Learning outcomes:

LSM3254 Ecology of Aquatic Environments

By the end of this lecture you should be able to discuss various aspects of the intertidal, including:

The intertidal: Rocky shores and soft sediments

Th tides The tid Coastal geomorphology and wave exposure. Physical factors, e.g. temperature, desiccation, salinity, and wave exposure (the vertical emersion gradient). Physical and biological interactions - vertical zonation. The particle size gradient.

Peter Todd Dept of Biological Sciences

Representative fauna. Feeding strategies.

SUPRALITTORAL

LITTORAL (intertidal)

SUBLITTORAL

Battle of the Origin of Bulges What causes the opposite bulge?

Most textbooks explain by the differences in centripetal and centrifugal forces across the Earth

High tide level

LAND
Low tide level

SEA

SEA

Battle of the Origin of Bulges What causes the opposite bulge?

Alternatively: Presence of the opposite bulge can be accounted for by the vector subtraction of the gravitational pull by the moon. Bulge is formed due to the tangential component of the resultant force from the Moons attraction. Not because water particles are pulled directly towards the moon!

7 days
NP

7 days

Sun

NP

Sun

7 days

7 days

Intertidal (or littoral) life along the coast is controlled by 2 very important gradients: The horizontal wave-action gradient g
(this determines what kind of substrate is available)

The horizontal wave-action gradient

The horizontal wave-action gradient is a complex combination of geomorphological and climatic/meteorological factors that determine whether an area of shore is sheltered or exposed (or in between)! In general, headlands tend to be exposed and rocky whereas bays are often sheltered and sandy/muddy.

The vertical emersion gradient

(this determines a large range of environmental parameters)

Horizontal wave action gradient

Wave get bent, or refracted, by features such as headlands and outcrops
McGraw Hill

Exposure to wave action

Prevailing wind direction and fetch are very important

Hitting a cliff can mean a wave gets reflected.

Wave-action gets focused on headlands and outcrops and diffused in bays

Maps of two indented coastlines to show variations in exposure to wave action

VP P

British Columbia Estuary Mapping System

Very Protected : Protected: Maximum wave fetch less than one kilometre; usually the location of all-weather anchorages, marinas and harbours. Maximum wave fetch less than 10 km; usually areas of provisional anchorages and low wave exposure except in extreme winds. Maximum wave fetch distances in the range of 10 to 50 km. Waves are low most of the time except during high winds. Maximum wave fetch distances between 50 and 500 km. Swells, generated in areas distant from the shore unit create relatively high wave conditions. During storms, extremely large waves create high wave exposures Maximum wave fetch distances greater than 500 km. High ambient wave conditions usually prevail within this exposure category, which is typical of open-Pacific type conditions

SP SE

Semi-protected: Semi-exposed :

Exposed:

Fetch not everything sea bottom also important

Measuring exposure to wave action

Measure how fast some erodible substance (e.g. balls of plaster of Paris clod cards) wears away. Measure some aspect of wave velocity (Helmuth and Denny, 2003.)

How exposed is Singapore?

Wiffle golf balls!

The Th rocky shore k h

The rocky shore

The intertidal is the most familiar of the marine systems because it is the most accessible. Rocky shores are particularly well studied due to extensive epifauna (as opposed to soft sediments).
Little specialist equipment is needed and it is (usually!) easy to return to the same spot for research.

Features of the rocky shore

Have few sediments as they are washed away. Organisms cannot easily burrow only a few rock-boring bivalves like piddocks (in wood, chalk and even sandstone).
Toothed shell that doesnt fully shut. Muscles are attached in a way that allows for a rotational grinding movement.

Most organisms live on the top of things = epifauna. However, this makes them vulnerable to the effects of exposure. Rocky shores can be uplifted/ing rock that has had little time erode. Hawaii a good example of young islands, with new land being created on a regular basis.

The vertical emersion gradient

The vertical emersion gradient is where the hydrosphere, atmosphere and lithosphere meet.
The physical conditions along the gradient from lower to the upper shore are extremely variable especially compared to the sea itself, which is a relatively constant environment It is a highly stressful environment for marine organisms primarily due to the time spent emersed, i.e. exposed to the air (as opposed to immersed).

Hawaii: volcanic activity not created by spreading or subduction but by a hot spot. The Pacific Plate has moved northwest across the hot spot creating the island chain the oldest being Kaui and the youngest and most active being Hawaii or the Big island

5,000,000 yrs old

The high degree of variability makes acclimatization difficult

The only stresses that may increase at lower shore levels are predation and light availability for plants/algae

Intertidal organisms have two main strategies dealing with these stresses: tolerate (sessile) or avoid (motile).
Low temp: Under typical New England winter conditions, as much as 5070% of the water content of invertebrates and algae freezes. Desiccation: Some barnacles can survive 28 days out of water and the rough periwinkle Littorina saxatilis, can survive over 42 days out of water. Salinity: Carcinus maenas controls osmotic pressure of internal organs, regardless of external conditions. Low O2: In the periwinkles Littorina neritoides (supralittoral fringe) the mantle cavity is modified as a simple lung Some crabs have lungs too lung. lungs too.

