Molluscs

A Shell-Collector’s Delight
1. Diversity: Molluscs include wormlike animals, giant squids, and species with various shell
configurations (single shell, two shells, eight shell plates, or no shell at all).
2. Shared Ancestry: Although some debate exists, several features suggest that molluscs form
a clade. Most molluscs share common characteristics.
3. Radula: Most molluscs possess a unique ribbon of teeth called a radula. This structure is
used for feeding, often scraping algae from hard surfaces.
4. Muscular Foot: Molluscs typically have a large muscular foot used for locomotion. The foot’s
shape and function vary across different mollusc classes.
5. Mantle: Molluscs have a tissue layer called the mantle. It plays multiple roles, including
secreting the shell and housing respiratory and sensory organs.
6. Diverse Adaptations: These common molluscan features are employed differently across
the eight mollusc classes. For instance:
o Radular teeth inject venom in some species.
o Some groups have lost the radula entirely.
o Snails crawl on their foot.
o Octopi divide their foot into muscular, prehensile arms.
7. Human Exploitation:
o Shells were historically used as currency on almost every continent. The money
cowry, a snail shell, served as widespread currency.
o Shells continue to be used in jewellery, buttons, and decoration worldwide.
o Pearls, produced by bivalves, are another valuable product.
o Most commonly, molluscs are consumed as food. Clams, oysters, scallops, mussels,
snails, abalone, squids, and octopi are among the commercially exploited molluscs.
o Humans consume different parts of molluscs, including the mantle, foot, and entire
body.

Mollusca
1. Diverse Group:
o Mollusca is one of the largest animal phyla after Arthropoda.
o It comprises over 90,000 living species and approximately 70,000 fossil species.
2. Distinctive Characteristics:
o The name “Mollusca” reflects one of their defining features: a soft body.
 o They develop via spiral mosaic cleavage and form a coelom by schizocoely.
o The ancestral larval stage is a trochophore, but development varies across different
classes.
3. Wide Range of Sizes:
o Molluscs come in various sizes, from almost microscopic to the giant squid
Architeuthis.
o These massive molluscs can grow up to 20 meters long, including their tentacles,
and weigh up to 900 kg (1980 pounds).
o While some giant clams (e.g., Tridacna gigas) reach 1.5 meters in length and weigh
over 250 kg, most molluscs have a maximum shell size of less than 10 cm.
4. Habitats and Lifestyles:
o Molluscs occupy a wide range of habitats:
o Their lifestyles vary:
▪ Bottom feeders, burrowers, borers, and pelagic forms
▪ Herbivorous grazers, predaceous carnivores, filter feeders, detritus feeders,
and parasites
5. Origin and Evolution:
o Molluscs originated in the sea according to fossil evidence.
o Most molluscs have remained in marine environments.
o Their evolution occurred primarily along shores, where food was abundant and
habitats varied.
6. Habitat Adaptations:
o Bivalves and gastropods are the only molluscs that moved into brackish and
freshwater habitats.
o Bivalves, being filter feeders, remained tied to aquatic surroundings.
o Slugs and snails (gastropods) were the only molluscs to invade land.
o Terrestrial snails are limited by their need for humidity, shelter, and calcium-rich
soil.
7. Threats to Molluscs:
o A healthy mollusc “fishery” relies on healthy oceans.
o Ocean acidification poses an unexpected threat to mollusc populations.
8. Ocean Acidification Mechanism:
o Increased atmospheric CO2 leads to chemical reactions in the oceans, lowering pH.
o As the ocean becomes more acidic, biologically available calcium decreases.
o Marine organisms, including molluscs, find it harder to form calcium skeletons.
o Calcium is crucial for healthy mollusc shells and coral tissue foundation.
9. Effects on Molluscs:
o Clams, oysters, mussels, and scallops produce thinner and weaker shells under low
pH conditions.
o Larval abalone struggles to develop normally and often cannot form proper shells.
o Oyster larval survivorship decreases with increasing ocean acidity.
o The Pacific Northwest oyster industry, with a $278 million revenue, faces concerns
due to acidification.
10. Cephalopods’ Resilience:
o Squid and octopus, lacking shells, appear less affected by ocean acidification.
o They represent a bright spot in future mollusc harvesting.
11. Coral Reef Ecosystem Impact:
o Direct effects of acidification on coral settlement and growth are observed.
o Neurological development of larval tropical reef fishes is also affected.
o Indirect damage to the coral reef ecosystem is equally concerning.
 o Changes in food webs, coral distribution, and algal abundance may impact molluscs
and other taxa not directly affected by acidification.
12. Class Aplacophora, Monoplacophora, and Scaphopoda:
o These classes have limited diversity.
o Members of class Polyplacophora (chitons) are common marine animals, especially
in the intertidal zone.
o Bivalves (class Bivalvia) have evolved many species, both marine and freshwater.
o Class Cephalopoda contains the largest and most intelligent of all invertebrates.
13. Class Gastropoda (Snails and Relatives):
o Most abundant and widespread of molluscs.
o Enormously diverse group.
o Basic body plan shared among molluscs.
o The coelom in molluscs is limited to a space around the heart, gonads, and part of
the kidneys.
o Unlike annelids, this coelom space is not used in locomotion.

FORM AND FUNCTION

1. Basic Body Plan:
o The mollusc body plan consists of two main portions:
▪ Head-foot: Contains feeding, cephalic sensory, and locomotor organs.
Primarily relies on muscular action.
▪ Visceral mass: Contains digestive, circulatory, respiratory, and reproductive
organs. Primarily relies on ciliary tracts.
2. Mantle and Mantle Cavity:
o Two folds of skin form a protective mantle.
o The mantle cavity lies between the mantle and body wall.
o The mantle cavity houses gills (ctenidia) or a lung.
o In some molluscs, the mantle secretes a protective shell over the visceral mass.
3. Diversity of Patterns:
o Modifications in the structures forming the head-foot and visceral mass result in the
great diversity observed in the Mollusca.
o Different classes of molluscs emphasize either the head-foot portion or the visceral
mass portion.

Head-Foot

• Well-Developed Heads: Most molluscs have well-developed heads that house their mouth
and specialized structures like sensory tentacles and photoreceptors.
• Photoreceptors: These light-sensing cells can be simple, but in cephalopods (such as squids
and octopuses), they are remarkably complex.
• Locomotor Organ: Typically located posterior to the mouth, the chief locomotor organ in
molluscs is the foot.

