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PHYLUM MOLLUSCA

• Molluscs are of one of the most diverse of all


invertebrate phyla which include the curious plate
chitons (amphineurans), the slugs and snails
(gastopods), tooth-shells (scaphopods), bivalves,
their possible progenitors (the extinct rostroconchs)
and finally the most complex and successful of all
molluscs: the cephalopods, which include the
modern squids, cuttlefish, octopuses and pearly
nautilus and the fossil ammonoids and belemnites.
Molluscs are mainly marines but a few bivalves and
gastropods have been successful in fresh water. One
group of gastropod, the pulmonates was ever able to
make transition from sea to land.
Common basic structures/features of
mollusca
• The shell which is secreted by a layer of tissue known as mantle
which directly underlines the shell. Throughout growth, calcium
carbonate is added to the edge of the shell from the mantle.
• The shell may either be external univalve, external bivalve,
internal or missing.
• In squids and cuttlefish and extinct belemnites, the shell has
been modified as an internal skeleton.
• In various gastropods e.g. marine nudibranch and terrestrial
slugs, and octopuses, the shell has been entirely lost. Visceral
mass located above the mantle cavity and contains the gut,
digestive glands, kidneys, the nervous, circulatory and muscular
systems
• Mantle cavity in which the gills reside, has been put to various
uses in different moluscan classes.
• The mantle cavity has an outlet to the sea through which waste
materials from the anus discharges as well as respired water.
This outlet has undergone a differential modification, most
markedly in the cephalopods where water taken into the
mantle cavity and squirted out as jet through a funnel, which is
the primary means of propulsion.
• The gills perform respiratory function except in the land
gastropods (pulmonates) where the internal surface of the
mantle is highly vascular and is modified as a lung. In bivalves,
the gills are also adapted for the gathering of suspended food
particles from the water.
• Visceral mass located above the mantle cavity and contains the
gut, digestive glands, kidneys, the nervous, circulatory and
muscular systems
CLASSIFICATION
The phylum mollusca have been classified into the following
classes:
1. Class Monoplacophora (Cambrian – Recent)
• A group of primitive marine molluscs with univalved limpet -
like shells.
• Have paired serially – repeated muscles, gills, nephridia
(excretory organs) as well as other internal organs.
• The foot is circular and central, and ringed by the mantle
cavity in which the gills lie.
• Are the only molluscan class with a true internal
segmentation
• Fossils such as Pilina and Tryblidium are known only from the
Palaeozoic.
• Neopilina, Vema and a few other general occur today mainly
in the deep sea.
2. Class Amphineura (Upper Cambrian - Recent)

• Marine molluscs having a bilaterally


symmetrical shell with seven or eight
calcareous plates.
• Have anterior mouth with radular, a posterior
mantle cavity with anus and gills and a ventral
foot.
• Are rare fossils but occur scattered through
the Phanerozoic e.g. chiton
3. Class Scaphopoda (Ordovician – Recent)

• Marine molluscs with small tapering curving shells


open at both ends.
• The anterior wider end with the mouth is
permanently embedded in sediment.
• The animals feed on small organisms using specially
adapted tentacle
• The anus is at the upper end and
• The gills are much reduced
• Recent species are more abundant than fossil ones
and
• Occur dominantly on the continental slope or shelf.
• Examples; Plagioglypta and Detalium
4. Class Bivalvia (Lamellibranchia or
Pelecypoda) (Ordovician - Recent)
• Have no definite head
• Soft parts are enclosed between paired but
unequilateral calcareous shells united by a toothed
dorsal hinge
• The valves can be shut by strong internal
musculature and
• Opened by the outward pressure of a springy
ligament along the hinge when the muscles relax
• The gills are large and modified for filter feeding
• The mantle cavity is connected to the outer
environment by siphons.
5. Class Rostronconchia (Lower Cambrian – Permian)

