MOLLUSCA
CC-I / UG FIRST YEAR
PHYLUM- MOLLUSCA
Mollusca : mollis i.e. soft
EVOLUTIONARY PERSPECTIVE
• This group has been very successful. It is the second largest
phylum after arthropoda.
• If success is measured by numbers of species, the molluscs are
twice as successful as vertebrates!
• The vast majority of the nearly 100,000 living species of
molluscs belongs to two classes: Gastropoda, the snails and
slugs; and Bivalvia, the clams and their close relatives.
RELATIONSHIPS TO OTHER ANIMALS
• Molluscs are protostomes. The similarities in the embryological
development of the molluscs and other protostomes, especially the
annelids (segmented worms), are striking.
• Some embryological stages, for example the trochophore larvae, are
virtually indistinguishable in molluscs and annelids.
• Certain adult structures of molluscs and annelids, for example the
excretory organs and their duct systems, are very similar in structure.
• Even though most zoologists accept the protosome affiliation of the
molluscs, the relationships between members of this phylum and
other protostomes is distant, and the ancestral evolutionary
pathways are speculative.
MOLLUSCAN CHARACTERISTICS
• Molluscs range in size and body form from the giant squid,
measuring 18 m in length, to the smallest garden slug, less than 1
cm long.
• In spite of this diversity, the phylum Mollusca (mol-luskah) (L.
molluscus, soft) is not difficult to characterize .
Characteristics of the phylum Mollusca include:
1. Body of two parts: head-foot and visceral mass
2. Mantle that secretes a calcareous shell and covers the visceral mass
3. Mantle cavity functions in excretion, gas exchange, elimination of
digestive wastes, and release of reproductive products.
4. Bilateral symmetry
5. Protostome characteristics, including trochophore larvae, spiral
cleavage, and schizocoelous coelom formation
6. Coelom reduced to cavities surrounding the heart, nephridia, and
gonads
7. Open circulatory system in all but one class (Cephalopoda)
8. Radula usually present and used in scraping food
Body of molluscs
• has three main regions: the head-foot, the visceral mass,
and the mantle.
• The headfoot is elongate with an anterior head, containing the mouth and
certain nervous and sensory structures, and an elongate foot, used for
attachment and locomotion.
• The visceral mass contains the organs of digestion, circulation,
reproduction, and excretion and is positioned dorsal to the head-foot.
• The mantle of a mollusc usually attaches to the visceral mass, enfolds most
of the body, and may secrete a shell that overlies the mantle.
The shell of a mollusc is secreted in three layers:
• The outer layer of the shell is called the periostracum. Mantle cells at
the mantle’s outer margin secrete this protein layer.
• The middle layer of the shell, called the prismatic layer, is the
thickest of the three layers and consists of calcium carbonate mixed
with organic materials. Cells at the mantle’s outer margin also secrete
this layer.
• The inner layer of the shell, the nacreous layer, forms from thin
sheets of calcium carbonate alternating with organic matter. Cells
along the entire epithelial border of the mantle secrete the nacreous
layer. Nacre secretion thickens the shell.
Periostracum
Prismatic layer
Nacreous layer
Mantle cavity
• Between the mantle and the
foot is a space called the
mantle cavity.
• The mantle cavity opens to the
outside
• It functions in gas exchange,
excretion, elimination of
digestive wastes, and release of
reproductive products.
Triploblastic – coelomate
• Triploblastic animals have tissues
derived from ectoderm, mesoderm,
and endoderm.
• A coelom is a body cavity completely
surrounded by mesoderm
• Coelom reduced and represented
mainly by pericardial cavity, gonadial
cavity and kidney.
Bilaterally symmetrical
• It is the arrangement of body parts such that a single plane,
passing between the upper and lower surfaces and through
the longitudinal axis of an animal, divides the animal into right
and left mirror images
Unsegmented body
• do not have their body divided into distinct segments (repeating
structures)
• Except- Monoplacophora
Protostome
• the first opening in the embryo becomes the
mouth
• characterized by
(a) spiral and determinate cleavage
(b) a mouth that forms from an embryonic
blastopore
(c) schizocoelous coelom formation
(d) a trochophore larva.
Trochophore larva
A trochophore is a type
of larva found in marine plankton.
It belongs to animals of several
related phyla, in the
superphylum Lophotrochozoa.
Digestive system
• Digestive system complete with a digestive gland or liver
(hepatopancreas); a rasping organ, the radula, usually present.