Radula

1. Structure: The radula is a ribbonlike membrane present in all molluscs except bivalves and
most solenogasters. It resembles a rasping, protrusible tongue.
2. Teeth: The radula bears a surface covered in tiny, backwards-pointing teeth. These teeth
serve various purposes, including scraping, piercing, tearing, and cutting.
 3. Function:
o Feeding: The radula’s primary function is to rasp fine particles of food material
from hard surfaces.
o Conveyor Belt: It also acts as a conveyor belt, continuously carrying food particles
toward the digestive tract.
4. Tooth Count and Replacement:
o Molluscs can have as few as a few teeth or as many as 250,000 teeth on their
radula.
o As the radula wears away anteriorly, new rows of teeth are continuously replaced
by secretion at its posterior end.
5. Species-Specific Patterns:
o The pattern and number of teeth in a row are specific to each species.
o These unique tooth arrangements are used in the classification of molluscs.
6. Specializations:
o Some molluscs exhibit interesting radular specializations. For example, certain
forms have adaptations for boring through hard materials or harpooning prey.

Molluscan foot

1. Locomotion and Attachment:

o The molluscan foot can be adapted for locomotion, attachment to a substratum, or
a combination of functions.
o It is typically a ventral, sole-like structure that facilitates creeping locomotion
through waves of muscular contractions.
2. Modifications:
o Various modifications exist:
▪ Attachment Disc of Limpets: Limpets have an attachment disc on their foot.
▪ Hatchet Foot of Bivalves: Bivalves possess a laterally compressed foot
resembling a hatchet for digging.
▪ Siphon in Squids and Octopuses: Squids and octopuses use a siphon for jet
propulsion.
▪ Secreted Mucus: Small molluscs that glide on cilia often use secreted mucus
for adhesion or as a slime tract.
▪ Hydraulic Extension in Snails and Bivalves: In snails and bivalves, the foot
extends hydraulically from the body by engorgement with blood.
▪ Burrowing Forms: Burrowing molluscs can extend their foot into mud or
sand, enlarge it with blood pressure, and use it as an anchor to draw the
body downward.
▪ Pelagic Forms: In free-swimming molluscs, the foot may be modified into
winglike parapodia or thin, mobile fins for swimming.

Visceral Mass

Mantle and Mantle Cavity

1. Mantle Structure:
o The mantle is a tissue that extends outward from the dorsal midline of the visceral
mass.
o Its outer surface secretes the shell in molluscs.
o The mantle also contains sensory receptors and may be involved in gas exchange.
2. Mantle Cape and Mantle Cavity:
 o As the mantle grows outward, it forms a cape-like structure around the body.
o The sides of this mantle cape wrap around the body, enclosing a space called the
mantle cavity.
o The mantle cavity plays a crucial role in a mollusc’s life.
3. Respiratory Organs and Waste Disposal:
o The mantle cavity usually houses respiratory organs (such as gills or lungs) that
develop from the mantle.
o Products from the digestive, excretory, and reproductive systems are emptied into
the mantle cavity.
o In aquatic molluscs, a continuous water current (maintained by surface cilia or
muscular pumping) brings oxygen and sometimes food into the mantle cavity.
o This same water current also flushes wastes and reproductive products out into the
environment.
4. Jet Propulsion in Cephalopods:
o In cephalopods (such as squids and octopuses), the muscular mantle and its cavity
are involved in jet propulsion for locomotion.
5. Protection and Gill Function:
o Many molluscs can withdraw their head or foot into the mantle cavity (which is
surrounded by the shell) for protection.
o The gills sit within the mantle cavity and extract oxygen from the water passing
through it.
o Blood inside the gills transports oxygen throughout the body.
6. Gill Structure:
o In its simplest form, a mollusc ctenidium (gill) consists of a long, flattened axis
extending from the wall of the mantle cavity.
o Leaflike gill filaments project from this central axis.
o Water is propelled by cilia between the gill filaments, and blood diffuses from an
afferent vessel to an efferent vessel.
o The countercurrent exchange mechanism ensures efficient oxygen uptake.

Shell

1. The molluscan shell, when present, is secreted by the mantle and is lined by it.
2. Variation in Shell Structure:
o There is great variation in shell structure among different molluscs.
o Freshwater molluscs usually have a thick shell that protects acids produced in water
due to leaf litter decay.
o In many marine molluscs, the shell is relatively thin, and in some cases, it is absent.

Internal Structure and Function

1. Gas Exchange and Respiratory Organs:

o Gas exchange occurs in specialized respiratory organs such as ctenidia (gills),
secondary gills, and lungs.
o The mantle surface also plays a role in gas exchange.
o Almost all molluscs have an open circulatory system.
2. Open Circulatory System:
o In an open circulatory system, blood is not entirely contained within blood vessels.
o It flows through vessels in some parts of the body and enters open sinuses in other
parts.
o This system is common in slow-moving animals.
 o Insects are an exception; they use the tracheal system for oxygen distribution.
3. Closed Circulatory System:
o In a closed circulatory system, blood moves to and from tissues within blood
vessels.
o Most cephalopods (such as squids and octopuses) have a closed circulatory system
with vessels and capillaries.
4. Digestive Tract:
o The digestive tract in molluscs is complex and highly specialized.
o Its structure varies according to the feeding habits of different molluscs.
o Extensive ciliary tracts aid in digestion.
5. Kidneys (Metanephridia):
o Most molluscs have a pair of kidneys called metanephridia.
o These nephridia open into the coelom through a structure called the nephrostome.
o The kidney ducts also serve for the discharge of eggs and sperm.
6. Nervous System:
o The molluscan nervous system consists of several pairs of ganglia connected by
nerve cords.
o It is generally simpler than that of annelids and arthropods.
o Neurosecretory cells in the nervous system produce a growth hormone and function
in osmoregulation in certain air-breathing snails.
7. Highly Specialized Sense Organs:
o Molluscs possess various types of highly specialized sense organs for detecting their
environment.

Reproduction and Life History

1. Dioecious and Hermaphroditic:

o Most molluscs exhibit dioecious (separate sexes) reproduction, although some are
hermaphroditic (possess both male and female reproductive organs).
2. Trochophore Larva:
o A free-swimming trochophore larva emerges from the egg in many molluscs.
o This larval form is characteristic of lophotrochozoans (a larger group that includes
molluscs).
3. Direct Metamorphosis:
o Some molluscs, like chitons, undergo direct metamorphosis from a trochophore
into a small juvenile.
o However, in groups like gastropods and bivalves, the trochophore stage leads to a
unique molluscan larval stage called a veliger.
4. Veliger Larva:
o The free-swimming veliger possesses the beginnings of a foot, shell, and mantle.
o In many molluscs, the trochophore stage occurs within the egg, and the veliger
hatches as the only free-swimming stage.
5. Exceptions:
o Cephalopods, certain freshwater bivalves, and some marine snails lack free-
swimming larvae.
o Instead, their juveniles hatch directly from eggs.

