|Researched by||Dr Harvey Tyler-Walters||Refereed by||Dr Stefan Kraan|
|Authority||(Linnaeus) Greville, 1830|
|Other common names||-||Synonyms||Alaria platyrhiza (Linnaeus) Greville, 1830|
Short cylindrical stipe (exceptionally up to 75 cm) continuing as a distinct midrib throughout the length of the narrow, ribbon-like, slightly wavy blade. Attached to substrate by claw-like holdfast termed haptera. The blade is yellowish, olive-green or rich brown in colour, supple to the touch and very flexible. Blade length varies seasonally but is usually between 30 cm - 1.5 m (exceptionally 4 m) in length. Blade may be tattered and torn by wave action sometimes leaving only the midrib at which point it may be confused with Chorda filum. Older plants may have flat, finger-like sporophylls, each up to 10 cm in length, growing from the stipe at the base of the blade. The sporophylls bear reproductive bodies called sori. When fertile the sori form a typical H-shaped figure on the sporophylls.
Other common names include wing kelp, honeyware, edible fucus, and bladder locks in England; dabberlocks and keys in Scotland; and murlins, ribini, and Cupog nag Cloc in Ireland (Guiry 2000). The species name Alaria esculenta literally means 'edible wings'. This species was originally described as Fucus esculentus Linnaeus, 1767. The class Phaeophyceae may alternatively be classified in the Phylum Heterokontophyta ( Hoek van den et al. 1995).
Alaria (Phaeophyceae, Alariaceae) is a common genus of kelps in the northern hemisphere. Fourteen species are currently recognised of which three (Alaria esculenta (L.) Greville, Alaria pylaii (Bory de Saint-Vincent) Greville, and Alaria grandifolia J. Agardh) are reported for the cold -temperate North Atlantic Ocean. Alaria esculenta, the type species described originally from the North Atlantic, exhibits a range of biogeographically correlated morphotypes suggesting the possibility of multiple specific or intraspecific entities or hybrids (Kraan pers. comm.; Kraan & Guiry 2000 in press). A key to the species of the genus Alaria is given by Widdowson (1971).
- none -
|Phylum||Ochrophyta||Brown and yellow-green seaweeds|
|Authority||(Linnaeus) Greville, 1830|
|Recent Synonyms||Alaria platyrhiza (Linnaeus) Greville, 1830|
|Typical abundance||High density|
|Male size range||Male size at maturity|
|Female size range||Large(>50cm)||Female size at maturity|
|Growth form||Straplike / Ribbonlike||Growth rate||20 cm/month|
|Body flexibility||High (greater than 45 degrees)||Mobility||Sessile|
|Characteristic feeding method||Autotroph|
|Typically feeds on||Not relevant|
|Sociability||Not relevant||Environmental position||Epifloral, Epilithic|
Bryozoa and several epiphytes including Litosiphon laminariae (Kraan pers. comm.).
|Is the species harmful?||No|
|Physiographic preferences||Open coast, Sea loch / Sea lough|
|Biological zone preferences||Lower eulittoral, Sublittoral fringe, Upper infralittoral|
|Substratum / habitat preferences||Artificial (man-made), Bedrock, Cobbles, Large to very large boulders, Pebbles, Small boulders|
|Tidal strength preferences||Moderately Strong 1 to 3 knots (0.5-1.5 m/sec.), Strong 3 to 6 knots (1.5-3 m/sec.), Weak < 1 knot (<0.5 m/sec.)|
|Wave exposure preferences||Exposed, Extremely exposed, Very exposed|
|Salinity preferences||Full (30-40 psu)|
|Depth range||0-8 m|
|Other preferences||-2 °C winter isotherm (as far as sea ice) and up to 16 °C summer isotherm.|
|Migration Pattern||Non-migratory / resident|
Alaria esculenta is present in the North Pacific and North Atlantic, where it is located north as far as the winter sea ice and as far south as the 16 °C summer isotherm, represented by the French coast of Brittany in the European North Atlantic (Luning, 1990). Its absence in the southern North Sea and English Channel is due to high summer surface temperatures of 16 °C, which it cannot survive (Munda & Luning, 1977; Widdowson, 1971; Sundene, 1962). Its distribution in the Arctic Sea is associated with the -2 °C February winter isotherm (Kraan pers. comm.).
|Reproductive type||Alternation of generations||Reproductive frequency||Annual episodic|
|Fecundity (number of eggs)||>1,000,000||Generation time||1 year|
|Age at maturity||8 - 14 months||Season||October - May|
|Life span||5-10 years|
|Larval/propagule type||-||Larval/juvenile development||Spores (sexual / asexual)|
|Duration of larval stage||1 day||Larval dispersal potential||10 -100 m|
|Larval settlement period|
This MarLIN sensitivity assessment has been superseded by the MarESA approach to sensitivity assessment. MarLIN assessments used an approach that has now been modified to reflect the most recent conservation imperatives and terminology and are due to be updated by 2016/17.
