Laminaria saccharina park on very sheltered lower infralittoral rock

Researched byDr Keith Hiscock Refereed byThis information is not refereed.
EUNIS CodeA3.3133 EUNIS NameLaminaria saccharina park on very sheltered lower infralittoral rock


UK and Ireland classification

EUNIS 2008A3.3133Laminaria saccharina park on very sheltered lower infralittoral rock
EUNIS 2006A3.3133Laminaria saccharina park on very sheltered lower infralittoral rock
JNCC 2004IR.LIR.K.Lsac.PkLaminaria saccharina park on very sheltered lower infralittoral rock
1997 BiotopeIR.SIR.K.Lsac.PkLaminaria saccharina park on very sheltered lower infralittoral rock


Silty rock with a Laminaria saccharina park (often the cape-form). Beneath the canopy, the bedrock and boulders are covered by coralline algal crusts and urchins such as Echinus esculentus and Psammechinus miliaris are present. Though present, foliose algae are less abundant than in the Laminaria hyperborea park (MIR.Lhyp.Pk) with the most common species being Phycodrys rubens and Delesseria sanguinea. The most conspicuous animals in this biotope are ascidians, particularly Ascidia mentula, Ciona intestinalis and Corella parallelogramma. (Information taken from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b). NB. Laminaria saccharina is now considered to be a synonym of Saccharina latissima.

Recorded distribution in Britain and Ireland

Widely recorded on the west coast of Scotland and in Shetland especially in sheltered sea loch and voe habitats. Otherwise identified from Portland Harbour.

Depth range

5-10 m, 10-20 m

Additional information


Listed By

Further information sources

Search on:


Habitat review


Ecological and functional relationships

Saccharina latissima is the most conspicuous species and dominates the biotope from the point-of-view of ecological relationships. The kelp fronds shade the understory algae and rock below and are likely to sweep the rock - both creating areas where other algae struggle to survive. The sea urchins Echinus esculentus and Psammechinus miliaris graze the rock below leaving, with the effects of frond-sweeping, extensive bare crustose coralline algae dominating the rock. However, large solitary tunicates colonize the rock and the algae, typifying situations of very low water movement where active suspension feeders thrive.

Seasonal and longer term change

Growths of ephemeral algae are likely during the summer together with fresh growth of perennial algal species. Associated fish such as two-spot gobies are likely to be present in higher abundance at the end of the summer than at the start. Seabed animal species in this biotope are not highly changeable.

Habitat structure and complexity

The biotope offers a wide range of surfaces for settlement and shelter of species. The bedrock is colonized by encrusting and foliose red algae with a variety of tubicolous animals and ascidian species attached. The holdfasts of Saccharina latissima offer refuges for a wide range of small mobile species such as worms and amphipods whilst the fronds may be colonized by encrusting bryozoans, hydroids and ascidians. The shelter afforded by algal fronds attracts small fish species. Complexity is increased if the rock is fissured or the biotope colonizing boulders where the underboulder habitat provides additional shelter and complexity.


Primary and secondary productivity are probably both high. Algae are consumed directly by urchins especially and also provide material for detritus feeders when they die and break-up. Much secondary productivity relies on the acquisition of suspended food by active suspension feeders especially ascidians.

Recruitment processes

The characterizing species in this biotope all have planktonic larvae and propagules and are mainly short-lived. There is therefore high recruitment and high turnover. However, species that require or prefer settlement on algal substrata will require presence of those substrata.

Time for community to reach maturity

The main characterizing species, Saccharina latissima, rapidly colonizes cleared areas of the substratum and Kain (1975) recorded that Saccharina latissima (studied as Laminaria saccharina) was abundant six months after the substratum was cleared so colonization should be rapid. However, whilst it most likely settles rapidly, the coralline algal species covering rock, represented by Lithophyllum incrustans, grows at a rate of only <7mm a year (Irvine & Chamberlain 1994) and will take much longer to reach significant cover.

Additional information


Preferences & Distribution

Recorded distribution in Britain and IrelandWidely recorded on the west coast of Scotland and in Shetland especially in sheltered sea loch and voe habitats. Otherwise identified from Portland Harbour.

Habitat preferences

Depth Range 5-10 m, 10-20 m
Water clarity preferences
Limiting Nutrients No information found
Salinity Full (30-40 psu)
Physiographic Enclosed coast / Embayment
Biological Zone Lower infralittoral
Substratum Bedrock, Large to very large boulders, Small boulders, Cobbles
Tidal Very Weak (negligible), Weak < 1 knot (<0.5 m/sec.)
Wave Sheltered, Very sheltered
Other preferences

Additional Information

Whilst the biotope has been recorded almost only in Scotland, it is most likely that suitable habitats have not been surveyed in other areas. The main characterizing species are found throughout Britain and Ireland.

Species composition

Species found especially in this biotope

    Rare or scarce species associated with this biotope


    Additional information

    No text entered

    Sensitivity reviewHow is sensitivity assessed?


    Saccharina latissima is the major canopy forming species and provides a variety of substrata for colonization whilst Lithophyllum incrustans is the major seabed substratum cover. Grazing is a key element of maintaining the community and the grazer identified as indicative of sensitivity is the sea urchin Echinus esculentus. Solitary sea squirts, Ciona intestinalis, represent the active suspension feeding elements of the fauna whilst the brittle star Ophiothrix fragilis represents passive suspension feeders.

