Described as inhospitable environment Exposure to the Elements e.g. Wind and Wave Action Lack of Shelter Ice (geographical location) But in some cases Very Rich Diversity of Organisms Rocky Shore Communities
Zonation (horizontal banding) Plants and Animals Universal Patterns Identified A dark belt of brown seaweeds, a white belt of barnacles and a black belt of lichens dominate the lower, middle and upper shore levels respectively. Rocky Shore Communities
Notice that the flora are dominated by brown seaweeds. At the bottom of the shore, exposed to air only during short periods of low water during Spring tides, are 'kelps' or laminarians: in this case, species of the genus Laminaria which can grow to several metres in length; a tough stipe supports a floppy frond. Above this, between MLW and MHW, the main seaweeds are 'wracks' or fucoids, especially species of the genus Fucus but also Ascophyllum nodosum and Pelvetia canaliculata. Seaweed size tends to grow smaller towards the top of the shore. This is in part a result of tougher conditions here, but also a result of interactions amongst species.
Rocky Shore Communities Where do red and green seaweeds fit into the picture of zonation? According to an old theory, the zonation of seaweeds was a response to the changing colour of light penetrating the sea. Thus green seaweeds were found at the top of the shore, where average water cover was small and light close to white; brown seaweeds at greater depth, adapted to green light; and red seaweeds at greatest depth, adapted to the prevailing blue colour of light penetrating here. However, a better explanation is this. Red seaweeds are less tolerant of dessication and less good at competing with brown seaweeds on shores on the west coast of Scotland, so they tend to be found as an understory, or in rock pools, or growing on other seaweeds. Green seaweeds such as Enteromorpha are not very resistant to dessication and are heavily grazed, but they can grow quickly, and seem to fluorish in regions of nutrient enrichment - e.g. where sewage pipes cross the beach.
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Rocky Shore Communities Communities vary from shore to shore despite vertical zonation Wave exposure Zonation more obvious where fewer plants e.g. barnacles Ballantines scale (figure)
Rocky Shore Communities Sheltered versus exposed (protection versus wave action) Increasing exposure – brown algae to barnacles, mussels and red algae In-between shores – mosaic of fucoids, barnacles and bare rock Pattern varies around the World though
Rocky Shore Communities Also differences in vertical zones with different exposures (Figure 3.2) Distributions of flora and fauna are not just the result of physical factors Wave forces are responsible for some organisms but not for all
Rocky Shore Communities Differences in environment e.g. water temperature and turbidity Some organisms exist in more than one type of environment, but show physical differences in size and form e.g. kelp, algae etc… Physical and biological factors are responsible
Rocky Shore Communities Wave action both influences structure and community composition by eliminating organisms that can not withstand the waves, but also alters the balance of competition, predation, and grazing pressures Difficult to devise a general model Other factors that need to be taken into account are: rock type and texture etc…. Localised features such as crevices, gullies, caves etc.
Rocky Shore Communities Littoral ecosystems Food webs Two examples provided by Little and Kitching (Figures 8.1a and b) Show how communities work – albeit simplified Provides a basis for questions to be asked Are communities stable or changeable?
Rocky Shore Communities
On sheltered shores algal-dominated communities are stable Moderately exposed shores are much more variable Exposed shores show little overall variation but small scale mosaic patterns change frequently
Rocky Shore Communities The changes that take place are deemed to be a combination of both physical and biological interactions Disturbance lies at the root of the change For example, wave action leads to physical removal Mobile ice in different climates; desiccation; Sand Scour etc… Result is often a mosaic community
Rocky Shore Communities Competition also influences the community structure Interspecies interactions affect succession and therefore structuring of communities Competition for space, food Balance between physical and biological factors
Rocky Shore Communities Also Predation, Grazing and Recruitment Paines hypothesis of Keystone Species that can free up space for others But... not all predators are dominant because of recruitment of prey; physiological limitations of predator; prey may have refuge from predator; resistance to predator (chemicals or behaviour patterns)
Rocky Shore Communities Grazing – different because it is a different process and involves different species Recruitment – miniature adults important in the density of populations e.g. larval supply
Rocky Shore Communities Macroalgae and Microalgae Dominate on some shores Brown, green and red Blue-green, diatoms etc. Microalgae more important in the food chain than macroalgae Microalgae have high rate of production
Rocky Shore Communities Distribution varies according to exposed shore etc. and also locally Exposed shore less evident than sheltered shore Variety of species always increases towards the sub-littoral zone Green algae less abundant: Enteromorpha, Ulva, Cladophora