Removal of the substratum would entail removal of the plants themselves, germlings and gametophytes. They can not re-attach once removed and would be swept away. Evidence from storm damage and experimental removal of adult Laminaria species in Australia indicates that kelp forest can re-grow within 14 months. These experiments did not remove the gametophyte 'seed' bank. New individuals of Alaria esculenta colonized within 10 m of the parent plants in Norway. However, given the potentially large number of spores and gametophytes it is likely that recolonization would occur rapidly and sporophytes may grow up to 20 cm/month under optimal conditions in the field and up to 10 cm/day in rope culture systems (Birkett et al., 1998b; Kain & Dawes, 1987).
Although smothering of the adult sporophyte may reduce photosynthetic activity it is unlikely to cause damage. However, juvenile sporophytes may be smothered and their growth inhibited. The germlings, zoospores and gametophytes are likely to be intolerant of smothering.
Increased sedimentation may result in smothering of adults (sporophytes), germlings and gametophytes (see above). Increased sediment deposition may increase sediment scour which may damage sporelings in particular. However, the most likely effect of increased siltation will be increased light attenuation and turbidity (see below).
Kelps are normally subtidal species and are likely to be intolerant of desiccation. Alaria esculenta may extend into the lower eulittoral in very exposed conditions. However, these marginal populations have a reduced age range in comparison to the subtidal populations due to the loss of plants resulting from sunshine at low tide. An increase in desiccation by 25% over a period of a year is likely to remove the population.
The lower limit of Alaria esculenta populations is probably controlled by competition from other kelp species such as Laminaria hyperborea, Laminaria digitata and foliose red algae. A decrease in emergence is likely to extend the population up shore. However, an increase is emergence is likely to result in loss of plants at the upper limit of its distribution. Lost individuals are replaced by recruitment. However, the loss of plants will open the canopy and allow slow growing germling to replace adult sporophytes or the maturation of gametophytes.
The presence of Alaria esculenta is associated with strong wave action and tidal flow. Therefore, it is likely to be intolerant of any reduction in wave exposure and flow rate, which is likely to subject it to competition from other Laminarians e.g. Laminaria digitata.
Alaria esculenta germlings tolerate up to 16 °C, above which growth is inhibited (Sundene, 1962). Sundene (1962) reported that Alaria esculenta was only found on shores where the August mean seawater surface temperature is 16 °C or lower (except in areas of extreme exposure). Munda & Lüning (1977) reported that temperatures of 16 -17 °C for a few weeks were lethal to Alaria esculenta sporophytes based on field experiments with specimens transplanted to Helgoland from north Iceland. They suggested that sporophytes were likely to survive above 16 °C for some days, but that longer duration was lethal. Temperature was, therefore, considered to be the main factor controlling its distribution in Europe (Sundene, 1962; Munda & Lüning, 1977; Widdowson, 1971; Lüning, 1990). Birkett et al. (1998b) further suggest that kelp are stenothermal (intolerant of temperature change) and that upper and lower lethal limits of kelp would be between 1-2 °C above or below the normal temperature tolerances. Given its distribution in the North Atlantic and Arctic Sea is associated with the -2 °C February winter isotherm (Kraan pers. comm.). This species is likely to be intolerant of change in temperature equivalent to either benchmark. Its southern distribution is likely to be affected by the position of the mean surface temperature isotherm in Europe and therefore climatic change. The temperature tolerances of the gametophyte stages are different to those of the adult.
