Green seaweeds (Enteromorpha spp. and Cladophora spp.) in shallow upper shore rockpools

28-11-2002
Researched byGeorgina Budd Refereed byThis information is not refereed.
EUNIS CodeA1.421 EUNIS NameGreen seaweeds (Enteromorpha spp. and Cladophora spp.) in shallow upper shore rockpools

Summary

UK and Ireland classification

EUNIS 2008A1.421Green seaweeds (Enteromorpha spp. and Cladophora spp.) in shallow upper shore rockpools
EUNIS 2006A1.421Green seaweeds (Enteromorpha spp. and Cladophora spp.) in shallow upper shore rockpools
JNCC 2004LR.FLR.Rkp.GGreen seaweeds (Enteromorpha spp. and Cladophora spp.) in shallow upper shore rockpools
1997 BiotopeLR.LR.Rkp.GGreen seaweeds (Enteromorpha spp. and Cladophora spp.) in upper shore rockpools

Description

Rockpools in the supralittoral, littoral fringe or upper eulittoral which are subject to variable salinity and widely fluctuating temperatures are characterized by the ephemeral green alga Ulva spp. or the filamentous green alga Cladophora spp. Due to the physical stress imposed on these upper shore pools, grazing molluscs are generally in lower abundance than eulittoral pools, allowing the green algae to proliferate under reduced grazing pressures. The rock surface is often covered by the black lichen Verrucaria maura. On more exposed shores crevices in the rock may contain small Mytilus edulis. The bright orange copepod Tigriopus fulvus is tolerant of large salinity fluctuations and may also occur in large numbers in these upper shore pools. (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

On rocky coasts around Britain and Ireland.

Depth range

Upper shore

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Habitat review

Ecology

Ecological and functional relationships

In rockpools high on the shore, the familiar flora and fauna of rockpools is lost, the community becomes greatly depleted consisting of forms that are highly adapted to the rigorous and almost estuarine conditions (Lewis, 1964) as physical factors are the dominant structuring force.
  • Amongst the fauna, crustaceans predominate. Large populations (720 x 103m²) of the copepod Tigriopus fulvus can occur in upper shore rockpools densely covered by the green alga Ulva intestinalis (Goss-Custard et al., 1979). Tigriopus fulvus is remarkably tolerant of extremes of salinity and temperature (Ranade, 1957). Ranade (1957) stated that Tigriopus fulvus could live normally between salinities of 4-90 psu. In laboratory experiments, Goss-custard et al. (1979) found the species to survive for 15 days in salinities ranging from 42-90 psu, but died after 84 hours in distilled water, and sank to the bottom in salinities greater than 90 psu in a state of apparent death. However, if transferred to seawater (35 psu) after some hours it could recover. In tests with a slowly rising temperature, the death point was 32°C at a salinity of 34 psu, but this rose to 41.8°C at a salinity of 90 psu. Thus high salinities enable Tigriopus fulvus to withstand high temperature, a feature useful for a species living in pools in a zone where insolation and evaporation may be considerable. Despite the instability of the high shore rockpool as a habitat, the copepod benefits from the lower abundance of predators, that are in greater abundance in lower shore rockpools (Dethier, 1980).
  • Ulva intestinalis provides shelter for the orange harpacticoid copepod, Tigriopus brevicornis, and the chironomid larva, Halocladius fucicola (McAllen, 1999). Ulva intestinalis is often the only seaweed found in supralittoral rockpools, and the copepod and chironomid species utilize the hollow thallus of Ulva intestinalis as a moist refuge from desiccation when the rockpools completely dry out. Several hundred individuals of Tigriopus brevicornis have been observed in a single thallus of Ulva intestinalis (McAllen, 1999).
  • There are three major sources of food available to the fauna of high shore rockpools: the thalli of Ulva sp. and other macroalgae, the epiphytic micro-organisms attached to the surface of the Ulva and the micro-organisms associated with the substratum (Clark, 1968).
  • The distribution of grazers, Melarhaphe neritoides and Littorina saxatilis extends into the upper littoral fringe, the former feeding on micro-algae and lichens, the latter grazing on macroalgae and the microalgal film on the rocks. Both winkles favour crevices, especially in dry weather, from which they can forage, but owing to the physical stresses of the upper shore, grazing molluscs are generally lower in abundance than in eulittoral pools allowing green algae to proliferate as a result of reduced grazing pressure.
  • A band of yellow and grey lichens (LR.YG) is usually found immediately above the zone of Verrucaria maura which occurs in this biotope. The fauna of the LR.YG biotope may extend into the LR.G biotope to exploit the lichen. For instance, lichens are fed on by fungivorous Cryptostigmata and other acarid mites and potentially by some lichen dwelling tardigrades (Gerson & Seaward, 1977) and the bristle tail Petrobius maritimus (Joosse, 1976), while rotifers have been reported to consume lichen ascospores (Gerson & Seaward, 1977).

Seasonal and longer term change

Rockpools constitute a distinct environment for which physiological adaptations by the flora and fauna may be required (Lewis, 1964). Physico-chemical parameters within rockpools fluctuate dramatically as a consequence of prolonged separation from the main body of the sea (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 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. However, 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 a period of 24 hours (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 affect 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. pH values >9 were recorded in rockpools on the Isle of Cumbrae (Morris & Taylor, 1983). At night changes occur in the opposite direction. Respiration utilizes much of the available oxygen and pH decreases.
Fluctuations especially in the abundance of green seaweeds is likely owing to the marked changes in salinity and temperature during the year. For instance, surface layers of Ulva may be bleached in the summer.

Habitat structure and complexity

Rockpools vary greatly in their physical features. Pools in bedrock may be shallow and well-lit or deep and shaded with overhanging sides and vertical surfaces. Algae growing within provide additional surface for colonization and for shelter. There is also a tendency for loose substrata (sand, stones, rocks) to accumulate in pools, the instability of which may cause abrasion and affect species diversity. Amongst rockpools, deep crevices may be found, around the entrance of which small mussels may cluster. Crevices also support their own specialized fauna with many air-breathing arthropods such as centipedes, millipedes, beetles, pseudoscorpions and primitive onchidellid pulmonates (see Lewis, 1964).

Productivity

Macroalgae and the microbial film of bacteria, blue-greens, diatoms, fungi and protozoans are the primary producers in this biotope. Accumulations of algal debris are also likely in high shore rockpools and such detrital material contributes to overall productivity. Information specific to the community was not found, but Workman (1983) gave an estimate of primary production by microalgal films on the high shore in the British Isles to be in the region of 60 g C/m²/yr, much of which will be utilized directly by grazers.