CLASSES OF MOLLUSCS
Molluscan classification

• Taxonomists recognized five classes of living molluscs for over 50 years:

o Amphineura
o Gastropoda
o Scaphopoda
o Bivalvia (also called Pelecypoda)
o Cephalopoda
• Discovery of Neopilina in the 1950s added another class:
o Monoplacophora
• Hyman proposed separate classes for solenogasters and chitons:
o Aplacophora (for solenogasters)
o Polyplacophora (for chitons)
• Aplacophora was subsequently divided into sister groups:
o Caudofoveata
o Solenogastres
• Characteristics of Caudofoveata and Solenogastres:
o Wormlike and shell-less
o Calcareous sclerites or spicules in their integument
o Reduced heads
o Lack nephridia
• Despite these similarities, there are important differences between these groups.

Class Aplacophora:
• Small group or marine Mollusca
• 2) about 320 species

Class Polyplacophora: Chitons

1. Diverse Group: Chitons belong to the class Polyplacophora and represent a diverse group of
marine mollusks. There are approximately 1000 currently described species.
2. Shell Structure: Chitons have a shell composed of nine separate shell plates or valves.
These plates overlap slightly at the front and back edges, allowing flexibility for locomotion
over uneven surfaces. When dislodged from rocks, chitons can even curl up into a
protective ball.
3. Feeding Behavior: Most chitons are stay-at-home organisms, venturing only short distances
for feeding. They primarily feed by scraping algae from rocks using their radula (a
specialized feeding structure). The radular teeth are reinforced with the iron-containing
mineral, magnetite.
 4. Unusual Predatory Behavior: While most chitons are herbivores, the species Placiphorella
velata stands out as an unusual predatory chiton. It uses a specialized head flap to capture
small invertebrate prey.
5. Attachment and Protection: Chitons cling tenaciously to rocks with their broad, flat foot. If
detached, they can roll up like armadillos for protection.
6. Mantle and Girdle:
o The mantle forms a girdle around the margin of the chiton’s shell plates.
o In some species, mantle folds cover part or all of the shell plates.
o The mantle cavity of polyplacophorans extends along the side of the foot.
o The gills are more numerous in chitons compared to other molluscan classes.
7. Gill Arrangement:
o Multiple gills hang down into the mantle cavity along part or all of the lateral pallial
groove.
o Each gill consists of a central axis with flattened filaments through which oxygen is
absorbed.
o Water enters the grooves anteriorly, flows across the gills, and leaves posteriorly.
8. Adaptations for Water Conservation:
o At low tide, the mantle margins can be tightly pressed to the substratum to
diminish water loss.
o In some circumstances, the mantle margins can be held open for limited air
breathing.
9. Additional Sensory Structures:
o A pair of osphradia (chemoreceptive sense organs) occupy the mantle grooves near
the anus of many chitons.
o Blood pumped by the three-chambered heart reaches the gills via the aorta and
sinuses.
o A pair of kidneys (metanephridia) carries waste from the pericardial cavity to the
exterior.
o Two pairs of longitudinal nerve cords connect in the buccal region.
10. Reproductive Features:
o Most are diecious.
o Trochophore larvae metamorphose directly into juveniles without an intervening
veliger stage.

Class Monoplacophora

1. Extinction and Rediscovery:

o Monoplacophora were long thought to be extinct.
o Living specimens of Neopilina were discovered in 1952 from the seafloor near the
west coast of Costa Rica.
o Approximately 25 species of monoplacophorans are now known.
2. Morphology and Characteristics:
 o Monoplacophorans are small mollusks.
o They have a low, rounded shell and a creeping foot.
o The mouth bears a characteristic radula.
o Superficially, they resemble limpets.
o Unlike most other molluscs, they exhibit serially repeated organs.
3. Repeating Structures:
o Monoplacophorans have:
▪ Three to six pairs of gills
▪ Two pairs of heart atria
▪ Three to seven pairs of metanephridia
▪ One or two pairs of gonads
▪ A ladderlike nervous system with 10 pairs of pedal nerves
o This serial repetition is also seen to a more limited extent in chitons.
4. Metameric Ancestry Hypothesis:
o The repeated structures raise questions about whether molluscs had a segmented
(metameric) ancestor.
o Current research suggests that Neopilina exhibits pseudometamerism, and molluscs
did not have a metameric ancestor.
o The Serialia hypothesis proposes that monoplacophorans and polyplacophorans are
sister taxa, where serially repeated structures evolved in their common ancestor.

Class Gastropoda

1. Diversity and Size:

o Gastropoda is the largest and most diverse class among molluscs.
o It comprises over 70,000 living species and more than 15,000 fossil species.
o The diversity is so immense that there is no single general term in our language that
can fully encompass it.
2. Habitats and Forms:
o Gastropods inhabit various environments, from marine to terrestrial.
o They include a wide range of forms:
▪ Snails
▪ Limpets
▪ Slugs
▪ Whelks
▪ Conchs
▪ Periwinkles
▪ Sea slugs
▪ Sea hares
▪ Sea butterflies
3. Adaptations and Locomotion:
o Gastropods are often sluggish and sedentary due to their heavy shells and slow
locomotion.
o Some are specialized for climbing, swimming, or burrowing.
4. Shell Characteristics:
o The shell, when present, is always univalve (of one piece).
o It may be coiled or uncoiled.
o Whorls start at the apex, with each successive whorl becoming larger and spiraling
around the central axis (columella).
o Shells can be right-handed (dextral) or left-handed (sinistral) based on the direction
of coiling.
 5. Operculum and Defense:
o Many snails possess an operculum, a plate made of tanned protein.
o The operculum covers the shell aperture when the body is withdrawn into the shell,
serving as a protective lid.
6. Size Range:
o Gastropods exhibit a wide size range:
▪ From microscopic forms to giant marine snails like Pleuroploca gigantea
(with shells up to 60 cm long).
▪ Sea hares (e.g., Aplysia) can reach up to 1 meter in length.
▪ Most gastropods, however, are between 1 and 8 cm in length.
▪ Some fossil species even exceed 2 meters in length.
7. Habitat Diversity:
o Gastropods inhabit a wide range of habitats:
▪ Marine gastropods occur in littoral zones, great depths, and some are even
pelagic.
▪ They adapt to brackish water, freshwater, and terrestrial environments.
▪ Despite challenges like soil mineral content, temperature extremes, dryness,
and acidity, they are widespread.
▪ Some gastropods occur at great altitudes and even in polar regions.
8. Versatile Habitats:
o Snails occupy diverse niches:
▪ In small pools, large bodies of water, and woodlands.
▪ Under rocks, mosses, and cliffs.
▪ On trees, underground, and even on the bodies of other animals.
o They have adapted to every mode of life except aerial locomotion.
9. Defense Mechanisms:
o Gastropods protect themselves through:
▪ Shells (their chief defence).
▪ Distasteful or toxic secretions.
▪ Secretive habits.
▪ Some species can deploy stinging cells from their cnidarian prey for defence.
▪ A few, like Strombus, can deliver an active blow with their foot (bearing a
sharp operculum).
10. Predators and Parasites:
o Despite defences, gastropods are preyed upon by birds, beetles, small mammals,
and fish.
o They serve as intermediate hosts for various parasites, especially trematodes.
o Larval stages of parasites often harm snails.

