This article describes the habitat of the rocky shores. It is one of the habitat sub-categories within the section dealing with biodiversity of marine habitats and ecosystems. It gives an overview about the biota that lives there, the problems and adaptations the habitat is facing with and the importance of it in the marine environment.
A rocky shore is an intertidal area that consists of solid rocks. It is often a biologically rich environment and can include many different habitat types like steep rocky cliffs, platforms, rock pools and boulder fields. Because of the continuously action of the tides, it is characterized by erosional features. Together with the wind, sunlight and other physical factors it creates a complex environment. Organisms that live in this area experience daily fluctuations in their environment. For this reason, they must be able to tolerate extreme changes in temperature, salinity, moisture and wave action to survive.
Sufficient loose or unconsolidated material and a suitable coastal environment to allow the sediments to accumulate are key features in the formation of beaches. The sediments can be terrigenous (land-derived), transported by rivers and streams or through erosion of coastal cliffs. In tropical regions or places with a lack of land-derived material, sediments may be biogenous and consists of broken corals or shells. Beaches on volcanic islands may be black and consists of broken lava and volcanic minerals. Rocky shores are usually steeper than sandy shores. These differences are the result of the different permeability of the sediments and the balance between waves that retreat and come up. Rocky shores have a high permeability and much of the upcoming wave percolates into the bottom. This reduces the outwash of sediment toward the sea. For this reason, coarse sediment can accumulate on the beach and can be transported further on the beach by stronger waves.
Each region on the coast has a specific group of organisms that form distinct horizontal bands or zones on the rocks. The appearance of dominant species in these zones is called vertical zonation. It is a nearly universal feature of the intertidal zone.
When the tide retreats, the upper regions become exposed to air. The organisms that live in this region are facing problems like gas exchange, desiccation, temperature changes and feeding. This upper region is called the supratidal or splash zone. It is only covered during storms and extremely high tides and is moistened by the spray of the breaking waves. Organisms are exposed to the drying heat of the sun in the summer and to extreme low temperatures in the winter. Because of these severe conditions, only a few resistant organisms live here. Common organisms are lichens. They are composed of fungi and microscopic algae living together and sharing food and energy to grow. The fungi trap moisture for both themselves and their algal symbiont. The algae on the other hand produce nutrients by photosynthesis. Green algae and cyanobacteria can also be found on the rocks of the North Atlantic coasts. They are capable of surviving on the moisture of the sea spray from waves. During the winter, they are found lower on the intertidal rocks. The algae growing higher on the rocks gradually die when the air temperature changes. At the lower edge of the splash zone, rough snails (periwinkles) graze on various types of algae. These snails are well adapted to life out of the water by trapping water in their mantle cavity or hiding in cracks of rocks. Other common animals are isopods, barnacles, limpets,…
The intertidal zone or littoral zone is the shoreward fringe of the sea bed between the highest and lowest limit of the tides. The upper limit is often controlled by physiological limits on species tolerance of temperature and drying. The lower limit is often determined by the presence of predators or competing species. Because the intertidal zone is a transition zone between the land and the sea, it causes heat stress, desiccation, oxygen depletion and reduced opportunities for feeding. At low tide, marine organisms face both heat stress and desiccation stress. The degree of this water loss and heating is determined by the body size and body shape. When body size increases, the surface area decreases so the water loss is reduced. Shape has a similar effect. Long and thin organisms dry up much faster than spherical organisms. Intertidal organisms can avoid overheating by evaporative cooling combined with circulation of body fluids. Higher-intertidal organisms are better adapted to desiccation than lower-intertidal organisms, because they encounter more hours of sun. The organisms are exposed directly to the air or they are enclosed in burrows. This results in oxygen depletion, so they can’t get rid of their metabolic waste. A solution for this problem is to reduce the metabolic rate.
The intertidal zone can be divided in three zones:
- High tide zone or high intertidal zone. This region is only flooded during high tides. Organisms that you can find here are anemones, barnacles, chitons, crabs,isopods, mussels, sea stars, snails,...
- Middle tide zone or mid-littoral zone. This is a turbulent zone that is (un)covered twice a day. The zone extends from the upper limit of the barnacles to the lower limit of large brown algae (e.g. Laminariales, Fucoidales). Common organisms are snails, sponges, sea stars, barnacles, mussels, sea palms, crabs,...