Recruitment processes

Flora:
Rockpools in the supralittoral, littoral fringe or upper eulittoral which are subject to variable salinity and widely fluctuating temperatures are characterized by the ephemeral green alga Ulva spp. or the filamentous green alga Cladophora spp.
Species of the genus Ulva are rapidly growing opportunists, favoured by the frequency and speed of their reproduction. The short lived plants reach maturity at a certain stage of development rather than relying on an environmental trigger. Ulva intestinalis can be found in reproductive condition at all times of the year, but maximum development and reproduction occur during the summer months especially towards the northern end of the distribution of the species (Burrows, 1991). The life history consists of an isomorphic (indistinguishable except for the type of reproductive bodies produced) alternation between haploid gametophytic and diploid sporophytic generations, but can be modified by environmental conditions (Burrows, 1959; Moss & Marsland, 1976; Reed & Russell, 1978).
The haploid gametophytes of Ulva produce enormous numbers of biflagellate motile gametes which cluster and fuse to produce a sporophyte (diploid zygote). The sporophyte matures and produces by meiosis large numbers of quadriflagellate zoospores that mature as gametophytes, and the cycle is repeated. Both gametes and spores may be released in such quantities into rock pools or slack water that the water mass is coloured green (Little & Kitching, 1996). Together spores and gametes are termed 'swarmers'. Swarmers are often released in relation to tidal cycles, with the release being triggered by the incoming tide as it wets the thallus. However, the degree of release is usually related to the stage of the spring/neap tidal cycle, so allowing regular periodicity and synchronization of reproduction (Little & Kitching, 1996). Christie & Evans (1962) found that swarmer release of Ulva intestinalis from the Menai Straits, Wales, peaked just before the highest tides of each neap-spring cycle. Mobility of swarmers belonging to Ulva intestinalis can be maintained for as long as 8 days (Jones & Babb, 1968). Algae such as Ulva intestinalis tend to have large dispersal shadows, with propagules being found far from the nearest adult plants, e.g. 35 km (Amsler & Searles, 1980).
Information on the ecology of reproduction and propagation of the genus Cladophora is limited. Reproduction is asexual, and achieved by the release of quadriflagellate zoospores and biflagellate isogametes formed in the terminal cells of fronds. The life history consists of an isomorphic (indistinguishable except for the type of reproductive bodies produced) alternation of gametophyte and sporophyte generations, the plants are dioecious (Burrows, 1991). Both zoospores and gametes can be found at most times of the year. Archer (1963) was unable to find any correlation between the time of reproduction, the state of tide or environmental conditions. Most species of Cladophora attach to the substratum by multicellular, branching rhizoids (van den Hoek, 1982); these basal holdfasts may serve as resistant structures from which new growths can arise.
Fauna:
Fraser (1936) describes the ecology and life-history of the copepod Tigriopus fulvus. The species mates throughout the year. Females of the species release a sex pheromone promoting sexual recognition and attraction in males (Lazzaretto et al., 1994). Females brood the fertilized eggs which may be released between 5-15 days after the appearance of the female's egg sac, the time being shorter in summer and longer in winter. A single female may produce numerous juveniles from several egg sacs without further mating. From the time of hatching a juvenile attains an adult form and the ability to reproduce in about two months (Fraser, 1936).
Internal fertilization occurs in all species of winkle (littorinids). Melarhaphe neritoides releases its eggs into the plankton, whilst the female Littorina saxatilis broods its eggs which hatch as live young. Although animals with planktonic larvae have a greater dispersive ability than those with direct development, the production of crawling, live young from egg capsules or brood pouches reduces reproductive losses and permits exploitation of locally favourable conditions. It can also lead to inbreeding and genetic isolation of populations. For instance, owing to dispersion in the plankton the population of Melarhaphe neritoides is genetically homogenous, which is reflected in their uniform colour. Littorina saxatilis, which bears live young and are variable in size and colour (Hawkins & Jones, 1992).

Time for community to reach maturity

To recruit, grow and reproduce in the unpredictable environment of high shore rockpools, the flora and fauna within need to be capable of rapid recruitment, early maturation and rapid growth in order to exploit the habitat, thus it is likely that the community would be considered mature in terms of species present and capable of reproduction within a few months.
For example, with the exception of Cladophora rupestris whose turfs may persist for many years, the macroalgal species, e.g. Ulva, Monostroma and Prasiola stipitata which are characteristic of this biotope are seasonal and short lived (ephemeral) algae, which recruit rapidly to available substrata. For instance, the thalli of Ulva intestinalis, which arise from spores and zygotes, grow within a few weeks into thalli that reproduce again, and the majority of the cell contents are converted into reproductive cells. The species is also capable of dispersal over a considerable distance. For instance, Amsler & Searles (1980) showed that swarmers of a coastal population of Ulva reached exposed artificial substrata on a submarine plateau 35 km away. Ulva is amongst the first multicellular algae to appear on substrata that have been cleared following a disturbance, e.g. following the Torrey Canyon oil spill in March 1967, species of the genus Ulva rapidly recruited to areas where oil had killed the herbivores that usually grazed on them, so that a rapid greening of the rocks (owing to a thick coating of macroalgae) was apparent by mid-May (Smith, 1968).

Additional information

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Preferences & Distribution

Recorded distribution in Britain and IrelandOn rocky coasts around Britain and Ireland.

Habitat preferences

Depth Range Upper shore
Water clarity preferences
Limiting Nutrients Nitrogen (nitrates), Phosphorus (phosphates)
Salinity Variable (18-40 psu)
Physiographic
Biological Zone Upper eulittoral, Infralittoral, Supralittoral
Substratum Bedrock
Tidal
Wave Exposed, Moderately exposed, Sheltered
Other preferences

Additional Information

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Species composition

Species found especially in this biotope

  • Tigriopus fulvus

Rare or scarce species associated with this biotope

-

Additional information

None entered

Sensitivity reviewHow is sensitivity assessed?

Explanation

Ulva intestinalis and Cladophora rupestris are important characterizing species of this biotope as they are tolerant and able to exploit the unstable conditions found in high shore rockpools. However, these macroalgae have been included as key structuring species, as they provide a habitat that supports an associated community. For instance, harpacticoid copepods feed upon Ulva intestinalis and utilize the hollow thalli as a moist refuge from desiccation when the rockpools completely dry out. Cladophora spp. also presents a substratum for colonization and a food resource. Tigriopus fulvus is included as an important characterizing species particularly well adapted to the temporal changes experienced in high shore rockpools.

