Coralline crust-dominated shallow eulittoral rockpools

13-11-2002
Researched byGeorgina Budd Refereed byThis information is not refereed.
EUNIS CodeA1.411 EUNIS NameCoralline crust-dominated shallow eulittoral rockpools

Summary

UK and Ireland classification

EUNIS 2008A1.411Coralline crust-dominated shallow eulittoral rockpools
EUNIS 2006A1.411Coralline crust-dominated shallow eulittoral rockpools
JNCC 2004LR.FLR.Rkp.CorCoralline crust-dominated shallow eulittoral rockpools
1997 BiotopeLR.LR.Rkp.CorCorallina officinalis and coralline crusts in shallow eulittoral rockpools

Description

Shallow rockpools throughout the eulittoral zone may be characterized by a covering of encrusting coralline algae on which Corallina officinalis often forms a dense turf. These 'coralline' pools have a striking appearance as they are dominated predominantly by red algae. Filamentous and foliose red algae found in these pools include Dumontia contorta, Mastocarpus stellatus and Ceramium rubrum [now Ceramium virgatum]. The green algae Cladophora rupestris and Ulva spp. can also occur. The pools may hold large numbers of grazing molluscs, particularly Littorina littorea (which often occurs in exceptionally high densities in upper shore pools), Patella vulgata and Gibbula cineraria. Gastropods may graze these pools to such an extent that they are devoid of any foliose red algae, and are reduced to encrusting coralline algae and large numbers of gastropods. Large brown algae are generally absent (compare with LR.FK), although small Halidrys siliquosa may be present. Within the pools, pits and crevices are often occupied by the anemone Actinia equina and small Mytilus edulis. Similar sized pools in the littoral fringe generally lack the encrusting coralline algae and are characterized by green algae (see LR.G). In Ireland, the sea urchin Paracentrotus lividus can dominate these shallow coralline pools (see LR.Cor.Par - Coralline crusts and Paracentrotus lividus in shallow eulittoral rockpools). In south-west Britain, the brown alga Bifurcaria bifurcata (LR.Cor.Bif - Bifurcaria bifurcata in shallow eulittoral rockpools) or Cystoseira spp. (LR.Cor.Cys - Cystoseira spp. in shallow eulittoral rockpools) can be regionally dominant. (Information taken from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b).

Recorded distribution in Britain and Ireland

Found on rocky shores on all coasts of Britain and Ireland.

Depth range

Mid shore, Upper shore

Additional information

In the extreme south west of Britain, another coralline algae, Jania rubens, which is similar to Corallina officinalis, may occur in large amounts, almost certainly within this biotope.

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Habitat review

Ecology

Ecological and functional relationships

The coralline algae are the dominant species in this biotope. To a great extent the rockpool biotope is an upward extension of ELR.Coff, although the rockpool biotope has its own characteristics.
  • Corallina officinalis and various lithothamnia are successful in the upper half of the eulittoral zone, especially in shallow, well-lit rockpools (Lewis, 1964). In this zone, some limitation on species develops and not all lower littoral species of the open rock surface can colonize upper shore in rockpools. For example, Fucus serratus can do so but Laurencia pinnatifida, Lomentaria articulata and Rhodymenia become much less plentiful, almost to the point of exclusion (Lewis, 1964).
  • Other filamentous and foliose red algae found in the pools include Dumontia contorta, Mastocarpus stellatus, Ceramium nodulosum and Chaetomorpha, Ectocarpus, Polysiphonia and Scytosiphon species. The green seaweeds Cladophora rupestris, Ulva spp. and Ulva lactuca can also occur in high abundance.
  • Seaweeds provide primary productivity either directly to grazing fish and invertebrates or indirectly, to detritivores and decomposers, in the form of detritus and drift algae or as dissolved organic material and other exudates.
The faunal communities of coralline turfs are described in detail by Hagerman (1968), Dommasnes (1968, 1969), Hicks (1985), Grahame & Hanna (1989), Crisp & Mwaiseje (1989), Bamber (1988) and Bamber & Irving (1993). (see ELR.Coff for details).
  • Corallina officinalis provides substratum for spirorbid worms (e.g. Spirorbis corallinae), epiphytes, periphyton, microflora (e.g. bacteria, blue green algae, diatoms and juvenile larger algae), and interstices between the fronds provide refuges from predation for a variety of small invertebrates.
  • Amphipods (e.g. Parajassa pelagica and Stenothoe monoculoides), isopods (e.g. Idotea pelagica and Jaera albifrons) and other mesoherbivores graze the epiphytic flora and senescent macroalgal tissue, which may benefit the macroalgal host, and may facilitate dispersal of the propagules of some macroalgal species (Brawley, 1992b; Williams & Seed, 1992). Mesoherbivores may also graze the macroalgae but do not normally adversely affect the canopy (Brawley, 1992b). Grazing is likely to be advantageous to encrusting corallines owing to the removal of epiphytes.
  • Foliose seaweeds are grazed by large numbers of molluscs, especially the winkle, Littorina littorea, the limpet, Patella vulgata and top shell, Gibbula cineraria. Littorinids show definite preferences for particular algal foods. Littorina littorea tends to prefer the green algae such as Ulva to perennial red algae (Little & Kitching, 1996). Thin filamentous or membranous seaweeds, such as Ulva, Ceramium and Polysiphonia, are likely to more edible than tougher leathery forms. Some red seaweeds such as Corallina officinalis and coralline crusts (Lithothamnion, Lithophyllum) protect their thalli with a coating of calcium carbonate and are probably relatively grazing resistant (Littler & Kauker, 1984). Ephemeral algal species may be able to escape herbivory in time and space, owing to the fact that they are less predictable for herbivores, occurring at different times and in different places, usually as a result of disturbance (Raffaelli & Hawkins, 1999). The chiton, Lepidochitona cinerea probably grazes the corallines directly.
  • Grazers of periphyton (bacteria, blue-green algae and diatoms) or epiphytic algae include harpacticoid copepods, small gastropods (e.g. Rissoa spp. and Littorina neglecta.
  • Within the pools, pits and crevices are likely to be occupied by the beadlet anemone, Actinia equina and small mussels, Mytilus edulis. The food of anemones consists of a wide variety of crustaceans, molluscs, worms, other invertebrates and even fishes, caught using nematocysts borne on its tentacles.
  • The barnacle Semibalanus balanoides may be found over the rock surface. It and small mussels, are preyed upon by the whelk, Nucella lapillus.

