SL 20303

MARINE ECOSYSTEM
Lecturer 7

Tidal flats
Adaptations to sandy beach life
Nutrient cycling

TIDAL FLATS AND MUDBANKS

Sandy beach are not the only open intertidal systems, as tidal
flats and mudbanks of many estuaries are so considered.
Intertidal mudbanks
tidal flats are different from beaches. Tidal flats often have sandy
areas, but most of a tidal flat is dark, sticky mud

Waves are the key difference between tidal flats and beaches.
Tidal flats are not like beaches because they are not affected by
waves. Waves change the size of sediment particles. A sample of
tidal flat mud contains different kinds and different sizes of
sediment particles

How??

ADAPTATIONS TO SANDY-BEACH LIFE

Most of the adaptations that distinguish sandy beach animals form

those of other marine habitats result from instability of the substratum
coupled with heavy wave action.

Burrowing behavior (which is displayed by animals inhabiting all types of

soft substrata) on high-energy sandy beach must be both rapid and
powerful in order not to be swept away by incoming swash and waves.
Faunal displays a high degree of mobility and ability to deal with the
swash climate.
This implies the necessity for mechanisms enabling the animals to
maintain their positions on the shore and to regain those positions once
lost.
The ability to orientate to environmental situations is developed to high
degree, not only among the aquatic faunas but particularly among the
air-breathing amphipods, isopods, and brachyurans inhibiting in the
upper part of the shore.
Although these semiterresterial forms can tolerate submersion in
seawater, they cannot risk being caught by the surf and swept out to
sea OR wander too far inland.

Combination of high degree of mobility and complex responses to

environmental cues has lead to typical sandy-beach animals
developing night-day and tidal rhythms of migration, which
maximize food source and possibly attenuate predation.
Heavy surf is one of the challenges faced by the sandy-beach
animals as prey are more difficult to locate and capture under
turbulent conditions.
Carrion is moved around in the surf and swash until it is deposited at
the edge of the tide where may be unavailable to many aquatic
scavengers.

In corporate with the highly erratic supply of carrion to the beach,

makes it essential for the scavengers to locate and reach their food
rapidly and to ensure that they are not parted from it until they
consumed as much of it as they can.
High rates of ingestion, efficient digestion and assimilation and
subsequent energy conservation are called for if the anima is to be
sure of surviving until the next unpredictable meal.

On sheltered sandy beaches (particularly in muddy sands)

respiratory adaptations are necessary to tolerate poor or
declining oxygen tensions, especially during ebb tides.
On high energy sands, oxygen tensions are normally high to a
depth of a meter or more below the sand surface.
Morphological adaptations are thus in evidence in some groups
where a thin elongated shape being advantageous.
Some tend to move up and down through the sand column.

1. LOCOMOTION

There are no truly sessile animals on intertidal sandy beaches and

locomotion plays a much larger part in the lives of the animals
than it does on other types of shores.
Examples:
Burrowing

Swimming is common among the Crustacea

Crawling, leaping also occur in other groups.

1.1 BURROWING
Involves two distinct phase: Penetration of the substratum followed
by burrowing movements.
Penetration is frequently facilitated by the thixotropic properties of
particulate substrata, so that the sand is liquefied by repeated
probing movements of the head (worms) or of the foot (in a
mollusk).

Using this method the sand may be penetrated without the

application of large forces.
Initial penetration can be very slow on very sheltered beaches
where the swash and waves will not easily dislodge the animal

It may takes anywhere from 20 seconds in Polychaeta to 15 minutes

in anemone.
For those in animals in the surf-swept beaches, the danger being
swept away is always present and the penetration needs to be fast
(a few seconds) and therefore larger forces may be applied.

