WELCOME TO ST ANDREWS ON TOUR: ORIELTON 2017

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WELCOME TO ST ANDREWS ON TOUR: ORIELTON 2017

STAFF ANDY BLIGHT (Capt) SAFETY Signature! GRANT BROWN JEFF GRAVES GINNY GREENWAY SUE HEALY FAITH JONES DAVID PATERSON DAVID SHUKER

Intertidal ecology: The classical ecological laboratory of the rocky coastline

Algae and animals are found in distinct bands (or zones) between low and high tide. Joseph Connell, Rocky Shore Biologist

The intertidal is the sea bed between the highest and lowest extent of the tides 1949 Stephenson & Stephenson s universal scheme of zonation Littoral fringe Eulittoral zone Sublittoral zone Lewis (1955)

Animals and plants have overlapping distributions 3 zones is over simplified Biological definitions often used but other influencing factors Therefore the Zonation concept is useful but we must be aware of its limitations Beware of Context dependency

Exposure

Littoral fringe Height up shore Littoral fringe Eulittoral zone Sublittoral zone Sheltered Eulittoral zone Sublittoral zone Exposed Degree of shelter

Degree of exposure

The effects of exposure on algal form Fucus vesiculosus Sheltered has long fronds, many bladders Exposed has no air bladders and is stunted Laminaria hyperborea blade density Durvillaea antarctica (New Zealand): the strongest kelp in the world?

TYPICAL ZONATION PATTERN: SHELTERED SHORE Pelvetia canaliculata Fucus spiralis Fucus vesiculosis Ascophyllum nodosum Fucus serratus Laminaria digitata Laminaria hyperborea Undaria pinnatifida

CREST PERIOD TIDESS TROUGH SPRING SPRING RANGE Chart datum: Lowest level NEAP NEAP

What controls algal vertical zonation on Rocky Shores? Hypothesis 1: Zonation controlled by dispersa and settlement of propagules Hypothesis 2: Exposure: Physically controlled and directly related to tidal level and physiological capabilities Hypothesis 3: Hypothesis 4: Inter-specific competition Predation/Grazing

Hypothesis 1: Zonation controlled by dispersal and settlement Most intertidal organisms produce propagules that enter the plankton and each new generation must recolonise the shore. Limited supply of propagules might influence dispersion? Laminaria Fucus 20 x 109 spores produced 9.8 x 105 colonise rocks 2 grow into adults 1 x 106 per plant per year Supply side ecology

Hypothesis 2: Physically controlled and related to tidal level Critical tidal levels (Doty 1946) There is no good evidence for this in the UK (Underwood 1978) Height above chart datum (m) % Emersion Different emersion periods (Water cover/weather) (Baker 1909/Schonbeck & Norton 1978) Coleman (1933) Light regime Thermal conditions (Dring 1982) (Newell 1979) Water seepage extends zone (Frank 1965)

Hypothesis 3: Inter-specific competition Dayton (1975) Kelps dominate lower regions, but when removed smaller opportunistic or fugitive species were able to move in Schonbeck and Norton (1980)/Norton s weeding experiments (1985)

Hypothesis 4: Grazing The dominant grazers on the rocky intertidal are gastropod molluscs, crustaceans, sea urchins and fish. Increasing evidence that it is grazing that prevents algae from extending their lower limits Himmelman & Steele (1971) demonstrated that when the urchin Strongylocentrotus droebachiensis was removed the alga Alaria esculenta was able to spread its range.

Take home messages 1. Zonation controlled by events after settlement 2. Physiological tolerance greater in littoral fringe spp 3. Limits of littoral fringe controlled by aerial exposure 4. Upper limits within eulittoral by physiology, competition and grazing 5. Lower limits of eulittoral by competition and grazing Upper limits of macroalgal distribution regulated by physical factors and lower limits by biological factors