- Low intertidal zone or lower littoral zone. This region is usually covered with water. It is only uncovered when the tide is extremely low. In contrast to the other zones, the organisms are not well adapted to long periods of dryness or to extreme temperatures. The common organisms in this region are brown seaweed,crabs, hydroids, mussels, sea cucumber, sea lettuce, sea urchins, shrimps, snails, tube worms,…
Tidal pools are rocky pools in the intertidal zone that are filled with seawater. They are formed where hard or resistant rocks occur and where wave action and other erosional processes have eroded parts of the exposed rocks. This leaves holes or depressions in the substrate where seawater can be collected at high tide. It is important that the rocks are hard enough to survive the regular wave actions, but be sufficiently weak (fractures, bedding differences) to be partly eroded. They can be small and shallow or deep. The smallest ones are usually found at the high intertidal zone, whereas the bigger ones are found near the water. When the tide retreats, the pool becomes isolated. Because of the regular tides, the pool is not stagnant and new water regularly enters the pool. This is necessary to avoid temperature stress, salinity stress, nutrient stress,… Pools that are located higher on the beach are not regularly renewed by tides. These pools are basically freshwater or brackish water communities. It has different characteristics in comparison with other coastal habitats. Several taxa are more abundant in pools than the surrounding environment. These taxa are members of the algae and gastropods. There is also a difference between high and low located pools for the composition. In low located pools, whelks, mussels, sea urchins and Littorina littorea are common. Periwinkles and Littorina rudis are found in high located pools. Other organisms that are commonly found in pools are flatworms, rotifers, cladocerans, copepods, ostracods, barnacles, amphipods, isopods, chironomid larvae and oligochaetes. Vertical zonation also has been documented in tidal pools.
The subtidal zone or sublittoral zone is the region below the intertidal zone and is continuously covered by water. This zone is much more stable than the intertidal zone. Temperature, water pressure and sunlight radiation remain nearly constant. Organisms do not dry out as often as organisms higher on the beach. They grow much faster and are better in competition for the same niche. More essential nutrients are acquired from the water and they are buffered from extreme changes in temperature.  
Problems and adaptations
In this section, the problems and the adaptations are discussed. The continuously changing environment makes that organisms have to be tolerant for these changes. Adaptations are a solution for these problems and are necessary to survive.
Intertidal organisms are regularly exposed to air and water. Air differs physically from seawater in diverse and important features. This influences the ability to exchange gas and their overall thermal balance with the surrounding environment. The chemical and gaseous constituents of the atmosphere have changed dramatically through geological times. Because of the much greater density of water, organisms become buoyant. In air, the gravity induces retraction of tentacles and other feeding organs. It also makes the body less resistant. For this reason, organisms need supporting structures when they are exposed to air. Attachment and body changes are also required. When exposed to the air, the organisms directly absorb solar radiation. The buffering capacity of water, because of the high rate of heat conductivity, disappears and the body temperature increases. In contrast to this, heat loss is much lower in air than in water. An adaptation to heating is the vaporization of internal water reserves.
Sunlight is another parameter that influences the organisms. When there is too much sunlight, organisms dry out and the capacity to capture light energy can be weakened. The light that is not used or dissipated can cause damage to subcellular structures. Too little sunlight reduces the growth and reproduction of the organism, because photosynthesis is reduced. Algae can avoid absorbing too much light by changing the complement or amount of pigments they produce. They also can rearrange the pigmented organelles within their cells. When free radicals are produced from an excess of light, they can be scavenged and deactivated.
The intertidal zone can experience extreme temperature changes. The organisms in this zone must be resistant to these changes to survive. Most of the marine organisms are ectothermic and need the warmth from the environment to survive. When the organisms are submerged, they are buffered against temperature changes, because the water is isothermal. When the organisms become exposed to the air, they can experience cool or warm temperatures. When the temperature is too low, the organisms must cope with physiological threats associated with cold stress. This can be the case in polar and temperate latitude coastal zones. The body fluids can then reach their freezing point and ice crystals develop. This causes damage to cell membranes and increasing the osmotic concentration of the remaining fluids. To avoid this cold stress, organisms can migrate to habitats that are more suitable. This can be a problem for sessile organisms. They can develop physiological and behavioral adaptations such as gaping shells (mussels). Some organisms have developed antifreeze proteins. Increasing the concentration of small osmolytes such as glycerol in the body fluids can decrease the freezing point. Another strategy is to control ice crystal formation. Organisms can control the speed and the exact location of the ice crystals. When the ice formation is intracellular, it is lethal but extracellular ice formation can be tolerated. When the temperature is too high, heat stress appears. Heat stress accelerates rates of metabolic processes. This can be avoided by evaporative cooling combined with circulation of body fluids.