Seasonal and longer term change

As communities in rockpools remain constantly submerged and the danger of desiccation is absent, it might be expected that rockpools form an easier environment in which to live for marine life than drying rock surfaces, and that species from regions lower on the shore would be able to extend much further up the shore. However, much of the lower shore open rock fauna is absent from rockpools. Rockpools constitute a distinct environment for which physiological adaptations by the flora and fauna may be required (Lewis, 1964). Conditions within rockpools are the consequence of prolonged separation from the main body of the sea, and physico-chemical parameters within them fluctuate dramatically (Huggett & Griffiths, 1986). In general, larger and deep rockpools low on the shore tend to correspond to the sublittoral habitat with a more stable temperature and salinity regime. In contrast, small and shallow pools higher on the shore are especially influenced by insolation, air temperature and rainfall, the effects of which become more significant towards the high shore, where pools may be isolated from the sea for a number of days or weeks (Lewis, 1964).
  • Weather conditions exert a considerable influence on temperature and salinity. Water temperature in pools follows the temperature of the air more closely than that of the sea. In summer, shallow pools or the surface waters of deeper pools are warmer by day, but may be colder at night, and in winter may be much colder than the sea (Pyefinch, 1943). In deeper pools, the vertical temperature gradation usually present in summer reverses during winter owing to density stratification, so that ice may form (Naylor & Slinn, 1958).
  • High air temperatures cause surface evaporation of water from pools, so that salinity steadily increases, especially in pools not flooded by the tide for several days. Alternatively, high rainfall will reduce pool salinity or create a surface layer of brackish/nearly fresh water for a period. The extent of temperature and salinity change is affected by the frequency and time of day at which tidal inundation occurs. If high tide occurs in early morning and evening the diurnal temperature follows that of the air, whilst high water at midday suddenly returns the temperature to that of the sea (Pyefinch, 1943). Heavy rainfall, followed by tidal inundation can cause dramatic fluctuations in salinity, and values ranging from 5-30 psu have been recorded in rockpools over a period of 24 hrs (Ranade, 1957). Rockpools in the supralittoral, littoral fringe and upper eulittoral are liable to gradually changing salinities followed by days of fully marine or fluctuating salinity at times of spring tide (Lewis, 1964).
  • Other physico-chemical parameters in rockpools demonstrate temporal change. The biological community directly affects oxygen concentration, carbon dioxide concentration and pH, and are themselves affected by changes in the chemical parameters. Throughout the day, algae photosynthesize and produce oxygen, the concentration of which may rise to three times its saturation value, so that bubbles are released. During photosynthesis algae absorb carbon dioxide and as concentrations fall, the pH rises. Morris & Taylor (1983) recorded pH values >9 in rockpools on the Isle of Cumbrae. At night changes occur in the opposite direction a respiration utilizes much of the available oxygen and pH decreases.
Corallina officinalis may be overgrown by epiphytes, especially during summer. This overgrowth regularly leads to high mortality of fronds due to light reduction (Wiedemann, pers. comm.). The ephemeral green seaweeds Ulva intestinalis and Ulva lactuca are likely to be more abundant in summer. In summer, corallines may be bleached and loose their pink pigment but in some species, e.g. Phymatolithon, this does not necessarily result in death of the plant and pigment may be re-synthesized (Little & Kitching, 1996).

Habitat structure and complexity

Bedrock forms the substratum of the biotope. Rockpools vary greatly in their physical features. Pools may be shallow and well-lit or deep and shaded with overhanging sides and vertical surfaces. Algae growing within provide additional surfaces for colonization and there is also a tendency for loose substrata (sand, stones, rocks) to accumulate in pools, the instability of which may affect species diversity. Within rockpools, crevices and pits may be found and exploited by species such the mussel Mytilus edulis and the beadlet anemone, Actinia equina, while the underside of stones and boulders support underboulder communities (see MLR.Fser.Fser.Bo for example).

Productivity

Little information concerning the productivity of coralline turf communities was found. The red algae, algal epiphytes and periphyton provide primary productivity to grazers, while their spores and phytoplankton provide primary productivity to suspension feeders. For instance, spore production by the encrusting 'coralline' algae, Lithophyllum incrustans may be up to 18 million m²/yr (Edyvean & Ford, 1986).

Secondary productivity of the invertebrate fauna may be high and coralline turf may support high abundances of invertebrates. For example, Choat & Kingett (1982) recorded the following numbers of epiphytic fauna: amphipods 1038 / 0.01m²; ostracods 219 / 0.01m².