The burrowing movements that follow penetration, once the

animal has gained adequate purchase in the sand, are
essentially similar in virtually all soft-bodied forms being based on
the alternate application of two types of anchorage, each of
which allows another part of the body to move into the sand.
First penetration where the anchor is formed by the dilation of
part of the posterior region of the bodies and prevents the
animals to be pushed out of the sand when it moves downwards.
The second, or terminal, anchor will be formed by the swelling of
the anterior or leading region (head of worm / distal portion of
the foot in a bivalve mollusk) so as to allow the posterior to be
drawn down.
In order to permit such changes in shape, soft bodied animals
have characteristically developed large fluid-filled cavities that
effect both changes in shape and the transfer of muscular
forces.

http://life.bio.sunysb.edu/marinebio/benthic.life.html

Bivalve mollusks are primitively infaunal, with a laterally compressed

and blade-like foot with which to thrust downward into mud or sand.
Like the burrowing polychaetes, they also show a preliminary
penetration phase followed by a series of digging cycles in which
the alternate formation of penetration and terminal anchors is in
evidence.
In the initial penetration phase, the animal lies on its side on the
sand, while the foot probes downward until sufficient purchase is
obtained to allow the shell to be drawn erect.
During each digging cycles, each cycles involves anchorage of the
shell (penetration anchor), while the foot thrusts downward by
contraction of the pedal transverse muscles in antagonism to the
distal retractor fibers.
The shell is then drawn downwards while dilation of the distal part of
the foot forms a terminal anchor.
The downward movement of the shell is facilitated by adduction of
the valves.

During shell adduction, the siphons are closed so that water can only
escape from the mantle cavity downward, through the pedal gape.
The jet of water liquefies and excavates sand from beneath the shell,
making its passage easier through the substratum.

http://life.bio.sunysb.edu/marinebio/benthic.life.html

In sand burrowing arthropods, on the other hand, the body has a

fixed shape imposed upon it by the exoskeleton and a burrow
must be excavated by the jointed appendages.
This is by no mean an disadvantage, and in general the aquatic
Crustacea burrow more rapidly than of any of the soft bodied
animals.
However, the use of jointed appendages also allows many semi
terrestrial Crustacea and insects to burrow rapidly into dry and
moist sand whereas soft bodied invertebrates can usually only
burrow into water saturated sand.

1.2 SURFING AND COPING WITH

SWASH
Surfing is a common form of locomotion among the resident
invertebrates of exposed sandy beaches, although it is much less in
evidence on relatively sheltered shores.
Surfing is normally associated with the tidal migrations up and down
the beach face, and at first sight appears to be a cheap form of
locomotion in terms of energy expenditure.

Bivalve, Donax, surfs with both its foot and siphon extended.
Most species of Donax surfing on every tide but the largest species
(Donax serra), shows a decreasing tendency to surf with increasing
size as the biggest individuals probably never leaving the sand.

Smaller individual surd intermittently, chiefly on spring tides, with

some migration at neaps daily tidal changes being largely ignored
by the adults.
Even where a species surfs on every tide, this does not imply that
every individual of the population does so.

Females carrying eggs do not surf under any circumstances,

however remain buried in the sand.
Crustaceans, being more mobile, regularly swim and surf in the
swash.

All sandy beach animals must be able to cope with swash.

They must move and orient and burrow in it, even if they are not
active migrators or surface forms.
The ability to handle swash is a key adaptation enabling
colonization of different beach types.

2.0 RHYTHMS OF ACTIVITY

Tidal rhythms of activity are by no means limited to sandy beach
animals.
All animals subjected to tidal rise and fall on all substrata must of
necessity suffer discontinuities of behavior imposed upon them by
the tidal cycle.
The implication of tidal ebb and flow on a soft substratum are,
however, different from those on a rocky shore.
In particular, the danger of desiccation is not an overriding
concern, as the animals can retreat below the surface of the
substratum or even below water table.

Intertidal filter-feeders cannot feed while the tide is out.

Carrion beached by the previous rising tide lies too far up the
slope to be available to the aquatic scavengers, whereas other
activities (such as reproductive behavior, which may depend on
the animals being submerged) are at a minimum.
Macrofaunal arthropods and mollusks, on the other hand tend to
leave the substratum and to show excursions up and down the
beach face with the tides, particularly on exposed beaches
showing a moderate tidal range.
Among Crustacea, such tidal migration are displayed by some
isopods, amphipods, mysid and decapods.

In many of these animals, an endogenous rhythm has been

either demonstrated or assumed, triggered by factors related to
the rise and fall of the tides.
Sandy beach mollusks on the other hand have not been shown
to possess internal clocks, and their activity rhythms are not
continued under constant conditions.