• The hepatopancreas is a digestive organ in arthropods and
mollusks that performs functions similar to the liver and
pancreas in mammals.
Radula
• The mouth of most molluscs possesses a
rasping structure called a radula, which
consists of a chitinous belt and rows of
posteriorly curved teeth.
• The radula overlies a fleshy, tonguelike
structure supported by a cartilaginous
odontophore.
• Muscles associated with the odontophore
permit the radula to be protruded from the
mouth.
• Muscles associated with the radula move the
radula back and forth over the odontophore.
• Food is scraped from a substrate and passed
posteriorly to the digestive tract.
Open circulatory
system
• Molluscan blood vessels do not form a closed system.
• Blood and other fluids empty into a large, fluid-filled hemocoel, through
which fluids move around the animal and deliver oxygen to the internal
organs. Eventually, the fluids reenter the blood vessels and are moved by
a heart.
• But cephalopods, which include squid, octopus, and cuttlefish, have a
closed circulatory system.
Respiration and excretion
• Respiration direct or by gills or lungs or both.
• Excretion by paired metanephridia (kidneys).
Nervous system
• Nervous system of paired ganglia, connectives and nerves.
Ganglia usually form a circum-centric ring.
• Sense organs includes eyes, statocysts and receptors for touch,
smell and taste.
• The gravity sensors of most molluscs are spherical organs called
statocysts.
• relatively complex
• most complex- Octopus
• 2 pairs of main nerve cords
• visceral- serve the internal organs
• pedal-control the foot
• brain encircles the esophagus
• sensory ability
• variety of organ systems
• large, complex eyes
■ larges eyes- large squid
Reproduction, fertilization and development
• Dioecious or monocious
• One or two gonads with gonoducts, opening into renal ducts
or to exterior
• External or internal fertilization
• Development direct or through larval stages
Habitat and food
• Terrestrial
• Aquatic- freshwater or marine
• Some are herbivores while others are carnivores.
CLASSIFICATION
• Molluscs are classified into the following classes according to
their symmetry and the characters of food, shell, mantle, gills,
nervous system, muscles and radula.
1. Monoplacophora 5. Pelecypoda/ Bivalvia
2. Amphineura 6. Cephalopoda
3. Scaphopoda
4. Gastropoda
1. CLASS MONOPLACOPHORA
• Members of the class Monoplacophora (Gr. monos, one + plak, plate
+ phoros, to bear) have an undivided, arched shell; a broad, flat foot;
and serially repeated pairs of gills and foot-retractor muscles.
• They are dioecious; however, nothing is known of their embryology.
• This group of molluscs was known only from fossils until 1952, when
a limpet-like monoplacophoran, named Neopilina, was dredged up
from a depth of 3,520 m off the Pacific coast of Costa Rica.
Neopilina
• Body Symmetry and Segmentation
• bilateral symmetry, with a body that is segmented. This segmentation is a
notable feature, distinguishing them from many other mollusk classes.
• Mantle and Shell Structure
• Mantle is dome-shaped and encircles the body as a circular fold. This structure
supports a single-piece shell. The shell’s single-valve nature is a defining
characteristic of this class.
• Head Features
• lacks eyes and tentacles, which differentiates it from several other mollusk
classes that possess these sensory organs.
• Foot and Muscle Arrangement
• The foot of these organisms is broad and flat, equipped with eight pairs of
pedal retractor muscles. This muscular arrangement allows for effective
movement and interaction with their environment.
• Gill and Respiratory Structure
• external gills, arranged in five pairs within pallial grooves.
• Excretory System
• six pairs of nephridia, two of which also function as gonoducts.
• Digestive System
• radula located within a radular sac, and an extensively coiled intestine, adapting them
for efficient nutrient absorption and digestion.
• Circulatory System
• Their circulatory system includes two pairs of auricles and a single ventricle, constituting
their heart. This structure ensures effective circulation of blood and nutrients
throughout the body.
• Reproductive Characteristics
• The species in this class are dioecious, meaning the sexes are separate. This distinction is
vital for their reproductive strategies and lifecycle.
• Representative Species
• Neopilina galatheae
2. CLASS AMPHINEURA
• Amphineura, derived from Greek ‘amphi’ (both) and ‘neuron’
(nerve), is a class of mollusks characterized by their distinct
nervous system and body structure.
• This class is divided into two subclasses:
1. Aplacophora
2. Polyplacophora
• Subclass 1: Aplacophora
• Body Structure: have a worm-like body covered by a mantle but lack
a distinct shell and foot.