• Extinct molluscs of bivalve – like appearance


but with one or more of the shell layers
continuous across the dorsal margin so that a
dorsal commissure is lacking.
• The juvenile shells are univalved and coiled.
6. Class Gastropoda (Cambrian – Recent)
• Snails of all kinds - marine, land and fresh water
– which creep along on a flattened ‘foot’
• A true head, with eyes and other sense organs
are present
• The single univalved shell is coiled planispirally or
more often helically.
• The internal organs are twisted by a 180o torsion
so that the mantle cavity faces anteriorily.
• Some secondarily shell-less groups that have lost
their torsion are also know.
7. Class Cephalopoda (Upper Cambrian – Recent)

• The most advanced of all molluscs


• Have external or internal chambered shells with the
chambers linked by a siphuncle and giving buoyancy
• Posses properly defined head with elaborate sense
organs and
• Move by jet propulsion of water from mantle cavity
• Examples, morden Nautilus, squids, octopuses,
ammonites, belemnites and extinct relatives of
Nautilus.
CLASS BIVALVIA

• All bivalves have a shell consisting of a pair of


calcareous valves between which the soft parts
of the body are enclosed.
• The majority of bivalves are marines, most are
benthic and live infaunally or epifaunally.
• Some genera have successfully colonised fresh
water habitats. Oysters, cockles and mussels are
living example.
• The valves are usually mirror images of each
other and are right and left with respect to the
body.
SHELL MORPHOLOGY
• The two valves on either side of the commissure (the
line of closure) are virtually mirror images of each other
(equivalves).
• Using Carastoderma as an example, an intact
Cerastoderma when examined from either end appears
heart-shaped, with the two valves symmetrical about
the vertical commissure and their umbones (the early –
formed part of the shell) close together and facing each
other.
• From the umbones radiate 22 to 28 strong ribs, crossed
by concentric growth lines which are records of the
former positions of the edge of the shell.
 
SHELL ORIENTATION

• The umbones are set slightly towards one end of the shell,
and it is this which is important in determining the
orientation of the shell and which valve is which.
• In standard orientation for Cerastoderma the umbones face
anteriorly and are closest to the anterior end; this is typical of
most though not all bivalves.
• If an intact Cerastoderma with both valves present is held in
the hand with the commissure vertical, the hinge horizontal
and the umbones directed away from the observer, the right
and left valves are immediately distinguished. In this
orientation the ligament which holds the valves together and
is instrumental in the opening of the shell, is nearest to the
observer and thus posterior.
GILL MORPHOLOGY
•  1. Protobranch: This gill type is small and leaf-like in
appearance and the fact that they occur in a primitive group
suggests that they are not far from the ancestral type. They
are found in the order Nuculoida.
• 2. Fillibranch: this gill type form lamellar sheets of individual
filaments in a W-shape.
• 3. Eulamellibranch: Tis gill type occurs in Cerastoderma and
is similar to filibranch, but here there are cross partitions
joining the filaments and making water filled cavity between
them.
• 4. Septibranch: This type of gill run transversely across the
mantle cavity,almost enclosing an inner chamber which
maintains only a small connection with the outer cavity. This
type is confined to a single super family of rock borers, the
poromyacea (subclass Anomalodesmata)
Modes of life of typical bivalve shells
Types of pelecypod dentition
Types of pelecypod ornamentation (1–19), Rib terminology (20), terminology to
describe rib cross section (21), and types of external rib sculpture (22)
CLASS GASTROPODA

• The class gastropoda are mostly marine, fresh water or


terrestrial animals ranging in size from 1mm to 10 cm,
although exceptionally they may reach 50 cm in length.
• Modern varieties include winkles, whelks, limpets,
snails and slugs.
• The majority of present gastropods and the ones
preserved have coiled shells.
• Gastropods all have a true head, usually equipped with
tentacles, eyes and other sense organs.
• The visceral part of the body largely resides inside the
shell which in such an example as Buccinum (fig.1)
• The head - foot, i.e. the protrusible part of the
body, can be withdrawn inside the shell by
retractor muscles, (the only attachment of the
soft part to the shell) and closed off from the
outside by a plate-like trapdoor (operculum).
• The mouth contains a rasping jaw (the
radula) in which its lower part, composed of a
multitude of tiny teeth which scrape against a
horny plate in the upper part of the mouth to
shred food material. Buccinum has a tubular
extension of the mouth (the proboscis) which is
present in the more advanced gastropods.
TORSION