Recruitment processes

Recruitment processes of some of the characterizing species of the biotope are given below:
  • Corallina officinalis has isomorphic sexual (gametophyte) and asexual (sporophyte) stages (see MarLIN review). Settled tetraspores develop into a perennial crustose base, from which the upright, articulate fronds develop. Sporelings formed within 48 hrs, a crustose base within 72 hrs, fronds being initiated after 3 weeks and the first intergeniculum (segment) formed within 13 weeks of settlement (Jones & Moorjani, 1973). Settlement and development of fronds is optimal on rough surfaces but settlement can occur on smooth surfaces (Harlin & Lindbergh, 1977; Wiedeman pers. comm.). Corallina officinalis settled on artificial substrata within 1 week of their placement in the intertidal in New England summer suggesting that recruitment is high (Harlin & Lindbergh, 1977).
  • Besides having a meristem, Lithophyllum incrustans has its conceptacles (reproductive organs) buried in its calcified thallus, and connected to the exterior by canals (Edyvean & Ford, 1986). Reproductive types (gametangial and tetrasporangial plants) occur from October to April but decline into summer although some conceptacles are present throughout the year (Irvine & Chamberlain 1994). It has been calculated that 1 mm x 1mm of reproductive thallus produces 17.5 million bispores per year with average settlement of only 55 sporelings/year (Edyvean & Ford 1984).
  • All the spores of red algae are non flagellate and dispersal is wholly a passive process (Fletcher & Callow, 1992). Spores vary in their sinking rate as determined by size and density. In general, due to the difficulties of re-entering the benthic boundary layer, it is likely that successful colonization is achieved under conditions of limited dispersal and/or minimum water current activity. Norton (1992) reported that although spores may travel long distances the reach of the furthest propagule does not equal its useful dispersal range, and most successful recruitment probably occurs within 10m of the parent plants. It is expected, therefore, that recruitment of foliose macroalgae in the biotope would occur from local populations and that establishment and recovery of isolated populations would be patchy and sporadic.
  • Littorina littorea is an iteroparous breeder with high fecundity (up to 100,000 for a large female (27 mm shell height)) that lives for several (at least 4) years. Littorina littorea can breed throughout the year but the length and timing of the breeding period are extremely dependent on climatic conditions. Littorina littorea sheds egg capsules directly into the sea and release is synchronized with spring tides on several separate occasions. Larval settling time or pelagic phase can be up to six weeks (Fish, 1972).
  • Recruitment of Patella vulgata fluctuates from year to year and from place to place (Bowman, 1981). Fertilization is external and the larvae are pelagic for up to two weeks before settling on rock at a shell length of about 0.2 mm. Winter breeding occurs only in southern England, in the north of Scotland it breeds in August and in north-east England in September.
  • Gammarid amphipods brood their embryos and offspring but are highly mobile as adults and probably capable of colonizing new habitats from the surrounding area (e.g. see Hyale prevostii review). Similarly isopods such as Idotea species and Jaera species brood their young. Idotea species are mobile and active swimmers and probably capable of recruiting to new habitats from the surrounding area by adult migration. Jaera albifrons, however, is small and may take longer to move between habitats and Carvalho (1989) suggested that under normal circumstances movement was probably limited to an area of less than 2 m. Hicks (1985) noted that epiphytic harpacticoid copepods lack planktonic dispersive larval stages but are active swimmers, which is therefore the primary mechanism for dispersal and colonization of available habitats. Some species of harpacticoids are capable to moving between low and mid-water levels on the shore with the tide, while in others, colonization rates decrease with increasing distance from the resident population. Overall immigration and in situ reproduction were thought to maintain equilibrium populations exposed to local extinction, although there may be local spatial variation in abundance (see Hicks, 1985).

Time for community to reach maturity

The epiphytic species diversity of the coralline turf in the rockpool is dependant on the Corallina officinalis cover and its growth form (Dommasnes, 1968, 1969; Seapy & Littler, 1982; Crisp & Mwaiseje, 1989). Corallina officinalis was shown to settle on artificial substances within 1 week of their placement in the intertidal in New England summer suggesting that recruitment is high (Harlin & Lindbergh, 1977). New fronds of Corallina officinalis appeared on sterilized plots within six months and 10% cover was reached with 12 months (Littler & Kauker, 1984). In experimental plots, up to 15% cover of Corallina officinalis fronds returned within 3 months after removal of fronds and all other epiflora/fauna (Littler & Kauker, 1984). Bamber & Irving (1993) reported that new plants grew back in scraped transects within 12 months, although the resistant crustose bases were probably not removed. New crustose bases may recruit and develop quickly but the formation of new fronds from these bases and recovery of original cover may take longer. Once a coralline turf has developed it will probably be colonized by epiphytic invertebrates such as harpacticoids, amphipods and isopods relatively quickly from the surrounding area. Therefore, the biotope would be recognizable once the coralline turf has regrown, which is likely to be quite rapid if the resistant crustose bases remain. Recruitment of red algae probably equally rapid, and once the algal turf has developed most of the epiphytic invertebrates would colonize quickly, although some species e.g. small brooding gastropods would take longer.

Additional information

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Preferences & Distribution

Recorded distribution in Britain and IrelandFound on rocky shores on all coasts of Britain and Ireland.

Habitat preferences

Depth Range Mid shore, Upper shore
Water clarity preferences
Limiting Nutrients Nitrogen (nitrates), Phosphorus (phosphates)
Salinity Full (30-40 psu)
Physiographic
Biological Zone Eulittoral, Upper eulittoral
Substratum Bedrock, Large to very large boulders, Small boulders
Tidal Weak < 1 knot (<0.5 m/sec.)
Wave Exposed, Moderately exposed, Sheltered, Very exposed
Other preferences Rockpools

Additional Information

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Species composition

Species found especially in this biotope

Rare or scarce species associated with this biotope

-

Additional information

The MNCR recorded ca 577 species in 176 records of this biotope, although not all species occurred in all records of the biotope (Connor et al., 1997b; JNCC, 1999).

Sensitivity reviewHow is sensitivity assessed?