Species indicative of sensitivity

Community ImportanceSpecies nameCommon Name
Key structuralCladophora rupestrisA green seaweed
Important characterizingTigriopus fulvusA copepod
Key structuralUlva intestinalisGut weed

Physical Pressures

 IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
High Very high Low Major decline Low
Removal of the substratum will result in the removal and loss of the biotope. Therefore, an intolerance of high has been recorded. The macroalgae within this biotope can rapidly colonize new substratum and grow rapidly, probably within a few months therefore a recoverability of very high has been recorded (see additional information below).
Intermediate Very high Low Decline Moderate
Ulva intestinalis is a filamentous seaweed without structural support for its thalli, therefore it is likely that entire plants would be smothered by an additional covering of 5 cm of sediment. Cladophora rupestris is a stout shrub like seaweed, whose fronds may grow up to 20 cm in height. A covering of sediment to a depth of 5 cm is likely to partially cover the seaweed, and at low tide the whole plant may be covered whilst lying limply on the rock or in shallow pools. Unless the sediment is removed by the incoming tide (which may be some time on the high shore where pools may be isolated from the main body of the sea for several days in succession), photosynthesis would be inhibited and fronds of macroalgae may begin to decay over the duration of one month. Spores, germlings and juveniles are likely to be highly intolerant of smothering by sediment (Vadas et al. 1992). An intolerance assessment of intermediate has been made to reflect the probable impact of smothering on germlings, thereby preventing recruitment for that period, and the inhibitory effects upon more mature specimens. On return to prior conditions, the macroalgae is likely to recover, either new growth will arise from the resistant multicellular branching rhizoids in the case of Cladophora rupestris (van den Hoek, 1982) that may remain in situ, or macroalgal species will recruit to cleared substrata via spores dispersed in the plankton. Smothering by impermeable material such as oil is likely to have a more severe and long lasting impact.
Not relevant Not relevant Not relevant Not relevant Not relevant
The effects of increased suspended sediment on macroalgae are likely to be indirect but include settlement of silt although, at the level of the benchmark, that is unlikely to cause smothering. Increased turbidity reduces light availability (see below). At the benchmark level an assessment of not relevant has been made and the effects of smothering and turbidity addressed elsewhere.
Tolerant Not sensitive* Not relevant High
The community is unlikely to be intolerant of a decrease in suspended sediment. The effects of increased light penetration owing to reduced turbidity are addressed elsewhere.
Intermediate Very high Low Minor decline Moderate
Ulva intestinalis is often very abundant on the high shore where desiccation stress is the primary factor controlling seaweed distribution, and may even be found above the tidal limits of the shore. Ulva intestinalis can survive several weeks of living in completely dried out rock pools, while becoming completely bleached on the uppermost layers, but remaining moist underneath the bleached fronds. Its ability to survive out of water for so long makes Ulva intestinalis an ideal refuge for copepods in supralittoral rockpools (McAllen, 1999). However, marine communities occurring at the highest level on the shore are living at the extreme of their physiological tolerance limits and so would not be likely to tolerate a further increase in desiccation, unless adequate refuge is available locally, e.g. in the case of littorinids, crevices in the rock, or, for other species, within pools. Increased desiccation would therefore result in the upper extent of species distributions being depressed. An intolerance assessment of intermediate has been made. On return to prior conditions, the community is likely to recover rapidly.
Tolerant Not relevant Not relevant Not relevant Low
The biotope is isolated from the main body of the sea for considerable and varying periods of time owing to its position high on the shore. At the benchmark level, it is unlikely that the community will demonstrate any significant additional stress attributable to an extra hour of emersion. An assessment of not sensitive has been made.
High High Intermediate Rise Low
A greater period of immersion will lessen the physico-chemical extremes that prevent the colonization of the 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.G community has been assessed to have a high intolerance to decreased emergence as it will allow the 'up-lift' of lower shore species into the biotope and will begin to change to another. 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
The community exists for the great majority of time in still water conditions. Upper shore rockpools may only flood with water due to wave action and would be unaffected by changes in tidal stream velocity that occur away from the shore. An assessment of not sensitive has been suggested.
Tolerant Not sensitive* Not relevant Low
The community exists for the great majority of time in still water conditions. Upper shore rockpools may only flood with water due to wave action and would be unaffected by changes in tidal stream velocity that occur away from the shore. An assessment of not sensitive has been suggested.
Tolerant Not relevant Not relevant Not relevant Moderate
At the benchmark level the community has been assessed not to be intolerant of increased temperature.
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. For example, the copepod Tigriopus fulvus is more tolerant of high temperatures at higher salinities (see ecological relationships). At a salinity of 34 psu, the death point of Tigriopus fulvus is reached at 32°C (Goss-Custard et al.,1979). Clark (1992) reviewed the influence of cooling water effluent on shore communities. Effects are usually restricted to the immediate vicinity of the outfall, but brown seaweeds were eliminated from a rocky shore heated to 27-30°C by a power station in Maine, whilst Ulva intestinalis increased significantly near the outfall (Vadas et al., 1976). Fortes & Lüning (1980) and Lüning (1984) reported that Cladophora rupestris could survive exposure to temperatures in the range 0 - 28°C for at least a week.
Tolerant Not sensitive* Not relevant Moderate
At the benchmark level the community has been assessed not to be intolerant of decreased temperature.
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. 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. Under extremely low temperatures, components of the community demonstrate tolerance. For instance, Kylin (1917) reported Ulva(as Enteromorpha species to be tolerant of a temperature of -20°C, whilst growth measurements of Cladophora rupestris led Cambridge et al. (1984) to conclude that the species was tolerant of temperatures of below -5°C. The copepod fauna may be less tolerant of near freezing temperatures. For instance, Davenport et al. (1997) stated that unlike other species of Tigriopus, Tigriopus fulvus could not withstand freezing temperatures, whilst Tigriopus brevicornis appears to be able to withstand exposure to low temperature (and to severe hypoxia) by entering a quiescent/dormant state during which its metabolic rate is significantly reduce (McAllen et al., 1999). At extremes of temperature some mortality would be expected.
Low Immediate Not sensitive No change Very low
The light attenuating effects of increased turbidity are likely to impact on the photosynthetic efficiency of macroalgal species and the microalgal film in the short term, but owing to the relative stillness of water in rockpools, suspended sediment causing increased turbidity is likely to settle out. At the benchmark level an intolerance assessment of low has been made. Optimal photosynthesis is likely is resume rapidly.
Tolerant* Not sensitive No change Moderate
As photoautotrophs, macroalgae and microalgal films are likely to benefit from reduced turbidity, as the light attenuating effects of turbid water reduce photosynthesis. An assessment of not sensitive* has been made.
Low Very high Very Low Rise Low
The biotope may occur on the upper shore in locations with varying wave exposures (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, high shore rockpools may remain isolated from the main body of the sea for several days or weeks in concession. 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). The upper limit of littorinids and Verrucaria may rise dramatically, so that the vertical extent of the shore may increase from a few metres in shelter to 30 m or more in extreme exposures. Hence over a year the upward extent of the LR.G biotope may increase following an increase of two ranks on the MNCR wave exposure scale. More frequent renewal of seawater at the lower extent of the biotope may allow other species less tolerant of temperature and salinity variation to establish and begin to change the biotope to another. An intolerance assessment of low has been suggested as increased competition for space and resources may occur. On return to prior conditions, recovery is likely to be very high as intolerant species decline.
Tolerant Not sensitive* Not relevant Low
The biotope may occur on the upper shore of locations with varying wave exposure (exposed to sheltered) (Connor et al., 1997b). As elevated wave exposure serves to 'up-lift' zones of animal and plant communities up the shore (see increased wave exposure), decreased wave exposure will presumably cause a reversal to some extent and the upward extent of species characteristic of the biotope dominate become lower. However, providing suitable habitats are available the LR.G biotope may develop in pools further down the shore no longer subject to frequent renewal of seawater. Therefore an assessment of not sensitive has been suggested.
Tolerant Not relevant Not relevant No change Moderate
The biotope is characterized by macroalgae and small crustaceans and molluscs unlikely to be disturbed by noise at the benchmark level. An assessment of not sensitive has been made.
Tolerant Not relevant Not relevant No change Moderate
The biotope is characterized by macroalgae and small crustaceans and molluscs unlikely to be disturbed by the visual presence of objects not normally found in the marine environment. An assessment of not sensitive has been made.
High Very high Low Minor decline Moderate
Ulva intestinalis and Cladophora rupestris are likely to be susceptible to abrasion as they are not of a resilient growth form and would easily be scraped from the substratum by dragging objects. Littorinids may be knocked from rocks by physical disturbance and unless damaged are likely to reattach. Small copepods would probably be able to avoid abrasive agents by seeking refuge. Intolerance has been assessed to be high. Ulva intestinalis and Cladophora rupestris are cosmopolitan species that reproduce rapidly enabling them to colonize available substrata, so recoverability has been assessed to be very high (see additional information below).
High Very high Low Decline Low
Ulva intestinalis typically forms a permanent attachment to suitable substrata. However, in some circumstances, the algae may becomes detached from the substratum, and buoyed-up by gas, it floats up to the surface and continues to grow in mats (e.g. Baeck et al., 2000) but in which case it would be lost from the biotope. Cladophora rupestris forms a permanent attachment to solid substrata. It is likely to be intolerant of displacement as once removed, mature plants are unable to reattach. Faunal species associated with the biotope are mobile. The intolerance of the biotope to displacement has been assessed to be high as the key structuring seaweed species, Ulva and Cladophora, would not be able to reattach. However, recoverability has been assessed to be very high (see additional information below).