Waves:

Wave action

Dislodge things Encourage scour by sand or shingle hi l Smother things Cause continuous rapid movement (impedes foraging, larval settlement, etc.)

But waves also:

Renew oxygen Deliver food and nutrients Disperse gametes Remove waste

A number of mechanisms limit the effects of wave stress

Wave shock!

Streamlined morphology Strong attachments and protection Living in groups Flexiblility

On wave-exposed shorelines, the major adaptive challenge is the force of waves hitting the shore

Nucella lapillus colour polymorphism

Variation in colour morphs too! The dog whelk (Nucella lapillus) has white and brown forms (plus intermediates). The white ones tend to be found in exposed areas with lots of sun while the darker morphs are found in more shady spots. spots

Predation and feeding

The effects of limpet grazing

Not much sediment so few deposit feeders. Lots of filter/suspension feeders but time available for feeding decreases higher up the shore. Many grazers scraping algae and bacteria etc. Also many p y predators, but all tend to seek shelter when , the tide is out. Terrestrial predators too e.g. birds and rats. Predation and competition are superimposed upon the physical characteristics. INTERACTIONS

Six months of no limpets = 100% cover of the green seaweed

Enteromorpha intestinalis

Interactions and vertical zonation

Rocky shore is limited by space not limited by food, especially for sessile organisms Two main strategies - get there first (good dispersal and colonising abilities) or take over: ) Taking over may involve buldozing neighbours (barnacles are good at this), blocking out light (various seaweeds), smother by growing over (e.g. musselbeds). The biological response to the environmental gradient between land and sea is vertical zonation The vertical distribution of any one species is controlled by a complex interaction between physical and biological factors.

Mytilus sp.

Pisaster ochraceus

Different levels/ zones on the rocky shore are occupied by different assemblages of algae and animals, each with a main abundance within a particular zone where conditions are favourable.

10

McGraw Hill

Intertidal vertical zonation is found throughout the world although species will vary. Easy to spot due to distinct bands.

The main physical factors controlling zonation are the tidal range and frequency, and how exposed the shore is. Greater tidal ranges result in more extensive intertidal zones. However, even in the absence of tides, a zone exists in which the sea laps against the shore or waves break and splash (the splash zone).

As a general rule the upper limit is determined by physical factors (emersion) whereas the lower limit is controlled by predation and competition.

The (rocky) intertidal can be split up into 3 to 4 principle zones

McGraw Hill

Often linked

Tidal range Usually based on a combination of physical and biological gradients

McGraw Hill

11

McGraw Hill

Soft sediments

Remember the horizontal wave-action gradient? In general, headlands tend to be exposed and rocky whereas bays are often sheltered and sandy/muddy. The type of community encountered is closely linked to the substrate/s present. In this lecture we will focus on soft sediments.

McGraw Hill

Soft sediment = soft bottom = anything that isnt rock or very hard = can burrow into it easily!
Shore types: Eroding (usually rock) or Depositing (usually occur in bays, inlets and estuaries) The geological history of an area determines the availability of sediment types (e.g. pebbles, sand and mud).

Longshore current result in longshore transport or beach drift

Note the zigzag effect

Most of these sediments are deposited by rivers and carried along the coast by longshore currents.

Coral reef associated sand has a different history!

12

The particle size gradient

Pebbles and coarse sand on exposed beaches (often steep) Muddy sand and mud as shelter increases (and thus water movement decreases) Fine silt only accumulates under the most sheltered conditions
Sediment composition (the relative amount of pebbles, sand, silt and clay) is directly related to water motion.

Think of what happens if you shake a jar full of different sand, pebbles and mud (silt and clay).

McGraw Hill

Wentworth Classification and the Phi Scale Grade name Particle size range (mm) >256 25664 644 42 21 10.5 0.50.25 0 5 0 25 0.250.125 0.1250.0625 0.06250.0039 <0.0039

Feel for yourself!

Phi units

Sediments are defined by their grain size

Boulder Cobble Pebble Granule Very coarse sand Coarse sand Medium Medi m sand Fine sand Very fine sand Silt mud Clay

beyond -8.0 -8.0-6.0 -6.0-2.0 -2.0-1.0 -1.00 01.0 1.02.0 10 20 2.03.0 3.04.0 4.08.0 beyond 8.0

13

Life in soft sediments

Sandy shores are always shifting (in fact, some move offshore altogether in the winter!). Muddy bottoms tend not to shift at all and therefore host a greater diversity of organisms (e.g. tubes and burrows remain in tact in mud, whereas they disintegrate relatively quickly in sand.) Pebble beaches really are lifeless as the churning of the pebbles by waves basically grinds everything to death. This also happens on coarse sand beaches - albeit to a lesser extent.

