|Basic Information||Biotope classification||Ecology||Habitat preferences and distribution||Species composition||Sensitivity||Importance|
Image Tom Mercer - Seaweeds in sediment (sand or gravel)-floored eulittoral rockpools. Image width ca 1 m.
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LR.FLR.Rkp.SwSed recorded () and expected () distribution in Britain and Ireland (see below)
The rockpool environment varies depending on factors such as pool depth, surface area, volume, orientation to sunlight, shading, internal topography, sediment content and type, together with wave exposure, shore height, and hence flushing rate, and the presence of absence of freshwater runoff. As a result, no two rockpools exhibit exactly the same physio-chemical conditions and exhibit large spatial variation in community structure, even between adjacent pools at the same shore height (Ganning, 1971; Metaxas & Scheibling, 1993; Metaxas et al., 1994). Any given rockpool is effectively unique (Metaxas & Scheibling, 1993). Therefore, while the organisms detailed below have been recorded in this rockpool biotope, not all, or in some case few, of the organisms, and hence the relationships between them, may occur in any one rockpool.
As communities in rockpools remain constantly submerged and the danger of desiccation is absent, it might be expected that rockpools provide 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, the rockpool environment differs from that of the surrounding emergent rock surfaces, so that not all species that thrive on the surrounding rock occur abundantly in rockpools and much of the lower shore open rock fauna is absent from rockpools (Lewis, 1964). Rockpools constitute a distinct environment for which physiological adaptations by the flora and fauna may be required (Lewis, 1964; Metaxas & Scheibling, 1993). The following description is based on reviews by Lewis (1964), Ganning (1971) and Metaxas & Scheibling (1993), the species listed in the MNCR database (JNCC, 1999) and additional references as cited. Macroalgae such as kelps, fucoids, red and green algae, erect and encrusting corallines provide primary productivity either directly to grazing invertebrates and fish or indirectly, to detritivores and decomposers, in the form of detritus and drift algae or as dissolved organic material and other exudates. Benthic microalgae and phytoplankton (e.g. diatoms) also add to primary productivity.
Where present, large macroalgae such as Halidrys siliquosa and laminarians (e.g. Laminaria digitata, Saccharina latissima and Saccorhiza polyschides) and fucoids (e.g. Fucus serratus, Fucus vesiculosus) shade the substratum (depending on density) so that understorey plants tend to be shade tolerant red algae. Understorey algae, by effectively restricting access to the substratum, may also inhibit or restrict recruitment of other species of macroalgae (Hawkins & Harkin, 1985; Hawkins et al., 1992).
Macroalgae compete for space with sessile invertebrates such as sponges, hydroids, ascidians and bryozoans.
Macroalgae provide substrata and refuges for a variety of invertebrates and epiphytic algae. The stipes and lamina of Laminaria spp. may support bryozoans (e.g. Membranipora membranacea or Electra pilosa) and grazing blue-rayed limpets (Helcion pellucida), while their holdfasts provide additional refuges for meiofauna and small invertebrates. If present, the stipes of Laminaria hyperborea may support numerous epiphytes such as Palmaria palmata, Phycodrys rubens and Cladophora rupestris (Goss-Custard et al., 1979). Where present, Halidrys siliquosa provide substratum for epiphytes, depending on location, including microflora (e.g. bacteria, blue green algae, diatoms and juvenile larger algae), Ulothrix and Ceramium sp., hydroids (e.g. Obelia spp.), bryozoans (e.g. Scrupocellaria spp.), and ascidians (e.g. Apilidium spp., Botryllus schlosseri, and Botrylloides leachi) (Lewis, 1964; Moss, 1982; Connor et al., 1997a).
The macroalgae provide refuges for small invertebrates, such as isopods, amphipods, ostracods and copepods. Corallina officinalis provides a substratum for small spirorbids e.g. Spirorbis corallinae, which is only found on Corallina officinalis. Increasing density of Spirorbis corallinae was shown to increase the species richness of the epiphytic fauna (Crisp & Mwaiseje, 1989). The invertebrate fauna of Corallina officinalis is detailed in £ELR.Coff£.