Chemical Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
High Very high Low Decline Low
Ulva intestinalis was assessed to have an intermediate intolerance to synthetic chemicals owing to evidence of adverse effects on the species viability and damage causing mortality (Moss & Woodhead, 1975; Scarlett et al., 1997; Smith 1968). Smith (1968) reported Cladophora rupestris to be amongst algae of unhealthy appearance following exposure to oil dispersants. The intolerance of the biotope has been assessed to be high as key structural species may be adversely affected. Assuming deterioration of contaminants, recovery of the key structuring species has been assessed to be very high (see additional information below).
Heavy metal contamination
Low High Low Minor decline Very low
The order of metal toxicity to algae varies, with the algal species and environmental conditions, but generally the order is Hg>Cu>Cd> Ag>Pb>Zn (Rice et al., 1973; Rai et al., 1981). Available evidence suggested that Ulva intestinalis was relatively tolerant of heavy metal exposure, only experiencing reduced growth (see full MarLIN review). 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 the diet, and evidence from experimental studies suggest that the diet is the most important source, e.g. Bryan et al., 1983). The intolerance of the community to heavy metal pollution has been assessed to be low as available evidence suggests effects upon species viability. Recoverability has been assessed to be high. Whilst the key structuring algal species may recolonize rapidly within six months, the littorinids associated with this biotope are likely to take longer to recover should the population become depleted owing to the fact that they have very localized recruitment with live young rather than a dispersive larval stage (see recruitment processes).
Hydrocarbon contamination
Intermediate Very high Low Decline Moderate
Bokn et al. (1993) examined the long term effects of the water-accommodated fraction (WAF) of diesel oil on rocky shore populations. Two doses (average hydrocarbon concentration in diesel oil equivalents; High: = 129.4 µ/mg/L and Low = 30.1µ/mg/L of WAF of diesel oil were delivered via sea water to established rocky shore mesocosms over a two year period, however there were no demonstrable effects in the abundance patterns of Cladophora rupestris, Ulva lactuca in the oil contaminated compared with the control mesocosms at the end of that period.

However, direct contact with viscous oil is likely to have a greater impact on the community. The toxic effects of oil on algae may be categorized as those associated with the coating of the fronds, e.g. coating by oil is likely to reduce CO2diffusion and light penetration to the plant, and those attributable to the uptake of hydrocarbons and subsequent disruption of cellular metabolism (Lobban & Harrison, 1997). For instance, Cullinane et al. (1975) summarised the damage caused to Cladophora rupestris following the crude oil spill in 1974 in Bantry Bay, Ireland. No damage was immediately apparent to Cladophora rupestris, but microscopic examination of material from rock pools at League Point showed complete bleaching of the terminal cells (only). Burrows (1991) indicated that following damage to the apical cells of fronds, that regeneration was possible. Ulva intestinalis also tends to recover very rapidly from oil pollution incidents. For instance, after the Torrey Canyon oil in 1967, grazing littorinid species were killed, and a dense flush of ephemeral green algae (Ulva and Blidingia) appeared on the rocky shore within a few weeks and persisted for up to one year (Smith, 1968).
Experience of and observations from oil spills such as the Sea Empress and Amoco Cadiz suggest that gastropod molluscs are highly intolerant of hydrocarbon pollution and it is likely that littorinid species would suffer mortality following oil contamination of this biotope. However, they are not very characteristic of the biotope and in the absence of their grazing pressure the green algae characteristic of the biotope may extend in distribution. Intolerance has been assessed to be intermediate. Recoverability has been assessed to be very high.
Radionuclide contamination
No information Not relevant No information Not relevant Not relevant
Insufficient
information.
Changes in nutrient levels
Tolerant* Not relevant Not sensitive* Rise Low
Nutrient enrichment of the water column, e.g. resulting from sewage discharge, can stimulate blooms of opportunistic algae, especially by those of the Chlorophyceae,Ulva, Cladophora and Ulva species (Knox, 1986). An assessment of not sensitive* has been made, as the species experience increased growth rates as a result of nutrient enrichment. Grazers in the biotope would benefit from a more abundance food supply.
Tolerant Not relevant Not relevant Not relevant High
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 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). The community has been assessed not to be intolerant of 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* Not relevant High
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 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). Values ranging from 5-30 psu have been recorded in a period of 24 hours (Ranade, 1957). The community has been assessed to be tolerant of 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 relevant Not relevant Not relevant Moderate
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 actually released as bubbles. The effect of which 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 have been recorded (Morris & Taylor, 1983). Algae in this biotope are unlikely to be adversely affected by decreased oxygen as they produce their own. The effect of oxygen saturation resulting from algal photosynthesis is lesser in high shore pools as they tend to contain fewer algae. The effect of severe hypoxia on the copepod Tigriopus brevicornis is for it to enter a quiescent/dormant state during which its metabolic rate is significantly reduced. It recovers on return to optimal conditions (McAllen et al., 1999). As inhabitants of littoral rockpools are subjected and therefore likely to be highly adapted to dramatic changes in physico-chemical conditions such as oxygen concentration, the community has been assessed not to be sensitive at the benchmark level.

