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

Novocrania anomala and Protanthea simplex on sheltered circalittoral rock



UK and Ireland classification


This biotope typically occurs in full to variable salinity conditions on very wave-sheltered circalittoral bedrock and boulder slopes subject to negligible tidal streams (this tends to be in the landward, very sheltered basins of fjordic sealochs). This biotope is characterized by often dense populations of the anemone Protanthea simplex, growing on the silty bedrock. The underlying rock surfaces are usually covered by encrusting red algae, the polychaete Spirobranchus triqueter, the brachiopods Novocrania anomala and Terebratulina retusa, the saddle oyster Pododesmus patelliformis and the polychaete Sabella pavonina. Scattered colonies of Alcyonium digitatum and the hydroid Bougainvillia ramosa may occasionally be recorded. A diverse range of ascidians including Ciona intestinalis, Ascidia mentula, Corella parallelogramma, Ascidia virginea, Polycarpa pomaria and Dendrodoa grossularia are also occasionally recorded. Echinoderms such as the common brittlestar Ophiothrix fragilis are frequently reported with their arms protruding from crevices in the rock, whilst the starfish Asterias rubens, Henricia oculata, and the sea urchin Echinus esculentus and Psammechinus miliaris are occasionally found on the boulder/rock surface. The whelk Buccinum undatum is often present but in very low numbers. The squat lobster Munida rugosa may be seen hiding in crevices. The hermit crab Pagurus bernhardus may also be recorded. (Information from JNCC, 2014).

Depth range

5-10 m, 10-20 m, 20-30 m, 30-50 m

Additional information

CR.LCR.BrAS.NeoPro includes two sub-biotopes (NeoPro.FS and NeoPro.VS) depending on the salintiy regime. Both are found in simialr physical habitats but Dendrodoa grossularia becomes common is the variable salinity sub-biotope (NeoPro.VS) and Protanthea simplex exhibits a higher abundance in the full salinty sub-biotope (NeoPro.FS) (JNCC, 2014).

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Habitat review


Ecological and functional relationships


Seasonal and longer term change


Habitat structure and complexity




Recruitment processes


Time for community to reach maturity


Additional information


Preferences & Distribution

Habitat preferences

Depth Range 5-10 m, 10-20 m, 20-30 m, 30-50 m
Water clarity preferencesPoor clarity / Extreme turbidity, Very high clarity / Very low turbidity, See additional information
Limiting Nutrients Data deficient
Salinity preferences Full (30-40 psu)
Physiographic preferences Enclosed coast / Embayment
Biological zone preferences Circalittoral
Substratum/habitat preferences Bedrock, Large to very large boulders, Small boulders
Tidal strength preferences Very Weak (negligible), Weak < 1 knot (<0.5 m/sec.)
Wave exposure preferences Extremely sheltered, Sheltered, Very sheltered
Other preferences

Additional Information

The two sub-biotopes included within this assessment are characterized by variable, reduced or low salinity which may influence biotope structure. The two sub-biotopes have some similarities although SCR.NeoPro.Den is more species rich and may occur in more open lochs (so far it has only been recorded from Loch Etive). The temperature preferences of the individual species selected to represent the biotope are quite different to the temperatures in which the biotope occurs in Britain and Ireland. For instance, Ciona intestinalis has a world-wide distribution and optimal growth occurs at between 15-20 degrees C, considerably higher than water temperatures on the west coast of Scotland. Protanthea simplex extends further north into colder waters. No information is available regarding limiting nutrients.

Species composition

Species found especially in this biotope

    Rare or scarce species associated with this biotope


    Additional information

    The biotope assessment also covers two sub-biotopes. Ciona intestinalis is not one of the characterizing species in SCR.NeoPro.CaTw. However, Ciona intestinalis is assumed to form a suitable surrogate, representing the sensitivity of the various other solitary ascidians in the sub-biotope.

    Sensitivity review


    There are no species in this biotope that can be classified as 'key', i.e. species that if lost would result in loss or degradation of the associated community. Three important characterizing species have been selected that are important for the classification of this biotope. The biotope name includes Neocrania anomala and Protanthea simplex so these have been included. The parent biotope complex is 'Brachiopod and solitary ascidian communities (sheltered rock)' Ciona intestinalis has been included as a representative solitary ascidian that is found with high frequency in the biotope. The very large solitary ascidian Ascidia mentula was also recorded in more than half the of the records of this biotope although less information was available on this species with which to assess sensitivity.