    Species indicative of sensitivity

    Community ImportanceSpecies nameCommon Name
    Important characterizingCiona intestinalisA sea squirt
    Key structuralEchinus esculentusEdible sea urchin
    Important characterizingLithophyllum incrustansEncrusting coralline alga
    Important characterizingOphiothrix fragilisCommon brittlestar
    Important characterizingSaccharina latissimaSugar kelp

    Physical Pressures

     IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
    High Moderate Moderate Major decline High
    Most of the species characteristic of this biotope are permanently attached to the substratum and would be removed upon substratum loss. For recoverability, see Additional Information.
    Intermediate High Low Decline Moderate
    Some species, especially Saccharina latissima, are likely to protrude above smothering material whilst some, such as Lithophyllum incrustans, will most likely survive under smothering material. Mobile species such as urchins and brittle stars will be able to migrate out of most smothering material. Others such as the active suspension feeders and low-growing foliose algae are likely to be killed by smothering. However, since keystone species are likely to survive an intolerance of intermediate has been indicated. For recoverability, see Additional Information.
    Intermediate High Low Minor decline Moderate
    Increase in suspended sediment is likely to have a significant effect in the low water movement regime in which this biotope lives. Settling silt may smother organisms or clog respiratory and feeding organs (especially sea squirts). However, many of the species in this biotope live in areas of high silt content and be able to survive. For effects on light penetration, see turbidity. For recoverability, see Additional Information.
    Tolerant High Not sensitive* Minor decline Moderate
    Decrease in suspended sediment levels is not likely to have a significant effect on this biotope although suspension and deposit feeders that gain nutrients from silt may be adversely affected. On the other hand, suspension feeders may be less affected by clogging by silt. For effects on light penetration, see turbidity.
    Intermediate High Low Minor decline Moderate
    The biotope is predominantly sublittoral but does extend onto the shore and therefore shows some ability to resist desiccation. On a sunny day at low water of spring tides, damage (bleaching) is likely to occur to the Saccharina latissima plants but not destroy them completely. Species living below the kelp fronds will be protected by them from the worst effects of desiccation. There may be a minor loss of species. For recoverability, see Additional Information.
    High High Moderate Decline Low
    The biotope is predominantly sublittoral and the dominant species (Saccharina latissima) and many of the subordinate species, especially solitary sea squirts, are unlikely to survive an increased emergence regime. Several mobile species such as sea urchins, brittle stars and feather stars are likely to move away. However, providing that suitable substrata are present, the biotope is likely to re-establish further down the shore within a similar emergence regime to that which existed previously.
    Not sensitive* Not relevant
    The biotope is sublittoral and so decrease in emergence is not relevant.
    Intermediate High Low Minor decline Low
    It is unlikely that species in the biotope will be killed by an increase in flow rate. Existing organisms are likely to persist although conditions will not be ideal. A few mobile species such as brittle stars might be swept away. However, in situations where the substratum on which Saccharina latissima occurs is of cobbles or pebbles, it is likely that kelp plants might cause sufficient drag for plants and attached organisms to be swept away. In that case, a different biotope is likely to develop.
    Low High Low Minor decline Low
    The biotope exists in areas with very little or no tidal flow.
    Low Very high Very Low No change Low
    The species characteristic of the biotope are well within the range of temperatures in which they occur geographically and are unlikely to be lost as a result of higher temperatures occurring in the long term. However, exposure to high temperatures for several days may produce stress in some components but recovery would be rapid.
    Low Very high Moderate Moderate
    The species characteristic of the biotope are well within the range of temperatures in which they occur geographically and are unlikely to be lost as a result of lower temperatures occurring in the long term. However, exposure to low temperatures for several days may result in some mortality. Records in Crisp (1964) suggest that the species in the biotope are likely to be of low susceptibility to cold although Psammechinus miliaris was adversely affected by the 1962/63 winter and Antedon bifida is believed to have been lost from the Menai Strait following the 1947 winter (D.J. Crisp pers. comm. to K. Hiscock).
    Low High Low Minor decline Low
    Several of the characteristic species are algae that rely on light for photosynthesis. Decrease in light penetration as a result of higher turbidity is unlikely to be fatal in the short term but in the long term will result in a reduction in downward extent and therefore overall extent of the biotope.
    Tolerant* Not sensitive Not relevant Moderate
    The biotope is characterized especially by algae which are likely to increase in downward extent if light penetration increases.
    High Moderate Moderate Major decline Moderate
    This is a fundamentally sheltered coast biotope with species that do not appear to occur in wave exposed situations. Increased wave action is likely to dislodge Saccharina latissima plants, especially if they are attached to cobbles, dislodge brittle stars and feather stars and interfere with feeding in solitary tunicates.
    Low High Low Minor decline Low
    Some small amount of wave action is most likely required to prevent stagnation occurring in this biotope. Stagnation would most likely result is some localized de-oxygenation. And some species in sheltered pockets would be lost.
    Tolerant Not relevant Not relevant No change High
    The macroalgae characterizing the biotope have no known sound or vibration sensors. The response of macroinvertebrates is not known.
    Tolerant Not relevant Not relevant No change High
    Macrophytes have no known visual sensors. Most macroinvertebrates have poor or short range perception and are unlikely to be affected by visual disturbance such as shading.
    Intermediate High Low Major decline High
    Saccharina latissima, other algae and the large solitary tunicates are likely to be especially intolerant of physical disturbance and to be removed from the substratum. Sea urchins, brittlestars, and feather stars are likely to be damaged. However, the main species covering rock, encrusting coralline algae, will survive increased abrasion including if cobbles are moved around. Overall, some keystone species are likely to be lost but some will remain and an intolerance of intermediate is suggested. For recoverability, see additional information below.
    Intermediate High Low Decline Low
    Although many of the species in the biotope are sessile and would therefore be killed if removed from their substratum, displacement will often be of the boulders or cobbles on which the community occurs in which case survival will be high. The 'Intermediate' ranking given here supposes that some individuals sessile organisms will be removed and die. Mobile organisms such as the echinoderms in the biotope are likely to survive displacement. Recovery rate assumes that the characteristic species of the biotope will remain, albeit in lower numbers.