Rocky Shore Communities Primary production of algae is substantial Eaten by grazers (but less of the larger sizes) Large plants end up as detritus Algal defences: growing out of reach; temporal escape (growing or fruiting when grazing pressure is least); deter grazers through structural adaptations; chemical
Rocky Shore Communities Brown Algae Fucoids and Laminarians or Kelps Different habitat and distribution Laminarians are affected by wave action, competition and sea urchins
Rocky Shore Communities Green Algae Enteromorpha for example occurs anywhere on the shore, but usually high-shore, Ulva on the lower shore, as is Cladophora Generally sheltered environments Very frequent and rapid reproduction aids colonization
Rocky Shore Communities Red Algae Very varied form: frondose and branching, filamentous, encrusting Live low on the shore (but some at top) Some live in tidal pools Like brown and green algae contain chlorophyll a, but also red pigments – phycoerythrins (absorb blue and green light)
Rocky Shore Communities Microalgae Provides most of the food that the limpets, winkles, topshells and other herbivores survive on Evidence of vertical zonation Different diatom assemblages on different substrate Effects of grazers at different times of the year
Rocky Shore Communities Grazers or Herbivores Live algae Dead algae (detritus) are detritivores Molluscan grazers most prominent on rocky shores Major control on algal vegetation Lower shore – sea urchins Mesograzers – amphipods and isopods overall Limpets and Winkles and many others specialise in intertidal life
Rocky Shore Communities Grazers may determine community structure Intertidal Subtidal Limpets: widespread, at all intertidal levels, adaptable, strong, resist wave attack and predators, demonstrate homing and territoriality, different feeding times Can be affected by desiccation Predators deterred by Stomping and Mushrooming
Rocky Shore Communities Winkles Common around the world Very adaptable Evidence of Zonation Preference for algal foods Others are: Topshells, Mesograzers (amphipods and isopods), Sea Urchins
Rocky Shore Communities Suspension Feeders Barnacles and bivalves (mussels) on wave- exposed shores Largely sessile Many others around the world Many man-made substrates – docks, ship hulls, offshore platforms Fouling communities
Rocky Shore Communities Rocky shores suspension feeders are sessile Depends on local conditions for supply of food and nutrients Tolerate conditions of flow, wave action, desiccation, Mussels, Barnacles, Polychaetes, Sea Anemones,
Rocky Shore Communities Interaction between species For example, efficiency of grazers e.g. limpets Competition But differences exist between sheltered and exposed shores
Rocky Shore Communities Habitat-modifying, dominant species such as algaes, seagrass, corals, and mussels are known as ''foundation species' or 'ecosystem engineers' because they provide habitat for a high diversity of species. This is achieved through increasing the heterogeneity of habitat, reducing the water flow, stabilising the substrate, increasing sedimentation, reducing light, and providing substrate for species to live on.
Rocky Shore Communities Another example of the effects of grazing is illustrated by following the events that occurred after the Torrey Canyon oil spill in England in Beaches were treated with 10, 000 tonnes of oil dispersants, which proved, however, more toxic than the oil to most sea-shore life. Most animals on the shore were killed, imcluding many Patella (limpets) which were normally present in large numbers. There followed a settlement of the green seaweeds Ulva and Enteromorpha spp. During the late summer and autumn of 1967, Fucus vesiculosus began to appear, and in places the large brown seaweeds Laminaria digitata and Himanthalia grew m further upshore than normal. Animal life was still reduced in number during 1968, but a few barnacles settle in areas free from Fucus. Patella settled and removed much of the Fucus in , and there was a return to the limpet-barnacle community by
Rocky Shore Communities BENEDDETTI-CECCHI, L. (2001). Variability in abundance of algae and invertebrates at different spatial scales on rocky shores. Marine Ecology Progress Series 215, BROEKHUYSEN, G.J. (1940). A preliminary investigation of the importance of desiccation, temperature and salinity as factors controlling the vertical distribution of certain intertidal marine gastropods in False Bay, South Africa. Transaction of the Royal Society of South Africa 28, GIBBONS, M.J. (1988). The impact of wave exposure on the meiofauna of Gelidium pristoides (Turner) Keutzing (Gelidiales: Rhodophyta). Estuarine, Coastal and Shelf Science 27, GIBBONS, M.J. & GRIFFITHS, C.L. (2000). A comparison of macrofaunal and meiofaunal distribution and standing stock across a rocky shore, with an estimate of their productivities. Marine Biology 93, POVEY, A. & KEOUGH, M.J. (1991). Effect of trampling on plant and animals populations on rocky shores. Oikos 61, SOUTHWARD, A.J. (1958). The zonation of plants and animals on rocky sea shores. Biological Reviews 33, P.G. Moore and R. Seed (editor), The Ecology of Rocky Coasts: Essays Presented to J.R. Lewis George A Knox. The Ecology of Seashores. CRC Press 2000 Brosnan, D.M. and Crumrine, L.L. (1994) Effects of human trampling on marine rocky shore communities. J. Exp. Mar. Biol. Ecol., 177,