Explanation

Corallina officinalis is the dominant characterizing species within the biotope and provides substratum and refuges for a diverse epifauna. Therefore, the community is dependant on the presence of Corallina officinalis and it has been included as key structuring. Encrusting coralline algae such as Lithophyllum incrustans are characteristic of the biotope. Gastropods Littorina littorea, Patella vulgata and Gibbula cineraria may graze pools in the eulittoral zone to such an extent that they are devoid of any foliose red algae and are reduced to encrusting coralline algae. They have been included as important structural species. Epiphytic grazers, such as amphipods, isopods small gastropods probably keep the turf free of epiphytic algae and are important structural species. Reference has been made to reviews of Hyale prevostii to represent the sensitivity of amphipods and small crustaceans.

Species indicative of sensitivity

Community ImportanceSpecies nameCommon Name
Key structuralCorallina officinalisCoral weed
Important structuralGibbula cinerariaGrey top shell
Important structuralHyale prevostiiAn amphipod
Important characterizingLithophyllum incrustansEncrusting coralline algae
Important structuralLittorina littoreaCommon periwinkle
Important structuralPatella vulgataCommon limpet

Physical Pressures

 IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
High High Moderate Major decline High
Removal of the substratum would result in loss of the coralline turf and its associated community. Therefore an intolerance of high has been recorded. Recoverability is likely to be high (see additional information below).
Intermediate High Low Decline Moderate
Corallina spp. accumulate more sediment than any other alga (Hicks, 1985). Significant sediment cover of the middle to lower intertidal in a South Californian shore, resulting from fresh water runoff, caused substantial decline in Corallina spp. cover as the fronds died back (Seapy & Littler, 1982). Although the fronds of Corallina officinalis may be intolerant of, rapid recovery will result from the resistant crustose bases (see additional information below). Encrusting coralline algae are frequently subject to cover by sediment and appear to survive well.
Faunal components of the biotope may be more intolerant of smothering than algae. If Littorina littorea cannot regain the surface then death may occur. Smothering of limpets by 5cm of sediment for one month is likely to interfere with locomotion, grazing and respiration. Mobile epifauna, such as amphipods and isopods are likely to be able to move through the sediment or debris to escape smothering. The species food source may be affected because smothering can reduce photosynthesis but Hyale prevostii for instance may migrate to alternative food sources on other parts of the shore. At the benchmark, level intolerance to smothering has been assessed to be intermediate as important structural species may be lost. Recoverability has been assessed to be high. For instance, Littorina littorea is an iteroparous breeder with high fecundity that lives for several (at least 4) years. Breeding can occur throughout the year. The planktonic larval stage is long (up to 6 weeks) although larvae do tend to remain in waters close to the shore so recruitment rates should be good.
Intermediate Very high Low Minor decline Low
Corallina species accumulate more sediment than any other alga (Hicks, 1985). Hence an increase in suspended sediment is likely to accumulate in the coralline turf. A significant increase may result in smothering (see above). An accumulation of sediment within the turf may attract more sediment dwelling interstitial invertebrates such as nematodes, harpacticoids and polychaetes, although in more wave exposed locations accumulation of sediment is likely to be minimal. Increased suspended sediment may also result in increased scour, which may adversely affect the fleshy red algae, and interfere with settling spores and recruitment if the factor is coincident with their major reproductive period. However, coralline algae, especially the crustose forms are thought to be resistant of sediment scour (Littler & Kauker, 1984), and will probably not be adversely affected at the benchmark level. Therefore, an increase in suspended sediment may reduce the epiphytic species diversity in the immediacy, and adversely affect the cover of fleshy red algae and an intolerance of intermediate has been recorded. Recoverability is likely to be very high as species are likely to remain in situ from which recruitment can occur.
Tolerant Not sensitive* No change Moderate
This community is unlikely to be dependant on suspended sediment. Although accumulated sediment within coralline turf habitats is likely to increase the species diversity of the epiphytic fauna, in very wave exposed locations, accumulated sediment in the habitat is likely to be minimal. A reduction in suspended sediment will probably reduce the risk of scour, and reduce food availability for the few suspension feeding species in the biotope (e.g. barnacles and spirorbids present). Therefore not sensitive has been recorded.
High Very high Low Decline Low
Corallina officinalis inhabits damp or wet gullies and rock pools and does not inhabit open rock in the upper shore. Fronds are highly intolerant of desiccation and do not recover from a 15%water loss, which might occur within 40-45 minutes during a spring tide in summer (Wiedemann, 1994). Occurrence of encrusting coralline algae seems to be critically determined by exposure to air and sunlight. Colonies survive in damp conditions under algal canopies or in pools but not on open rock where desiccation effects are important. An intolerance assessment of high has been made owing to the fact that should a rockpool community dominated by coralline algae 'loose its water' and be subject to continuous emersion for an hour and hence desiccation it is likely that the key structuring species will suffer loss. Mobile epifauna are likely to seek shelter elsewhere and species such as Patella vulgata found throughout the intertidal zone are likely to tolerate a desiccation event to some extent. On return to prior conditions, recoverability of the key structuring species has been assessed to be very high (see additional information below).
Intermediate Very high Low Decline Low
Bleached corallines were observed 15 months after the 1964 Alaska earthquake which elevated areas in Prince William Sound by 10 m. Similarly, increased exposure to air caused by upward movement of 15 cm due to nuclear tests at Armchitka Island, Alaska adversely affected Corallina pilulifera (Johansen, 1974). However, the upper extent of LR.Coff is determined by the availability of rock pools and wet gullies. An increase in emergence and concomitant increase in desiccation on the open shore is of lesser importance where moisture is provided by standing water and shade. An increase in emergence will, however, expose the rockpool to increased evaporation, rainfall, and heating or cooling, so that some adverse effects on the fauna and flora are likely. Therefore, an intolerance of intermediate has been recorded. Recovery of the coralline algal community is likely to be very high (see additional information below).
High High Intermediate Rise Low
A greater period of immersion will lessen the physico-chemical extremes that prevent the colonization of the mid to upper shore by species more typical of the lower shore. Desiccation stress will be lessened and temperature and salinity changes less severe. Thus the LR.Cor community has been assessed to have a high intolerance to decreased emergence as it will probably allow the 'up-lift' of lower shore species into the biotope and will it begin to change to another biotope. On return to prior conditions, it is likely that species which entered the biotope would be lost owing to intolerance, and recoverability has been assessed to be high to indicate that colonizing species may persist for a period but disappear as the biotope community re-stabilizes.
Not relevant Not relevant Not relevant Not relevant Not relevant
Water flow rate in this biotope is typically only that of the ebb and flood tide speed, which hardly affects intertidal habitats and is far exceeded by the strength of wave action. An increase in water flow rate is therefore considered not relevant.
Low Very high Moderate Minor decline Very low
Wave action is far more important than tidal flow so not relevant has been recorded.
Intermediate Very high Low Decline Low
In general, the water temperature of rockpools follows that of the air more closely than that of the sea and throughout any 24 hour period dramatic changes in temperature may be observed. For instance, Pyefinch (1943) plotted diurnal changes in a pool lying above mean high water during July. When the pool was out of contact with the sea, water temperature increased by 5 °C from 14 to 19 °C over a three hour period, and decreased suddenly to 14 °C within 1.5 hours when the incoming tide reached it. Hence, the community that inhabits such environments needs to be especially tolerant of acute temperature changes. Mobile species (e.g. littorinids, isopods and intertidal fish) will exhibit physiological and behavioural adaptations to temperature change and will probably migrate to deeper water within the rockpool if possible, or migrate to the surrounding rock surface to escape extreme temperatures, while fixed species such as macroalgae will need to tolerate the temperature change (see Newell, 1979).