    Chemical Pressures

    High Moderate Moderate Decline Moderate
    Several of the species characteristic of the biotope are reported as having high intolerance to synthetic chemicals. For instance, Cole et al. (1999) suggested that herbicides such as Simazine and Atrazine were very toxic to macrophytic algae. Hoare & Hiscock (1974) noted that almost all red algal species and many animal species were absent from Amlwch Bay in North Wales adjacent to an acidified halogenated effluent. Red algae have also been found to be sensitive to oil spill dispersants (O'Brien & Dixon, 1976; Grandy quoted in Holt et al. 1995). Recovery is likely to occur fairly rapidly - see Additional Information.
    Heavy metal contamination
    Low High Low Minor decline Very low
    Sporophytes of Saccharina latissima have a low intolerance to heavy metals, but the early life stages are more intolerant. The effects of copper, zinc and mercury on Saccharina latissima (studied as Laminaria saccharina) have been investigated by Thompson & Burrows (1984). They observed that the growth of sporophytes was significantly inhibited at 50 µg Cu /l, 1000 µg Zn/l and 50 µg Hg/l. Zoospores were found to be more intolerant and significant reductions in survival rates were observed at 25 µg Cu/l, 1000 µg Zn/l and 5 µg/l. Little is known about the effects of heavy metals on echinoderms. Bryan (1984) reported that early work had shown that echinoderm larvae were intolerant of heavy metals, e.g. the intolerance of larvae of Paracentrotus lividus to copper (Cu) had been used to develop a water quality assessment. Kinne (1984) reported developmental disturbances in Echinus esculentus exposed to waters containing 25 µg / l of copper (Cu). Sea-urchins, especially the eggs and larvae, are used for toxicity testing and environmental monitoring (reviewed by Dinnel et al. 1988). Taken together with the findings of Gomez & Miguez-Rodriguez (1999) above it is likely that echinoderms are intolerant of heavy metal contamination. Overall, the biotope is may show some minor change following heavy metal contamination at the level of the baseline.
    Hydrocarbon contamination
    Intermediate High Low Decline Low
    Red algae have been found to be intolerant of oil and oil spill dispersants (O'Brien & Dixon 1976; Grandy quoted in Holt et al. 1995). Foliose red algae in the biotope may be subject to bleaching and death and, in lower shore/shallow sublittoral situations, encrusting calcareous algae are likely to be bleached and by 'fresh' oil. However, observations following the Sea Empress oil spill (Chamberlain, 1997) suggest that regeneration from below the destroyed area of crustose corallines repairs damage and recovery occurred within a year. Holt et al. (1995) report that Saccharina latissima (studied as Laminaria saccharina) has been seen to show no discernible effects from oil spills. Feather stars and sea urchins have both been observed to be killed by oil or oil and dispersant. For recoverability, see additional information below. Whilst some keystone (grazing) species might be killed, the overall character of the biotope will most likely remain and an intolerance of 'Intermediate' has been indicated.
    Radionuclide contamination
    No information Not relevant No information Insufficient
    Not relevant
    Changes in nutrient levels
    Low High Low Minor decline Low
    Evidence is equivocal. For Saccharina latissima (studied as Laminaria saccharina), Conolly & Drew (1985) found that plants at the most eutrophic site in a study on the east coast of Scotland where nutrient levels were 25% higher than average exhibited a higher growth rate. However, Read et al. (1983) reported that, after removal of a major sewage pollution in the Firth of Forth, Saccharina latissima (studied as Laminaria saccharina) became abundant where previously it had been absent. Increased nutrients may increase the abundance of ephemeral algae and result in smothering or changing the character of the biotope. Any recovery is likely to be high as species are unlikely to be completely lost: see Additional Information.
    Tolerant Not relevant Not relevant Not relevant Moderate
    The biotope occurs in full salinity conditions and so increase in salinity from variable or low would not adversely affect it.
    Low High Low Minor decline Moderate
    The biotope occurs in situations that are naturally subject to fluctuating or low salinities: it grows in areas where freshwater run-off dilutes near-surface waters and most components are likely to survive reduced salinity conditions. For instance, Saccharina latissima (studied as Laminaria saccharina) can survive in salinities of 8 psu although growth is retarded below 16 psu (Kain 1979). Delesseria sanguinea is also tolerant of salinities as low as 11 psu in the North Sea. The brittle star Ophiothrix fragilis occurs in salinities of 16 psu and even down to 10 psu (Wolff 1968) and the feather star Antedon bifida is typically present in situations of high freshwater outflow such as at the entrance to the Tamar (own observations). However, some species in the biotope such as the sea urchin Echinus esculentus are unlikely to survive in lower salinity and may perish. Most characteristic species are likely to survive. Species that are lost are likely to have planktonic larvae and recolonize rapidly.
    Intermediate Moderate Moderate Decline Moderate
    The biotope occurs in areas where still water conditions occur and therefore some hypoxia is likely. However, in severe conditions, death of constituent species is likely. Cole et al. (1999) suggest possible adverse effects on marine species below 4 mg/l and probable adverse effects below 2mg/l. For instance, death of a bloom of the phytoplankton Gyrodinium aureolum in Mounts Bay, Penzance in 1978 produced a layer of brown slime on the sea bottom. This resulted in the death of fish and invertebrates, including Echinus esculentus, a characterizing species, presumably due to anoxia caused by the decay of the dead dinoflagellates (Griffiths et al. 1979). For recoverability, see Additional Information.

    Biological Pressures

    Low High Low No change Moderate
    There is little information on microbial pathogen effects on the characterizing species in this biotope. However, Saccharina latissima may be infected by the microscopic brown alga Streblonema aecidioides. Infected algae show symptoms of Streblonema disease, i.e. alterations of the blade and stipe ranging from dark spots to heavy deformations and completely crippled thalli (Peters & Scaffelke, 1996). Infection can reduce growth rates of host algae.Echinus esculentus is susceptible to 'Bald-sea-urchin disease', which causes lesions, loss of spines, tube feet, pedicellariae, destruction of the upper layer of skeletal tissue and death. It is thought to be caused by the bacteria Vibrio anguillarum and Aeromonas salmonicida. Bald sea-urchin disease was recorded from Echinus esculentus on the Brittany Coast. Although associated with mass mortalities of Strongylocentrotus franciscanus in California and Paracentrotus lividus in the French Mediterranean it is not known if the disease induces mass mortality (Bower 1996). However, no evidence of mass mortalities of Echinus esculentus associated with disease have been recorded in Britain and Ireland. It is likely that microbial pathogens will have only a minor possible impact on this biotope.
    Tolerant Not relevant Not relevant No change Moderate
    This assessment of intolerance relates to known non-native species in October 2001. Although non-native species may colonize the biotope they are unlikely to significantly displace or affect native species.
    Intermediate High Low Minor decline Moderate
    Extraction of Saccharina latissima may occur but the plant rapidly colonizes cleared areas of the substratum: Kain (1975) recorded that Saccharina latissima (studied as Laminaria saccharina) was abundant six months after the substratum was cleared so recovery should be rapid. Associated species are unlikely to be affected by removal of Saccharina latissima unless protection from desiccation on the lower shore is important. Echinus esculentus may also be collected. The collection of Echinus esculentus for the curio trade was studied by Nichols (1984). He concluded that the majority of divers collected only large specimens that are seen quickly and often missed individuals covered by seaweed or under rocks, especially if small. As a result, a significant proportion of the population remains.

