Mollusca overview

There are more than 100,000 described species of Mollusc alive today and 35,000
fossil species, they range in size from just 2 mm to over 20 m in the giant squid, and
occupy all major habitats.

They are believed to have originated in a Precambrian marine environment where they

occupied the shore area offering them a great abundance and diversity of food and
habitats. It is these varied conditions which probably led to their great diversity.

Later bivalves and gastropods moved from salt to brackish then fresh water habitats.

And eventually some gastropods left the water and invaded the land, but were limited
by their need for humidity and calcium.

Some molluscs have multiple pairs of organs, e.g. Monoplacophora, and so were

thought to have evolved from a segmented ancestor making them closely related to
the Annelids. This is no longer thought to be the case. It is now believed that the
molluscan ancestor was probably more like a flatworm that developed some sort of
protective calcareous covering over its dorsal surface. This covering would have
limited respiration, and it is thought that the evolution of gills may have occurred at
the same time. Two extinct phyla, Hyolitha and Wiwaxiida have protective coverings
that may be similar to the ancestral molluscan shell.

Most molluscs are slow-movers. Movement is by muscular ripples down a flat, sticky
pad. However the cephalopods (octopus, squid) move by a form of jet propulsion by
squirting water out from their mantle cavity.

The mollusc shell

The shell is usually formed of three distinct layers of different forms of calcium
carbonate, from the outside the first layer is the periostracum, then a
thick prismatic layer, and finally layers of nacre.

The mantle forms a skirt, and retractor muscles can pull the shell down over the foot
when the animal is disturbed. The mantle cavity usually holds the gills or lungs, and
the products from the digestive, excretory and reproductive systems are usually
emptied into the mantle cavity.
The mollusc radula

The radula (above) is found in most molluscs. It is best seen when aquatic snails graze
the algae of aquarium walls.

It is a moveable ribbon or strap on which there are backwardly pointing chitinous

teeth, usually arranged in rows (see above ). There can be as many as 250,000 teeth.
The backwards and forwards movement of the radula rasps at the surface, breaking off
food particles. As the teeth wear the ribbon is moved forwards, so there is a constant
replacement of teeth.

The action of the radula is as follows -

 The mouth opens

 The supporting cartilage moves forwards bringing the radula in contact with the
surface
 The teeth scrape the surface
 Food particles move up into the mouth as it closes
 Cartilage and radula are withdrawn (see diagrams left ).

This action is repeated rhythmically. The pattern and number of teeth in a row can be
used in identification as they are different in each species.

Bivalvia

Bivalvia overview
The Bivalves are the mussels, clams, scallops, shipworms, piddocks and oysters. There are over
15,000 species world wide (32 British freshwater species). They are laterally compressed with a
pair of shell valves hinged at the dorsal end (see drawings below). The head is greatly reduced;
they have no radula or tentacles, and most are without eyes, although some have eyes at the
margins of the mantle (see Pecten maximus).
They are mainly sedentary filter feeders, have paired gills, and range in size from 1 mm to over 1
m. The giant clam, Tridacna gigas, is the largest bivalve. It can be over 1.5 m across and weigh
over 225 kg. The oldest clam ever found was estimated to be around 400 years old, and some
freshwater mussels and clams can live to be over 100 years old.

Edible oysters and other bivalves have been an important food source of man since prehistoric
times.

Bivalves are mainly marine, with a few freshwater species. The sexes are separate, although some
may be hermaphrodite.The two shells are held together by a dorsal hinge, the ligament can look
like glue oozing out between the two hinges. The shells are held slightly open at rest, but can be
kept closed by a pair of powerful adductor muscles (see above), which work in opposition to the
hinge ligament. When you eat a scallop it is the adductor muscle you are eating.

Bivalves sense gravity using microscopic sensors (see diagram below). A hollow chamber is
lined with nerve cells bearing sensory hairs. These can detect when a mineral weight (statocyst,
see below) is touching them, and so give information about the animal's orientation.