    Species indicative of sensitivity

    Community ImportanceSpecies nameCommon Name
    Important characterizingCiona intestinalisA sea squirt
    Important characterizingProtanthea simplexSealoch anemone

    Physical Pressures

     IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
    High Moderate Moderate Major decline Low
    All three important characterizing species are highly intolerant of substratum loss. The slower growing and longer lived Neocrania anomala will probably be the limiting factor in the recovery of the biotope although both Neocrania anomala and Protanthea simplex have moderate recoverability.
    High Moderate Moderate Major decline Moderate
    Both Neocrania anomala and Protanthea simplex are highly intolerant of smothering. Ciona intestinalis, being also recorded from areas of modified substratum and high siltation is more tolerant but still intermediately intolerant. The slower growing and longer lived Neocrania anomala will probably be the limiting factor in the recovery of the biotope although both Neocrania anomala and Protanthea simplex have moderate recoverability.
    Low Very high Very Low Minor decline Moderate
    All three of the selected important characterizing species have low intolerance to siltation. Neocrania anomala and Protanthea simplex have very high recoverability from siltation.
    Not relevant Not relevant Not relevant Not relevant High
    The deeper water location (greater than 10 m) of this biotope means that desiccation is highly unlikely to be a relevant factor.
    Not relevant Not relevant Not relevant Not relevant High
    The deep water location (greater than 10 m) of this biotope means that exposure to an emergence regime is highly unlikely to be a relevant factor.
    High Moderate Moderate Major decline Low
    The biotope occurs in the landward basins of fjordic sea lochs where water flow rate is likely to be low. Increases in flow rate may cause high intolerance in Neocrania anomala and intermediate intolerance in Protanthea simplex. The slower growing and longer lived Neocrania anomala will probably be the limiting factor in the recovery of the biotope.
    High Moderate Moderate Major decline Low
    Protanthea simplex has high intolerance to long term chronic temperature increases. The biotope has a restricted distribution along the west coast of Scotland. Long term increases in temperature will cause a decrease in available suitable habitat. Ciona intestinalis may be intermediately intolerant of short term acute decreases in temperature and Neocrania anomala may be intermediately intolerant of short term acute increases in temperature. Intolerance in the lower salinity biotopes where Protanthea simplex is absent may be different. The moderate recoverability of Protanthea simplex from temperature change is likely to be the slowest part of biotope recovery.
    Tolerant Not relevant Not relevant No change Low
    None of the selected important characterizing species from this biotope are intolerant of changes in turbidity.
    Not relevant Not relevant Not relevant Not relevant Moderate
    The selected important characterizing species in this biotope may have high or intermediate intolerance to wave exposure. However, the circalittoral and highly sheltered location of the biotope means that changes in wave exposure are extremely unlikely so the factor has been assessed as not relevant.
    Tolerant Not relevant Not relevant No change High
    Neocrania anomala has low intolerance to noise vibrations but overall the biotope is unlikely to be sensitive to disturbance by noise.
    Tolerant Not relevant Not relevant No change High
    Neocrania anomala has a shadow reflex that causes the valves to clamp shut giving a low intolerance to visual presence but overall, the biotope is unlikely to be sensitive to visual presence.
    High Moderate Moderate Major decline High
    Erect epifaunal species are particularly vulnerable to physical disturbance. Hydroids and bryozoans are likely to be removed or damaged by bottom trawling or dredging (Holt et al., 1995). Veale et al. (2000) reported that the abundance, biomass and production of epifaunal assemblages decreased with increasing fishing effort. Hydroid and bryozoan matrices were reported to be greatly reduced in fished areas (Jennings & Kaiser, 1998 and references therein). Damage to emergent epifauna was the first sign of damage from scallop dredging on horse mussel beds (see Modiolus modiolus) (Service & Magorrian, 1997; Service, 1988; Magorrian & Service, 1998). For example, Protanthea simplex and Ciona intestinalis were both considered to be highly intolerant of physical disturbance and abrasion (see species reviews). The growth form and more durable nature of the valves of Neocrania anomala suggested an intermediate intolerance. Given the likely intolerance of epifaunal communities, an overall intolerance of high has been suggested.