The light penetration influences the maximum depth at which kelps species can grow. Dring (1982) reported that laminarians grow at depths at which the light levels are reduced to 1% of incident light at the surface. This varies with the turbidity of the sea water from 100 m in the Mediterranean to only 6-7 m in the silt laden German Bight. Increased turbidity due to coastal engineering, dredging, cooling water plumes been reported to result in the loss of local kelp forest. Suspended material in vicinity of sewage outfalls have been reported to result in reduced the depth range and the fewer new plants under the canopy. However, Alaria esculenta is excluded from deep waters by competition from other kelp species and is unlikely to be light limited (except Rockall populations). Although likely to be intolerant of extreme increase of light attenuation it is unlikely to be affected by a long term change of turbidity of one level on the water clarity scale.
Alaria esculenta is characteristic of wave exposed coasts and favoured by increasing exposure. However, a decrease in wave exposure is likely to subject it to competition with Laminaria spp., which are likely to replace it as the dominant species as exposure decreases
|Tolerant||Not relevant||Not sensitive||High|
Plants have no known sound or vibration receptors
|Tolerant||Not relevant||Not sensitive||High|
Macroalgae are not known to react to the rapid changes in light and shade that would be associated with movement and have no known visual receptors.
Alaria esculenta is adapted to exposed and very wave exposed coasts. The force of wave action in these habitats is significant source of physical abrasion, which is mitigated by the flexibility of the frond. However, abrasion, for instance from a vessel stranding or equivalent disturbance, is likely to snag, damage, and remove fronds. Therefore, an intolerance of intermediate has been recorded. Recoverability is likely to be high.
Alaria esculenta is permanently attached to the substratum. If removed if can not re-attach (except in experimental conditions) and will be lost from the population. However, the population will recover fairly rapidly from the losses due the presence of germlings, large number of spores and gametophytes.
Atrazine was lethal to young sporophytes of Laminaria hyperborea at 1 mg/l and caused growth suppression at 10 µg/l (Hopkin & Kain, 1978). Mixed detergents, herbicides (dalapon 2,4-D) were not toxic at the levels tested. Cole et al. (1999) report the following as very toxic to macrophytes: atrazine; simazine; diuron; and linuron (herbicides). It is likely therefore that Laminariales such as Alaria esculenta are intolerant of atrazine and some other herbicides. PCBs inhibited growth, gametogenesis and sporophyte recruitment in Macrocystis pyrifera at 5 µg/litre and, therefore, may have similar sub-lethal effects on Alaria esculenta
Mercury and copper were found to be lethal at 50 µg/l and 100 µg/l respectively and toxic at 50 and 10 µg/ l respectively in Laminaria hyperborea. Zinc and cadmium were lethal at 5 mg/l and 10 mg/l respectively. Therefore, it is likely that Laminariales such as Alaria esculenta are intolerant of copper at benchmark level and so intolerance is assessed as intermediate. Zinc, cadmium and mercury especially are likely to cause sub-lethal effects.
Mucilaginous slime coating on kelp fronds is thought to protect them from coatings of oil. Hydrocarbons in solution reduce photosynthesis and may be algicidal. Reduction in photosynthesis depends on the type of oil, its concentration, length of exposure, method used to prepare oil-water mixture and irradiance in experimental trials (Lobban & Harrison, 1994). The sublittoral fringe and lower eulittoral populations of Alaria esculenta would be most vulnerable to oiling. Subtidal populations being only exposed to oil emulsions or oil adsorbed to particles. Kelps are relatively insensitive to dispersants (Birkett et al. 1998). Three days exposure to 1% diesel emulsion reduced photosynthesis completely in young Macrocytsis plants. Laminaria digitata exposed to diesel oil at 130 µg/litre reduced growth by 50% in a two year experiment. No growth inhibition was noted at 30 µg/l and the plants recovered completed in oil-free conditions. Overall, laminarians such as Alaria esculenta are probably relatively tolerant of oiling and an intolerance of low has been recorded.
|No information||No information||No information||Not relevant|
No information found.
All kelp species are efficient absorbers of nutrients (nitrates and phosphates) and can take up and store excess nutrients. In exposed sites the turnover of fresh seawater suggests that nutrients are continuously replenished. Eutrophication is associated with loss of perennial macrophytes, a reduction in the depth range and replacement by mussels or opportunistic algae species (Fletcher, 1996; Birkett et al., 1998). Increased nutrients may increase growth of epiphytes and plankton, resulting in reduced light penetration for photosynthesis and a subsequent reduction in the depth at which kelp could grow. However, nutrients are often added to macrophyte cultures to increase productivity. Therefore a rank of intermediate intolerance has been given to represent the likely indirect effects on turbidity and competition.