Bivalve locomotion
Locomotion is achieved by extending the foot (see below), which then swells as blood is pumped
into it and acts as an anchor in the sediment, the foot muscle is then shortened as the animal pulls
itself towards it (see Yoldia limatula below). Some can swim by clapping their valves together.
Below is Yoldia limatula. The foot has a flattened sole which can be folded up to push into mud,
then opened to anchor as the rest of the body is pulled downwards.

Bivalvia - cockles

The common cockle (Ceratoid duple) shown above is found just below the surface at low tide.

When covered by water it extends two papillae siphons just above the surface of the sand or mud
and filter feeds by ciliary action. Much of the material taken in is fine sand, so before cooking
cockles must be kept in clean sea water so that the fine sand particles are ejected by the cilia.
The Royal cloak shell, Gloria alliums is above. It is in the Pectin family, and is found in the
Indo-Pacific region.

Tantulocardia aculeate, the Prickly cockle

The Prickly cockle, also known as the Spiny cockle, above, is found in European and West
African waters in mud and sand down to 350 metres deep. It has thorns or teeth running down
the20 or so ribs of the shell, and can grow as big as 115 mm across. It is fairly common. The
outside of the shell is usually pale brown, while the inside is white. Predators of this cockle
include starfish.

Heart cockle, Corundum Marissa

The Heart cockle is found in the Indo-Pacific, and as its name suggests the shell is heart-shaped
when viewed from above. It lives among sand and coral debris in a hollow it excavates. It grows
to around 4 - 6 cm across. The shell is very thin, so much so that light can penetrate through the
shell allowing green algae to live and photosynthesise inside the shell. The Heart cockle has a
symbiotic relationship with the algae and it benefits from the metabolites the algae produces,
while the algae can grow in an environment safe from grazing.

Bivalvia - clams
Above is the Fluted giant clam, or Scaled clam, Tridana squamosa. The fluted giant
clam is found on sandy bottoms around coral reefs at 15 - 20 m deep in the Indo-
Pacific, and has been introduced to Hawaii. The fluted edges are called scutes, and are
used by crabs and other small invertebrates as shelter. It grows up to 40 cm across.It
gets most of its food from a symbiotic relationship with a single celled algae that
colonizes its mantle. The clam extends the mantle during the day for the algae to
photosynthesise. This species is popular in large aquaria. It has been collected and
traded for human consumption, but is now bred in captivity.

Giant clam
Below right is the giant clam, Tridacna gigas, the largest bivalve. Clams in the
Tridacna genus usually have a symbiotic relationship with algae which live on the
mantle tissue and provide an extra source of nutrition.

Sand gaper clam, Mya arenaria

The Sand gaper clam, also known as the Soft shell clam is above. It is found buried in
sand, gravel or mud near the shore or in estuaries, from 50 cm to 192 m deep around
the U. K. and Irish coast. It is also found on the North American Atlantic coast where
it is caught for food, and the Pacific coast where it is classed as an invasive species.

Its siphon is so long in order to reach the surface, that it cannot fit inside its shell (see
the drawing on the right). A fully grown specimen can reach over 20 cm long. Its shell
is thin and easily damaged giving it one of its common names. Its main predators are
humans, sea otters, crabs and sea gulls.

Quahog, Hard-shelled clam, Mercenaria mercenaria

The Quahog (see the photograph above) is native to the east coast of North America
from Prince Edward Island down the the Yucatan Peninsula, where it burrows into
mud and sand in waters up to 10 m deep. More recently the quahog has been
introduced to parts of northern Europe.

Quahog reproduction is sexual and external. Sexual maturity is reached after one year.
At first the sexual ratio is strong skewed towards males, with around 98% of juveniles
being male. As the population ages many of these males change to females, so at
reproductive maturity there are roughly equal numbers of each sex. Like other clams
the Quahog is a filter feeder. Its predators include sea stars, whelks, crabs, snails, bird
and humans. The entire quahog body is edible, not just the adductor muscle. Most
commercially caught quahogs are between 4 and 8 years old.

It was famously used by native North Americans to make their currency, which they
called wampum. The whole shell was used for this, but the purple parts were the most
valuable. The shell was cut into small pieces, and these pieces were made into beads,
then treaded on to necklaces, belts and clothes.