Biological Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
Not relevant Not relevant Not relevant Not relevant Not relevant
Insufficient
information.
Not relevant Not relevant Not relevant Not relevant Not relevant
No known alien species are reported to adversely affect the biotope. An assessment of not relevant has been made.
Not relevant Not relevant Not relevant Not relevant Not relevant
It is extremely unlikely that any of the species indicative of sensitivity would be targeted for extraction and we have no evidence for the indirect effects of extraction of other species on this biotope.
Not relevant Not relevant Not relevant Not relevant Not relevant

Additional information

Recoverability
It is likely that Ulva and Cladophora species will have a considerable capacity for recovery. Both species are widespread around the British Isles and Ireland, and may be found in reproductive condition all year round. Numerous motile swarmers (gametes and spores) are released and in the water column they can be dispersed over considerable distances. In addition to recruitment by swarmers, new growth of Cladophora rupestris may arise from the resistant multicellular branching rhizoids (van den Hoek, 1982) that may remain in situ. Recoverability has therefore been assessed to be very high. For instance, after the Torrey Canyon tanker oil spill in mid March 1967, recolonization by sporelings of Ulva and Cladophora species had occurred by the end of April (Smith, 1968). Recovery of the copepod Tigriopus fulvus would be expected to be rapid (presuming a residual or localized population remained from which to recruit) as the species is in reproductive condition all year round and reaches sexual maturity within two months. It also can produce more than one brood from one fertilization. These aforementioned species are characteristic of the biotope, which would be recognized upon their probably rapid re-establishment. Other components of the community, such as the littorinids and other grazers, are of lower abundance owing to physical conditions and are not considered to be characterizing species. However, in their total absence the biotope would be considered to be impoverished. Owing to recruitment of live young in a localized area without a dispersive larval stage recovery of such species may take longer.

Importance review

Policy/Legislation

Habitats Directive Annex 1Reefs

Exploitation

No text entered.

Additional information

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Bibliography

  1. Abou-Aisha, K.M., Kobbia, I., El Abyad, M., Shabana, E.F. & Schanz, F., 1995. Impact of phosphorus loadings on macro-algal communities in the Red Sea coast of Egypt. Water, Air, and Soil Pollution, 83 (3-4), 285-297.

  2. Airoldi, L., 2003. The effects of sedimentation on rocky coast assemblages. Oceanography and Marine Biology: An Annual Review, 41,161-236
  3. Airoldi, L. & Hawkins, S.J., 2007. Negative effects of sediment deposition on grazing activity and survival of the limpet Patella vulgataMarine Ecology Progress Series, 332, 235-240.
  4. Albrecht, A. & Reise, K., 1994. Effects of Fucus vesiculosus covering intertidal mussel beds in the Wadden Sea. Helgoländer Meeresuntersuchungen, 48 (2-3), 243-256.
  5. Albrecht, A.S., 1998. Soft bottom versus hard rock: Community ecology of macroalgae on intertidal mussel beds in the Wadden Sea. Journal of Experimental Marine Biology and Ecology229 (1), 85-109.
  6. Alströem-Rapaport, C., Leskinen, E. & Pamilo, P., 2010. Seasonal variation in the mode of reproduction of Ulva intestinalis in a brackish water environment. Aquatic Botany, 93 (4), 244-249.
  7. Amsler, C.D. & Searles, R.B., 1980. Vertical distribution of seaweed spores in a water column off shore of North Carolina. Journal of Phycology, 16, 617-619.
  8. Archer, A.A., 1963. A new approach to the taxonomy of the branched members of the Cladophoraceae in the British Isles. , Ph.D. thesis, Liverpool University.
  9. Arnold, D.C., 1957. The response of the limpet, Patella vulgata L., to waters of different salinities. Journal of the Marine Biological Association of the United Kingdom, 36, 121-128.
  10. Baden, S.P., Pihl, L. & Rosenberg, R., 1990. Effects of oxygen depletion on the ecology, blood physiology and fishery of the Norway lobster Nephrops norvegicus. Marine Ecology Progress Series, 67, 141-155.
  11. Baeck, S., Lehvo, A. & Blomster, J., 2000. Mass occurrence of unattached Enteromorpha intestinalis on the Finnish Baltic Sea coast. Annales Botanici Fennici, 37, 155-161.
  12. Bokn, T.L., Moy, F.E. & Murray, S.N., 1993. Long-term effects of the water-accommodated fraction (WAF) of diesel oil on rocky shore populations maintained in experimental mesocosms. Botanica Marina, 36, 313-319.
  13. Bonner, T. M., Pyatt, F. B. & Storey, D. M., 1993. Studies on the motility of the limpet Patella vulgata in acidified sea-water. International Journal of Environmental Studies, 43, 313-320.
  14. Bowman, R. S. (1985). The biology of the limpet Patella vulgata L. in the British Isles: Spawning time as a factor determining recruitment sucess. in Moore and Seed (eds) The Ecology of Rocky Coasts, Hodder and Stoughton, London, pages 178{193.
  15. Bowman, R. S. and Lewis, J. R. (1986). Geographical variation in the breeding cycles and recruitment of Patella spp. Hydrobiologia, 142:41-56.
  16. Bowman, R.S. & Lewis, J.R., 1977. Annual fluctuations in the recruitment of Patella vulgata L. Journal of the Marine Biological Association of the United Kingdom, 57, 793-815.
  17. Bryan, G.W. & Gibbs, P.E., 1983. Heavy metals from the Fal estuary, Cornwall: a study of long-term contamination by mining waste and its effects on estuarine organisms. Plymouth: Marine Biological Association of the United Kingdom. [Occasional Publication, no. 2.]
  18. Bryan, G.W., 1984. Pollution due to heavy metals and their compounds. In Marine Ecology: A Comprehensive, Integrated Treatise on Life in the Oceans and Coastal Waters, vol. 5. Ocean Management, part 3, (ed. O. Kinne), pp.1289-1431. New York: John Wiley & Sons.
  19. Burrows, E.M., 1959. Growth form and environment in Enteromorpha. Botanical Journal of the Linnean Society, 56, 204-206.
  20. Burrows, E.M., 1991. Seaweeds of the British Isles. Volume 2. Chlorophyta. London: British Museum (Natural History).
  21. Cabral-Oliveira, J., Mendes, S., Maranhão, P. & Pardal, M., 2014. Effects of sewage pollution on the structure of rocky shore macroinvertebrate assemblages. Hydrobiologia, 726 (1), 271-283.
  22. Cambridge, M., Breeman, A.M., van Oosterwijk, R. & van den Hoek, C., 1984. Temperature responses of some North American Cladophora species (Chlorophyceae) in relation to their geographic distribution. Helgoländer Wissenschaftliche Meeresuntersuchungen, 38, 349-363.
  23. Carlson, R.L., Shulman, M.J. & Ellis, J.C., 2006. Factors Contributing to Spatial Heterogeneity in the Abundance of the Common Periwinkle Littorina Littorea (L.). Journal of Molluscan Studies, 72 (2), 149-156.
  24. Chandrasekara, W.U. & Frid, C.L.J., 1998. A laboratory assessment of the survival and vertical movement of two epibenthic gastropod species, Hydrobia ulvae, (Pennant) and Littorina littorea (Linnaeus), after burial in sediment. Journal of Experimental Marine Biology and Ecology, 221, 191-207.
  25. Christie , A.O. & Evans, L.V., 1962. Periodicity in the liberation of gametes and zoospores of Enteromorpha intestinalis Link. Nature, 193, 193-194.
  26. Clark, M.E., 1968. The ecology of supralittoral rockpools with special reference to the copepod fauna. , Ph.D. Thesis, University of Aberdeen, Scotland.
  27. Clark, R.B., 1992. Marine pollution, 3rd edition. Oxford: Clarendon Press.
  28. Clark, R.B., 1997. Marine Pollution, 4th ed. Oxford: Carendon Press.
  29. Cole, S., Codling, I.D., Parr, W., Zabel, T., 1999. Guidelines for managing water quality impacts within UK European marine sites [On-line]. UK Marine SACs Project. [Cited 26/01/16]. Available from: http://www.ukmarinesac.org.uk/pdfs/water_quality.pdf