Kelps are found in full salinity and thought to be stenohaline, i.e. intolerant of salinity change. Alaria esculenta sporophytes grew poorly at salinities below 25 psu (Sundene 1961). Although its may penetrate into the lower eulittoral in is probably highly intolerant of long term reductions in salinity (Birkett et al., 1998).
|No information||Not relevant||No information||Not relevant|
No information was found concerning on the effects of de-oxygenation in macrophytes.
Galls on the blade of Laminaria hyperborea and spot disease are associated with the endophyte Streblonema sp. although the causal agent is unknown (bacteria, virus or endophyte). Resultant damage to the blade and stipe may increase losses in storms. The endophyte inhibits spore production and therefore recruitment and recoverability. Streblonema sp. has been reported growing on Alaria esculenta but no further information was found (Lein et al., 1991).
The Japanese kelp Undaria pinnatifida (wakame) has recently spread to the south coast of England from Brittany where it was introduced for aquaculture. It is presently restricted to man made structures but could spread in the ballast water of commercial or recreational boats and shipping. Its potential competition with other kelps in the UK, including Alaria esculenta requires further study (Birkett et al., 1998).
There is considerable material on the effects of harvesting kelp species (Birkett et al. 1998; Guiry & Blunden, 1991) but little evidence concerning the effects of harvesting on Alaria esculenta populations. However, evidence from other kelp species suggest that the macrophytes can recover within 3-4 years, although the effects on the rest of the community is poorly studied. In canopy clearance experiments, Alaria esculenta may appear early in the succession suggesting that it would recover more rapidly.
Extraction of sea urchin predators such as lobsters and sea otters in Nova Scotia was suggested as a possible cause of catastrophic sea urchin infestation and loss of kelp forest. Removal of grazing species may reduce competition with urchins leading to a increase in their population and subsequent loss of kelp species. However, present evidence indicates that periodic good recruitment to the urchin population may have a greater effect. It seem likely that each of these factors influences urchin populations to different extents in different areas. These effects are poorly studied in UK populations of kelp. An increased urchin or other grazer population may increase time taken for the population to recover, since they are likely to remove young sporophytes.
The ecophysiology and chemical tolerances of Alaria esculenta is poorly studied. Most of the information presented here is based on work on other Laminariales. The effects of pollutants on macrophytes, including Laminariales, is reviewed by Loban and Harrison (1997).
- no data -
|National (GB) importance||-||Global red list (IUCN) category||-|
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.). Available from: http://www.ukmarinesac.org.uk/publications.htm
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.
Dieck, T.I., 1993. Temperature tolerance and survival in darkness of kelp gametophytes (Laminariales: Phaeophyta) - ecological and biogeographical implications. Marine Ecology Progress Series, 100, 253-264.
Dring, M.J., 1982. The Biology of Marine Plants. London: Edward Arnold.
Druehl, L., 1988. Laminaria aquaculture in British Columbia. In Proceedings of the Fourth Alaska Aquaculture Conference, November 18-21, 1987, Sitka, Alaska USA, (Alaska Sea Grant Report, No. 88-4.), pp. 3-10.
Fish, J.D. & Fish, S., 1996. A student's guide to the seashore. Cambridge: Cambridge University Press.
Fletcher, R.L., 1996. The occurrence of 'green tides' - a review. In Marine Benthic Vegetation. Recent changes and the Effects of Eutrophication (ed. W. Schramm & P.H. Nienhuis). Berlin Heidelberg: Springer-Verlag. [Ecological Studies, vol. 123].
Guiry, M.D. & Blunden, G., 1991. Seaweed Resources in Europe: Uses and Potential. Chicester: John Wiley & Sons.
Guiry, M.D. & Hession, C., 1996. Eat up your seaweed. Ireland of the Welcomes, 45, 22-25.
Guiry, M.D. & Nic Dhonncha, E., 2000. AlgaeBase. World Wide Web electronic publication http://www.algaebase.org, 2000-01-01
Guiry, M.D., 1997. Research and development of a sustainable Irish seaweed industry. Occasional Papers in Irish Science and Technology, No. 14, Went Memorial Lecture 1996., Royal Dublin Society, Dublin, 11pp.