Amphipods, isopods (e.g. Idotea granulosa) 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 also graze the macroalgae but do not normally adversely affect the canopy (Brawley, 1992b).
Grazers of periphyton (bacteria, blue-green algae and diatoms) or epiphytic algae include harpacticoid copepods, the limpets Patella vulgata and Patella ulyssiponensis, the blue-rayed limpet Helcion pellucidum, and gastropods such as Gibbula cineraria, Gibbula umbilicalis, Littorina saxatilis, Littorina littorea, Littorina obtusata and Rissoa spp. Limpets and littorinids also graze macroalgal sporelings and green algae especially.
Coralline algae are probably relatively grazing resistant (Littler & Kauker, 1984) and few species graze the corallines directly except perhaps chitons and limpets of the genus Tectura.
Grazing by littorinids and gammarid amphipods has been shown to significantly affect macroalgal abundance and diversity. For example in cage experiments in littoral fringe pools, Parker et al. (1993) found that gammarid amphipods significantly reduced the erect macroalgal canopy, while littorinids grazed microalgae and macroalgal sporelings, and prevented the establishment of erect and encrusting algal canopies. Both groups reduced the species richness of the algal canopy (Parker et al., 1993). In tidepools in Nova Scotia, Chapman (1990) and Chapman & Johnson (1990) reported that grazers (especially littorinids) reduced the abundance of Fucus spp. sporelings and juveniles but increased the abundance of ephemeral algae, while having no effect on the encrusting red alga Hildenbrandia rubra (Metaxas & Scheibling, 1993). Conversely, Lubchenco (1978) noted that the addition of littorinids to mid-shore pools in Massachusetts decreased the abundance of dominant Ulva spp. (as Enteromorpha spp.) in favour of Chondrus crispus (Metaxas & Scheibling, 1993) a less palatable red alga. Wolfe & Harlin (1988a) noted that Rhode Island tidepools with the highest littorinid densities had the lowest abundance of macroalgae. Similarly, removal of the limpet Patella vulgata from high tidal pools at Lough Ine resulted in an increased abundance of Ulva (as Enteromorpha) sp. (Goss-Custard et al., 1979). Where present, suspension feeders include barnacles (e.g. Semibalanus balanoides), the mussel Mytilus edulis, hydroids, tubeworms ( e.g. Spirorbis spp. and Pomatoceros spp.), ascidians, bryozoans and sponges. However, the abundance of barnacles and mussels in rockpools is usually low (Lewis, 1964), presumably due to heavy predation on juveniles by the dog whelk Nucella lapillus and crabs (e.g. Carcinus maenas and Cancer pagurus).
The sediment provides habitat for deposit feeding annelids e.g. Arenicola marina and terebellids.
Invertebrate predators include turbellarians and nemerteans feeding on small invertebrates such as copepods and small gastropods. Lower to mid shore rockpools provide refugia for dog whelks Nucella lapillus feeding on barnacles and small mussel within the rockpool and /or leaving the rockpool to forage at high tide. Similarly, crabs such as Carcinus maenas and Cancer pagurus are generalist predators of gastropods (e.g. littorinids) and bivalves as well as scavengers. Passive carnivores include sea anemones such as Anemonia viridis, Actinia equina and Urticina felina. In addition, intertidal fish such as the shanny Lipophrys pholis and gobies prey on small invertebrates such as copepods, amphipods and isopods.
As with grazing, predation pressure is potentially higher in mid to low shore rockpools, since predators can continue feeding irrespective of the state of the tide. For example, in New South Wales, whelks were shown to reduce the abundance of barnacles, tubeworms and limpets (Fairweather, 1987; Metaxas & Scheibling, 1993). Dethier (1984) concluded that harpacticoid copepod abundance in mid to low shore pools was low due to the presence of predators such as fish and to a lesser extent anemones. The reduced abundance of barnacles and mussels observed in rockpools (Lewis, 1964) is probably partly due to increased predation pressure.
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.