    The recolonization of epifauna on vertical rock walls was investigated by Sebens (1985, 1986). He reported that rapid colonizers such as encrusting corallines, encrusting bryozoans, amphipods, and tubeworms recolonized within 1-4 months. Ascidians such as Dendrodoa carnea, Molgula manhattensis and Aplidium spp. achieved significant cover in less than a year, and, together with Halichondria panicea, reached pre-clearance levels of cover after 2 years. A few individuals of Alcyonium digitatum and Metridium dianthus colonized within 4 years (Sebens, 1986). Large sponges and sea anemones would probably take longer to reach pre-clearance levels. Therefore, a recoverability of moderate has been recorded.

    High Moderate Moderate Major decline High
    Neocrania anomala individuals are cemented to the substratum and cannot reform an attachment if displaced from the substratum. Ciona intestinalis has some limited ability to reform attachments following displacement. The slower growing and longer lived Neocrania anomala will probably be the limiting factor in the recovery of the biotope.

    Chemical Pressures

    No information No information No information Insufficient
    Not relevant
    Heavy metal contamination
    No information No information No information Insufficient
    Not relevant
    Hydrocarbon contamination
    No information No information No information Insufficient
    Not relevant
    Radionuclide contamination
    No information No information No information Insufficient
    Not relevant
    Changes in nutrient levels
    Tolerant Not relevant Not relevant No change Moderate
    There is some evidence that increased levels of organic nutrients is of benefit to Ciona intestinalis populations (Naranjo et al., 1996). Dissolved organic matter can also form a nutrition component for other species such as sponges. It is unlikely that changes in nutrient levels will have much effect on the biotope.
    Intermediate High Low Decline Low
    The biotope SCR.NeoPro contains Protanthea simplex which has intermediate intolerance to decreases in salinity. The sea loch anemone may only survive in fully saline waters as it is not found in the two similar sub-biotopes where salinity is variable, reduced or low. Reductions in salinity in the biotope may cause this species to be lost, changing the biotope. The two sub-biotopes A4.3142 and SCR.NeoPro.CaTw are characterized by low/reduced and variable salinity respectively. In SCR.NeoPro.Den, increases in salinity may cause high intolerance. Recoverability of SCR.NeoPro based on the recoverability of Protanthea simplex from change in salinity is high. However, in the sub-biotopes, where Protanthea simplex is absent recoverability may be different.
    Intermediate High Low Decline Low
    Protanthea simplex has intermediate intolerance to decreases in oxygen concentration. In the sub-biotopes where the sea loch anemone does not occur changes in oxygenation may have different effects. Brachiopods can sustain anaerobic metabolism for 3-5 days although at low oxygen concentrations, activity may be reduced. It is likely that characterizing species other than the three selected for the assessment are intolerant of decreases in oxygenation. Cole et al. (1999) suggest possible adverse effects on marine species below 4 mg/l and probable adverse effects below 2mg/l. Recoverability of SCR.NeoPro based on the recoverability of Protanthea simplex and Neocrania anomala from change in oxygenation is likely to be high. However, in the sub-biotopes, where Protanthea simplex is absent, recoverability may be different.

    Biological Pressures

    No information No information No information Insufficient
    Not relevant
    No information No information No information Insufficient
    Not relevant
    Not relevant Not relevant Not relevant Not relevant Low
    It is extremely unlikely that any of the species indicative of sensitivity would be targeted for extraction and we have no evidence for the indirect effects of extraction of other species on this biotope.
    Not relevant Not relevant Not relevant Not relevant Low

    Additional information



    1. Álvarez, F. & Emig, C., 2000. Brachiopoda from the Luso-Iberian zone. I. Biology and ecology. In The Millennium Brachiopod Congress, London, 2000. Abstracts.

    2. Aneiros, F., Rubal, M., Troncoso, J.S. & Bañón, R., 2015. Subtidal benthic megafauna in a productive and highly urbanised semi-enclosed bay (Ría de Vigo, NW Iberian Peninsula). Continental Shelf Research, 110, 16-24.