    An intermediate intolerance has been suggested to reflect the possibility that either of these two species may experience some loss. Given the majority of each is likely to remain however, recovery has been assessed as high.

    Low High Low Minor decline Moderate

    Additional information

    SIR.Lsac.Pk is likely to be naturally disturbed and damaged during storms where the substratum is of mobile cobbles. None of the species present are likely to be long-lived or slow growing and recolonization will be fairly rapid. The main characterizing species, Saccharina latissima, rapidly colonizes cleared areas of the substratum and Kain (1975) recorded that Saccharina latissima (studied as Laminaria saccharina) was abundant six months after the substratum was cleared so recovery should be rapid. However, the main group covering rock, encrusting coralline algae represented by Lithophyllum incrustanswhich grows at a rate of <7mm a year (Irvine & Chamberlain, 1994) will take much longer to cover rocks. Most other characterizing species have a planktonic larva and/or are mobile and so can migrate into the affected area. Development of a balance between grazing species and algae may be of critical importance to recovery. Because some species might not have recovered full abundance within five years and there are likely still to be changes in the algae-grazers balance after five years, recoverability is likely to be only moderate after catastrophic loss but high if only a portion of the biotope is lost.

    Importance review


    Habitats Directive Annex 1Reefs, Large shallow inlets and bays
    UK Biodiversity Action Plan Priority


    No exploitation of this biotope is known.

    Additional information



    1. Lüning, K., 1979. Growth strategy of three Laminana species (Phaeophyceae) inhabiting different depth zones in the sublittoral region of Hegloland (North Sea). Marine Ecological Progress Series, 1, 195-207.
    2. Pihl, L., Wennhage, H. & Nilsson, S., 1994. Fish assemblage structure in relation to macrophytes and filamentous epiphytes in shallow non-tidal rocky-and soft-bottom habitats. Environmental Biology of Fishes, 39 (3), 271-288.
    3. Wernberg, T. & Thomsen, S., 2005. Miniview: What affects the forces required to break or dislodge macroalgae? European Journal of Phycology, 40 (2), 139-148.
    4. Andersen, G.S., 2013. Patterns of Saccharina latissima recruitment. Plos One, 8 (12). pages?

    5. Andersen, G.S., Pedersen, M.F. & Nielsen, S.L., 2013. Temperature, Acclimation and Heat Tolerance of photosynthesis in Norwegian Saccharina latissima (Laminariales, Phaeophyceae). Journal of Phycology, 49 (4), 689-700.

    6. Andersen, G.S., Steen, H., Christie, H., Fredriksen, S. & Moy, F.E., 2011. Seasonal patterns of sporophyte growth, fertility, fouling, and mortality of Saccharina latissima in Skagerrak, Norway: implications for forest recovery. Journal of Marine Biology

    7. Arzel, P., 1998. Les laminaires sur les côtes bretonnes. Évolution de l'exploitation et de la flottille de pêche, état actuel et perspectives. Plouzané, France: Ifremer.
    8. Bartsch, I., Vogt, J., Pehlke, C. & Hanelt, D., 2013. Prevailing sea surface temperatures inhibit summer reproduction of the kelp Laminaria digitata at Helgoland (North Sea). Journal of Phycology, 49 (6), 1061-1073.
    9. Bartsch, I., Wiencke, C., Bischof, K., Buchholz, C.M., Buck, B.H., Eggert, A., Feuerpfeil, P., Hanelt, D., Jacobsen, S. & Karez, R., 2008. The genus Laminaria sensu lato: recent insights and developments. European Journal of Phycology, 43 (1), 1-86.
    10. Bekkby, T. & Moy, F.E., 2011. Developing spatial models of sugar kelp (Saccharina latissima) potential distribution under natural conditions and areas of its disappearance in Skagerrak. Estuarine Coastal and Shelf Science, 95 (4), 477-483.

    11. Bernstein, B.B., Williams, B.E. & Mann, K.H., 1981. The role of behavioral responses to predators in modifying urchins' (Strongylocentrotus droebachiensis) destructive grazing and seasonal foraging patterns. Marine Biology, 63 (1), 39-49.
    12. Birkett, D.A., Maggs, C.A., Dring, M.J. & Boaden, P.J.S., 1998b. Infralittoral reef biotopes with kelp species: an overview of dynamic and sensitivity characteristics for conservation management of marine SACs. Natura 2000 report prepared by Scottish Association of Marine Science (SAMS) for the UK Marine SACs Project., Scottish Association for Marine Science. (UK Marine SACs Project, vol V.).,
    13. Bishop, G.M., 1985. Aspects of the reproductive ecology of the sea urchin Echinus esculentus L. Ph.D. thesis, University of Exeter, UK.

    14. Bolton, J.J. & Lüning, K., 1982. Optimal growth and maximal survival temperatures of Atlantic Laminaria species (Phaeophyta) in culture. Marine Biology, 66, 89-94.

    15. Bonsdorff, E. & Vahl, O., 1982. Food preferences of the sea urchins Echinus actus and Echinus esculentus. Marine Behaviour and Physiology, 8 (3), 243-248.

    16. Boolootian, R.A.,1966. Physiology of Echinodermata. (Ed. R.A. Boolootian), pp. 822-822. New York: John Wiley & Sons.

    17. Borum, J., Pedersen, M.F., Krause-Jensen, D., Christensen, P.B. & Nielsen, K., 2002. Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland. Marine Biology, 141, 11-19.
    18. Bower, S.M., 1996. Synopsis of Infectious Diseases and Parasites of Commercially Exploited Shellfish: Bald-sea-urchin Disease. [On-line]. Fisheries and Oceans Canada. [cited 26/01/16]. Available from:

    19. 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.
    20. Buck, B.H. & Buchholz, C.M., 2005. Response of offshore cultivated Laminaria saccharina to hydrodynamic forcing in the North Sea. Aquaculture, 250 (3-4), 674-691.