The temperature of pools fluctuates with air temperature and sunlight, and tend to warm throughout the day, especially if in direct sunlight (Daniel & Boyden, 1975; Goss-Custard et al., 1979). Shaded pools exhibit less dramatic changes in temperature (Daniel & Boyden, 1975). For example, the temperature of an high shore pool exposed to direct sunlight rose quickly in the morning to a maximum of 25 °C, while a shaded high shore pool only rose by 1 °C, even though air temperatures reached 20 °C (Daniel & Boyden, 1975). In addition, deeper pools may become stratified, with warmer water near the surface and cooler near the bottom (Daniel & Boyden, 1975), primarily due to sunlight. They noted that pool temperatures remained almost constant at night and suggested that pool temperatures would fluctuate slowly during the day under overcast conditions. In deeper pools, the vertical temperature gradation present in summer, may reverse during winter owing to density stratification, so that ice may form (Naylor & Slinn, 1958). Morris & Taylor (1983) reported warmer water at the bottom of the pools and cooler at the surface, which they attributed to cooling of the surface water by wind. Examples of temperature ranges reported for mid to low shore pools include annual maxima and minima of 1-25 °C and 2-22 °C (Morris & Taylor, 1983), a diurnal range of 24 °C (day) and 13 °C (night) for a mid shore pool (Daniel & Boyden, 1975), and surface water temperature ranges of 14-19.25 °C and 15.5-20.75 °C in mid shore pools (Pyefinch, 1943). Smaller (or shallow pools) are more affected by climatic change in temperature than larger and/or deeper ones (Ganning, 1971). Morris & Taylor (1983) noted that temperature showed the greatest seasonal variation of all the physical parameters examined. In summer, the minimum recorded temperatures were greater than the maximum temperatures recorded in winter, and the daily temperature ranges were greater in summer than in winter, in both high and low shore pools (Morris & Taylor, 1983).
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. In addition, freezing of surface water increases the salinity of the underlying water (Ganning, 1971; Metaxas & Scheibling, 1993). Daniel & Boyden (1975) and Morris & Taylor (1983) reported little variability in salinity over one tidal cycle, and Ganning (1971) suggested that changes in salinity were of limited importance. However, heavy rain resulted in a layer of low salinity water on the surface of pools. Morris & Taylor (1983) stated that a low salinity layer of 2-10 mm was normal but after one storm the low salinity layer increased in depth, eventually resulting in a homogeneous pool of brackish water. Morris & Taylor (1983) reported an annual salinity range in mid to low shore pools of 26-36.5 ppt.
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). 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 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 (Pyefinch, 1943; Ganning, 1971; Daniel & Boyden, 1975; Goss-Custard et al., 1979; Morris & Taylor, 1983; Metaxas & Scheibling, 1993). In summer, rockpools are likely to be supersaturated with oxygen during the day (Pyefinch, 1943). For example, the greatest range of oxygen saturation of 101.7% occurred in a seaweed dominated, sediment floored pool, which reached over 190%saturation on some days (Pyefinch, 1943). Daniel & Boyden (1975) noted that a mid shore, seaweed dominated pool reached 194% saturation (ca 15 mg O2/l) but that oxygenation was also marked in shaded pools. A pool with dense fauna exhibited a maximum saturation of 210% (Pyefinch, 1943). 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 as respiration utilizes much of the available oxygen and pH decreases. Daniel & Boyden (1975) reported oxygen depletion at night, with mid to low shore pools reduced to 8-44% saturation. They noted that the crab Carcinus maenas leaves the pools at night, and that other species with the ability to air-breathe could also do so, e.g. limpets, littorinids, and the shanny Lipophrys pholis. They also observed that shrimps gathered at the edge of high shore pools at night, presumably to take advantage of the better oxygenated surface layer (Daniel & Boyden, 1975). Daily fluctuation in oxygen concentration and pH also vary seasonally, and with the height of the pool on the shore or pool depth (Daniel & Boyden, 1975; Morris & Taylor, 1983; Metaxas & Scheibling, 1993). Morris & Taylor (1983) noted variation in oxygen concentration with pools with pool depth and proximity to pool algae. Again, inundation by the rising tide causes sudden changes in oxygenation, either sudden drops during the day or increases at night.