    3. Bay-Nouailhat, W., 2007. Description de Sarcodictyon roseum. [cited 17.03.2016]. Available from:

    4. Blum, J.C., Chang, A.L., Liljesthröm, M., Schenk, M.E., Steinberg, M.K. & Ruiz, G.M., 2007. The non-native solitary ascidian Ciona intestinalis (L.) depresses species richness. Journal of Experimental Marine Biology and Ecology, 342 (1), 5-14.

    5. Bradshaw, C., Veale, L.O., Hill, A.S. & Brand, A.R., 2000. The effects of scallop dredging on gravelly seabed communities. In: Effects of fishing on non-target species and habitats (ed. M.J. Kaiser & de S.J. Groot), pp. 83-104. Oxford: Blackwell Science.

    6. Butman, C.A., 1987. Larval settlement of soft-sediment invertebrates: the spatial scales of pattern explained by active habitat selection and the emerging role of hydrodynamical processes. Oceanography and Marine Biology: an Annual Review, 25, 113-165.

    7. Caputi, L., Crocetta, F., Toscano, F., Sordino, P. & Cirino, P., 2015. Long-term demographic and reproductive trends in Ciona intestinalis sp. A. Marine Ecology, 36 (1), 118-128.

    8. Carlgren, O., 1921. Actiniaria. Pt. 1. Danish Ingolf Expedition, Vol. V, No. 9., pp. 31.  Copenhagen: Bianco Luno.

    9. Carlgren, O., 1949. A survey of the Ptychodactiaria, Corallimorpharia and Actiniaria. Kungliga Svenska Vetenskapsakadamiens Handlingar, Series 4, 1, 16-110.

    10. Carver, C., Mallet, A. & Vercaemer, B., 2006. Biological synopsis of the solitary tunicate Ciona intestinalis. Canadian Manuscript Report of Fisheries and Aquatic Science, No. 2746, v + 55 p. Bedford Institute of Oceanography, Dartmouth, Nova Scotia.

    11. 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.

    12. Chia, F.S. & Spaulding, J.G., 1972. Development and juvenile growth of the sea anemone Tealia crassicornis. Biological Bulletin, Marine Biological Laboratory, Woods Hole, 142, 206-218.

    13. 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:

    14. 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.

    15. Connor, D.W., Dalkin, M.J., Hill, T.O., Holt, R.H.F. & Sanderson, W.G., 1997a. Marine biotope classification for Britain and Ireland. Vol. 2. Sublittoral biotopes. Joint Nature Conservation Committee, Peterborough, JNCC Report no. 230, Version 97.06., Joint Nature Conservation Committee, Peterborough, JNCC Report no. 230, Version 97.06.

    16. Davies, C.E. & Moss, D., 1998. European Union Nature Information System (EUNIS) Habitat Classification. Report to European Topic Centre on Nature Conservation from the Institute of Terrestrial Ecology, Monks Wood, Cambridgeshire. [Final draft with further revisions to marine habitats.], Brussels: European Environment Agency.

    17. 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.

    18. Dumont, C., Gaymer, C. & Thiel, M., 2011. Predation contributes to invasion resistance of benthic communities against the non-indigenous tunicate Ciona intestinalis. Biological Invasions, 13 (9), 2023-2034.

    19. Dybern, B.I., 1965. The life cycle of Ciona intestinalis (L.) f. typica in relation to the environmental temperature. Oikos, 16, 109-131.

    20. Dybern, B.I., 1967. The distribution and salinity tolerance of Ciona intestinalis (L.) f. typica with special reference to the waters around southern Scandinavia. Ophelia, 4 (2), 207-226.

    21. Dybern, B.I., 1969. Distribution and ecology of ascidians in Kviturdvikpollen and Vågsböpollen on the west coast of Norway. Sarsia, 37 (1), 21-40.

    22. Ellington, W.R., 1982. Metabolic responses of the sea anemone Bunodosoma cavernata (Bosc) to declining oxygen tensions and anoxia. Physiological zoology, 240-249.

    23. Emig, C.C., 1997. Ecology of inarticulated brachiopods. In Treatise of Invertebrate Paleontology (Kaesler, RL; editor). Part H, 473-495. Geological Society of America and the University of Kansas Press.