Form and Function

Torsion

1. Larval Stages:
o Gastropod development begins with a trochophore larval stage.
o This is followed by a veliger larval stage, during which the shell first forms.
o The veliger larva typically has two ciliated velar lobes used for swimming, and the
developing foot becomes visible.
2. Initial Body Orientation:
o Initially, the mouth is anterior and the anus is posterior.
o However, the relative positions of several body parts change during development.
3. Torsion Process:
o Torsion is a crucial process in gastropod development.
o It occurs in two steps:
▪ First, an asymmetrical foot retractor muscle contracts, pulling the shell and
enclosed viscera (organs) 90 degrees counterclockwise relative to the head.
▪ This movement brings the anus from the posterior to the right side of the
body.
▪ Interestingly, the movement of the shell is independent of visceral
movements.
▪ The shell rotates between 90 and 180 degrees, establishing its position for
adulthood.
4. Mantle Cavity and Anus Movement:
o Previously, it was assumed that the mantle cavity, housing both the gills and the
anus, moved with the anus during the first 90 degrees of torsion.
o However, studies reveal that the mantle cavity develops on the right side of the
body near the anus but is initially separate.
o Eventually, the anus and mantle cavity move farther to the right, and the mantle
cavity remodels to encompass the anus.
5. Digestive Tract Changes:
o In a slower and more variable process, the digestive tract moves laterally and
dorsally.
o As a result, the anus lies above the head within the mantle cavity
6. Torsion Process:
o After torsion, the anus and mantle cavity open above the mouth and head.
o The left gill, kidney, and heart atrium shift to the right side, while the original right-
side structures move to the left.
o The nerve cords form a figure eight pattern.
7. Protective Adaptations:
o Due to the space available in the mantle cavity, the gastropod’s sensitive head end
can now be withdrawn into the protection of the shell.
o The tougher foot and, if present, the operculum, form a barrier to the outside.
8. Ontogenetic Torsion vs. Evolutionary Torsion:
o The described developmental sequence is called ontogenetic torsion.
o Evolutionary torsion refers to the changes that produced the modern torted
gastropod body from the ancestral untorted form.
o Traditionally, it was assumed that morphological changes in ontogenetic torsion
represented the sequence of evolutionary changes.
9. New Hypothesis:
o Recent studies propose a different scenario:
▪ The ancestral gastropod likely had two lateral mantle cavities, similar to
those in Neopilina and chitons.
▪ A single mantle cavity over the head may have arisen when the left lateral
mantle cavity was lost.
▪ The right cavity expanded toward the middle of the body after the first 90
degrees of torsion.
10. Asynchronous Displacements:
o Careful study of ontogenetic torsion reveals that asynchronous displacements of
the shell, visceral mass, and anus are possible.
o Some features move together in certain taxa.
11. Reinterpretation of Torsion:
o Torsion is now seen as a conserved anatomical stage:
 ▪ The shell has moved to the adult position.
▪ The anus and mantle cavity are on the right side of the body.
12. Detorsion and Anus Position:
o In some gastropods, varying degrees of detorsion occur in groups like
opisthobranchs and pulmonates.
o The anus opens to the right side or even posteriorly.
o However, both of these groups were derived from torted ancestors.
13. Curious Arrangement and Evolutionary Pressures:
o Torsion results in an unusual arrangement where the anus opens over the head and
gills.
o This creates the possibility of wastes being washed back over the gills and head (a
phenomenon known as fouling).
o Scientists wonder what strong evolutionary pressures led to this peculiar
realignment of body structures.
14. Proposed Explanations:
o Several explanations have been proposed, although none are entirely satisfying.
o For instance, sense organs in the mantle cavity (osphradia) might better sample
water when turned in the direction of travel.
15. Importance of Torsion:
o The consequences of torsion and the need to avoid fouling have been crucial in the
subsequent evolution of gastropods.
o However, to fully explore these consequences, we must also understand another
unusual feature of gastropods—coiling.

Gastropod coiling and its consequences:

1. Coiling and Torsion:

o Coiling, or spiral winding of the shell and visceral mass, is distinct from torsion.
o While coiling may occur during the larval stage alongside torsion, the fossil record
indicates that coiling originated earlier in gastropods than torsion did.
o All living gastropods have descended from coiled, torted ancestors, although some
have lost these characteristics.
2. Early Gastropod Shells:
o Early gastropods had a bilaterally symmetrical planospiral shell where all whorls lay
in a single plane.
o However, this planospiral shell was not very compact because each whorl had to lie
completely outside the preceding one.
3. Solution: Conispiral Shape:
o The conispiral shape solved the compactness problem:
▪ Each succeeding whorl is at the side of the preceding one.
▪ However, this shape was unbalanced.
4. Better Weight Distribution:
o To achieve better weight distribution, gastropods shifted the shell upward and
posteriorly.
o The shell axis became oblique to the longitudinal axis of the foot.
o The main body whorl’s weight and bulk pressed on the right side of the mantle
cavity.
5. Loss of Right-Side Organs:
o As a consequence, the gill, atrium, and kidney on the right side were lost in most
living gastropods.
o This led to a condition of bilateral asymmetry.
 6. Avoiding Fouling:
o Some modern species have secondarily returned to a planospiral shell form.
o Loss of the right gill likely adapted to the mechanics of carrying a coiled shell.
o Most modern prosobranchs display this condition, which allows them to avoid the
fouling problem caused by torsion.
o Water enters the left side of the mantle cavity and exits the right side, carrying
wastes from the anus and nephridiopore located near the right side.

Gastropod feeding habits:

1. Herbivorous Gastropods:
o Most gastropods are herbivorous, feeding on algae and other plant material.
o They use their specialized feeding structure, the radula, to rasp particles from hard
surfaces.
o Herbivores exhibit different feeding strategies:
▪ Grazers: These gastropods feed by grazing on surfaces, scraping algae.
▪ Browsers: Some gastropods selectively browse on specific types of algae.
▪ Planktonic Feeders: Certain species feed on plankton suspended in the
water.
2. Specific Examples:
o Haliotis (Abalone): Holds seaweed with its foot and breaks off pieces using its
radula.
o Land Snails: Forage at night for green vegetation.
o Bullia and Buccinum: Scavengers living on dead and decaying flesh.
o Melongena: Feeds on clams, especially the razor clam (Tagelus), by thrusting its
proboscis between the gaping shell valves.
3. Carnivorous Strategies:
o Some snails are carnivores:
▪ Urosalpinx cinerea (Oyster Borers): Drill holes through oyster shells using
their radula.
▪ They alternate between rasping and chemical activity to create neat round
holes.
▪ Once inserted through the hole, they feed on the soft flesh using their
radula.
4. Specialized Predators:
o Cyphoma gibbosum and related species live on gorgonians in shallow, tropical coral
reefs.
o These snails, commonly called flamingo tongues, have a brightly coloured mantle
that envelops the shell during normal activity.
o When disturbed, they can quickly withdraw their mantle into the shell aperture.
5. Venomous Adaptation:
o A gland charges the radular teeth with a highly toxic venom.
o When a Conus senses prey, a single radular tooth slides into position at the tip of its
proboscis.
o Upon striking the prey, the proboscis expels the tooth like a harpoon, and the
venom immediately paralyzes the prey.
o This adaptation prevents escape of swiftly moving prey.
6. Digestion Strategies:
o Gastropods employ various digestion methods:
▪ Ciliary feeders: Stomachs act as sorting regions, and most digestion occurs
intracellularly in digestive glands.
 ▪ Others macerate food using the radula and sometimes a gizzard, followed
by extracellular digestion in the stomach or digestive glands.