Lüning (1990) reported that Corallina officinalis from Helgoland survived one week exposure to temperatures between 0 °C and 28 °C. An abrupt increase in temperature of 10 °C caused by the hot, dry 'Santa Anna' winds (between January -and February) in Santa Cruz, California resulted in die back of several species of algae exposed at low tide (Seapy & Littler, 1984). Although fronds of Corallina spp. dramatically declined, summer regrowth resulted in dense cover by the following October, suggesting that the crustose bases survived. Severe damage was noted in Corallina officinalis as a result of desiccation during unusually hot and sunny weather in summer 1983 (an increase of between 4.8 and 8.5 °C) (Hawkins & Hartnoll 1985). Littler & Kauker (1984) suggested that the crustose base was more resistant of desiccation or heating than fronds.

Most of the other species within the biotope are distributed to the north and south of Britain and Ireland and unlikely to be adversely affected by long-term temperature change. But Hawkins & Hartnoll (1985) suggested that typical understorey red algae were susceptible to hot dry weather and that occasional damaged specimens of Palmaria palmata, Osmundea pinnatifida and Mastocarpus stellatus were observed after the hot summer of 1983.

It is likely that Corallina officinalis fronds are intolerant of abrupt short term temperature increase although they may not be affected by long term chronic change and the crustose bases are probably less intolerant of than fronds. Epifaunal species will decline due to loss of coralline turf cover. Similarly, acute increases in temperature will probably reduce the cover of the characterizing red algae. The rockpool community is probably tolerant of a short term 5 °C change in temperature (see benchmark) and under normal circumstances the body of water protects species from the immediate effects of desiccation. But a long term increase in average temperature may expose the community to extremes of temperature above the tolerance limits, resulting in loss of some species, especially red algae and a reduced abundance of coralline algae. Therefore, an intolerance of intermediate has been recorded. Recoverability is probably very high (see additional information below).