Ganning (1971) noted that the temperature of sediment at the bottom of pools showed little variation in temperature, and noted that diurnal fluctuations of greater than 1 °C were rarely observed in bottom sediments of pools deeper than 0.5 m, although a temperature of fall 0.5-1 °C was observed at the water/sediment interface. The presence of sediment in pools increases the risk of scour, which is likely to vary seasonally, increasing in winter storms. In addition, anoxic conditions within the sediment could potentially cause significant and sudden decreases in oxygen levels if the sediment was stirred up by wave action, although no evidence of this effect was found.
Seasonal change in communities
Tidepool community structure has been show to vary markedly over time, particularly with season (Metaxas & Scheibling, 1993). However, most studies have examined tidepools overseas, or different tidepools communities to those that occur in this biotope (Dethier, 1984; Wolf & Harlin, 1988a,b; Kooistra et al., 1989; Metaxas et al., 1994) so that the species concerned are very different. Seasonal changes in macroalgal cover and diversity were associated with changes in water temperature, light intensity and day length, and reduced grazing pressure from littorinids in winter, although the community types were relatively stable over time in Rhode Island pools (Wolfe & Harlin, 1988a,b). Metaxas et al. (1994) noted that sheet forming algae (e.g. Ulva lactuca) were found throughout the year, filamentous forms (e.g. Cladophora and Ceramium) were present in late spring and summer (although other studies found them to be present from late spring to late autumn) while thick leathery and encrusting forms did not vary seasonally. However, macroalgal diversity was lowest in summer and autumn, especially in mid shore pools, probably due to grazing. Metaxas et al. (1994) also noted that the abundance of mussels, littorinids and whelks in Rhode Island peaked in summer due to recruitment but varied significantly between pools.
Red algae exhibit seasonal variation in growth and reproduction and red algal turf declines in abundance during the winter months, partly due to die back and abrasion during winter storms. Although protected from wave action in deep pools, macroalgae will be particularly susceptible to damage and abrasion by wave action and winter storms in shallow sediment filled pools. For example, maximum growth of Furcellaria lumbricalis occurs in March/April (Austin, 1960b) and release of carpospores and tetraspores occurs in December/January (Bird et al., 1991). Reproductive bodies are present on the gametophytes of Ahnfeltia plicata between July and January and mature carposporophytes occur between October and July (Maggs & Pueschel, 1989). However, in the Bristol Channel, Bamber & Irving (1993) noted that the biomass of Corallina officinalis increased steadily through spring and summer and began to decline after July. Mastocarpus stellatus (as Gigartina stellata) was reported have a perennial holdfast, losing many erect fronds in winter, which grow back in spring (Dixon & Irvine, 1977). Osmundea pinnatifida also shows seasonal variation in growth, expanding its perennial holdfast in June to September, and producing erect fronds from October onwards reaching a maximum in February to May (Maggs & Hommersand, 1993). 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 depending on grazing pressure. In summer, erect and encrusting corallines may be bleached (especially in shallow pools) 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).
In deep pools the underlying rock is likely to be covered by encrusting corallines. Large macroalgal species (e.g. kelps and fucoids) may dominate the surface of the pool. Their depth within the pool (vertical zonation) is limited by self-shading so that only corallines and red algae occur beneath them. The interface between the bottom sediment and the rock surface is likely to support only sand resistant red algae and fauna, e.g. sand-tolerant algae such as Furcellaria lumbricalis, Polyides rotundus, Ahnfeltia plicata, Rhodothamniella floridula, and the anemone Urticina felina. The sediment may support infauna such as lugworm (e.g. Arenicola marina), the sand mason worm Lanice conchilega, terebellids and meiofauna. The upper limit of some species of algae within the pool may be limited by the summer surface water temperatures, and or desiccation after evaporation (e.g. corallines). Grazing intensity due to littorinids may also affect the abundance of fleshy macroalgae, so that the pools may be dominated by less palatable red algae (e.g. Chondrus crispus and Mastocarpus stellatus). Vertical surfaces within deep pools, and crevices or overhangs present, are likely to be dominated by encrusting fauna e.g. the sponges Halichondria panicea and Hymeniacidon perleve, tubeworms and anemones. The surface of larger stones and pebbles may support tubeworms and the holdfasts of kelps or Chorda filum. The holdfasts of kelps and fucoids, and fronds of filamentous species and erect corallines provide refuges for small invertebrates (e.g. amphipods, isopods and small gastropods) or meiofauna (e.g. copepods) (see ecological relationships above). In addition, the shade of macroalgae provide refuges for shrimps (e.g. Palaemon spp.) and intertidal fish (e.g. blennies and gobies), while crevices and underboulder habitats provide additional refuges for crabs.