• Skin and Radula: Their skin is embedded with calcareous spicules,
and they possess a simple radula.
• Examples: Representative examples include Neomenia,
Nematomenia, and Chaetoderma.
Neomenia Nematomenia
• Subclass 2: Polyplacophora
• Body and Head: Polyplacophora species are dorso-ventrally
flattened with a small head. They lack eyes and tentacles.
• Shell and Radula: These mollusks have a radula, mantle, foot, and
external gills. The posterior mantle cavity is a significant feature.
Their shell consists of eight calcareous dorsal plates.
chiton (Tonicella lineata)
3. CLASS SCAPHOPODA
• Members of the class Scaphopoda (Gr. skaphe,
boat + podos, foot) are called tooth shells or tusk
shells.
• The over three hundred species are all burrowing
marine animals that inhabit moderate depths.
• Their most distinctive characteristic is a conical
shell that is open at both ends.
• The head and foot project from the wider end of
the shell, and the rest of the body, including the
mantle, is greatly elongate and extends the
Dentalium
length of the shell.
• Scaphopods live mostly buried in the
substrate with head and foot oriented down
and with the apex of the shell projecting
into the water above.
• Incurrent and excurrent water enters and
leaves the mantle cavity through the
opening at the apex of the shell.
• Functional gills are absent, and gas
exchange occurs across mantle folds.
• Scaphopods have a radula and tentacles,
which they use in feeding on foraminiferans.
• Sexes are separate, and trochophore and
veliger larvae are produced.
4. CLASS GASTROPODA
(gaster, gut + podos, foot)
• Gastropods are the most species-
rich and widely distributed
molluscs, with nearly 70,000
living species.
• Gastropoda is the largest and
most varied molluscan class.
• Its members occupy a wide
variety of marine, freshwater, and
terrestrial habitats.
• Snails, whelks, limpets, slugs,
nudibranchs (sea slugs), and
abalones are all gastropods.
• Most species move by gliding on their muscular foot, but in a few
species—the sea butterflies and heteropods—the foot is a swimming
organ with which the animal moves through open ocean waters.
• Nudibranchs have lost their protective shell over the course of evolution.
Nudibranchs Sea butterflies
• Sometimes their brilliant coloration is aposematic, meaning it serves
to warn potential predators of toxicity.
(sea slugs)
Brilliant
colours as
warning
• Other nudibranch species exhibit camouflaged coloration.
• Shell present or absent, univalve and usually coiled.
• Shelled gastropods have one-piece shells.
• The only molluscs that live in terrestrial environments—land snails
and slugs— are gastropods.
• In these terrestrial species, the mantle tissue is modified into a highly
vascularized lung.
Cowrie
TORSION
• One of the most significant modifications of the molluscan body
form in the gastropods occurs early in gastropod development.
• Torsion is a 180°, counterclockwise twisting of the visceral
mass, mantle, and mantle cavity.
• Torsion positions the gills, anus, and openings from the excretory
and reproductive systems just behind the head and nerve cords,
and twists the digestive tract into a U shape.
Torsion (coiling) of body mass at sometime
in development.
After torsion, the digestive tract is looped,
A pretorsion gastropod larva. The
and the mantle cavity opens near the head.
posterior opening of the mantle cavity
The foot is drawn into the shell last, and the
and the untwisted digestive tract.
operculum closes the shell opening.
Significance
• The adaptive significance of torsion is speculative; however, three
advantages are plausible.
1. First, without torsion, withdrawal into the shell would proceed with the
foot entering first and the more vulnerable head entering last.
With torsion, the head enters the shell first, exposing the head less to
potential predators. In some snails, a proteinaceous covering, called an
operculum, on the dorsal, posterior margin of the foot enhances protection.
When the gastropod draws the foot into the mantle cavity, the operculum
closes the opening of the shell, thus preventing desiccation when the snail is
in drying habitats.
2. A second advantage of torsion concerns an anterior opening
of the mantle cavity that allows clean water from in front of the
snail to enter the mantle cavity, rather than water
contaminated with silt stirred up by the snail’s crawling.
3. The twist in the mantle’s sensory organs around to the head
region is a third advantage of torsion because it makes the snail
more sensitive to stimuli coming from the direction in which it
moves.
c) A hypothetical adult ancestor, (d) Modern adult gastropods have an
showing the arrangement of anterior opening of the mantle cavity
internal organs prior to torsion. and the looped digestive tract.
Detorsion
• Some gastropods undergo detorsion, in which the embryo
undergoes a full 180° torsion and then untwists approximately
90°.