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

    25. Fent, K., 1996. Ecotoxicology of organotin compounds. Critical reviews in toxicology, 26 (1), 3-117.

    26. Glantz, M.H., 2005. Climate variability, climate change and fisheries. Cambridge: Cambridge University Press.

    27. 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.

    28. Gulliksen, B., 1977. Studies from the U.W.L. "Helgoland" on the macrobenthic fauna of rocks and boulders in Lübeck Bay (western Baltic Sea). Helgoländer wissenschaftliche Meeresunters, 30, 519-526.

    29. Hammond, L., 1983. Experimental studies of salinity tolerance, burrowing behavior and pedicle regeneration in Lingula anatina (Brachiopoda, Inarticulata). Journal of Paleontology, 1311-1316.

    30. Hartnoll, R.G., 1998. Circalittoral faunal turf biotopes: an overview of dynamics and sensitivity characteristics for conservation management of marine SACs, Volume VIII. Scottish Association of Marine Sciences, Oban, Scotland. [UK Marine SAC Project. Natura 2000 reports.]

    31. Havenhand, J. & Svane, I., 1989. Larval behaviour, recruitment, and the role of adult attraction in Ascidia mentula O. F. Mueller: Reproduction, genetics and distributions of marine organisms. 23rd European Marine Biology Symposium. Olsen and Olsen, 127-132.

    32. Havenhand, J.N. & Svane, I., 1991. Roles of hydrodynamics and larval behaviour in determining spatial aggregation in the tunicate Ciona intestinalis. Marine Ecology Progress Series, 68, 271-276.

    33. 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.

    34. Heip, C.H., Keegan, B.F. & Lewis, J.R., 1985. Long-Term Changes in Coastal Benthic Communities. In Proceedings of a Symposium, held in Brussels, Belgium, December 9–12, 1985: Springer Science & Business Media.

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

    36. 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.

    37. 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.

    38. Hiscock, K., 1985. Littoral and sublittoral monitoring in the Isles of Scilly. September 22nd to 29th, 1984. Nature Conservancy Council, Peterborough, CSD Report, no. 562., Field Studies Council Oil Pollution Research Unit, Pembroke.

    39. 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.

    40. Jackson, A. 2000. Novocrania anomala, A brachiopod. 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. [17.03.16] Available from:

    41. Jackson, A., 2008. Ciona intestinalis. A sea squirt. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [On-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 16/12/15] Available from:

    42. Jackson, A. 2008b. Protanthea simplex, Sealoch anemone. 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. [17.03.2016] Available from:

    43. James, M.A., Ansell, A.D., Collins, M.J., Curry, G.B., Peck, L.S. & Rhoda, M.C., 1992. Biology of living brachiopods. Advances in Marine Biology, 28, 175-387.

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

    45. JNCC (Joint Nature Conservation Committee), 1999. Marine Environment Resource Mapping And Information Database (MERMAID): Marine Nature Conservation Review Survey Database. [on-line]

    46. Kerby, J.L., Richards‐Hrdlicka, K.L., Storfer, A. & Skelly, D.K., 2010. An examination of amphibian sensitivity to environmental contaminants: are amphibians poor canaries? Ecology Letters, 13 (1), 60-67.

    47. Kocak, F. & Kucuksezgin, F., 2000. Sessile fouling organisms and environmental parameters in the marinas of the Turkish Aegean coast. Indian journal of marine sciences, 29 (2), 149-157.

    48. Lambert, C.C. & Lambert, G., 1998. Non-indigenous ascidians in southern California harbors and marinas. Marine Biology, 130 (4), 675-688.

    49. Laupsa, M., 2015. Spawning, settlement and growth of Ciona intestinalis in Øygarden, Hardangerfjorden and Kvitsøy. Master's thesis. University of Bergen.

    50. 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.

    51. Long, J.A. & Stricker, S.A., 1991. Brachiopoda. In Reproduction of marine invertebrates, Vol. VI. Echinoderms and Lophophorates. (ed. A.C. Giese, J.S. Pearse & V.B. Pearse). California: The Boxwood Press.

    52. Mackie, G. & Bone, Q., 1976. Skin impulses and locomotion in an ascidian tadpole. Journal of the Marine Biological Association of the United Kingdom, 56 (03), 751-768.