  30. Connor, D.W., Allen, J.H., Golding, N., Howell, K.L., Lieberknecht, L.M., Northen, K.O. & Reker, J.B., 2004. The Marine Habitat Classification for Britain and Ireland. Version 04.05. Joint Nature Conservation Committee, Peterborough. www.jncc.gov.uk/MarineHabitatClassification,
  31. Connor, D.W., Brazier, D.P., Hill, T.O., & Northen, K.O., 1997b. Marine biotope classification for Britain and Ireland. Vol. 1. Littoral biotopes. Joint Nature Conservation Committee, Peterborough, JNCC Report no. 229, Version 97.06., Joint Nature Conservation Committee, Peterborough, JNCC Report No. 230, Version 97.06.
  32. Corradi, M.G., Gorbi, G. & Zanni, C., 2006. Hypoxia and sulphide influence gamete production in Ulva sp. Aquatic Botany, 84 (2), 144-150.
  33. Crisp, D.J. (ed.), 1964. The effects of the severe winter of 1962-63 on marine life in Britain. Journal of Animal Ecology, 33, 165-210.
  34. Cullinane, J.P., McCarthy, P. & Fletcher, A., 1975. The effect of oil pollution in Bantry Bay. Marine Pollution Bulletin, 6, 173-176.
  35. Daly, M.A. & Mathieson, A.C., 1977. The effects of sand movement on intertidal seaweeds and selected invertebrates at Bound Rock, New Hampshire, USA. Marine Biology, 43, 45-55.
  36. Davenport, J. & Davenport, J.L., 2005. Effects of shore height, wave exposure and geographical distance on thermal niche width of intertidal fauna. Marine Ecology Progress Series, 292, 41-50.
  37. Davenport, J., Barnett, P.R.O. & McAllen, R.J., 1997. Environmental tolerances of three species of the harpacticoid copepod genus Tigriopus. Journal of the Marine Biological Association of the United Kingdom, 77, 3-16.
  38. Davies, C.E. & Moss, D., 1998. European Union Nature Information System (EUNIS) Habitat Classification. Report to European Topic Centre on Nature Conservation from the Institute of Terrestrial Ecology, Monks Wood, Cambridgeshire. [Final draft with further revisions to marine habitats.], Brussels: European Environment Agency.
  39. Davies, M.S., 1992. Heavy metals in seawater: effects on limpet pedal mucus production. Water Research, 26, 1691-1693.
  40. Davies, S.P., 1970. Physiological ecology of Patella IV. Environmental and limpet body temperatures. Journal of the Marine Biological Association of the United Kingdom, 50 (04), 1069-1077.
  41. Dethier, M.N., 1980. Tidepools as refuges: predation and the limits of the harpacticoid copepod Tigriopus californicus (Baker). Journal of Experimental Marine Biology and Ecology, 42, 99-111.
  42. Diaz, E.R., Kraufvelin, P. & Erlandsson, J., 2012. Combining gut fluorescence technique and spatial analysis to determine Littorina littorea grazing dynamics in nutrient-enriched and nutrient-unenriched littoral mesocosms. Marine Biology, 159 (4), 837-852.
  43. Dodds, W.K. & Gudder, D.A., 1992. The ecology of Cladophora. Journal of Phycology, 28, 415-427.
  44. Ekaratne, S.U.K. & Crisp, D.J., 1984. Seasonal growth studies of intertidal gastropods from shell micro-growth band measurements, including a comparison with alternative methods. Journal of the Marine Biological Association of the United Kingdom, 64, 183-210.
  45. Evans, R.G., 1948. The lethal temperatures of some common British littoral molluscs. The Journal of Animal Ecology, 17, 165-173.
  46. Fortes, M.D. & Lüning, K., 1980. Growth rates of North Sea macroalgae in relation to temperature, irradiance and photoperiod. Helgolander Meeresuntersuchungen, 34, 15-29.
  47. Fraser, J.H., 1936. The occurrence, ecology and life-history of Tigriopus fulvus (Fischer). Journal of the Marine Biological Association of the United Kingdom, 20, 523-536.
  48. Fretter, V. & Graham, A., 1994. British prosobranch molluscs: their functional anatomy and ecology, revised and updated edition. London: The Ray Society.
  49. Gerson, U & Seaward, M.R.D., 1977. Lichen - invertebrate associations. In Lichen ecology (ed. M.R.D. Seaward), pp. 69-119. London: Academic Press.
  50. Goss-Custard, S., Jones, J., Kitching, J.A. & Norton, T.A., 1979. Tide pools of Carrigathorna and Barloge Creek. Philosophical Transactions of the Royal Society. Series B: Biological Sciences, 287, 1-44.
  51. Grenon, J.F. & Walker, G., 1981. The tenacity of the limpet, Patella vulgata L.: an experimental approach. Journal of Experimental Marine Biology and Ecology, 54, 277-308.
  52. Hawkins, S. J. & Jones, H. D., 1992. Rocky Shores. London: Immel.
  53. Hawkins, S.J. & Southward, A.J., 1992. The Torrey Canyon oil spill: recovery of rocky shore communities. In Restoring the Nations Marine Environment, (ed. G.W. Thorpe), Chapter 13, pp. 583-631. Maryland, USA: Maryland Sea Grant College.
  54. Hayden, H.S., Blomster, J., Maggs, C.A., Silva, P.C., Stanhope, M.J. & Waaland, J.R., 2003. Linnaeus was right all along: Ulva and Enteromorpha are not distinct genera. European Journal of Phycology, 38, 277-294.
  55. Hoare, R. & Hiscock, K., 1974. An ecological survey of the rocky coast adjacent to the effluent of a bromine extraction plant. Estuarine and Coastal Marine Science, 2 (4), 329-348.