Tolerant Not sensitive* No change Moderate
In general, the water temperature of rockpools follows that of the air more closely than that of the sea and dramatic temporal changes in temperature may be observed. Under extremely low temperatures, components of the community demonstrate tolerance. Lüning (1990) reported that Corallina officinalis from Helgoland survived 0 °C when exposed for one week. New Zealand specimens were found to tolerate -4 °C (Frazer et al., 1988). Lüning (1990) suggested that most littoral algal species were tolerant of cold and freezing. For example, the photosynthetic rate of Chondrus crispus recovered after 3hrs at -20 °C but not after 6hrs (Dudgeon et al., 1990). The photosynthetic rate of Mastocarpus stellatus higher on the shore fully recovered from 24hrs at -20 °C. Littorina littorea is a hardy intertidal species and can tolerate long periods of exposure to the air and consequently wide variations in temperature. Adult snails can easily tolerate sub-zero temperatures and the freezing of over 50% of their extracellular body fluids (English & Storey, 1998). Adults of Patella vulgata are also largely unaffected by short periods of extreme cold. Ekaratne & Crisp (1984) found adult limpets continuing to grow over winter when temperatures fell to -6 °C, and stopped only by still more severe weather. However, loss of adhesion after exposure to -13 °C has been observed with limpets falling off rocks and therefore becoming easy prey to crabs or birds (Fretter & Graham, 1994). However, in the very cold winter of 1962-3 when temperatures repeatedly fell below 0 °C over a period of 2 months large numbers of Patella vulgata were found dead (Crisp, 1964). Nevertheless, the community has been assessed not sensitive to decreased temperature at the benchmark level.
Tolerant Not relevant Not relevant Not relevant Low
The biotope essentially occur in shallow waters where light attenuation due to increase turbidity is probably low. Red algae and coralline algae especially are known to be shade tolerant and are common components of the understorey on seaweed dominated shores. Therefore, a decrease in light intensity is unlikely to adversely affect the biotope.
Tolerant Not sensitive* No change Moderate
An increase in light intensity is unlikely to adversely affect the biotope.
Tolerant Not relevant Not relevant Not relevant Low
The biotope may occur on the upper shore in locations with varying wave exposures (very exposed to sheltered) (Connor et al., 1997b). The effects of wave exposure upon rockpool communities high on the shore are likely to depend on tidal amplitude as within a shore, and where the tidal amplitude is significant, the time for which organisms are subjected to wave action will vary along the intertidal gradient. For instance, during neap tide periods, mid and higher shore rockpools may remain isolated from the main body of the sea for several days or weeks in succession. During such times, wave action is unlikely to be of direct influence other than generating a spray, whilst during periods of tidal immersion wave action may directly affect the community. The changes in community composition that occur with increased wave exposure are accompanied by striking changes in the vertical levels of zones on the shore. In north-west Europe, all the zones become greater in vertical extent as wave exposure increases, and thus are found at greater heights above chart datum (Little & Kitching, 1996). In balance, over a year it is likely that as a result of increased wave action the vertical extent of the biotope up the shore will increase with no major changes in community composition occurring, so an assessment of not sensitive has been made.
Low Very high Moderate Minor decline Very low
The biotope may occur on the upper shore in locations with varying wave exposures (very exposed to sheltered) (Connor et al., 1997b). A decrease in wave action (directly effective only when the biotope is immersed) may mean that siltation and smothering of epifauna may occur and remain in the long term (see smothering above). Overall, only limited adverse effects are likely and an intolerance assessment of low has been made. However, in situations where the biotope occurs on the upper shore due to wave splash, and reduction in wave exposure may result in loss these examples of the biotope.
Tolerant Not relevant Not relevant No change Low
None of the species in this biotope are know to respond to noise or vibration at the benchmark level, an assessment of not sensitive has been made.
Tolerant Not relevant Not relevant No change Moderate
The mobile invertebrates are probably capable of responding to localized shading, experienced by the approach of a predator. But their visual acuity is likely to be low and they are unlikely to respond to visual disturbance at the benchmark level.
Intermediate High Low Minor decline Low
Abrasion by an anchor or mooring may remove some fronds of the foliose red algae and scrape coralline turf, although most species would grow back from their remaining holdfasts. Trampling may be more damaging (see A1.122) but is likely to be less significant in rockpool than on open rock surfaces. Therefore an intolerance assessment of intermediate has been made.
High High Moderate Decline Moderate
The majority of the characterizing fauna, such as the limpets, winkles, top shells isopods, amphipods and harpacticoid copepods are either highly mobile or would be able to re-attach to the substratum if undamaged, and are unlikely to be adversely affected by displacement. But the dominant macroalgae are permanently attached to the substratum and if removed will be lost, resulting in loss of the biotope overall. If their holdfasts and bases are also removed then recovery will be prolonged but still relatively rapid

Chemical Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
High High Moderate Major decline High
Smith (1968) reported that oil and detergent dispersants from the Torrey Canyon spill affected high water plans of Corallina officinalis more than low shore plants and some plants were protected in deep pools. In areas of heavy spraying, however, Corallina officinalis was killed (Smith, 1968). Regrowth of fronds had begun within two months after spraying ceased (Smith, 1968). O'Brien & Dixon (1976) suggested that red algae were the most sensitive group of algae to oil or dispersant contamination, possibly due to the susceptibility of phycoerythrins to destruction. They also reported that red algae are effective indicators of detergent damage since they undergo colour changes when exposed to relatively low concentration of detergent. However, Smith (1968) reported that red algae such as Chondrus crispus, Mastocarpus stellatus, and Laurencia pinnatifida were amongst the algae least affected by detergents. Laboratory studies by Grandy (1984) on the effects of oil and dispersants on several red algal species concluded that they were all sensitive to oil/dispersant mixtures, with little difference between adults, sporelings, diploid or haploid life stages.

Cole et al. (1999) suggested that herbicides were (not surprisingly) very toxic to algae and macrophytes. Hoare & Hiscock (1974) noted that with the exception of Phyllophora species, all red algae including encrusting coralline forms, were excluded from the vicinity of an acidified halogenated effluent discharge in Amlwch Bay, Anglesey and that intertidal populations of Corallina officinalis occurred in significant amounts only 600m east of the effluent. Chamberlain (1996) observed that although Lithophyllum incrustans was quickly affected by oil during the Sea Empress spill, recovery occurred within about a year. The oil was found to have destroyed about one third of the thallus thickness but regeneration occurred from thallus filaments below the damaged area.

Most pesticides and herbicides were suggested to be very toxic for invertebrates, especially crustaceans (amphipods isopods, mysids, shrimp and crabs) and fish (Cole et al., 1999).

intolerance to synthetic chemicals has been assessed to be high, owing to the likely loss of key structural and important characterizing as well as other red foliose algae from the rockpools. On return to prior conditions, recoverability has been assessed to be high (see additional information below).