Example: The tidal migratory responses of Donax, appear to be

dictated entirely by changing physical conditions during ebb
and flow. They also appeared to display some endogenous
rhythmicity, with jumping behavior being strong during high tide
periods and least during low tide.
For some isopods, little swimming takes place during neap tides,
the animals mostly remaining buried in the sand or leaving it only
for short periods.
Swimming activity is also marked at night than during the day.

3.0 CHOICE OF HABITATS

At least once during their lives, a majority of intertidal animals
must choose their habitat and such choice may be critical at the
end of the larval development or at the metamorphosis (as is the
case with the macrofauna of high energy beaches) the choice
have to be made repeatedly as their migratory behavior or
unpredictable conditions (such as storms) carry the animals into
less favorable sites.
Once carried out to the sea, tidal rhythms and endogenous
clocks are of no use in regaining their positions on the shore and
they must rely entirely on orientation cues to reach favor area.
Responses to water currents, to patterns of polarized light, and to
changes in hydrostatic pressure may all be important in this
regards although other factors, which have not yet been
investigated, may also be involved.

The two most important factors defining the beach habitat for
benthic macrofauna are sand texture and swash flow over the
beach face.
Sand particle size is perhaps the most critical factor and exerts a
considerable influence on burrowing.
Crustaceans and mollusks have shown a clear response to sand
particle size in terms of burrowing performance and prefer fine to
medium sands than coarser sands.
In most species, there are size differences in burrowing, small
individuals burying faster than larger individuals.

There are also temperature effects, invertebrates burrowing faster

at higher temperatures within tolerance limits.
Thus, sand particle size in the habitat is a critical factor affecting
the most important feature of sandy beach animals: mobility and
burrowing ability.

Similar to the macrofaunal, the distribution of some species of the

meiofaunal is governed partly by a choice of particle size range.
However, grain size and factors dependent on it may interact
with variables such as oxygen tension and pore space to
determine the distribution of many members of the meiofauna.
Indeed, the apparent preferences of many species for particular
grades of sand may be due mainly to other related factors, such
as water circulation and availability of oxygen.

4.0 NUTRITION
The absence of attached macrophytes on intertidal sands dictates a
predominance of filter feeders and scavengers among the resident
invertebrate macrofauna.
the physiological and behavioral adaptations of carnivorous scavengers
are likely linked to the highly erratic nature of their food supply and
unpredictability as to what type of carrion will be available, whereas the filter
feeders rely on a more constant but poorer diet.
The scavengers will not only accept a wide variety of food but will typically
turn predator when the opportunity arises and they also tent to have more
than one method of feeding.
i.e: some crabs are both scavengers and predators, displaying a remarkable
ability to locate, dig up and capture their prey.
Yet they will also scoop up the few millimeters of sand by means of their chelae
and in order to ingest meiofauna, microflora or organic matter in general.

The invertebrates might be facing some challenges in securing

the food source as the carrion is typically stranded in the swash
zone and consequently moved by the waves.
They will take every precaution steps not to be parted from it:
If the food is soft, the whelks proboscis is thrust deeply into it so that
it is anchored to the food.

If the food is a bivalve or some other animals with hard shell, then
some tent to have the foot as efficient suction cup that renders the
animals and its food inseparable by the waves.
If the food is very small (a fragment of jellyfish), they will drag it to the
bottom of the sand and consumes it there.

5.0 RESPIRATION
The respiratory adaptations of aquatic animals inhabiting low energy sandy
beaches are different from those found of surf-swept beaches (or indeed on
rocky shores) and resemble the adaptations of animals burrowing into
marine muds being associated mainly with the incidence of low oxygen
tensions, particularly during ebb tide.
On high-energy intertidal sands, where oxygen tensions are high, most of
the macrofaunal species are very much more active than those of lowenergy shores the higher activity being especially associated with tidal
migrations.

Yet the amount of food availability may be lower than is found on sheltered
shores and it is thus not surprising to find that the accent here appears to be
on reduction of metabolic rate and on other ways of conserving energy.

An important method of conserving energy is to reduce, where possible,

the metabolic effects of acute increases in environmental factors such as
temperature.
This may be achieved behaviorally by avoiding high temperature which
spends the daylight hours deep within its relatively cool burrow.