    53. Magorrian, B.H. & Service, M., 1998. Analysis of underwater visual data to identify the impact of physical disturbance on horse mussel (Modiolus modiolus) beds. Marine Pollution Bulletin, 36, 354-359.

    54. Manuel, R.L., 1988. British Anthozoa. London: Academic Press.[Synopses of the British Fauna, no. 18.]

    55. Marin, M.G., Bresan, M., Beghi, L. & Brunetti, R., 1987. Thermo-haline tolerance of Ciona intestinalis (L. 1767) at different developmental stages. Cahiers de Biologie Marine, 28, 45-57.

    56. Mazouni, N., Gaertner, J. & Deslous-Paoli, J.-M., 2001. Composition of biofouling communities on suspended oyster cultures: an in situ study of their interactions with the water column. Marine Ecology Progress Series, 214, 93-102.

    57. MBA (Marine Biological Association), 1957. Plymouth Marine Fauna. Plymouth: Marine Biological Association of the United Kingdom.

    58. McCammon, H.M., 1972. Establishing and Maintaining Articulate Brachiopods in Aquaria. Journal of Geological Education, 20 (3), 139-142.

    59. McCammon, H.M., 1973. The ecology of Magellania venosa, an articulate brachiopod. Journal of Paleontology, 266-278.

    60. McDonald, J., 2004. The invasive pest species Ciona intestinalis (Linnaeus, 1767) reported in a harbour in southern Western Australia. Marine Pollution Bulletin, 49 (9), 868-870.

    61. Meadows, P.S. & Campbell, J.I., 1972. Habitat selection by aquatic invertebrates. Advances in Marine Biology, 10, 271-382.

    62. Mercier, A., Pelletier, É. & Hamel, J.-F., 1998. Response of temperate sea anemones to butyltin contamination. Canadian Journal of Fisheries and Aquatic Sciences, 55 (1), 239-245.

    63. Mercier, A., Pelletier, E. & Hamel, J.-F., 1996. Toxicological response of the symbiotic sea anemone Aiptasia pallida to butyltin contamination. Marine Ecology Progress Series, 14 (1), 133-146.

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

    65. Millar, R.H., 1966. Tunicata Ascidiacea. Oslo, Universitetsforlaget.

    66. 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.

    67. Monniot, C. & Monniot, F., 1994. Additions to the inventory of eastern tropical Atlantic ascidians; arrival of cosmopolitan species. Bulletin of Marine Science, 54 (1), 71-93.

    68. 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.

    69. Naylor. P., 2011. Great British Marine Animals, 3rd Edition. Plymouth. Sound Diving Publications.

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

    71. Nielsen, C., 1991. The development of the brachiopod Crania (Neocrania) anomala (OF Müller) and its phylogenetic significance. Acta Zoologica, 72 (1), 7-28.

    72. Nomaguchi, T.A., Nishijima, C., Minowa, S., Hashimoto, M., Haraguchi, C., Amemiya, S. & Fujisawa, H., 1997. Embryonic thermosensitivity of the ascidian, Ciona savignyi. Zoological Science, 14 (3), 511-515.

    73. Nyholm, K-G., 1959. On the development of the primitive actinian Protanthea simplex, Carlgren. Zoologiska Bidrag Fran Uppsala, Band 33 1958-1962, 69-78.

    74. Pérès, J.M., 1943. Recherches sur le sang et les organes neuraux des Tuniciers. Annales de l’Institut Oceanographique (Monaco), 21, 229-359.

    75. Petersen, J. & Riisgård, H.U., 1992. Filtration capacity of the ascidian Ciona intestinalis and its grazing impact in a shallow fjord. Marine Ecology-Progress Series, 88, 9-17.

    76. Picton, B.E. & Morrow, C.C., 2015. Ascidia mentula O F Müller, 1776. In Encyclopedia of Marine Life of Britain and Ireland. [cited 26/01/16]. Available from:

    77. Prestrud, P., Strøm, H. & Goldman, H.V., 2004. A catalogue of the terrestrial and marine animals of Svalbard. Norsk Polarinstitutt.