• The mantle cavity thus opens on the right side of the body,
behind the head.
SHELL COILING
• Shell coiling is a distinctive feature of gastropod molluscs, such as snails,
where their shells spiral around a central axis.
• The opening, or aperture, of the shell is usually to one side of the axis.
• The direction of coiling can be either dextral (right-handed) or sinistral (left-
handed).
LOCOMOTION
• Nearly all gastropods have a flattened foot that is often ciliated,
covered with gland cells, and used to creep across the substrate.
• The smallest gastropods use cilia to propel themselves over a
mucous trail.
• Larger gastropods use waves of muscular contraction that move
over the foot.
• The foot of some gastropods is modified for clinging, as in abalones
and limpets, or for swimming, as in sea butterflies and sea hares.
5. CLASS BIVALVIA / PELECYPODA / LAMELLIBRANCHIA
(bis, twice + valva, leaf )
CHARACTERS
• Body enclosed in a bivalve shell and laterally compressed.
• No head, tentacles, eyes, jaws and radula.
• Foot often hatchet-shaped and extending between mantle
lobes.
• Mostly filter-feeding.
• Usually dioecious, veliger or glochidium larva.
• Mostly marine, a few freshwater.
• Clams, oysters, scallops,
and mussels are all familiar
bivalves.
• The 30,000 living species
are found in both marine
and freshwater
environments.
• Bivalves have a very small
head and a hinged, two-
part shell that extends over
the sides of the body as
well as the top.
• Many clams use their foot to burrow into mud and sand.
• Bivalves feed by taking in water through an opening called an incurrent
siphon and filtering food from the water with their large gills, which are
also the main sites of gas exchange.
• Water and gametes exit through the excurrent siphon.
• Fertilization takes place in open water in most species.
SHELL AND ASSOCIATED STRUCTURES
• The two convex halves of the shell
are called valves.
• Along the dorsal margin of the shell
is a proteinaceous hinge and a series
of tongue-and-groove modifications
of the shell, called teeth, that
prevent the valves from twisting.
• The oldest part of the shell is the
umbo, a swollen area near the shell’s
anterior margin.
• Although bivalves appear to have two
shells, embryologically, the shell forms
as a single structure.
• The shell is continuous along its dorsal
margin, but the mantle, in the region of
the hinge, secretes relatively greater
quantities of protein and relatively little
calcium carbonate.
• The result is an elastic hinge ligament.
• The elasticity of the hinge ligament
opens the valves when certain muscles
relax.
• Adductor muscles at either end of
the dorsal half of the shell close the
shell.
• Anyone who has tried to force apart
the valves of a bivalve mollusc knows
the effectiveness of these muscles.
• This is important for bivalves
because their primary defense
against predatory sea stars is to
tenaciously refuse to open their
shells.
• The bivalve mantle attaches to the
shell around the adductor muscles
and near the shell margin.
• If a sand grain or a parasite lodges between the shell and the mantle,
the mantle secretes nacre around the irritant, gradually forming a pearl.
• The Pacific oysters, Pinctada margaritifera and Pinctada mertensi, form
the highest-quality pearls.
PEARL FORMATION
• Pearl is a concretion(a soilidified mass) formed by molluscs .
• It consists of nacre or mother of pearl . The nacre is formed of two
substances , namely a calcium carbonate which is in the form of
argonite or calcite and an albuminoid substance called conchiolin.
• Kokichi Mikimoto of Toba ( Japan ) is known to be the father of
pearl industry .
• He discovered a method to induce foreign particle between the
mantle and the shell of the pearl oyester and thus stimulated
pearl formation .
• The pearl of oysters produce pearl as an adaptation against outside material .
• When a foreign material enter the body such as a particle of sand or a small parasite ,
it adheres with the mantle .
• The mantle epithelium starts secreting concentric layers of nacre around the foreign
material .
• The completed structure is called pearl .
• Most precious oysters are found in pearl
oysters of the genus Pinctada.
• Important species for pearls are P.
vulgaris , P. chemnitzi , P. margaritifera ,
P. anomioides and P. atropurpurea found
in Indian waters .
Tridacna gigas - Giant clam
6. CLASS CEPHALOPODA
(L. cephalic, head +Gr. podos, foot)
• The class Cephalopoda (L. cephalic, head + Gr. podos, foot)
includes the octopuses, squid, cuttlefish, and nautili.
Nautilus Cuttle fish
• They are the most complex
molluscs and, in many ways, the
most complex invertebrates.