Internal Form and Function

1. Gill Arrangements:
o Most marine gastropods respire through a single gill located in the mantle cavity.
This gill is supplied with oxygen by a current of water that flows through the mantle
cavity.
o The water current follows a U-shaped path, ensuring that waste products are
flushed away from the anus (which is positioned above the animal’s head) to
prevent fouling.
o Some ancient lineages of gastropods have bipectinate gills, resembling a bird’s
feather. These gills have narrow filaments projecting on either side of a central stalk.
o In certain gastropods like abalones and keyhole limpets, there are two gills, which is
believed to be the arrangement in the earliest fossil gastropods.
o Other gastropods have lost one gill, and the remaining gill has evolved for efficient
water circulation through the mantle cavity.
o Unipectinate gills are more common, firmly anchored to the mantle wall with a
single row of filaments. These allow gastropods to thrive in muddy or sandy
environments.
2. Pulmonates (Terrestrial and Freshwater Gastropods):
o Pulmonates lack gills but have a simple lung. Their mantle contains a highly vascular
area that serves as the lung.
o The lung opens to the outside through a small opening called a pneumostome.
o Aquatic pulmonates must surface periodically to expel gas bubbles from their lungs.
o To inhale, they curl the edge of the mantle around the pneumostome, forming a
siphon.
3. Other Features:
o Most gastropods have a single nephridium (kidney).
o Their circulatory and nervous systems are well-developed.
o Sense organs include eyes or simple photoreceptors, statocysts, tactile organs, and
chemoreceptors.
o The simplest type of gastropod eye is a cuplike indentation in the skin, lined with
pigmented photoreceptor cells.
o Some gastropods have an eyecup containing a lens covered with a cornea.
o An osphradium, located at the base of the incurrent siphon, serves as a
chemosensory area in some forms, although its function remains unknown in
others.
4. Sexual Types:
o Gastropods exhibit both dioecious (separate male and female individuals) and
monoecious (hermaphroditic) reproductive systems.
o In monoecious species, there is an exchange of spermatozoa or spermatophores
during copulation.
5. Courtship and Excitement:
o Many gastropods engage in courtship ceremonies.
o Some terrestrial pulmonates (land snails) enhance excitement before copulation by
ejecting a dart from a specialized sac into their partner’s body.
6. Egg Deposition:
o After copulation, each partner deposits its eggs in shallow burrows in the ground.
 o Primitive gastropods discharge ova and sperm into seawater, where fertilization
occurs.
o Embryos then hatch as free-swimming trochophore larvae.
7. Fertilization Methods:
o Internal fertilization is common in most gastropods.
o Fertilized eggs may be:
1. Emitted singly to float among plankton.
2. Laid in gelatinous layers attached to a substratum.
3. Enclosed in tough egg capsules or a variety of egg cases.
o Offspring generally emerge as veliger larvae or directly as young snails.
8. Variations:
o Some species, including many freshwater snails, are ovoviviparous, brooding their
eggs and young in their oviduct.

Class Bivalvia (Pelecypoda)

1. Name and Characteristics:

o Bivalvia, also known as Pelecypoda (meaning “hatchet-footed”), consists of molluscs
with two shells.
o The group includes various species such as mussels, clams, scallops, oysters, and
shipworms.
o Bivalves exhibit a wide size range:
▪ From tiny pea clams measuring 1 to 2 mm in length.
▪ To giant South Pacific clams (Tridacna), which can exceed 1 m in length and
weigh up to 225 kg (500 pounds).
2. Feeding and Habitat:
o Most bivalves are sedentary filter feeders.
o They rely on cilia on their gills to create water currents that bring in food particles.
o Unlike gastropods, bivalves lack a head, a radula, and significant cephalization.
o Bivalves inhabit various environments:
▪ Marine habitats are common.
▪ Some live in brackish water, while others thrive in streams, ponds, and
lakes.

Form and Function

Shell

1. Shell Structure and Hinge Ligament:

o Bivalves have laterally compressed bodies, and their two shells (valves) are held
together dorsally by a hinge ligament.
o The hinge ligament causes the valves to gape ventrally.
o Adductor muscles work in opposition to the hinge ligament, drawing the valves
together.
2. Umbo and Growth:
o The umbo is the oldest part of the shell.
o Growth occurs in concentric lines around the umbo.
3. Pearl Production:
o Pearl production is a by-product of a protective mechanism.
o When a foreign object (such as a grain of sand or parasite) becomes lodged between
the shell and mantle, the animal responds.
o The mantle secretes many layers of nacre (mother-of-pearl) around the irritating
object.
o Pearls are formed in this process.
4. Cultured Pearls:
o To culture pearls, particles of nacre (usually taken from the shells of freshwater
bivalves) are inserted between the shell and mantle of a certain species of oyster.
o These oysters are kept in enclosures for several years.
o The Japanese extensively use the oyster genus Meleagrina for pearl culture.

Body and Mantle

1. Visceral Mass and Muscular Foot:

o The visceral mass (the internal organs) is suspended from the dorsal midline.
o The muscular foot is attached to the visceral mass anteroventrally.
2. Ctenidia (Gills):
o The ctenidia (gills) hang down on each side.
o Each gill is covered by a fold of the mantle.
3. Modified Mantle Edges:
o The posterior edges of the mantle folds are modified to form:
▪ Dorsal excurrent openings: These allow water and waste products to exit.
▪ Ventral incurrent openings: These allow water with food particles to enter.
4. Muscular Siphons (In Some Marine Bivalves):
o In certain marine bivalves, the mantle is drawn out into long muscular siphons.
o These siphons serve several purposes:
▪ Allowing the clam to burrow into mud or sand.
▪ Extending the siphons to reach the water above for respiration and feeding.