A common error on the part of marine biologists have been to rely on the
measurement of the rate of oxygen consumption as an index of total
metabolism, thus ignoring the anaerobic components of respiration (which
may be considerable in some cases).
Among sandy beach animals one might expect the anaerobic
component to be highest in animals from very sheltered shores (where
hypoxia is common condition), and least from those from high energy
shores where fully oxygen saturated conditions normally occur.

6.0 ENVIRONMENTAL TOLERANCE

It is clear that the majority of intertidal animals have tolerance
levels of natural variables that exceed those necessary for
survival in their particular habitats.
Some species may regularly encounter conditions close to their
upper lethal limits and their behavior and physiology of the
animal is needed.
Ocypodid crabs have evaporative cooling where the water
branchial cavity being replaced from time to time entering the
burrow, by plunging into the sea or by absorption from the
substratum.

Evaporative cooling may also take place in some aquatic

crustacean while in air.

Meiofauna living in the upper layers of the beach may also in

occasion be near their upper limits of tolerance, although in most
cases migration downward through the sand column will relieve
the situation.
Differences in tolerance levels are displayed not only between
intertidal and subtidal forms between animals inhabiting different
zones on the shore.
The correspondence among zonal level, the relative importance
of aerial respiration, and resistance to desiccation, temperature,
and salinity changes are now almost as well documented for
sandy-beach species as for those on rocky shores.
Differences in thermal tolerances among meiofaunal organisms
are also be expected in relation to tidal level and mean depth
within the sand column.

Rates of locomotion are also temperature dependent as low

temperatures jeopardize the ability to undertake migratory
excursions successfully and prove dangerous to the life of the
animal.
Low temperatures significantly decrease the rates of burrowing in
sandy beach bivalves and gastropods, increasing the risk of the
animals being swept away by the waves before an adequate
purchase has been achieved in the substratum.
Low temperatures may also prevent migratory behavior
altogether.

7.0 AVOIDANCE OF PREDATORS

Sandy-beach invertebrates are particularly vulnerable to
predation, and it is not surprising to find a wide variety of
mechanisms whereby the animals avoid predators.

Deep burrowing is clearly one device that renders them

unavailable to birds and fish.
During feeding, the siphon of bivalve is exposed and bitten-off
siphons form an important part of the diet of some fishes and a
few birds.
The loss of one or both siphons does not seriously incapacitate
the animal for any length of time, as the siphons are rapidly
regenerated.

Some deeper-than-normal burrowing activities occur in response to

certain chemical substances which the animals are quite sensitive.
This is in marked contrast to the animals response to pollutants and may
be a reaction to the scent of predators.

Tidal migration of some invertebrates of exposed sandy beaches, keeping

the animals in the swash zone, also reduce predation by making it difficult
for birds or fishes to get at them.
Some animals have developed, in addition, escape responses.
I,e: some crabs rely on an impressive threat display the chelae being
held open and aloft, pointing at the predator, while the crab burrows
rapidly backward and disappear into the sand.

NUTRIENT CYCLING
Nitrogen and phosphorus are the most important nutrient elements in the
majority of marine systems, with nitrogen more often limiting to plant growth
than phosphorus.
In beach/surf zone ecosystems, nitrogen requirements may be considered to
be those of the surf-zone phytoplankton and benthic diatoms.
There are various forms of nitrogen input to the beaches and surf zones.
Groundwater seepage represents one of the most important and ubiquitous
nitrogen sources.

Rain also introduce a small amount of nitrogen.

Sandy beaches have long been considered active in nutrient

recycling through their role in mineralizing organic materials
emanating from the sea and have been labeled great digestive
and incubating systems.
All animals recycle nutrients in the dorm of urine and feces.
The interstitial system is considered more important than the
macrofauna in nutrient recycling.
As water is filtered through the sand, the interstitial biota consumes
the dissolved and particulate organic matter and recycles the
nutrients.

Nitrification is the dominant process in well oxygenated sands, most

organic nitrogen being mineralized to nitrate.

Nitrogen may be lost to beach/surf zone areas by export to

adjacent coaster waters or by accumulation in sediment.

Losses to the sea may occur via active migration of fishes and
zooplankton out of the surf zone or by the passive transport of
DON, PON and plankton out to sea, alongshore and into
estuaries.
These exports may be significant where water turnover rates are
high and surplus material is produced.

In general, the ecosystem must have losses to balance the

surplus of nitrogen.