    21. Burrows, E.M. & Pybus, C., 1971. Laminaria saccharina and marine pollution in North-East England. Marine Pollution Bulletin, 2, 53-56.
    22. Burrows, E.M., 1958. Sublittoral algal population in Port Erin Bay, Isle of Man. Journal of the Marine Biological Association of the United Kingdom, 37, 687-703.
    23. Burrows, E.M., 1971. Assessment of pollution effects by the use of algae. Proceedings of the Royal Society of London, Series B, 177, 295-306.
    24. Chapman, A.R.O., 1981. Stability of sea urchin dominated barren grounds following destructive grazing of kelp in St. Margaret's Bay, Eastern Canada. Marine Biology, 62, 307-311.
    25. Chavanich, S. & Harris, L.G., 2004. Impact of the non-native macroalga Codium fragile (sur.) hariot ssp. tomentosoides (van goor) silva on the native snail Lacuna vincta (montagu, 1803) in the gulf of maine. Veliger, 47 (2), 85-90.

    26. Cie, D.K. & Edwards, M.S., 2011. Vertical distribution of kelp zoospores. Phycologia, 50 (4), 340-350.

    27. Cole, S., Codling, I.D., Parr, W. & Zabel, T., 1999. Guidelines for managing water quality impacts within UK European Marine sites. Natura 2000 report prepared for the UK Marine SACs Project. 441 pp., Swindon: Water Research Council on behalf of EN, SNH, CCW, JNCC, SAMS and EHS. [UK Marine SACs Project.],
    28. Comely, C.A. & Ansell, A.D., 1988. Invertebrate associates of the sea urchin, Echinus esculentus L., from the Scottish west coast. Ophelia, 28, 111-137.

    29. Conolly N.J. & Drew, E.A., 1985. Physiology of Laminaria. III. Effect of a coastal eutrophication on seasonal patterns of growth and tissue composition in Laminaria digitata and L. saccharina. Marine Ecology, Pubblicazioni della Stazione Zoologica di Napoli I, 6, 181-195.
    30. Cosse, A., Potin, P. & Leblanc, C., 2009. Patterns of gene expression induced by oligoguluronates reveal conserved and environment‐specific molecular defence responses in the brown alga Laminaria digitata. New Phytologist, 182 (1), 239-250.
    31. Davison, I.R. & Pearson, G.A., 1996. Stress tolerance in intertidal seaweeds. Journal of Phycology, 32 (2), 197-211.

    32. Davison, I.R., Greene, R.M. & Podolak, E.J., 1991. Temperature acclimation of respiration and photosynthesis in the brown alga Laminaria saccharina. Marine Biology, 110, 449-454.
    33. Dayton, P.K. & Tegner, M.J., 1984. Catastrophic storms, El-Nino, and patch stability in a southern-california kelp community. Science, 224 (4646), 283-285.
    34. Devinny, J. & Volse, L., 1978. Effects of sediments on the development of Macrocystis pyrifera gametophytes. Marine Biology, 48 (4), 343-348.

    35. Drobyshev, V.P., 1971. Acclimatisation of marine algae when maintained in media of differing salinities. Ékologiya, 1, 96-98.

    36. Druehl, L.D., 1967. Distribution of two species of Laminaria as related to some enviromental factors 1. Journal of Phycology, 3(2), 103-108.

    37. Druehl, L.D., 1970. The pattern of Laminariales distribution in the northeast Pacific. Phycologia, 9 (3), 237-247.

    38. Engelen, A.H., Leveque, L., Destombe, C. & Valer, M., 2011. Spatial and temporal patterns of recovery of low intertidal Laminaria digitata after experimental spring and autumn removal. Cahiers De Biologie Marine, 52 (4), 441-453.

    39. Estes, J.A. & Duggins, D.O., 1995. Sea otters and kelp forests in Alaska: generality and variation in a community ecological paradigm. Ecological Monographs, 65, 75-100.

    40. Frieder, C., Nam, S., Martz, T. & Levin, L., 2012. High temporal and spatial variability of dissolved oxygen and pH in a nearshore California kelp forest. Biogeosciences, 9 (10), 3917-3930.

    41. Gaylord, B., Reed, D.C., Raimondi, P.T. & Washburn, L., 2006. Macroalgal spore dispersal in coastal environments: Mechanistic insights revealed by theory and experiment. Ecological Monographs, 76 (4), 481-502.

    42. Gayral, P. & Cosson, J., 1973. Exposé synoptique des données biologiques sur la laminaire digitée Laminaria digitata. Synopsis FAO sur les pêches, no. 89.
    43. Gerard, V., 1982. In situ water motion and nutrient uptake by the giant kelp Macrocystis pyrifera. Marine Biology, 69 (1), 51-54.

    44. Gerard, V.A. & Du Bois, K.R., 1988. Temperature ecotypes near the southern boundary of the kelp Laminaria saccharina. Marine Biology, 97, 575-580.
    45. Gerard, V.A. & Mann, K.H., 1979. Growth and production of Laminaria longicruris ( Phaeophyta) populations exposed to different intensities of water movement 1. Journal of Phycology, 15 (1), 33-41.

    46. Gerard, V.A., 1987. Hydrodynamic streamlining of Laminaria saccharina Lamour. in response to mechanical stress. Journal of Experimental Marine Biology and Ecology, 107, 237-244.
    47. Gorgula, S.K. & Connell, S.D., 2004. Expansive covers of turf-forming algae on human-dominated coast: the relative effects of increasing nutrient and sediment loads. Marine Biology, 145 (3), 613-619.

    48. Gorman, D., Bajjouk, T., Populus, J., Vasquez, M. & Ehrhold, A., 2013. Modeling kelp forest distribution and biomass along temperate rocky coastlines. Marine Biology, 160 (2), 309-325.
    49. Gunnill, F., 1985 Population fluctuations of seven macroalgae in southern California during 1981-1983 including effects of severe storms and an El Nino. Journal of Experimental Marine Biology and Ecology, 85, 149-164.

    50. Hall-Spencer, J.M. & Moore, P.G., 2000a. Impact of scallop dredging on maerl grounds. In Effects of fishing on non-target species and habitats. (ed. M.J. Kaiser & S.J., de Groot) 105-117. Oxford: Blackwell Science.
    51. Harder, D.L., Hurd, C.L. & Speck, T., 2006. Comparison of mechanical properties of four large, wave-exposed seaweeds. American Journal of Botany, 93 (10), 1426-1432.