Locomotion

1. Muscular Foot and Anchoring:

o Bivalves initiate movement by extending a slender muscular foot between their
valves.
o They pump blood into their foot, causing it to swell and act as an anchor in the mud
or sand.
o Longitudinal muscles then contract to shorten the foot, pulling the animal forward.
2. Scallops and File Shells:
 o Some bivalves, such as scallops and file shells, swim with a jerky motion.
o They achieve this by clapping their valves together, creating a sort of jet propulsion.
3. Mantle Edges and Swimming Direction:
o The mantle edges can direct the stream of expelled water.
o As a result, bivalves can swim in virtually any direction.

Gills

1. Mantle and Gills:

o Bivalves perform gaseous exchange through both their mantle and gills.
o The gills are highly modified for filter-feeding.
2. Gill Structure:
o Bivalve gills are derived from primitive ctenidia (gill structures).
o Filaments on each side of the central axis greatly lengthen during evolution.
o As long filaments fold back toward the central axis, they develop a somewhat
elongated W shape.
o Filaments lying beside each other become joined by ciliary junctions or tissue
fusions.
3. Platelike Lamellae and Water Tubes:
o These modified filaments form platelike lamellae.
o Inside the lamellae, there are many vertical water tubes.
o Water enters the incurrent siphon, propelled by ciliary action.
o It then enters the water tubes through pores between the filaments in the lamellae.
o Finally, water proceeds dorsally into a common suprabranchial chamber and leaves
through the excurrent aperture.

Feeding

1. Filter Feeding:
o Most bivalves are filter feeders.
o Respiratory currents bring both oxygen and organic materials to the gills.
o Ciliary tracts direct these currents to the tiny pores of the gills.
2. Mucus Secretion and Particle Entrapment:
o Gland cells on the gills and labial palps secrete copious amounts of mucus.
o This mucus entangles particles suspended in water as it passes through the gill
pores.
o The mucous masses slide down the outside of the gills toward food grooves at the
lower edge of the gills.
3. Particle Sorting and Ingestion:
o Heavier sediment particles fall from the ctenidia (gills).
o Smaller particles travel along the food grooves toward the labial palps.
o The labial palps, being grooved and ciliated, sort the particles.
o Tasty particles encased in the mucous mass are directed into the mouth.
4. Variations in Feeding Strategies:
o Some bivalves, like Nucula and Yoldia, are deposit feeders.
▪ They have long proboscides attached to the labial palps.
▪ These proboscides can be protruded onto sand or mud to collect food
particles.
o Shipworms burrow in wood and feed on particles they excavate.
o Symbiotic bacteria live in a special organ in some bivalves and produce cellulase to
digest wood.
 o Giant clams gain much of their nutrition as adults from the photosynthetic products
of symbiotic dinoflagellates living in their mantle tissue.
o Septibranchs, another group of bivalves, draw small crustaceans or bits of organic
debris into the mantle cavity by a sudden inflow of water created by the pumping
action of a muscular septum in the mantle cavity.

Internal Structure and Function

1. Digestive System:
o The floor of the stomach in filter-feeding bivalves is folded into ciliary tracts. These
tracts help sort a continuous stream of particles.
o Most bivalves have a cylindrical style sac that opens into the stomach. This sac
secretes a gelatinous rod called a crystalline style.
o The crystalline style projects into the stomach and is kept whirling by cilia in the
style sac.
o Rotation of the style serves several purposes:
▪ It helps dissolve the surface layers of the style.
▪ It frees digestive enzymes, especially amylase, contained in the style.
▪ It assists in rolling the mucous food mass.
o Dislodged particles are sorted, and suitable ones are either directed to the digestive
gland or engulfed by amebocytes.
o Further digestion occurs intracellularly.
2. Circulatory System:
o Bivalves have a three-chambered heart located in the pericardial cavity.
o The heart consists of two atria and a ventricle.
o It beats slowly, ranging from 0.2 to 30 times per minute.
o Part of the blood is oxygenated in the mantle and returns to the ventricle through
the atria.
o The rest of the blood circulates through sinuses, passes through a vein to the
kidneys, and then to the gills for oxygenation before returning to the atria.
3. Excretory System:
o A pair of U-shaped kidneys (nephridial tubules) lies just ventral and posterior to the
heart.
o The glandular portion of each tubule opens into the pericardium.
o The bladder portion empties into the suprabranchial chamber.
4. Nervous System and Sense Organs:
o The nervous system consists of three pairs of widely separated ganglia connected
by commissures and a system of nerves.
o Sense organs are poorly developed and include:
▪ Statocysts in the foot.
▪ Oshradia (of uncertain function) in the mantle cavity.
▪ Tactile cells.
▪ Sometimes simple pigment cells on the mantle.
▪ Scallops (Aequipecten, Chlamys) have a row of small blue eyes along each
mantle edge. Each eye has a cornea, lens, retina, and pigmented layer.
▪ Tentacles on the margin of the mantle of Aequipecten and Lima have tactile
and chemoreceptor cells.

Reproduction and Development

1. Sexes and Gamete Discharge:

 o Bivalves usually have separate sexes.
o Gametes (sperm and eggs) are discharged into the suprabranchial chamber to be
carried out with the excurrent flow.
o Remarkably, an oyster may produce 50 million eggs in a single season.
2. Fertilization:
o In most bivalves, fertilization is external.
o The embryo undergoes several developmental stages:
▪ Trochophore
▪ Veliger
▪ Spat (juvenile shelled form)
3. Freshwater Clams and Glochidia:
o In most freshwater clams, fertilization is internal.
o Eggs drop into the water tubes of the ctenidia (gills).
o There, they are fertilized by sperm entering with the incurrent flow.
o The developing larvae reach a stage called glochidium, which is a specialized veliger.
o Glochidia need to attach themselves to specific fish hosts and live parasitically for
several weeks to complete their development.
4. Mussel Larval Tactics:
o Various mussel species employ unique tactics to get their larvae in contact with
suitable fish hosts.
o Some simply discharge glochidia into the water column.
▪ If they come into contact with a suitable passing fish or amphibian, they
attach to the gills or skin and complete their development.
o Other species use the mantle flap of brooding females.
▪ This mantle flap holds the glochidia in a gelatinous packet called a
conglutinate.
▪ The conglutinate’s size and shape are unique to each mussel species.
▪ The mantle flap acts as a lure to bring potential host species into contact
with the glochidia.
▪ For example, the conglutinate of a gravid female pocketbook mussel
(Lampsilis ovata) grows to resemble a small fish.
▪ The mantle flap is then wriggled like a fishing lure to attract nearby bass or
other sunfish.
▪ These fish serve as hosts for the glochidia.
▪ When a hungry bass strikes the mantle, it gets a mouthful of glochidia,
which promptly attach to the fish’s gills.
▪ After encysting on a suitable host to complete development, the juveniles
detach and sink to the substratum to begin independent lives.
▪ This larval “hitchhiking” helps distribute organisms whose locomotion is
very limited while also preventing larvae from being swept downstream out
of lakes.