Heavy metal contamination
Intermediate High Low Decline Moderate
Little information was found concerning the effects of heavy metals on turf forming and encrusting coralline algae. However, Bryan (1984) suggested that the general order for heavy metal toxicity in seaweeds is: organic Hg> inorganic Hg > Cu > Ag > Zn> Cd>Pb.
Most of the information available suggests that adult gastropod molluscs are rather tolerant of heavy-metal toxicity (Bryan, 1984). Winkles may absorb metals from the surrounding water by absorption across the gills or from their diet, and evidence from experimental studies on Littorina littorea suggest that diet is the most important source (Bryan et al., 1983). The species has been suggested as a suitable bioindicator species for some heavy metals in the marine environment. Bryan et al. (1983) suggested that the species is a reasonable indicator for Ag, Cd, Pb and perhaps As. In the Fal estuary Patella vulgata occurs at, or just outside, Restronguet Point, at the end of the creek where metal concentrations are in the order: Zinc (Zn) 100-2000 µg/l, copper (Cu) 10-100µg/l and cadmium (Cd) 0.25-5µg/l (Bryan & Gibbs, 1983). However, in the laboratory Patella vulgata was found to be intolerant of small changes in environmental concentrations of Cd and Zn by Davies (1992). At concentrations of 10µg/l pedal mucus production and levels of activity were both reduced, indicating a physiological response to metal concentrations. Exposure to Cu at a concentration of 100 µg/l for one week resulted in progressive brachycardia (slowing of the heart beat) and the death of limpets. Zn at a concentration of 5500 µg/l produced the same effect (Marchan et al., 1999).
An intolerance assessment of intermediate has been made as evidence suggests that important grazers in the biotope would be adversely affected by heavy metal pollution of the biotope. Recruitment patterns of such species suggest that recoverability would be high.
Hydrocarbon contamination
High High Moderate Major decline Moderate
Where exposed to direct contact with fresh hydrocarbons, encrusting coralline algae appear to have a high intolerance. Crump et al. (1999) described "dramatic and extensive bleaching" of 'Lithothamnia' following the Sea Empress oil spill. Observations following the Don Marika oil spill (K. Hiscock, pers. comm.) were of rockpools with completely bleached coralline algae. However, Chamberlain (1996) observed that although Lithophyllum incrustans was affected in a short period of time by oil during the Sea Empress spill, recovery occurred within about a year. The oil was found to have destroyed about one third of the thallus thickness but regeneration occurred from thallus filaments below the damaged area.

Following the Torrey Canyon oil spill in 1967, oil and detergent dispersants affected high shore specimens of Corallina officinalis more than low shore specimens. Plants in deep pools were afforded some initial protection, although probably later affected by contaminated runoff. In areas of heavy spraying, however, Corallina officinalis was killed. (Smith 1968). Intolerance to hydrocarbon pollution has been assessed to be high, as key structural and important characterizing coralline algal species will be lost and the biotope not be recognized in their absence. Recoverability of the key structural and important characterizing turf forming and encrusting coralline algae has been assessed to be high (see additional information below).

Radionuclide contamination
No information Not relevant No information Not relevant Not relevant
Insufficient
information.
Changes in nutrient levels
Low Very high Very Low No change Low
Corallines seem to be tolerant and successful in polluted waters. Kindig & Littler (1980) demonstrated that Corallina officinalis var. chilensis in South California showed equivalent or enhanced health indices, highest productivity and lowest moralities (amongst the species examined) when exposed to primary or secondary sewage effluent. Little difference in productivity was noted in chlorinated secondary effluent or pine oil disinfectant. However, specimens from unpolluted areas were less tolerant, suggesting physiological adaptation to sewage pollution (Kindig & Littler 1980). Also, increased nutrients may result in overgrowth of coralline algae by other epiphytic algae. Grazers in the biotope may benefit from increase availability of food resources. An intolerance assessment of low has been made as key structural and important characterizing species may experience reduced viability. On return to prior conditions, recovery has been assessed to be very high as, although the biotope will recover to typical abundances, some species may remain abnormally abundant for a period whilst the biotope community re-stabilizes.
Tolerant Not relevant Not relevant No change Moderate
Conditions within rockpools are the consequence of prolonged separation from the main body of the sea, and physico-chemical parameters within them fluctuate dramatically (Huggett & Griffiths, 1986). Small and shallow pools are especially influenced by insolation, air temperature and rainfall, the effects of which become more significant towards the high shore, where pools may be isolated from the sea for a number of days or weeks (Lewis, 1964). Rockpools in the mid eulittoral are likely to experience lesser changes in salinity than those of the supralittoral, littoral fringe and upper eulittoral which are liable to gradually changing salinities followed by days of fully marine or fluctuating salinity at times of spring tide (Lewis, 1964). The community has been assessed to be not sensitive to increased salinity at the benchmark level because it represents a lesser change in salinity than the community might normally be expected to experience and the community persists owing to the tolerance of species to short-term acute changes.
Tolerant Not sensitive* No change Moderate
Conditions within rockpools are the consequence of prolonged separation from the main body of the sea, and physico-chemical parameters within them fluctuate dramatically (Huggett & Griffiths, 1986). Small and shallow pools are especially influenced by insolation, air temperature and rainfall, the effects of which become more significant towards the high shore, where pools may be isolated from the sea for a number of days or weeks (Lewis, 1964). Rockpools in the mid eulittoral are likely to experience lesser changes in salinity than those of the supralittoral, littoral fringe and upper eulittoral which are liable to gradually changing salinities followed by days of fully marine or fluctuating salinity at times of spring tide (Lewis, 1964). Salinity values ranging from 5-30 psu have been recorded in upper shore rockpools over a period of 24 hrs (Ranade, 1957). The community has been assessed to be not sensitive to decreased salinity at the benchmark level because it represents a lesser change in salinity than the community might normally be expected to experience and the community persists owing to the tolerance of species to short-term acute changes.
Low Very high Very Low Minor decline Low
Oxygenation of the rockpool is usually high due to photosynthetic activity and frequent water exchange. During the day, algae within rockpools produce oxygen by photosynthesis, and oxygen concentrations may rise to three times the saturation value, so that it is released as bubbles. The effect of oxygen production is to increase the pH of water in the pool owing to utilization of carbon dioxide. At night, when photosynthesis has ceased, algal respiration may utilize much of the available oxygen and minimum values of 1-5 % saturation of oxygen have been recorded in situ (Morris & Taylor, 1983). The algal component of the biotope may be intolerance of reduced oxygen concentration in darkness when they can only respire. However, corallines may be more tolerant than most algae due to their low rates of respiration (see Littler & Kauker 1984 for values).