• During developments, gastropods twisted their dorsal part (including


the shell) through 180o relative to the head, as a result of which large
velum - a flat bilobed organ covered with cilia of gastropod larva) can
be withdrawn inside the shell in case of danger through a process
called torsion.
• The advantage conferred by this process ie being able to tuck away
the velum was so great that the resultant displacement and
asymmetry of the internal organs was a minor price to pay for the
protective advantage to the gastropod when in the larval state and
retained in the adult.
• The main problem that gastropods acquired through torsion was
that, excreta from the anus (located in the mantle cavity) would be
expelled just over the head where the mouth is located.
• Also the orientation of the shell relative to the head - foot mass is
altered, so that in the snail, for example the shell is sited posteriorly
rather than anteriorly above the animal’s head.
REMEDIAL MEASURES
• In many gastropods there are devices that
cope with torsion by separating the inhalant
and exhalent water.
• Exhalent siphons are present in some
gastropods
• and in others there is an indentation (slit
band) in the shell, forming a channel through
which the exhaled water is carried dorsally
away from the shell and head regions below.
CLASSIFICATION
• 1. Subclass prosobranchia
  Order archeogastropoda
  Order caenogastropoda

• 2. Subcalsss opsisthobranchiata
 
• 3. Subclass pulmonata
Prosobranchiata

• Mantle cavity faces forward (anterior)


• Developed special deflecting devices to
separate and divert the inhalant and exhalent
streams
• This subclass is further subdivided into the 2
orders:
• Archaeogastropods (L. Cambrian – Recent)
• Caenogastropods (Ordovician – Recent)
The archaeogastropods

• The name means ancient gastropods; and they are the


parent stock of all subsequent groups.
• They are closest to the hypothetical archimollusc, with the
possession of paired gills which, in more advanced forms
are reduced to one.
• This reduction allows the anus to move further in the
exhalent direction and so helps prevent fouling.
• The exhalent current still has to be deflected from the
head, however, especially since that is where the inhalant
stream enters.
• It is therefore taken out through a hole in the shell or a
deep cleft in the aperture. The presence of a hole (trema)
or the trace of a slit in the growth lines (slit band or
selenizone) is easily recognisable in fossils.
The Caenogastropods
• These are the recent gastropods they deflect
their exhalent currents as best as possible.
• Their most important modification is the
extension of the inhalant siphon often within a
tube of shell material called the siphonal
canal.
• The idea is to separate inhalant and exhalent
currents and prevent recirculation.
• This group has been very successful and it
could be due to their more efficient gill system
that is less sensitive to fouling.
Opistonbranchiata (Carboniferous – Recent)
• Mantle cavity in rear position
• Members of this subclass undergo
detorsion, reversing the original process

Pulmonata (Mesozoic – Recent)


• No gills and instead breathe air directly by
the use of mantle wall highly charged with
blood vessels for easy gaseous exchange.
MODE OF LIFE

• Gastropods live by grazing on algae and higher plants in shallow waters or


in moist conditions on land. Carnivorous habits are seen among
pulmonates, but only rarely.

• Also rare carnivorous like conch may deliver powerful strings – sufficiently
powerful to paralyse its prey .Many divers in tropical areas have drowned
as a result of paralysis brought on such stings.

• Prosobranchs are known to be effective hunters. Certain species may ingest


soft parts of bivalves like clams by prising the valves apart using the
muscular foot and employing their own apertural margin as a wedge.

• Filter feeding is a mode of life that is usually adapted by fixed, cemented or


burrows dwelling forms.