    78. Printrakoon, C. & Kamlung-ek, A., 2013. Socioeconomic study and economic value of living fossil, Lingula sp. in mangrove ecosystem in Trat Province, Thailand. Chinese Journal of Population Resources and Environment, 11 (3), 187-199.

    79. Radolović, M., Bakran-Petricioli, T., Petricioli, D., Surić, M. & Perica, D., 2015. Biological response to geochemical and hydrological processes in a shallow submarine cave. Mediterranean Marine Science, 16 (2), 305-324.

    80. Ramsay, A., Davidson, J., Bourque, D. & Stryhn, H., 2009. Recruitment patterns and population development of the invasive ascidian Ciona intestinalis in Prince Edward Island, Canada. Aquatic Invasions, 4 (1), 169-176.

    81. Ramsay, A., Davidson, J., Landry, T. & Stryhn, H., 2008. The effect of mussel seed density on tunicate settlement and growth for the cultured mussel, Mytilus edulis. Aquaculture, 275 (1), 194-200.

    82. Renborg, E., Johannesson, K. & Havenhand, J., 2014. Variable salinity tolerance in ascidian larvae is primarily a plastic response to the parental environment. Evolutionary ecology, 28 (3), 561-572

    83. Riisgård, H.U., Jürgensen, C. & Clausen, T., 1996. Filter-feeding ascidians (Ciona intestinalis) in a shallow cove: implications of hydrodynamics for grazing impact. Journal of Sea Research, 35 (4), 293-300.

    84. Robbins, I., 1984a. The regulation of ingestion rate, at high suspended particulate concentrations, by some phleobranchiate ascidians. Journal of Experimental Marine Biology and Ecology, 82 (1), 1-10.

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

    86. Rowley, S.J., 2008. A sea squirt (Ascidia mentula). 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 26/01/16]. Available from:

    87. Rudwick, M.J.S., 1970. Living and fossil brachiopods. London: Hutchinson University Library

    88. Ruppert, E.E. & Barnes, R.D., 1994. Invertebrate zoology (6th ed.). Fort Worth, USA: Saunders College Publishing.

    89. Sabbadin, A., 1957. Il ciclo biologico di Ciona intestinalis (L.), Molgula manhattensis (De Kay) e Styela plicata (Lesueur) nella Laguna Veneta.

    90. Scheltema, R.S., 1974. Biological interactions determining larval settlement of marine invertebrates. Thalassia Jugoslavica, 10, 263-296.

    91. Sebens, K.P., 1981. Recruitment in a Sea Anemone Population: Juvenile Substrate Becomes Adult Prey. Science, 213 (4509), 785-787.

    92. Sebens, K.P., 1985. Community ecology of vertical rock walls in the Gulf of Maine: small-scale processes and alternative community states. In The Ecology of Rocky Coasts: essays presented to J.R. Lewis, D.Sc. (ed. P.G. Moore & R. Seed), pp. 346-371. London: Hodder & Stoughton Ltd.

    93. Sebens, K.P., 1986. Spatial relationships among encrusting marine organisms in the New England subtidal zone. Ecological Monographs, 56, 73-96.

    94. Seeley, R., 2006. Sealife survey. MarLIN news, 9.

    95. Service, M. & Magorrian, B.H., 1997. The extent and temporal variation of disturbance to epibenthic communities in Strangford Lough, Northern Ireland. Journal of the Marine Biological Association of the United Kingdom, 77, 1151-1164.

    96. Service, M., 1998. Recovery of benthic communities in Strangford Lough following changes in fishing practice. ICES Council Meeting Paper, CM 1998/V.6, 13pp., Copenhagen: International Council for the Exploration of the Sea (ICES).

    97. Shick, J.M., 2012. A functional biology of sea anemones: Springer Science & Business Media.

    98. Shumway, S., 1978. Respiration, pumping activity and heart rate in Ciona intestinalis exposed to fluctuating salinities. Marine Biology, 48 (3), 235-242.

    99. Shumway, S.E., 1978. Activity and respiration of the sea anemone, Metridium senile (L.) exposed to salinity fluctuations. Journal of Experimental Marine Biology and Ecology, 33, 85-92.