    52. Harker, M., Berkaloff, C., Lemoine, Y., Britton, G., Young, A.J., Duval, J.-C., Rmiki, N.-E. & Rousseau, B., 1999. Effects of high light and desiccation on the operation of the xanthophyll cycle in two marine brown algae. European Journal of Phycology, 34 (1), 35-42.

    53. Hawkins, S.J. & Harkin, E., 1985. Preliminary canopy removal experiments in algal dominated communities low on the shore and in the shallow subtidal on the Isle of Man. Botanica Marina, 28, 223-30.
    54. Hawkins, S.J. & Hartnoll, R.G., 1985. Factors determining the upper limits of intertidal canopy-forming algae. Marine Ecology Progress Series, 20, 265-271.
    55. Heinrich, S., Valentin, K., Frickenhaus, S., John, U. & Wiencke, C., 2012. Transcriptomic analysis of acclimation to temperature and light stress in Saccharina latissima (Phaeophyceae). Plos One, 7 (8), e44342.

    56. Hurd, C.L., 2000. Water motion, marine macroalgal physiology, and production. Journal of Phycology, 36 (3), 453-472.
    57. Isaeus, M., 2004. Factors structuring Fucus communities at open and complex coastlines in the Baltic Sea. Department of Botany, Botaniska institutionen, Stockholm.

    58. Johnston, E., Marzinelli, E., Wood, C., Speranza, D. & Bishop, J., 2011. Bearing the burden of boat harbours: Heavy contaminant and fouling loads in a native habitat-forming alga. Marine Pollution Bulletin, 62 (10), 2137-2144.

    59. Kain, J.M., 1975a. Algal recolonization of some cleared subtidal areas. Journal of Ecology, 63, 739-765.

    60. Kain, J.M., 1979. A view of the genus Laminaria. Oceanography and Marine Biology: an Annual Review, 17, 101-161.
    61. Karsten, U., 2007. Research note: salinity tolerance of Arctic kelps from Spitsbergen. Phycological Research, 55 (4), 257-262.

    62. Kinne, O., 1977. International Helgoland Symposium "Ecosystem research": summary, conclusions and closing. Helgoländer Wissenschaftliche Meeresuntersuchungen, 30(1-4), 709-727.

    63. Kirk, J., 1976. Yellow substance (gelbstoff) and its contribution to the attenuation of photosynthetically active radiation in some inland and coastal south-eastern Australian waters. Marine and Freshwater Research, 27 (1), 61-71.

    64. Kirst, G., 1990. Salinity tolerance of eukaryotic marine algae. Annual review of plant biology, 41 (1), 21-53.

    65. Kirst, G.O. & Wiencke, C., 1995. Ecophysiology of polar algae. Journal of Phycology, 31 (2), 181-199.

    66. Krumhansl, K.A. & Scheibling, R.E., 2011. Detrital production in Nova Scotian kelp beds: patterns and processes. Marine Ecological Progress Series, 421, 67-82.

    67. Krumhansl, K.A. & Scheibling, R.E., 2012. Detrital subsidy from subtidal kelp beds is altered by the invasive green alga Codium fragile ssp fragile. Marine Ecology Progress Series, 456, 73-85.

    68. Krumhansl, K.A. & Scheibling, R.E., 2012. Detrital subsidy from subtidal kelp beds is altered by the invasive green alga Codium fragile ssp fragile. Marine Ecology Progress Series, 456, 73-85.

    69. Krumhansl, K.A., 2012. Detrital production in kelp beds. degree of Doctor of Philosophy, Department of Biology, Dalhousie University, Halifax, Nova Scotia.

    70. Krumhansl, K.A., Lee, J.M. & Scheibling, R.E., 2011. Grazing damage and encrustation by an invasive bryozoan reduce the ability of kelps to withstand breakage by waves. Journal of Experimental Marine Biology and Ecology, 407 (1), 12-18.

    71. Lüning, K., 1990. Seaweeds: their environment, biogeography, and ecophysiology: John Wiley & Sons.

    72. Lüning, K. & Dring, M., 1979. Continuous underwater light measurement near Helgoland (North Sea) and its significance for characteristic light limits in the sublittoral region. Helgoländer Wissenschaftliche Meeresuntersuchungen, 32 (4), 403-424.

    73. Lauzon-Guay, J.-S. & Scheibling, R., 2007. Seasonal variation in movement, aggregation and destructive grazing of the green sea urchin (Strongylocentrotus droebachiensis) in relation to wave action and sea temperature. Marine Biology, 151 (6), 2109-2118.

    74. Lawrence, J.M., 1975. On the relationships between marine plants and sea urchins. Oceanography and Marine Biology: An Annual Review, 13, 213-286.

    75. Lee, J.A. & Brinkhuis, B.H., 1988. Seasonal light and temperature interaction effects on development of Laminaria saccharina (Phaeophyta) gametophytes and juvenile sporophytes. Journal of Phycology, 24, 181-191.
    76. Levin, P.S., Coyer, J.A., Petrik, R. & Good, T.P., 2002. Community-wide effects of nonindigenous species on temperate rocky reefs. Ecology, 83(11), 3182-3193.

    77. Lewis, G.A. & Nichols, D., 1980. Geotactic movement following disturbance in the European sea-urchin, Echinus esculentus (Echinodermata: Echinoidea). Progress in Underwater Science, 5, 171-186.

    78. Ling, S.D., Johnson, C.R., Frusher, S.D. & Ridgeway, K.R., 2009. Overfishing reduces resilience of kelp beds to climate-driven catastrophic phase shift. Proceedings of the National Academy of Sciences USA, 106, 22341-22345.

    79. Lobban, C.S. & Harrison, P.J. (eds.), 1994. Seaweed Ecology and Physiology. Cambridge, uk: Cambridge University Press, pp. 366.