Bivalvia - oysters

Above is the edible American east coast oyster, Crassostrea virginica. The lower

valve is usually larger than the upper valve.

Below is an oyster shell where instead of forming a pearl round a piece of trapped

grit a small fish has been trapped and preserved forever as a fish-shaped pearl. An
oyster can produce as many as 50 million eggs a year.
Below is the American thorny oyster, Spondylus americanus, which is found in the
Caribbean, and the Atlantic coast from North Carolina down to Brazil between 9 and
45 m deep. It can reach 10 cm in diameter. The lower shell is flat and attached to the
substrate. It tends to remain in the same place. In the past it was widely traded in
South America as it was used for decoration.

Spondylus princeps, Pacific thorny oyster

Spondylus princeps, Pacific thorny oyster, also known as the Spiny oyster,
Chrysanthemum shells, on the right is found from the Gulf of California down to Peru
at around 50 m deep.

It cements the bottom half of its shell to a reef or other solid substrate, and sponges are
attached to live and grow on the spiky top half od the shell. This provides camouflage.
Fully grown individuals can reach up to 150 mm across. The outer colouring varies
from pink through to red and orange.

In the past it has been used as offerings to gods, and as currency by native south
Americans. In fact native indians call it "Food of the Gods". It was also used
decoratively in jewellery - as it still is today.

Above is the Black-lipped pearl oyster, Pinctada margaritifera. It is native to the

Indo-Pacific, but found in many other areas as it is raised commercially for pearls. In
the wild it is found on tropical coral reefs down to 75 m deep. It produces back and
grey "Black South Sea Pearls" of very high quality. Its common name comes from the
inner black rim of the shell that you can see in the photograph on the right. Externally
the shell colour varies from dark grey to brown and green, and often with white spots.
Fully grown adults can be as large as 25 cm across.
Above is the Hammer shell or Blackhammer oyster, Malleus malleus, it is found in
the Indo-Pacific on reefs and in sea grass attached to structures, or else free-living and
partly buried in sediment at depths from 1.0 15 m. Sometimes it is found in large
colonies.

Why the shell takes in such a strange shape is unknown. It can grow up to 20 cm long
and is collected for food; the shell is also used to make lime, and cut up to be used in
jewellery and decoration.

Bivalvia - scallops etc

Scallops
Pecten maximus, the great scallop (above) can grow as large as 15 cm across. Along the edges of
its mantle is has finger-like sensory processes, and between many of these are eyes. During its
early life it is attached to the substrate by a byssus (strong hair-like filaments), but later in life it
is free living and is one of the species of bivalves which can "swim" by clapping its valves
(shells) together.
Above is the Noble scallop, Chlamys senatoria nobilis from Japan. Its shell colour is controlled
genetically, and can be orange, purple or brown, or a combination of these colours.

Below is the Leather donax, Hecuba scortum, found in the littoral zone, usually in mud and
sand in Central Indo-Pacific and South China Sea. Its length can reach 9 cm. The inner surface is
a beautiful smooth, glossy violet/pink
Bassinia disjecta, Wedding cake venus

Wedding cake venus above is found in the South Australian waters. It is believed that the ridges
on the shell help protect it against predatory snails.

Shipworms and piddocks

Shipworm and piddocks are the boring bivalves

This group have evolved to use the edge of their valves to bore into wood and even
into rock. The anterior end of the shell is usually serrated to aid the boring action.
Teredo norvegica, the common shipworm, above
Teredo norvegica became known as the shipworm because its shell is greatly reduced,
(it lies towards the bottom of the image), and does not contain all of its body. The rest
of the body lies in a chalky lining secreted as it burrows, so it does look worm-like.

At one the time it was not understood how shipworms could have evolved before man
started building boats and structures in the sea. It takes millions of years for animals to
evolve and man has been building boats for only a few thousand years. However now
it is known that timber is quite plentiful in the oceans especially near great river
estuaries and after storms, so the evolution of the shipworm did not need the presence
of boats.