Rockpool species also display zonation patterns, similar to the emergent species. For example, brown algae and corallines are usually dominant in mid to low shore pools, while green algae tend to dominant high shore pools (Metaxas & Scheibling, 1993). Kooistra et al. (1989) noted vertical zonation within pools and found that macroalgal communities could be allocated to different depths within pools in the lower or higher parts of the shore. However, the communities studied in Brittany differed markedly from those found in this biotope. Similarly, zonation patterns have also been reported in flatworms, rotifers, oligochaetes, cladoceans, copepods, ostracods, barnacles, amphipods, isopods, chironomid larvae and fish (see Metaxas & Scheibling, 1993). Littorina littorea, mussels, whelks, limpets and sea urchins tend to dominate in lower shore pools, while other littorinids dominate higher on the shore (Metaxas & Scheibling, 1993). Nevertheless, Metaxas et al. (1994) noted that horizontal spatial variability between pools within the same shore height appeared to be as great as variability along the intertidal gradient, and suggested that the physical setting of the pool may be of primary importance in determining the macroalgal abundance. Dethier (1984) examined the effect of natural disturbance rockpool communities in the coast of Washington State. She noted that disturbance such as heat stress in summer and wave action in winter occurred regularly (ca 1.6 times per year per pool). The observed disturbances affected dominant species, so that no one dominant species could occupy all the pools within the tidal range at any one time. None of the tidal pool assemblages observed were stable over many generations and disturbances resulted in a mosaic of species assemblages within pools in any one region (Dethier, 1984).
Information specific to the community was not found but Workman (1983) gave an estimate of primary production by microalgal films on the lower shore in the British Isles to be in the region of 100 g C/m²/yr, much of which will be utilized directly by grazers, while primary productivity for fucoids on sheltered shores was estimated to be 1250 g C/m²/yr (Hawkins et al., 1992) and for encrusting corallines to be 1000 g C/m²/yr (Dawes et al., 1991; Raffaelli & Hawkins, 1999). Ganning & Wulff (1970) reported primary productivity values in terms of gross photosynthesis of between 2 and 3.5 O2/m3/ hr in brackish water rock pools dominated by green algae. Overall, deep rockpools with abundant macroalgae are likely to be highly productive mesocosms on the shore. However, shallower pools, with only sparse macroalgal cover due to sediment scour are likely to be far less productive.
Dethier (1984) noted that few rockpool populations, even of dominant species, remained static over time, based on long-term observations over several years. On the coast of Washington State, partial disturbance (a reduction in abundance or cover) resulted in relatively rapid recovery of the community for example; encrusting corallines recovered in over 2 months and erect corallines attained 87% of their original cover in 2 years. The red algae Rhodomela sp. exhibited 39% recovery from total loss after 2 years and Cladophora sp. exhibited 77% recovery after total loss. Dethier (1984) concluded that disturbance was a factor that resulted in a mosaic of different communities in rockpools within an area and that, at any point in time, separate rockpool communities were probably in different stages of recovery.
This review can be cited as follows:
Tyler-Walters, H. 2005. Seaweeds in sediment (sand or gravel)-floored eulittoral rockpools. Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 25/01/2015]. Available from: <http://www.marlin.ac.uk/habitatecology.php?habitatid=326&code=2004>