    80. Lüning, K. & Dring, M.J., 1975. Reproduction, growth and photosynthesis of gametophytes of Laminaria saccharina grown in blue and red light. Marine Biology, 29, 195-200.
    81. Lüning, K., 1980. Critical levels of light and temperature regulating the gametogenesis of three laminaria species (Phaeophyceae). Journal of Phycology, 16, 1-15.
    82. Lüning, K., 1988. Photoperiodic control of sorus formation in the brown alga Laminaria saccharina. Marine Ecology Progress Series, 45, 137-144.
    83. Lyngby, J.E. & Mortensen, S.M., 1996. Effects of dredging activities on growth of Laminaria saccharina. Marine Ecology, Publicazioni della Stazione Zoologica di Napoli I, 17, 345-354.
    84. Müller, R., Laepple, T., Bartsch, I. & Wiencke, C., 2009. Impact of oceanic warming on the distribution of seaweeds in polar and cold-temperate waters. Botanica Marina, 52 (6), 617-638.
    85. Markham, J.W. & Munda, I.M., 1980. Algal recolonisation in the rocky eulittoral at Helgoland, Germany. Aquatic Botany, 9, 33-71.
    86. Mikhaylova, T.A., 1999. The initial stages of experimental forming of Laminaria communities in the White Sea. Botanicheskii Zhurnal (St. Petersburg), 84 (3), 56-66.

    87. Molfese, C., Beare, D. & Hall-Spencer, J.M., 2014. Overfishing and the Replacement of Demersal Finfish by Shellfish: An Example from the English Channel. Plos One, 9 (7).

    88. Moy, F., Alve, E., Bogen, J., Christie, H., Green, N., Helland, A., Steen, H., Skarbøvik, E. & Stålnacke, P., 2006. Sugar Kelp Project: Status Report No 1. SFT Report TA-2193/2006, NIVA Report 5265 (in Norwegian, with English Abstract), 36 pp.

    89. Moy, F.E. & Christie, H., 2012. Large-scale shift from sugar kelp (Saccharina latissima) to ephemeral algae along the south and west coast of Norway. Marine Biology Research, 8 (4), 309-321.

    90. Nagel, K., Schneemann, I., Kajahn, I., Labes, A., Wiese, J. & Imhoff, J.F., 2012. Beneficial effects of 2,4-diacetylphloroglucinol-producing pseudomonads on the marine alga Saccharina latissima. Aquatic Microbial Ecology, 67 (3), 239-249.

    91. Netalgae, 2012. Seaweed industry in Europe. (24/04/2014).
    92. Nichols, D., 1984. An investigation of the population dynamics of the common edible sea urchin (Echinus esculentus L.) in relation to species conservation management. Report to Department of the Environment and Nature Conservancy Council from the Department of Biological Sciences, University of Exeter.
    93. Nielsen, M., Krause-Jensen, D., Olesen, B., Thinggaard, R., Christensen, P. & Bruhn, A., 2014a. Growth dynamics of Saccharina latissima (Laminariales, Phaeophyceae) in Aarhus Bay, Denmark, and along the species’ distribution range. Marine Biology, 161 (9), 2011-2022.

    94. Nimura, K., Mizuta, H. & Yamamoto, H., 2002. Critical contents of nitrogen and phosphorus for sorus formation in four Laminaria species. Botanica Marina, 45, 184-188.

    95. Norton, T.A., 1978. The factors influencing the distribution of Saccorhiza polyschides in the region of Lough Ine. Journal of the Marine Biological Association of the United Kingdom, 58, 527-536.
    96. Norton, T.A., 1978. The factors influencing the distribution of Saccorhiza polyschides in the region of Lough Ine. Journal of the Marine Biological Association of the United Kingdom, 58, 527-536.
    97. Norton, T.A., 1978. The factors influencing the distribution of Saccorhiza polyschides in the region of Lough Ine. Journal of the Marine Biological Association of the United Kingdom, 58, 527-536.
    98. O’Brien, J.M., Scheibling, R.E. & Krumhansl, K.A., 2015. Positive feedback between large-scale disturbance and density-dependent grazing decreases resilience of a kelp bed ecosystem. Marine Ecology Progress Series, 522, 1-13.

    99. Oates, B.R., 1985. Photosynthesis and amelioration of desiccation in the intertidal saccate alga Colpornema peregrina. Marine Biology, 89, 109-119.

    100. Oates, B.R., 1986. Components of photosynthesis in the intertidal saccate alga Halosaccion americanum (Rhodophyta, Palmariales). Journal of Phycology, 22, 217-223.

    101. Parke, M., 1948. Studies on British Laminariaceae. I. Growth in Laminaria saccharina (L.) Lamour. Journal of the Marine Biological Association of the United Kingdom, 27, 651-709.
    102. Parker, H., 1981. Influence of relative water motion on the growth, ammonium uptake and carbon and nitrogen composition of Ulva lactuca (Chlorophyta). Marine Biology, 63 (3), 309-318.

    103. Parker, H., 1982. Effects of simulated current on the growth rate and nitrogen metabolism of Gracilaria tikvahiae (Rhodophyta). Marine Biology, 69 (2), 137-145.

    104. Pérez, R., 1971. Écologie, croissance et régénération, teneurs en acide alginique de Laminaria digitata sur les cotes de la Manche. Revue des Travaux de l'Institut des Peches Maritimes, 35, 287-346.
    105. Peteiro, C. & Freire, O., 2013. Biomass yield and morphological features of the seaweed Saccharina latissima cultivated at two different sites in a coastal bay in the Atlantic coast of Spain. Journal of Applied Phycology, 25(1), 205-213.

    106. Peteiro, C., Sánchez, N., Dueñas-Liaño, C. & Martínez, B., 2014. Open-sea cultivation by transplanting young fronds of the kelp Saccharina latissima. Journal of Applied Phycology, 26 (1), 519-528.

    107. Peters, A.F. & Schaffelke, B., 1996. Streblonema (Ectocarpales, Phaeophyceae) infection in the kelp Laminaria saccharina in the western Baltic. Hydrobiologia, 326/327, 111-116.
    108. Reed, D.C., Rassweiler, A. & Arkema, K.K., 2008. Biomass rather than growth rate determines variation in net primary production by giant kelp. Ecology and evolution, 89, 2493-2505

    109. Robins, P.E., Neill, S.P., Giménez, L., Jenkins, S.R. & Malham, S.K., 2013. Physical and biological controls on larval dispersal and connectivity in a highly energetic shelf sea. Limnology and Oceanography, 58(2), 505-524.