  56. Hruby, T. & Norton, T.A., 1979. Algal colonization on rocky shores in the Firth of Clyde. Journal of Ecology, 67, 65-77.
  57. Huggett, J. & Griffiths, C.L., 1986. Some relationships between elevation, physico-chemical variables and biota of intertidal rockpools. Marine Ecology Progress Series, 29, 198-197.
  58. Hyslop B.T. & Davies, M.S., 1998. Evidence for abrasion and enhanced growth of Ulva lactuca L. in the presence of colliery waste particles. Environmental Pollution, 101 (1), 117-121.
  59. Hyslop, B.T., Davies, M.S., Arthur, W., Gazey, N.J. & Holroyd, S., 1997. Effects of colliery waste on littoral communities in north-east England. Environmental Pollution, 96 (3), 383-400.
  60. 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,
  61. Jones, W.E. & Babb, M.S., 1968. The motile period of swarmers of Enteromorpha intestinalis (L.) Link. British Phycological Bulletin, 3, 525-528.
  62. Joosse, E.N.G., 1976. Littoral apterygotes (Collembola and Thysanura). In Marine insects (ed. L. Cheng), pp. 151-186. Amsterdam: North-Holland Publishing Company.
  63. Kain, J.M., & Norton, T.A., 1990. Marine Ecology. In Biology of the Red Algae, (ed. K.M. Cole & Sheath, R.G.). Cambridge: Cambridge University Press.
  64. Kamer, K. & Fong, P., 2001. Nitrogen enrichment ameliorates the negative effects of reduced salinity on green macroalga Enteromorpha intestinalis. Marine Ecology Progress Series, 218, 87-93.
  65. Kennison, R.L. & Fong, P., 2013. High amplitude tides that result in floating mats decouple algal distribution from patterns of recruitment and nutrient sources. Marine Ecology Progress Series, 494, 73-86.
  66. Kitching, J.A. & Thain, V.M., 1983. The ecological impact of the sea urchin Paracentrotus lividus (Lamarck) in Lough Ine, Ireland. Philosophical Transactions of the Royal Society of London, Series B, 300, 513-552.
  67. Kraufvelin, P., 2007. Responses to nutrient enrichment, wave action and disturbance in rocky shore communities. Aquatic Botany, 87 (4), 262-274.
  68. Kylin, H., 1917. Kalteresistenze der Meerealen. Bericht der Deutschen Botanischen Gesellschafter, 35, 370-384.
  69. Lüning, K., 1990. Seaweeds: their environment, biogeography, and ecophysiology: John Wiley & Sons.