Class Scaphopoda
 1. Description:
o Scaphopoda, commonly called tusk shells or tooth shells, are benthic marine
mollusks.
o They are found from the subtidal zone to over 6000 m in depth.
o Their body is slender, covered with a mantle, and enclosed in a tubular shell open
at both ends.
o The scientific name Scaphopoda means “shovel foot,” referring to the head used for
burrowing in marine mud and sediments.
2. Shell Characteristics:
o Scaphopod shells are curved, tubular, and shaped like elephant tusks.
o Most species are 2.5 to 5 cm long, but they can range from 4 mm to 25 cm long.
o Fossils show specimens up to 30 cm long.
3. Burrowing and Respiration:
o The foot, protruding through the larger end of the shell, is used to burrow into mud
or sand.
o The narrow posterior end of the shell remains exposed to the water above.
o Gills are absent, and gaseous exchange occurs in the mantle.
o Most food consists of detritus and protozoa from the substratum.
o Captacula (long tentacles) extending from the head help capture food and convey it
to the nearby mouth.
o Sexes are separate, and the larva is a trochophore.

Class Cephalopoda

Cephalopoda (squids, octopuses, nautiluses, devilfish, and cuttlefish):

1. Diverse Group:
o Cephalopoda includes various marine creatures such as squids, octopuses,
nautiluses, devilfish, and cuttlefish.
o All of them are active predators.
2. Feeding and Predation:
o Cephalopods are predaceous, primarily feeding on small fishes, molluscs,
crustaceans, and worms.
o Their arms are highly mobile and used to capture and handle food.
o They swiftly seize prey and bring it to the mouth.
3. Salivary Glands and Digestion:
o Octopuses and cuttlefishes have salivary glands that secrete venom for
immobilizing prey.
o Their strong, beaklike jaws grasp prey, and the radula tears off pieces of flesh.
 o Salivary gland secretions begin the digestive process, allowing partially liquefied
food to be ingested.
o Extracellular digestion occurs in the stomach and cecum.

Form and Function

Shell

1. Cuttlefishes, Squids, and Octopuses:

o Cuttlefish: Cuttlefishes have a small, curved shell entirely enclosed by the mantle.
o Squids: Most of the shells in squids has disappeared, leaving only a thin,
proteinaceous strip called a pen, which is also enclosed by the mantle.
o Octopus: In octopuses, the shell has disappeared entirely.

Locomotion

1. Jet Propulsion:
o Cephalopods swim using jet propulsion.
o They forcefully expel water from the mantle cavity through a ventral funnel (or
siphon).
o The funnel is mobile and can be pointed forward or backwards to control direction.
o The force of water expulsion determines their speed.

2. Squids and Cuttlefishes:

o Squids:
▪ Streamlined body built for speed.
▪ Excellent swimmers.
▪ Lateral fins serve as stabilizers but are held close to the body for rapid
swimming.
o Cuttlefishes:
▪ Swim more slowly than squids.

3. Nautilus:
o Active at night.
o Its gas-filled chambers keep the shell upright.
o Moves surprisingly well, although not as fast as squids.
 4. Octopus:
o Globular body with no fins.
o Can swim backward by spurting jets of water from its funnel.
o Better adapted to crawling over rocks and coral, using suction discs on its arms to
pull or anchor itself.
o Some deepwater octopods have webbed arms and swim similarly to medusae
(jellyfish).

Internal Features

The active habits of cephalopods are reflected in their internal anatomy, particularly their
respiratory, circulatory, and nervous systems.

cephalopod gills and circulatory systems:

1. Gills:
o Nautiloids Exception: Except for nautiloids, cephalopods have one pair of gills.
o Ciliary Propulsion: Unlike other organisms, cephalopod gills lack cilia because ciliary
propulsion wouldn’t circulate enough water for their high oxygen requirements.
o Radial Muscles: Instead, radial muscles in the mantle wall compress the wall,
enlarging the mantle cavity and drawing water inside.
o Expelling Water: Strong circular muscles contract and forcibly expel water through
the funnel (ventral siphon).
o One-Way Valves: A system of one-way valves prevents water from being taken in
through the funnel or expelled around the mantle margin.
2. Circulatory System:
o Ancestral Molluscs: The open circulatory system of ancestral molluscs would be
inadequate for cephalopods.
o Closed Network: Cephalopods have a closed network of vessels for their circulatory
system.
o Capillaries and Gills: Capillaries conduct blood through the gill filaments.
o Systemic Circulation: Unlike vertebrates, where blood goes directly to the gills or
lungs, the molluscan plan of circulation places the systemic circulation before the
blood reaches the gills.
o Branchial Hearts: To address this functional problem, cephalopods developed
accessory or branchial hearts at the base of each gill. These hearts increase the
pressure of blood flowing through the capillaries there.

Nervous and Sensory Systems

1. Elaborate Nervous and Sensory Systems:

o Cephalopods exhibit more sophisticated nervous and sensory systems compared to
other molluscs.
o Their brain is the largest among invertebrates.
 o The brain consists of several lobes containing millions of nerve cells.

2. Squids:
o Squids possess giant nerve fibers, which are among the largest known in the animal
kingdom.
o These fibers are activated when the squid is alarmed.
o They initiate maximal contractions of the mantle muscles, enabling a speedy
escape.
3. Complex Eyes:
o Except for Nautilus, cephalopods have highly complex eyes.
o Eye components include the cornea, lens, chambers, and retina.
o Statocysts control eye orientation; they are larger and more complex than in other
molluscs.
o Cephalopod eyes are held in a constant relation to gravity, ensuring that the slit-
shaped pupils remain horizontal.
o Their eyesight is excellent, with visual acuity underwater far surpassing our own.
4. Learning and Discrimination:
o Cephalopods can be taught to discriminate between shapes (e.g., square and
rectangle).
o They remember such discrimination for a considerable time.
o Observational learning occurs: When one octopus observes another being rewarded
for a correct choice, the observer learns and consistently makes the same selection.
5. Tactile Exploration:
o Octopods use their arms for tactile exploration.
o They can discriminate between textures by feel but not necessarily between shapes.
o Their arms are well supplied with both tactile and chemoreceptor cells.
6. Sense of Hearing:
o Interestingly, cephalopods seem to lack a sense of hearing.