• Many gastropods are passive floaters on seaweed or specially constructed


rafts, but the pteropods (subclass opisthobranchiata) are free swimmers.
STRATIGRAPHICAL VALUE

• Mollusc especially gastropods are useful in the


subdivision of the tertiary strata particularly
where sedimentation was fresh water or
shallow marine – as zonal indices.
• Gastropods are useful in the identification of
environmental conditions from the more recent
past. Archaeologists use gastropods to
determine the environment of early man.
Many of the snails then living are still alive
today and are known to have very strict
preferences in terms of climate, rainfall, type of
vegetation and so on.
CLASS CEPHALOPODA
CHARACTERISTICS
• They are entirely marine and are the most highly evolved of
all molluscs.
• They include the modern nautilus, the Argonauts, squids
and octopuses, and the extinct ammonoids and belemnites.
• Cephalopods are distinguished by having a properly
developed head with a good brain and elaborate sensory
organs.
• A modified foot to a ring of muscular tentacles around a
month in a well developed head region equipped with
highly efficient eyes.
• A sophisticated nervous systems.
• An ability to move rapidly by expulsion of water from a
muscular funnel beneath the head.
CHARACTERISTICS

• An ink sac which inject a smoky fluid into the water when the animal is
threatened, under cover of which they can make their own jet propelled escape.
Ink sac, must have been a protective device evolved at later stage in
cephalopods evolution.

• A strong beak like jaws with which they feed.

• A univalve shell which may either be external (nautilus) internal (belemnites) or


missing (Octopuses) and is usually composed mainly of aragonite. It may either
be wholly chambered when external, or partially chambered when internal

• Chambers filled with gas which may be used to make the animal buoyant. In
those forms having an external shell, the animal lives in the most recently
formed chambers.

• A siphuncle, a thin thread of body tissue situated within a tube which perforate
the septa connecting the body chamber with the protoconch
CLASSIFICATION OF CEPHALOPODS
• There have been considerable difficulties in classifying cephalopods, especially in
erecting large natural categories.

• A provisional classification based on gill morphology which may be inferred but


not confirmed in fossils, broadly divided cephalopods into two divisions: (Owen,
1832)

• 1. Tetrabranchiata: Forms with four gills present or postulated within the mantle
cavity. This includes the following subclasses.
• Endoceratoidea (Ordovician - ? Silurian)
• Actinoceratoidea (Ordovician - Carboniferous)
• Nautiloidea (Cambrian - Recent.)
• Bactritiodea (Ordovician - Permian.)
• Ammoniodea (Devonian - Cretaceous)

• 2. Dibranchiate: Forms with two gills within the mantle cavity. This includes
subclass coleoidea
CLASSIFICATION OF CEPHALOPODS

• From the above subclasses, only nautiloidea, ammonoidea and


coleoidea belong to the important fossil forming groups.

• Nautiloidea and ammonoidea have external chambered shells, while


the last one has an internal chambered shell.

• The grouping of nautiloids and ammonoids to the same


tetrabranchiata is based largely on consideration of the similar basic
soft parts of the two. The bases of which, an extinct ammonoids
reconstruction have been made (fig 1).

• Yet there are some recent discoveries that indicate that the biological
affinities of ammonoids are in many ways closer to coleoids
(diabranchiate cepalopods) such as squids, cuttle fish and octopuses
than nautilus
(b) Ammonoid morphology reconstruction on
(a) Nautilus pompilus shell sectioned the basis of the anatomy of nautilus, showing
subcentrally marginal siphuncle and inferred organisation of
soft parts
APTICHI AND ANAPTCHI

• Aptychi are paired calcitic plates commonly found


in association with Mesozoic ammonites, which
superficially resemble bivalves fig. 1c.
• Aptychi are usually scattered on bedding planes
where ammonites are abundant, and where the
aragonitic shells of ammonites are crushed as in
so many Jurassic shells
• Anaptychi consist of a single plate of chitin or
other organic material, normally flattened and in
a butterfly shape.
• Anaptychi don’t fit the aperture properly and
thus presumably were not opercula.
(c) Aptychus of Hildoceras
SUBCLASS NAUTILOIDEA