  70. Lazzaretto, I., Franco, F. & Battaglia, B., 1994. Reproductive behaviour in the harpacticoid copepod Tigriopus fulvus. Hydrobiologia, 292-293, 229-234.
  71. Le Quesne W.J.F. 2005. The response of a protandrous species to exploitation, and the implications for management: a case study with patellid limpets. PhD thesis. University of Southampton, Southampton, United Kingdom.
  72. Lersten, N.R. & Voth, P.D., 1960. Experimental control of zoid discharge and rhizoid formation in the green alga Enteromorpha. Botanical Gazette, 122, 33-45.
  73. Lewis, J.R., 1964. The Ecology of Rocky Shores. London: English Universities Press.
  74. Lewis, S., Handy, R.D., Cordi, B., Billinghurst, Z. & Depledge, M.H., 1999. Stress proteins (HSPs): methods of detection and their use as an environmental biomonitor. Ecotoxicology, 8, 351-368.
  75. Lewis, S., May, S., Donkin, M.E. & Depledge, M.H., 1998. The influence of copper and heat shock on the physiology and cellular stress response of Enteromorpha intestinalis. Marine Environmental Research, 46, 421-424.
  76. Little, C. & Kitching, J.A., 1996. The Biology of Rocky Shores. Oxford: Oxford University Press.
  77. Little, C., Partridge, J.C. & Teagle, L., 1991. Foraging activity of limpets in normal and abnormal tidal regimes. Journal of the Marine Biological Association of the United Kingdom, 71, 537-554.
  78. Littler, M.M., Martz, D.R. & Littler, D.S., 1983. Effects of recurrent sand deposition on rocky intertidal organisms: importance of substrate heterogeneity in a fluctuating environment. Marine Ecology Progress Series. 11 (2), 129-139.
  79. Lobban, C.S. & Harrison, P.J., 1997. Seaweed ecology and physiology. Cambridge: Cambridge University Press.
  80. Lüning, K., 1984. Temperature tolerance and biogeography of seaweeds: the marine algal flora of Helgoland (North Sea) as an example. Helgolander Meeresuntersuchungen, 38, 305-317.
  81. Marchan, S., Davies, M.S., Fleming, S. & Jones, H.D., 1999. Effects of copper and zinc on the heart rate of the limpet Patella vulgata (L.) Comparative Biochemistry and Physiology, 123A, 89-93.
  82. Marshall, D.J. & McQuaid, C.D., 1989. The influence of respiratory responses on the tolerance to sand inundation of the limpets Patella granularis L.(Prosobranchia) and Siphonaria capensis Q. et G.(Pulmonata). Journal of Experimental Marine Biology and Ecology, 128 (3), 191-201.
  83. Marshall, D.J. & McQuaid, C.D., 1993. Effects of hypoxia and hyposalinity on the heart beat of the intertidal limpets Patella granvlaris (Prosobranchia) and Siphonaria capensis (Pulmonata). Comparative Biochemistry and Physiology Part A: Physiology, 106 (1), 65-68
  84. Martinez, B., Pato, L.S. & Rico, J.M., 2012. Nutrient uptake and growth responses of three intertidal macroalgae with perennial, opportunistic and summer-annual strategies. Aquatic Botany, 96 (1), 14-22.
  85. Martins, I., Oliveira, J.M., Flindt, M.R. & Marques, J.C., 1999. The effect of salinity on the growth rate of the macroalgae Enteromorpha intestinalis (Chlorophyta) in the Mondego estuary (west Portugal). Acta Oecologica, 20 (4), 259-265.
  86. McAllen, R., 1999. Enteromorpha intestinalis - a refuge for the supralittoral rockpool harpacticoid copepod Tigriopus brevicornis. Journal of the Marine Biological Association of the United Kingdom, 79, 1125-1126.
  87. McAllen, R., Taylor, A.C. & Davenport, J., 1999. The effects of temperature and oxygen partial pressure on the rate of oxygen consumption of the high-shore rock pool copepod Tigriopus brevicornis. Comparative Biochemistry and Physiology A, 123, 195-202.
  88. Morris, S. & Taylor, A.C. 1983. Diurnal and seasonal variations in physico-chemical conditions within intertidal rock pools. Estuarine, Coastal and Shelf Science, 17, 339-355.
  89. Moss, B. & Marsland, A., 1976. Regeneration of Enteromorpha. British Phycological Journal, 11, 309-313.
  90. Moss, B.L. & Woodhead, P., 1975. The effect of two commercial herbicides on the settlement, germination and growth of Enteromorpha. Marine Pollution Bulletin, 6, 189-192.
  91. Naylor, E. & Slinn, D.J., 1958. Observations on the ecology of some brackish water organisms in pools at Scarlett Point, Isle of Man. Journal of Animal Ecology, 27, 15-25.
  92. Niesenbaum R.A., 1988. The ecology of sporulation by the macroalga Ulva lactuca L. (chlorophyceae). Aquatic Botany, 32, 155-166.
  93. Pedersen, M.F., Borum, J. & Fotel, L. F., 2009. Phosphorus dynamics and limitation of fast and slow-growing temperate seaweeds in Oslofjord, Norway. Marine Ecology Progress Series, 399, 103-115
  94. 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/
  95. Pinn, E.H. & Rodgers, M., 2005. The influence of visitors on intertidal biodiversity. Journal of the Marine Biological Association of the United Kingdom, 85 (02), 263-268.
  96. Povey, A. & Keough, M.J., 1991. Effects of trampling on plant and animal populations on rocky shores. Oikos61: 355-368.
  97. Pyefinch, K. A., 1943. The intertidal ecology of Bardsey Island, North Wales, with special reference to the recolonization of rock surfaces, and the rock pool environment. Journal of Animal Ecology, 12, 82-108.
  98. Raffaelli, D. & Hawkins, S., 1999. Intertidal Ecology 2nd edn.. London: Kluwer Academic Publishers.
  99. Rai, L., Gaur, J.P. & Kumar, H.D., 1981. Phycology and heavy-metal pollution. Biological Reviews, 56, 99-151.
  100. Ranade, M.R., 1957. Observations on the resistance of Tigriopus fulvus (Fischer) to changes in temperature and salinity. Journal of the Marine Biological Association of the United Kingdom, 36, 115-119.
  101. Reed, R.H. & Russell, G., 1979. Adaptation to salinity stress in populations of Enteromorpha intestinalis (L.) Link. Estuarine and Coastal Marine Science, 8, 251-258.
  102. Ribeiro, P.A., Xavier, R., Santos, A.M. & Hawkins, S.J., 2009. Reproductive cycles of four species of Patella (Mollusca: Gastropoda) on the northern and central Portuguese coast. Journal of the Marine Biological Association of the United Kingdom, 89 (06), 1215-1221.
  103. Rice, H., Leighty, D.A. & McLeod, G.C., 1973. The effects of some trace metals on marine phytoplankton. CRC Critical Review in Microbiology, 3, 27-49.
  104. Robles, C., 1982. Disturbance and predation in an assemblage of herbivorous Diptera and algae on rocky shores. Oecologia, 54 (1), 23-31.
  105. Scarlett, A., Donkin, M.E., Fileman, T.W. & Donkin, P., 1997. Occurrence of the marine antifouling agent Irgarol 1051 within the Plymouth Sound locality: implications for the green macroalga Enteromorpha intestinalis. Marine Pollution Bulletin, 38, 645-651.
  106. Sfriso, A., Marcomini, A. & Pavoni, B., 1987. Relationships between macroalgal biomass and nutrient concentrations in a hypertrophic area of the Venice Lagoon. Marine Environmental Research, 22 (4), 297-312.
  107. Shanks, A.L. & Wright, W.G., 1986. Adding teeth to wave action- the destructive effects of wave-bourne rocks on intertidal organisms. Oecologia, 69 (3), 420-428.
  108. Smith, G.M., 1947. On the reproduction of some Pacific coast species of Ulva. American Journal of Botany, 34, 80-87.
  109. Smith, J.E. (ed.), 1968. 'Torrey Canyon'. Pollution and marine life. Cambridge: Cambridge University Press.
  110. Southward, A.J. & Southward, E.C., 1978. Recolonisation of rocky shores in Cornwall after use of toxic dispersants to clean up the Torrey Canyon spill. Journal of the Fisheries Research Board of Canada, 35, 682-706.
  111. Southward, A.J., Hawkins, S.J. & Burrows, M.T., 1995. Seventy years observations of changes in distribution and abundance of zooplankton and intertidal organisms in the western English Channel in relation to rising sea temperature. Journal of Thermal Biology, 20, 127-155.
  112. Storey, K.B., Lant, B., Anozie, O.O. & Storey, J.M., 2013. Metabolic mechanisms for anoxia tolerance and freezing survival in the intertidal gastropod, Littorina littorea. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 165 (4), 448-459.
  113. Sverdrup, H.U., Johnson, M.W. & Fleming, R.H., 1942. The Oceans. New York: Prentice Hall.
  114. Tolhurst, L.E., Barry, J., Dyer, R.A. & Thomas, K.V., 2007. The effect of resuspending sediment contaminated with antifouling paint particles containing Irgarol 1051 on the marine macrophyte Ulva intestinalis. Chemosphere, 68 (8), 1519-1524.
  115. UKTAG, 2014. UK Technical Advisory Group on the Water Framework Directive [online]. Available from: http://www.wfduk.org
  116. Vadas, R.L., Johnson, S. & Norton, T.A., 1992. Recruitment and mortality of early post-settlement stages of benthic algae. British Phycological Journal, 27, 331-351.
  117. Vadas, R.L., Keser, M. & Rusanowski, P.C., 1976. Influence of thermal loading on the ecology of intertidal algae. In Thermal Ecology II, (eds. G.W. Esch & R.W. McFarlane), ERDA Symposium Series (Conf-750425, NTIS), Augusta, GA, pp. 202-212.
  118. Van den Hoek, C., 1982. The distribution of benthic marine algae in relation to the temperature regulation of their life histories. Biological Journal of the Linnean Society, 18, 81-144.
  119. Vaudrey, J.M.P., Kremer, J.N., Branco, B.F. & Short, F.T., 2010. Eelgrass recovery after nutrient enrichment reversal. Aquatic Botany, 93 (4), 237-243.
  120. Vermaat J.E. & Sand-Jensen, K., 1987. Survival, metabolism and growth of Ulva lactuca under winter conditions: a laboratory study of bottlenecks in the life cycle. Marine Biology, 95 (1), 55-61.
  121. Wells, E., Best, M., Scanlan, C. & Foden, J., 2014. Opportunistic Macroalgae Blooming. Water Framework Directive- development of classification tools for ecological assessment., Water Framework Directive-United Kingdom Technical Advisory Group (WFD-UKTAG),

Citation

This review can be cited as:

Budd, G.C. 2002. Green seaweeds (Enteromorpha spp. and Cladophora spp.) in shallow upper shore 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/246

Last Updated: 28/11/2002