Particle size gradient (mix of particles also important)

Particle size gradient and organisms

Surface dwelling species are present at both ends of the particle size gradient Infauna (organisms live in the substrate) are restricted to smaller particle sizes (Non-transient) macrobiota are absent from the middle section of the gradient but surface may have a few microscopic species.

Sandy vs muddy shores

Exposed, sandy beaches Prone to high t P t hi h temperatures and desiccation t dd i ti Little fluctuation in salinity Well oxygenated Reduced organic matter and hence limited bacterial activity

Sheltered mudflats Rarely (if ever) dries out (thus not so hot) Greater fluctuation in salinity Poorly oxygenated Increased organic matter and thus greater bacterial activity

Not a comfortable living space!

14

Temperatures
The midday heat of a summer sun may raise the surface temperature of the sand much higher than the returning sea. M dfl t are not affected so much as they are buffered by Mudflats t ff t d h th b ff d b standing water.

Oxygen availability
No sunlight below a few mm and therefore no photosynthesis. What oxygen there is gets used up by the animal respiration. Sand contains little organic matter (it is washed away) and therefore g ( y) feels clean. Mud is the opposite! Muddy bottoms especially bad as they have more organic material but less oxygen getting in. Deeper down water conditions become anoxic (no oxygen at all) and thus anaerobic respiration found. Sandy beaches can also have this layer just deeper down (1m +). Chemocline. Other organisms also have to adapt to low levels of oxygen:
Some pump (oxygen-rich) water through their borrows Some use siphons to suck water from the surface Others have properly adjusted to low levels of oxygen (rather than just avoiding them) through special hemoglobins and reduced metabolism. A few have symbiotic bacteria

The marine-freshwater gradient of salinity

Coarse sandy beaches - sufficiently rapid drainage of water - little opportunity for standing water to evaporate or be diluted by rainwater On mudflats - often a great deal of standing water thus heavy rainfall or evaporation at low tide may considerably change its salinity.

Sandy vs muddy shores

The fauna of mud and sand

Most organisms on soft sediments are either too small to be seen with the naked eye or they are buried so they dont attract the same amount of attention as rocky shores, for instance. instance Many animals borrow in the sediment to keep from being washed away, i.e. infauna. Interstitial fauna are microscopic organisms that live among the tiny spaces between grains and are represented by. Most phyla are ep esented a e represented.
Note: interstitial and meiobenthos and meiofauna are often used interchangeably (Some say the latter are technically those organisms between 0.5mm and 62m, however other sources say meiofauna is 100-1000m, 21000m, etc.)

15

Tardigrades Copepods

Ciliates Cnidarians

Polychaetes

Macrofauna (infauna and epifauna)

Gastrotrichs Molluscs

Burrowing

Bivalves Use their muscular foot Shell sculpture or ribbing thought to increase friction in one direction Worms Elongated body Use penetration and terminal anchors Shrimps/crustaceans B Burrow head fi t with th i appendages h d first ith their d Urchin Heart urchin has spatulate spines for burrowing

All burrowing organisms are bioturbators

McGraw Hill

16

Much the same basic idea for both molluscs and worms

Types of feeding

Predators, e.g. Snails (Polinices moon snail), crabs, worms, fishes and birds. Suspension feeders, e.g. bivalves, worms, etc. Deposit feeders, e.g. amphipods, sea cucumbers, worms and snails.

suspension feeder

deposit feeder

Deposit feeding

17

Deposit feeding

McGraw Hill

Hydrobia

Gorbushin A.M. 1997 Field evidence for trematode induced gigantism in Hydrobia spp. (Gastropoda: Prosobranchia). J. Mar. Biol. Ass. U.K., 77, 785-800.

In summary:
Tide are a result of the sun and moons gravity. Rocky shores are found on more exposed areas of coast.Soft sediments shores are found on sheltered areas of coast. coast Along the shoreline life is only found ON rock or IN relatively fine sediments Organisms living in the intertidal have to survive a wide rage of stresses. Intertidal organisms have various adaptations to cope with this harsh environment. Generally, the upper limit of marine life on the rocky shore is determined by h i l factors ( b physical f t (emersion) whereas th l i ) h the lower di t ib ti i distribution is controlled by predation and competition. Organisms living soft sediments must adapt to this shifting environment. Most soft sediment organisms have the ability to burrow and deposit and suspension feeding is common.

18