Salinity stress can occur in the external medium and in surface films. The concentration of the fluids determines whether or not the organism will lose water. When the osmolality of the cell is lower than the surrounding medium, the cell loses water from the internal fluids to the environment (hyperosmotic stress). When the intracellular osmolality is higher than the environment, there is an influx of water into the cell from the environment (hypoosmotic stress). Multicellular organisms respond to this salinity stress by compartmentalization. This buffers the cells from sharp changes in the osmotic environment. When the tissue has an immediate contact with the external medium, a solution can be to regulate intercellular osmotic pressure by actively excreting salts or water. Another solution is to change the internal osmolality. This can be done by incorporating ions or compatible solutes in the internal fluids.
Organisms are threat by desiccation during emersion at low tides or when they are positioned in the high intertidal zones. Dehydratation due to evaporative water loss is the most common mechanism. Highly mobile organisms can avoid the desiccation by migrating to a region that is more suitable. Less mobile organisms restrict various activities (reduced metabolism) and attach more firmly to the substrate. Physiological features to tolerate water loss are desiccation-resistant egg cases, reduction in water permeability of membranes, accumulation of metabolic end products, reduction of metabolic and developmental rates, maintenance of intracellular osmolytes and gene expression for production of protective macromolecules.
A wide variety of strategies to escape from predation exists. The first strategy is calcification. It makes it more difficult for the predator to eat these organisms. This strategy is applied by algae. It makes them tougher and less nutritious. A second one is the production of chemicals, usually produced as secondary metabolites. These chemicals can be produced all the time such as toxins, but other chemicals are only produced in response to stimuli (inducible defence). Another way to avoid predation is to have two distinct anatomical forms within one life cycle. This can be e.g. an alternation between a crusty form when the predator is present and a more delicate form (e.g. blade) when the predator is absent. Also the shape of the body can be a distinct evolutionary advantage. Bioluminescence is another strategy to avoid predators. Many intertidal and subtidal predators visually forage. The light is used for warning, blinding, making scare, misleading or attracting the predator. A commonly used form of protection against predation is camouflage. This can be visually or chemically. Visual camouflage means that the prey becomes invisible to the predator by using the same colors as the environment. Chemical camouflage is the passive adsorption of chemicals. The predator does not smell the prey anymore, because the smell is masked.
One way to protect organisms from waves is permanent attachment. But this strategy can not be used by organisms that have to move to feed themselves. These organisms have to make a compromise between mobility and attachment. Another way to be protected is to burrow themselves into the sediment. But an alternative is to seek protected habitats. Attachment can be done by different structures. Bivalves usually use threads (byssal threads) to attach to rocky surfaces or to other organisms. But it can also be done by a foot. Another one is cementation. This is the case for bivalves such as oysters, scallops and some other forms. They lay on their side, with the lower valve cemented firmly to the bottom. This can be combined by reduction or enlargement of certain muscles.  
Why are rocky shores important?
- Providing a home for a lot of organisms
- Nursery area for many fish and crustacean species
- Shelter in areas where seaweeds break the waves power
- Providing food for fishes
- Algal beds important food source for rare and threatened species like sea turtles
- Feeding ground at low tide for wading birds
- Stabilization inshore sediment
- http://www.marbef.org – Sprung M.
- Knox G.A. 2001. The ecology of seashores. CRC Press LLC. p. 557
- Karleskint G. 1998. Introduction to marine biology. Harcourt Brace & Company. p.378
- Levinton J.S. 1995. Marine biology: function, biodiversity, ecology. Oxford university press. p.420
- Denny M.W. Gaines S.D. 2007. Encyclopedia of tidepools & rocky shores. University of California Press. p. 705
- Levinton J.S. 1995. Marine biology: function, biodiversity, ecology. Oxford University Press. p. 420
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