefaniak, L.M., Auster, P.J. & Babb, I.G., 2014. Loss of an erect sponge on a rock reef in Long Island Sound (north-west Atlantic). Marine Biodiversity Records, 7, e115.

  106. Storr, J.F. 1976. Ecological factors controlling sponge distribution in the Gulf of Mexico and the resulting zonation. In Aspects of Sponge Biology (ed. F.W. Harrison & R.R. Cowden), pp. 261-276. New York: Academic Press.

  107. Tillin, H. & Tyler-Walters, H., 2014. Assessing the sensitivity of subtidal sedimentary habitats to pressures associated with marine activities. Phase 2 Report – Literature review and sensitivity assessments for ecological groups for circalittoral and offshore Level 5 biotopes. JNCC Report No. 512B,  260 pp. Available from:

  108. Tjensvoll, I., Kutti, T., Fosså, J.H. & Bannister, R., 2013. Rapid respiratory responses of the deep-water sponge Geodia barretti exposed to suspended sediments. Aquatic Biology, 19, 65-73.

  109. Tyler-Walters, H., 2005c. Bugula turbinata an erect bryozoan. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 30.03.16] Available from:

  110. Tyler-Walters, H. & Ballerstedt, S., 2007. Flustra foliacea Hornwrack. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. Available from:

  111. Vacelet, J., 1994. Control of the severe sponge epidemic—Near East and Europe: Algeria, Cyprus, Egypt, Lebanon, Malta, Morocco, Syria, Tunisia, Turkey. Yugoslavia. Technical Report–the struggle against the epidemic which is decimating Mediterranean sponges FI: TCP/RAB/8853. Rome, Italy. 1–39 p,  pp.

  112. Van Dolah, R.F., Wendt, P.H. & Nicholson, N., 1987. Effects of a research trawl on a hard-bottom assemblage of sponges and corals. Fisheries Research, 5 (1), 39-54.

  113. Wapstra, M. & van Soest, R.W.M., 1987. Sexual reproduction, larval morphology and behaviour in demosponges from the southwest of the Netherlands. Berlin: Springer-Verlag

  114. Ward-Paige, C.A., Risk, M.J., Sherwood, O.A. & Jaap, W.C., 2005. Clionid sponge surveys on the Florida Reef Tract suggest land-based nutrient inputs. Marine Pollution Bulletin, 51 (5), 570-579.

  115. Webster, N.S., 2007. Sponge disease: a global threat? Environmental Microbiology, 9 (6), 1363-1375.

  116. Wulff, J., 1995. Effects of a hurricane on survival and orientation of large erect coral reef sponges. Coral Reefs, 14 (1), 55-61.

  117. Wulff, J., 2006. Resistance vs recovery: morphological strategies of coral reef sponges. Functional Ecology, 20 (4), 699-708.

  118. Zahn, R., Zahn, G., Müller, W., Kurelec, B., Rijavec, M., Batel, R. & Given, R., 1981. Assessing consequences of marine pollution by hydrocarbons using sponges as model organisms. Science of The Total Environment, 20 (2), 147-169.


This review can be cited as:

Readman, J.A.J., 2016. [Flustra foliacea] and colonial ascidians on tide-swept exposed circalittoral mixed substrata. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 18-06-2018]. Available from:

Last Updated: 07/07/2016