    110. Roleda, M.Y. & Dethleff, D., 2011. Storm-generated sediment deposition on rocky shores: Simulating burial effects on the physiology and morphology of Saccharina latissima sporophytes. Marine Biology Research, 7 (3), 213-223.

    111. Roleda, M.Y., Dethleff, D. & Wiencke, C., 2008. Transient sediment load on blades of Arctic Saccharina latissima can mitigate UV radiation effect on photosynthesis. Polar Biology, 31 (6), 765-769.

    112. Saier, B. & Chapman, A.S., 2004. Crusts of the alien bryozoan Membranipora membranacea can negatively impact spore output from native kelps (Laminaria longicruris). Botanica Marina, 47 (4), 265-271.

    113. Sanderson, J., Dring, M., Davidson, K. & Kelly, M., 2012. Culture, yield and bioremediation potential of Palmaria palmata (Linnaeus) Weber & Mohr and Saccharina latissima (Linnaeus) adjacent to fish farm cages in northwest Scotland. Aquaculture, 354, 128-135.

    114. Scheibling, R.E. & Gagnon, P., 2006. Competitive interactions between the invasive green alga Codium fragile ssp tomentosoides and native canopy-forming seaweeds in Nova Scotia (Canada). Marine Ecology Progress Series, 325, 1-14.

    115. Scheibling, R.E., Hennigar, A.W. & Balch, T., 1999. Destructive grazing, epiphytism, and disease: the dynamics of sea urchin-kelp interactions in Nova Scotia. Canadian Journal of Fisheries and Aquatic Sciences, 56 (12), 2300-2314.

    116. Sivertsen, K. & Bjorge, A., 2015. On the brink of the Arctic: Unusual intertidal sub-Arctic kelp associations in the Porsangerfjord, North Norway. Marine Biology Research, 11 (4), 405-413.

    117. Sjøtun, K. & Schoschina, E.V., 2002. Gametophytic development of Laminaria spp. (Laminariales, Phaeophyta) at low temperatures. Phycologia, 41, 147-152.
    118. Smale, D.A. & Wernberg, T., 2013. Extreme climatic event drives range contraction of a habitat-forming species. Proceedings of the Royal Society B-Biological Sciences, 280 (1754).

    119. Smale, D.A., Burrows, M.T., Moore, P., O'Connor, N. & Hawkins, S.J., 2013. Threats and knowledge gaps for ecosystem services provided by kelp forests: a northeast Atlantic perspective. Ecology and evolution, 3 (11), 4016-4038.
    120. Spurkland, T. & Iken, K., 2011a. Salinity and irradiance effects on growth and maximum photosynthetic quantum yield in subarctic Saccharina latissima (Laminariales, Laminariaceae). Botanica Marina, 54, 355-365.

    121. Spurkland, T. & Iken, K., 2011b. Kelp Bed Dynamics in Estuarine Environments in Subarctic Alaska. Journal of Coastal Research, 133-143.

    122. Steneck, R.S., Vavrinec, J. & Leland, A.V., 2004. Accelerating trophic-level dysfunction in kelp forest ecosystems of the western North Atlantic. Ecosystems, 7 (4), 323-332.

    123. Stickle, W.B. & Diehl, W.J., 1987. Effects of salinity on echinoderms. In Echinoderm Studies, Vol. 2 (ed. M. Jangoux & J.M. Lawrence), pp. 235-285. A.A. Balkema: Rotterdam.

    124. Strong, J.A. & Dring, M.J., 2011. Macroalgal competition and invasive success: testing competition in mixed canopies of Sargassum muticum and Saccharina latissima. Botanica Marina, 54 (3), 223-229.

    125. Sundene, O., 1964. The ecology of Laminaria digitata in Norway in view of transplant experiments. Nytt Magasin for Botanik, 11, 83-107.
    126. Ursin, E., 1960. A quantitative investigation of the echinoderm fauna of the central North Sea. Meddelelser fra Danmark Fiskeri-og-Havundersogelser, 2 (24), pp. 204.

    127. Van den Hoek, C. & Donze, M., 1967. Algal phytogeography of the European Atlantic coasts. Blumea, 15 (1), 63-89.
    128. Van den Hoek, C., Mann, D.G. & Jahns, H.M., 1995. Algae: an introduction to phycology: Cambridge University Press.

    129. Wang, X., Broch, O.J., Forbord, S., Handa, A., Skjermo, J., Reitan, K.I., Vadstein, O. & Olsen, Y., 2014. Assimilation of inorganic nutrients from salmon (Salmo salar) farming by the macroalgae (Saccharina latissima) in an exposed coastal environment: implications for integrated multi-trophic aquaculture. Journal of Applied Phycology, 26 (4), 1869-1878.

    130. Weile, K., 1996. Baseline study of Laminaria populations in Øresund. Doc. nr. 95/120/1E. By VRI/Toxicon AB for Øresundskonsortiet

    131. Wernberg, T., Smale, D.A., Tuya, F., Thomsen, M.S., Langlois, T.J., de Bettignies, T., Bennett, S. & Rousseaux, C.S., 2013. An extreme climatic event alters marine ecosystem structure in a global biodiversity hotspot. Nature Climate Change, 3 (1), 78-82.

    132. Wheeler, W.N., 1980. Effect of boundary layer transport on the fixation of carbon by the giant kelp Macrocystis pyrifera. Marine Biology, 56, 103–110.

    133. Wilce, R., 1965. Studies in the genus Laminaria. III. A revision of the north Atlantic species of the Simplices section of Laminaria. Bot. gothoburg., 3, 247-256.

    134. Wolff, W.J., 1968. The Echinodermata of the estuarine region of the rivers Rhine, Meuse and Scheldt, with a list of species occurring in the coastal waters of the Netherlands. The Netherlands Journal of Sea Research, 4, 59-85.
    135. Yarish, C., Penniman, C.A. & Egan, B., 1990. Growth and reproductibe responses of Laminaria longicruris (Laminariales, Phaeophyta) to nutrient enrichment. Hydrobiologia, 204, 505-511.


    This review can be cited as:

    Hiscock, K. 2001. Laminaria saccharina park on very sheltered lower infralittoral 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. Available from:

    Last Updated: 27/11/2001