• The anterior portion of their foot
has been modified into a circle of
tentacles or arms used for prey
capture, attachment, locomotion,
and copulation.
• The foot is also incorporated into a
funnel associated with the mantle
cavity and used for jetlike
locomotion.
• The molluscan body plan is further
modified in that the cephalopod
head is in line with the visceral
mass.
• Cephalopods have a highly
muscular mantle that encloses all
of the body except the head and
tentacles.
• The mantle acts as a pump to
bring large quantities of water
into the mantle cavity.
SHELL
• Ancestral cephalopods probably
had a conical shell.
• The only living cephalopod that
possesses an external shell is the
nautilus.
• In all other cephalopods, the shell is
reduced or absent.
• In cuttlefish, the shell is internal and
laid down in thin layers, leaving
small, gas-filled spaces that
increase buoyancy.
• The shell of a squid is reduced to an
internal, chitinous structure.
• The shell is absent in octopuses.
LOCOMOTION
• As predators, cephalopods
depend on their ability to move
quickly using a jet-propulsion
system.
• The mantle of cephalopods
contains radial and circular
muscles.
• When circular muscles contract,
they decrease the volume of the
mantle cavity and close collar-
like valves to prevent water from
moving out of the mantle cavity
between the head and the
mantle wall.
• Water is thus forced out of a
narrow funnel.
• Muscles attached to the funnel
control the direction of the
animal’s movement.
• Radial mantle muscles bring
water into the mantle cavity
by increasing the cavity’s
volume.
• Posterior fins act as stabilizers
in squid and also aid in
propulsion and steering in
cuttlefish.
• “Flying squid’’ (family Onycoteuthidae)
have been clocked at speeds of 30
km/hr.
• Octopuses are more sedentary animals.
• They may use jet propulsion in an
escape response, but normally, they
crawl over the substrate using their
tentacles.
FEEDING AND DIGESTION
• Cephalopods locate their prey
by sight and capture prey with
tentacles that have adhesive
cups.
• In squid, the margins of these
cups are reinforced with tough
protein and sometimes possess
small hooks.
• All cephalopods have jaws and a
radula.
• The jaws are powerful, beaklike
structures for tearing food, and
the radula rasps food, forcing it
into the mouth cavity.
• Cuttlefish and nautili feed on small invertebrates on the ocean floor.
• Octopuses are nocturnal hunters and feed on snails, fish, and crustaceans.
• Octopuses have salivary glands that inject venom into prey.
• Squid feed on fishes and shrimp, which they kill by biting across the back
of the head.
CIRCULATORY SYSTEM
• Cephalopods, unlike other molluscs, have a closed
circulatory system.
• Blood is confined to vessels throughout its circuit around the
body.
• Capillary beds connect arteries and veins, and exchanges of
gases, nutrients, and metabolic wastes occur across capillary
walls.
NERVOUS SYSTEM
• The cephalopod nervous system is unparalleled in any other
invertebrate.
• Cephalopod brains are large, and their evolution is directly related to
cephalopod predatory habits.
• The brain forms by a fusion of ganglia.
• Large areas are devoted to controlling muscle contraction (e.g.,
swimming movements and sucker closing), sensory perception, and
functions such as memory and decision making.
• Research on cephalopod brains has provided insight into human brain
functions.
EYES
• The eyes of octopuses,
cuttlefish, and squid are
similar in structure to
vertebrate eyes.
• (This similarity is an
excellent example of
convergent evolution.)
• Cephalopods can form
images, distinguish shapes, Cephalopod Eye. The eye is immovable in a supportive and
and discriminate some protective socket of cartilages. It contains a rigid, spherical
colors. lens. An iris in front of the lens forms a slit-like pupil that can
open and close in response to varying light conditions. Note
that the optic nerve comes off the back of the retina.
Cephalopods have pigment cells called chromatophores.
• When tiny muscles attached to these
pigment cells contract, the
chromatophores quickly expand and
change the color of the animal.
• Color changes, in combination with
ink discharge, function in alarm
responses.
• In defensive displays, color changes
may spread in waves over the body to
form large, flickering patterns.
• Color changes may also help
cephalopods to blend with their
background.
• Color changes are also involved with courtship displays.
• Some species combine chromatophore displays with bioluminescence.
• All cephalopods possess an ink gland that opens just behind the anus.
• Ink is a brown or black fluid containing melanin and other chemicals.
• Discharged ink confuses a predator, allowing the cephalopod to escape.
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