Cephalopods’ remarkable neural adaptations contribute to their agility, intelligence, and survival
strategies.

cephalopod communication and signalling:

1. Social Behavior and Communication:

o Nautiloids and Deep-Water Cephalopods: Little is known about the social behaviour
of nautiloids or deep-water cephalopods.
o Studied Forms: Inshore and littoral cephalopods, such as Sepia, Sepioteuthis, Loligo,
and Octopus, have been extensively studied.
2. Predominant Means of Communication:
o While cephalopods have well-developed tactile senses and some chemical
sensitivity, visual signals are the primary mode of communication.
o These visual signals involve a variety of movements of the arms, fins, and body, as
well as colour changes.
3. Color Changes and Chromatophores:
o Chromatophores are specialized cells in the skin containing pigment granules.
o Tiny muscle cells surround each elastic chromatophore.
o When these muscles contract, they pull the cell boundary outward, causing the
chromatophore to expand greatly.
o As the cell expands, the pigment becomes dispersed, altering the animal’s colour
pattern.
 o Relaxation of the muscles allows chromatophores to return to their original size,
concentrating the pigment again.
4. Elaborate Colour and Pattern Changes:
o Under nervous and hormonal control, cephalopods achieve an elaborate system of
colour and pattern changes:
▪ General darkening or lightening.
▪ Flushes of pink, yellow, or lavender.
▪ Formation of bars, stripes, spots, or irregular blotches.
o These colours serve various purposes, including:
▪ Danger signals.
▪ Protective coloring.
▪ Courtship rituals.
▪ Other adaptive behaviours.
5. Instantaneous Message Transmission:
o By assuming different colour patterns on different body parts, a squid can
simultaneously transmit three or four different messages to different individuals
and in different directions.
o The cephalopod can instantaneously change any or all of these messages.
6. Colour Vision:
o Despite their ability to match colourful backgrounds, cephalopods likely lack colour
vision.
o In an experiment with equal-intensity background colours, they were unable to
match colours.
o Their photoreceptors are rhabdomere cells, not rods and cones like in vertebrates.
o Most cephalopods have only one visual pigment, making colour detection unlikely.
o An exception is the bioluminescent firefly squid, which has three visual pigments,
similar to humans.
7. Luminescent Organs:
o Many deep-water cephalopods have evolved elaborate luminescent organs.
o The wonderful color patterns we see on cephalopod skin are visible to other
cephalopods as polarized light patterns.
o Cephalopods use this ability to enhance discrimination of translucent prey and fish
with silvery reflective scales.
8. Ink Defense Mechanism:
o Most cephalopods (except nautiloids) use ink as a protective device.
o The ink sac empties into the rectum and contains sepia, a dark fluid with melanin
pigment.
o When alarmed, the cephalopod releases a cloud of ink:
▪ The ink may hang in the water as a blob or be contorted by water currents.
▪ The animal quickly departs, leaving the ink as a decoy to confuse predators.

In summary, cephalopods employ a sophisticated visual language through colour and movement,
conveying a wealth of information rapidly.

Reproduction

1. Sexual Dimorphism:
o Cephalopods exhibit separate sexes.
o Spermatozoa are encased in spermatophores and stored in a sac that opens into
the mantle cavity.
2. Male Reproductive Organ:
 o Hectocotylus: One arm of adult males is modified as an intromittent organ.
o The hectocotylus is used to pluck a spermatophore from the male’s own mantle
cavity and insert it into the mantle cavity of a female near the oviduct opening.
3. Male Behavior and Color Displays:
o Color Displays: Before copulation, males often undergo color displays, apparently
directed against rival males.
4. Egg Fertilization and Attachment:
o Eggs are fertilized as they leave the oviduct.
o They are usually attached to stones or other objects.
5. Parental Care:
o Some octopods tend to their eggs.
o Argonauta (paper nautilus) females secrete a fluted “shell” or capsule in which eggs
develop.
6. Embryonic Development:
o The large, yolky eggs undergo meroblastic cleavage.
o During embryonic development, the head and foot become indistinguishable.
o The ring around the mouth, which bears the arms (tentacles), may be derived from
the anterior part of the foot.
7. Juvenile Stage:
o Juveniles hatch from eggs; there is no free-swimming larva stage in cephalopods .

PHYLOGENY AND ADAPTIVE DIVERSIFICATION

1. Ancient Origins:
o Fossils attributed to Mollusca appear in geological strata as old as the early
Cambrian period.
o A Cambrian fossil radula was recently found in Alberta, Canada.
2. Protostomes and Trochozoa:
o Molluscs are protostomes, sharing features like spiral cleavage, mesoderm from
the 4d blastomere, and a trochophore larva.
o They are allied with annelids in the lophotrochozoan subgroup Trochozoa.
3. Relationships Among Lophotrochozoans:
o Opinions differ on the exact relationships among lophotrochozoans.
o Some suggest that molluscs and annelids are sister taxa, but the branching order is
not depicted.
4. Annelids vs. Molluscs:
o Annelid worms have a developmental pattern similar to that of molluscs.
o However, the annelid body is metameric, composed of serially repeated segments.
o In contrast, there are no true segments in molluscs.
5. Coelom and Protostomes:
o Both annelids and molluscs are coelomate protostomes.
o However, the coelom is greatly reduced in molluscs compared to annelids .
6. Segmented Ancestor Hypothesis:
o The hypothesis that annelids and molluscs shared a segmented ancestor is
strengthened if the repeated body parts present in Neopilina (class
Monoplacophora) and in some chitons can be considered evidence of metamerism.
o However, morphological and developmental studies indicate that these parts are
not remnants of an ancestral metameric body.
7. Evolution of Repeated Parts:
 o Analysis of molecular characters from a wide range of molluscs, including a
monoplacophoran, suggests that the evolution of repeated parts (such as gills and
muscles) is complex.
o The cladogram from this study places monoplacophorans as the sister taxon to
chitons, uniting the two taxa with repeated body parts in a clade called Serialia.
o Serialia is phylogenetically nested within a clade of unsegmented molluscs,
indicating that the repeated structures are derived molluscan features, not
ancestral features.
8. Recent Molecular Phylogenies:
o Recent molecular phylogenies do not support the Serialia hypothesis.
o Instead, these studies support clade Aculifera, uniting caudofoveates and
solenogasters.
o These two wormlike taxa appear to be derived forms that lost the shell plates and
other features.
9. Conchifera and Cephalopods:
o Conchifera is a clade of shelled molluscs.
o Relationships among members of Conchifera differ with specific studies.
o The placement of cephalopods is especially variable:
▪ Previous studies placed cephalopods as sister to gastropods.
▪ Newer work shows them as the sister taxon to monoplacophorans.
▪ One study places cephalopods outside Conchifera as the sister taxon to
Aculifera.
10. Habitat Adaptations:
o Molluscs exhibit diversity due to their adaptation to different habitats and modes
of life.
o Their feeding methods range from sedentary filter-feeding to active predation.
11. Radular Diversity:
o There is an enormous variety in radular structure and function, especially among
gastropods.
12. Versatile Mantle:
o The glandular mantle is a highly adaptable molluscan structure.
o Besides secreting the shell and forming the mantle cavity, it can be modified into
gills, lungs, siphons, and apertures.
o It sometimes functions in locomotion, feeding processes, or as a sensory organ.
13. Evolution of Shells:
o The shell has undergone various evolutionary adaptations.
o A recent study reveals that the cap-shaped “limpet” shell of gastropods evolved
independently 54 times across fossil and modern taxa.