• Nautilus is the only living cephalopod genus with a


coiled external shell. The nautilus shell, some 200
cm in diameter, is planispiral and ornamented
externally with a radial colours banding of irregular
and bilateral symmetrical orange – brown stripes.
• The shell is divided into internal gas-filled chambers
(or camerae) by septa, concave towards the
aperture;
• The animal resides in the last chambers (body
chamber) and moves forward each time a new
septum is secreted.
• A single tube (the siphuncle) passes through the centre of
each septum and connect the chambers
• Each septum meets the inner wall of the external shell along
a slightly curved line (suture line)
• Fossil nautilus and its relatives are usually preserved with the
shell dissolved and the chambers filled with spar or matrix.
• In such cases the position of each septum is marked by a
suture line
• Growth is very rapid in living nautilus, new septa being
emplaced about every two weeks on average.
• The living animal itself is separated by a fluid cushion from
the last septum.
• The soft parts can be considered as two separate units:
• The body, which is fully enclosed by the mantle and contains
the viscera and the mantle cavity with its contents
• The head-foot, a cartilage – supported structure with thirty-eight
tentacles surrounding the mouth with its horny parrot – like
jaws.
• The eyes are placed laterally on the head – foot.
• The dorsal part of the head-foot, above the tentacles, has the
form of a hood which normally extends some way up the shell.
• This has a tough warty outer skin, so that when the tentacles are
withdrawn for protection the hood closes the aperture,
presenting a largely impenetrable and uninviting surface to any
predator.
• Below the tentacles is the hyponome or funnel, a long tubular
structure which can be turned in any direction.
• An empty nautilus shell (fig 1) shows short backward – pointing
septal necks piercing each septum ventrally.
• The siphunle consist of septal necks and the permeable strands
between them (siphonal tube).
SUBCLASS AMMONOIDEA
• This group is known in vernacular as ammonites and is amongst the
most abundant and well known fossils.
• They were derived from nautiloids probably during the early
Devonian and from the Carboniferous until the Cretaceous when
they cased to live.
• The suture line normally follows a complex pattern; each suture
marks the junction of a septum with the inner surface of the shell
wall.
• The shell is planispirally coiled.
• The siphuncle is situated near the outer margin (venter)
• One order, the upper Devonian clymeniida, is typified by a dorsal
suture running along the inner margin of the shell (dorsum); in this
group, the septal necks are retrochoanitic (backwardly pointing), as
in nautiloids and very long.
• This contract with the most advanced ammonoid condition, where
the septal necks are short and prochoanitic (directed forwards).
ORNAMENTATION
• Whereas the shell of coiled nautiloids are often
unornamented or have only a feeble external
sculpture, ammonoid shells are frequently ribbed,
and the ribs may have knobs, tubercles or spines;
in very compressed form there may be a keel.
• In such general as the Jurassic Kosmoceras such
developments are carried to an extreme, and in
addition lateral rostrums lappets are present on
either side of a compressed aperture fig. 2
SUTURES IN AMMONOIDS
• Goniaties: are Palaeozoic ammonoids with sharply angular
and generally zigzag sutural without any accessory
crenulations (Goniatitic sutures) fig 4a
• Not all ammonoids wish such sutures are goniatites
• There are some unrelated Mesozoic genera with similar
sutures
• Ceratites: are Triassic ammonoids with frilled lobes, though
the saddles are entire fig 5b (certitic sutures)
• Some curious Cretaceous general (pseudoceratities) have
similar sutures
• Ammonites: are Mesozoic ammonites with finely
subdivided and complex lobes and saddles, (ammonitic
sutures) fig 4c
• Though there are some Permian ammonoids with sutures
also of this kind
HETEROMORPHS IN AMMONITES
• At certain periods in geological history, some
groups of ammonites evolved shells of highly
aberrant form.
• Such shells are known as heteromorphs.
• Some appeared during the late Triassic,
• Some in the Jurassic and there was a more
extensive development of heteromorphs during
the late Cretaceous.
• Heteromorphs may be loosely coiled with the
whorls wholly or partially separated, fig 5 e.g.
Choristocera (Trias), Spiroceras (Jur), and
Lytocriocera (Cret)

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