Hardy, F.G. & Guiry, M.D., 2003. A check-list and atlas of the seaweeds of Britain and Ireland. London: British Phycological Society
Hayward, P., Nelson-Smith, T. & Shields, C. 1996. Collins pocket guide. Sea shore of Britain and northern Europe. London: HarperCollins.
Hiscock, S., 1979. A field key to the British brown seaweeds (Phaeophyta). Field Studies, 5, 1- 44.
Hopkin, R. & Kain, J.M., 1978. The effects of some pollutants on the survival, growth and respiration of Laminaria hyperborea. Estuarine and Coastal Marine Science, 7, 531-553.
Indergaard, M. & Minsaas, J., 1991. Animal and Human Nutrition. In Seaweed Resources in Europe: Uses and Potential, (Ed. M.D. Guiry & G. Blunden). Chichester: John Wiley & Sons.
JNCC (Joint Nature Conservation Committee), 1999. Marine Environment Resource Mapping And Information Database (MERMAID): Marine Nature Conservation Review Survey Database. [on-line] http://www.jncc.gov.uk/mermaid
Kain, J.M. & Dawes, C.P., 1987. Useful European seaweeds: past hopes and present cultivation. Hydrobiologia, 151/152, 173-181.
Kain, J.M., Holt, T.J. & Dawes, C.P., 1990. European Laminariales and their cultivation. In Economically important marine plants of the Atlantic: their biology and cultivation, (ed. C. Yarish, C.A. Penniman and P. van Patten). Connecticut Sea Grant College Program. Groton.
Kraan S. & Guiry M.D., 2001. Are North Atlantic Alaria esculenta and A. grandifolia (Alariaceae, Phaeophyceae) conspecific? European Journal of Phycology, 36(1), 35-42
Lein, T.E, Sjotun, K. & Wakili, S., 1991. Mass - occurrence of a brown filamentous endophyte in the lamina of the kelp Laminaria hyperborea (Gunnerus) Foslie along the south western coast of Norway Sarsia, 76, 187-193.
Levring, T., Hoppe, H.A. & Schmid, O.J., 1969. Marine Algae: a survey of research and utilization. Hamburg: Cram, de Gruyter & Co. [Botanica Marina Handbooks, Vol. 1.]
Lewallen, E. & Lewallen, J., 1996. Sea vegetable gourmet cookbook and wildcrafter's guide. California: Mendocino Sea Vegetable Company.
Lobban, C.S. & Harrison, P.J., 1997. Seaweed ecology and physiology. Cambridge: Cambridge University Press.
Madlener, J.C., 1977. The sea vegetable book. New York: Clarkson N. Potter.
Mai, K.S., Mercer, J.P. & Donlon, J., 1996. Comparative studies of on the nutrition of 2 species of Abalone, Haliotis tuberculata and Haliotis discus Hanni-Ino. 5. The role of poly-unsaturated fatty acids in macroalgae in Abalone nutrition. Aquaculture, 139, 77-89.
Munda, I.M. & Luning, K., 1977. Growth performance of Alaria esculenta off Helgoland. Helgolander Wissenschaftliche Meeresuntersuchungen, 29, 311-314.
Newton, L., 1931. A handbook of the British seaweeds. London: British Museum (Natural History).
Nisizawa, K., Noda, H., Kikuchi, R., Watanabe, T., 1987. The main seaweed foods in Japan. Hydrobiologia, 151/152, 5-29.
Norton, T.A., 1992. Dispersal by macroalgae. British Phycological Journal, 27, 293-301.
Picton, B.E. & Costello, M.J., 1998. BioMar biotope viewer: a guide to marine habitats, fauna and flora of Britain and Ireland. [CD-ROM] Environmental Sciences Unit, Trinity College, Dublin., http://www.itsligo.ie/biomar/
Stein, F., Sjotun, K., Lein, T.E. & Rueness, J., 1995. Spore dispersal in Laminaria hyperborea (Laminariales, Phaeophyceae) Sarsia, 80, 47-53.
Sundene, O., 1962. The implications of transplant and culture experiments on the growth and distribution of Alaria esculenta. Nytt Magasin for Botanik, 9, 155-174.
Widdowson, T.B., 1971. A taxonomic revision of the genus Alaria Greville. Syesis, 4, 11-49.
Yamanaka, R. & Akiyama, K., 1993. Cultivation and utilization of Undaria pinnatifida (wakame) as food. Journal of Applied Phycology, 5, 249-253.
This review can be cited as:
Last Updated: 29/05/2008