The larva settles on a piece of timber and starts to burrow. As it burrows the foot grips
the side of the tunnel and the siphons are protected by two calcareous flaps. Food is
obtained through the siphons, but the shipworm has the ability to digest the cellulose
of the wood it drills into. It can cause considerable damage to wooden ships.

The oval piddock, Zirphaea crispata

The oval piddock, Zirphaea crispata, above, bores into rock. As with the shipworm,
the foot holds the animal in place while the rocking backwards and forwards of the
valves rubs away the rock. The valves do get worn down by this action, but every now
and then the mantle edge creeps over the edge of the valve and deposits another layer
of chalk.

Pholas dactylus, the European piddock.

The European piddock is found in the N. E. Atlantic, North Sea and English Channel,
right. It bores into soft rocks, e.g. chalk, sandstone and clay. It can be up to 12 cm
long, excluding the siphon which can add another 12 - 24 cm to its length. In the dark
it phosphoresces a blue/green light.

Below is a drawing of Pholas sp. showing the rock-cutting teeth and the siphon.

Mussels

Mussels attach themselves to surfaces by a thin guy rope called a byssus (see below).
These are placed on the anchoring surface by the foot (see below), and enable the
mussel to withstand battering from the sea.
The scar inside an empty mussel shell shows where the adductor muscle pulling the
two shells together anchors.

Mussels are both freshwater (see below) and marine, and are filter feeders. Water
enters through the siphon. Compare the very muscular foot of the common cocklewith
that of the mussel. The mussel uses its foot much less. Mussels often live in large
groups and can cause problems for coastal power stations by blocking the pipes of
their cooling systems.

Horse mussel, Modiolus modiolus

The Horse mussel, Modiolus modiolus, above, is usually found deeper than the low
water line, most commonly at 5 - 7 metres deep. It can be found all around the U. K.
coast, but is more common in the north and west. It is also found along the Atlantic
coast of North America from the Arctic down to Florida, and on the Pacific coast from
the Arctic down to California. It lives partly buried in sediment, but can attach itself to
solid substrate.
It can grow up to 20 cm long and live for up to 50 years, making it the largest
European mussel. Small mussels are preyed on by crabs and starfish, but those larger
than 6 cm have few predators.

Horse mussels are important as they help stabilise mobile sea beds, thereby creating
habitat for many other animals. Trawling and dredging destroy these habitats. The
mussel beds or reefs are held together by the byssus threads of live and dead mussels
that have tangled with dead mussel shells and any other detritus.

Date mussel, Lithophaga lithophaga

The Date mussel,Lithophaga lithophaga, above, is also known as the Date shell. It is

found in the N. E. Atlantic, the Mediterranean and the Red Sea.

As can be seen in the photograph on the right it bores into rocks aided by secretions
from its pallial gland. Previously the mussel was considered a delicacy, but because
collecting the mussel usually involved destroying the rocky habitat, the collection in
many countries has now been banned or restricted.

Above is Margaritifera margaritifera, the pearl mussel, a freshwater mussel. Fully

grown specimens can reach 150 mm long, and can live fore 190 years. Large pearls
are rare; most pearls are around 2 mm in diameter. It is found in large, fast rivers with
sand or gravel beds in which it buries itself. Recently it has declined through pearl
collecting.

Above is Anodonta cygnea, the swan mussel. The shell when fully grown can be 15
cm long and olive brown. It is found in firm mud in canal bottoms, slow river and
lakes.

The eggs are produced from June to August, as many as 500 000 per individual. They
are kept in the outer gills which now become brood pouches, and are fertilised by
sperm carried on a water current from a nearby male.

After 9 months they leave the brood pouch and become parasites of fish for 3 months.
When a miniature mussel drops off the fish and, hopefully, lands in mud.

Above is Pisidium sp., the pea-shell cockle or Pea mussel. It has just one siphon. It
is common in sand and mud of all types of fresh water including marshes, ditches and
mountain tarns.
It is hermaphrodite and viviparous producing fully developed young. It is very small,
just 3 - 5 mm, and buff coloured.