Littorina littorea can endure long periods of oxygen deprivation. The snails can tolerate anoxia by drastically reducing their metabolic rate (down to 20% of normal) (MacDonald & Storey, 1999). However, this reduces feeding rate and thus the viability of a population may be reduced. Patella vulgata was assessed to have an intermediate intolerance to low oxygen concentrations (see MarLIN review). At the benchmark level, the community has been assessed to have a low intolerance to reduced oxygenation of the water. Mobile species are likely to leave unfavourable locations, whilst immobile species are likely to suffer an energy expenditure, e.g. owing to changes in metabolism, in order to survive. On return to optimal conditions, recoverability has been assessed to be very high.

Biological Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
No information Not relevant No information Not relevant Not relevant
Several coralline and non-coralline species are epiphytic on Corallina officinalis. Irvine & Chamberlain (1994) cite tissue destruction caused by Titanoderma corallinae. However, no information on pathogenic organisms in the British Isles was found. In Rhodophycota, viruses have been identified by means of electron microscopy (Lee, 1971) and they are probably widespread. However, nothing is known of their effects on growth or reproduction in red algae and experimental transfer from an infected to an uninfected specimen has not been achieved (Dixon & Irvine, 1977). Overall, insufficient information was found to make an assessment
Low Very high Very Low Minor decline Low
The non-native wireweed Sargassum muticum may occur extensively in examples of this biotope. But the biotope persists, probably because of the small area of basal attachment of Sargassum. Sargassum probably competes with other macroalgae for light and nutrients so an intolerance of low has been recorded.
Intermediate Very high Low Decline Low
Corallina officinalis was used in Europe as a vermifuge although it no longer seems to be collected for this purpose (Guiry & Blunden, 1991). Corallina officinalis is collected for medical purposes; the fronds are dried and converted to hydroxyapatite and used as bone forming material (Ewers et al., 1987). It is also sold as a powder for use in the cosmetic industry. An European research proposal for cultivation of Corallina officinalis is pending as of May 2000 (Wiedemann pers. comm.). Littorina littorea may also be collected. Both Chondrus crispus and Mastocarpus stellatus, although not species indicative of sensitivity, are collected as 'carragheen' by hand picking and racking in Europe (Guiry & Blunden, 1991). However, as long as holdfasts remain recovery will probably be rapid. Overall, intermediate intolerance has been suggested to reflect the fact that some species may experience a small decline. Recovery is likely to be very high (see additional information).
Not relevant Not relevant Not relevant Not relevant Not relevant

Additional information

Recoverability:
Corallina officinalis probably has good recruitment and settled on artificial substances within 1 week of their placement in the intertidal in New England summer (Harlin & Lindbergh, 1977). New fronds of Corallina officinalis appeared on sterilized plots within six months and 10% cover was reached with 12 months (Littler & Kauker, 1984). Bamber & Irving (1993) reported that new plants grew back in scraped transects within 12 months, although the resistant crustose bases were probably not removed. Similarly, in experimental plots, up to 15% cover of Corallina officinalis fronds returned within 3 months after removal of fronds and all other epiflora/fauna but not the crustose bases (Littler & Kauker, 1984). Although new crustose bases may recruit and develop quickly the formation of new fronds from these bases and recovery of original cover may take longer, and it is suggested that a population is likely to recover within a few years.

Chamberlain (1996) observed that although Lithophyllum incrustans was quickly affected by oil during the Sea Empress spill, recovery occurred within about a year. The oil was found to have destroyed about one third of the thallus thickness but regeneration occurred from thallus filaments below the damaged area. A recoverability of high is therefore suggested. If colonies were completely destroyed new growth would be slow and, because of low growth rates, recoverability would be low. If death occurred, recoverability will be slow. Spores will settle and new colonies will arise rapidly on bare substratum but growth rate is slow (2-7 mm per annum - see Irvine & Chamberlain 1994).

Recolonization of Patella vulgata on rocky shores is rapid as seen by the appearance of limpet spat 6 months after the Torrey Canyon oil spill reaching peak numbers 4-5 years after the spill (Southward & Southward, 1978). The epifauna are mainly composed of mobile species that will recruit quickly from surrounding habitats, and will therefore, recover quickly once the coralline turf has developed.

Importance review

Policy/Legislation

Habitats Directive Annex 1Reefs
UK Biodiversity Action Plan Priority

Exploitation

Corallina officinalis was used in Europe as a vermifuge although it no longer seems to be collected for this purpose (Guiry & Blunden, 1991). Corallina officinalis is collected for medical purposes; the fronds are dried and converted to hydroxyapatite and used as bone forming material (Ewers et al., 1987). It is also sold as a powder for use in the cosmetic industry. A European research proposal for cultivation of Corallina officinalis is pending as of May 2000 (Wiedemann, pers. comm.). Both Chondrus crispus and Mastocarpus stellatus are collected as 'carragheen' by hand picking and racking in Europe (Guiry & Blunden, 1991).

Additional information

-

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Citation

This review can be cited as:

Budd, G.C. 2002. Coralline crust-dominated shallow eulittoral rockpools. 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: http://www.marlin.ac.uk/habitat/detail/240

Last Updated: 13/11/2002