    100. Smith, J.E. (ed.), 1968. 'Torrey Canyon'. Pollution and marine life. Cambridge: Cambridge University Press.

    101. Stanley, J.A., Wilkens, S., McDonald, J.I. & Jeffs, A.G., 2016. Vessel noise promotes hull fouling. In The Effects of Noise on Aquatic Life II: Springer, pp. 1097-1104.

    102. Steele-Petrović, H.M., 1975. An explanation for the tolerance of brachiopods and relative intolerance of filter-feeding bivalves for soft muddy bottoms. Journal of Paleontology, 552-556.

    103. Steele-Petrović, H.M., 1976. Brachiopod food and feeding processes. Palaeontology, 19 (3), 417-436

    104. Steele-Petrović, H.M., 1979. The physiological differences between articulate brachiopods and filter-feeding bivalves as a factor in the evolution of marine level-bottom communities. Palaeontology, 22 (1), 101-134.

    105. Svane, I., 1984. Observations on the long-term population dynamics of the perennial ascidian, Ascidia mentula O F Müller, on the Swedish west coast. The Biological Bulletin, 167 (3), 630-646.

    106. Svane, I. & Havenhand, J.N., 1993. Spawning and dispersal in Ciona intestinalis (L.) Marine Ecology, Pubblicazioni della Stazione Zoologica di Napoli. I, 14 , 53-66.

    107. Thayer, C.W., 1986. Are brachiopods better than bivalves? Mechanisms of turbidity tolerance and their interaction with feeding in articulates. Paleobiology, 12 (02), 161-174.

    108. Therriault, T.W. & Herborg, L.-M., 2008. Predicting the potential distribution of the vase tunicate Ciona intestinalis in Canadian waters: informing a risk assessment. ICES Journal of Marine Science: Journal du Conseil, 65 (5), 788-794.

    109. 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:

    110. Tunnicliffe, V. & Wilson, K., 1988. Brachiopod populations: Distribution in fjords of British Columbia(Canada) and tolerance of low oxygen concentrations. Marine ecology progress series. 47 (2), 117-128.

    111. Van Ofwegen, L., 2015. Sarcodictyon roseum. World Register of Marine Species: [17/03/16]. Available from

    112. Van Ofwegen, L., Grasshoff, M. & Van der Land, J., 2001. Octocorallia (excl. Pennatulacea). European register of marine species: a check-list of the marine species in Europe and a bibliography of guides to their identification. Collection Patrimoines Naturels, 50, 104-105.

    113. Veale, L.O., Hill, A.S., Hawkins, S.J. & Brand, A.R., 2000. Effects of long term physical disturbance by scallop fishing on subtidal epifaunal assemblages and habitats. Marine Biology, 137, 325-337.

    114. Verhoeven, J. & Van Vierssen, W., 1978a. Structure of macrophyte dominated communities in two brackish lagoons on the island of Corsica, France. Aquatic Botany, 5, 77-86.

    115. Whittingham, D.G., 1967. Light-induction of shedding of gametes in Ciona intestinalis and Morgula manhattensis. Biological Bulletin, Marine Biological Laboratory, Woods Hole, 132, 292-298.

    116. Wood. C., 2005. Seasearch guide to sea anemones and corals of Britain and Ireland. Ross-on-Wye: Marine Conservation Society.

    117. WoRMS, 2015. World Register of Marine Species. (11/04/2007).

    118. Yamaguchi, M., 1975. Growth and reproductive cycles of the marine fouling ascidians Ciona intestinalis, Styela plicata, Botrylloides violaceus, and Leptoclinum mitsukurii at Aburatsubo-Moroiso Inlet (Central Japan). Marine Biology, 29 (3), 253-259.

    119. 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.

    120. Zhang, J. & Fang, J., 1999. Study on the oxygen consumption rates of some common species of ascidian. Journal of fishery sciences of China, 7 (1), 16-19.

    121. Zhang, J., Fang, J. & Dong, S., 1999. Study on the ammonia excretion rates of four species ascidian. Marine Fisheries Research, 21 (1), 31-36.


    This review can be cited as:

    Readman, J.A.J. & Jackson, A. 2016. [Novocrania anomala] and [Protanthea simplex] on sheltered circalittoral rock. 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 19-06-2018]. Available from:

    Last Updated: 31/03/2016