M
Mollusk
Vimala K John1 and Akash Gautam2
1
Research and PG Department of Zoology, St:
Thomas’ College, Thrissur, India
2
Centre for Neural and Cognitive Sciences,
University of Hyderabad, Hyderabad, India
Synonyms
Mollusca
Definition
Mollusks are the members of a diverse group of
invertebrates with a thin fleshy soft unsegmented
body, divisible into a head, visceral mass, and
foot. Members of this phylum exhibit an “organ
system” grade of the organization (Anderson
2001), e.g., snails, octopus, squids, oysters, chi-
tons, mussels, etc.
Etymology
The term mollusk was coined by the French nat-
uralist Lamarck. This term is originated from the
Latin word molluscu, meaning “thin shelled,”
derived from the root Latin word Mollis, meaning
“soft.” Cuvier first used the term in 1798 to
describe the animals with reduced shells, such as
© Springer Nature Switzerland AG 2021
J. Vonk, T. K. Shackelford (eds.), Encyclopedia of Animal Cognition and Behavior,
https://doi.org/10.1007/978-3-319-47829-6_1203-1
cuttlefish, snails, squids, etc. The study of mol-
lusks is known as malacology, whereas mollusks’
shell study is called conchology. Due to the vast
diversity of species and many fossil forms, they
are considered the largest marine phylum and
comprise about 23% of all marine organisms
(Amsler et al. 2001). They constituted the
second-largest phylum of non-chordate after
arthropods and were believed to appear during
the Cambrian period (Ruppert and Barnes 1994).
Present Taxonomical Status
The phylum Mollusca is considered one of the
most conspicuous of the invertebrate group,
consisting of about 220,000 living species. The
fossil record also indicates a long and extensive
history (Telford and Budd 2011). Mollusks are
one of the largest metazoan phyla, which includes
about 130,000 described extant species and about
70,000 described fossil species (Haszprunar et al.
2008). As per Ponder and Lindberg (2008), Mol-
lusk fauna’s total living species diversity com-
prises 20,000–30,000. According to modern
taxonomists, the following classes include the
mollusk phylum: the Gastropoda (slugs, snails,
etc.), the Bivalvia or pelecypods (mussels,
clams, etc.), the Cephalopoda (sepia, cuttlefish,
octopus, etc.), the Polyplacophora (chitons), the
Scaphopoda (dentalium), the Monoplacophora
(univalves), and the Aplacophora (Barth and
Broshears 1982).
2 Mollusk
Habit, Habitat, and Ecological
Importance
Mollusks’ habitat ranges from the deepest ocean
to intertidal/abyssal zone, freshwater, and lands.
A few members form a significant part of fresh-
water fauna, and the others include terrestrial hab-
itat. They are found on rocky, sandy, or muddy
substrata, and some members are burrowing,
some are crawling, few members become attached
to a substratum, or some are free-swimming
forms. Among all classes, Gastropoda is the
most important and comprises more than 80% of
all mollusk species (Fortunato 2015). Cephalo-
pods such as squids, cuttlefish, etc. are considered
the most neurologically advanced invertebrates
(Amsler et al. 2001).
Mollusks are bilaterally symmetrical and slug-
gish invertebrates (Apte 2015). Their size ranges
from microscopic organisms to 20 m long. They
are considered important ecosystem engineers to
frame and protect the aquatic habitat’s bottom
(Fortunato 2015). These animals play an essential
role in stabilizing many marine ecosystems and
keeping the algal and bacterial levels balanced.
They play a crucial role in recycling nutrients, soil
generation, and water filtration due to their abun-
dance. Most bivalve members contribute to the
organic turnover in the intertidal (littoral) zones
of marine and freshwater ecosystems because of
their active filtering. However, this filtering activ-
ity may also seriously interfere with the various
invertebrate larval populations or planktons found
suspended and free swimming in water. They also
form an essential trophic level in the aquatic food
chain and act as good bioindicators for the eco-
logical quality in all types of aquatic habitats
(Avila et al. 2000, 2004). Most members form
food sources to a wide variety of other taxonomic
groups like humans, birds, fish, mammals, and
other invertebrates.
Mollusks play a crucial role in the lives of
humans. They form a rich source of protein,
good fat, and minerals (e.g., lime from the shells
is used in poultry food and building material). In
many regions, mollusk shells are used to manu-
facture ornaments and jewelry. This phylum plays
a significant role in the pearl industry. Pearl
oysters (Pinctada margaritifera and P. mertensi),
which inhabit the Indian and Pacific Ocean’s
warmer parts along the coast of China, India, Sri
Lanka, and Japan, form the most valuable natural
pearls. Pearls are produced by most bivalves
(pelecypods), including freshwater calms and
marine oysters. When a small foreign object
such as a particle of sand or parasite lodges
between the shell and mantle, the foreign object
becomes a nucleus around in which concentric
layers of nacreous are laid by mantle to form a
pearl (Avila et al. 2000).
A few members of this group are used as a
medicine, especially for sore throat and cough.
A recent research shows that a wide variety of
secondary metabolites from mollusks possess
anticancer, anti-inflammatory, and antimicrobial
abilities. Most of these compounds are in clinical
trials. The extracts from the shells and bioactive
compounds of some mollusks act as growth inhib-
itors of cancer cells in mice (Beckendorf et al.
2005; Cohan et al. 2003).
Some mollusks, such as slug, shipworm, etc.,
act as pests and destroy garden and agriculture
cultivations by eating leaves and cutting up roots
and stems. Moreover, a few members of gastro-
pod species are intermediate hosts for disease-
causing parasites such as liver fluke (phylum
Platyhelminthes, class Trematoda), which causes
fascioliasis or liver rot in cattle. The zebra mussel
Dreissena polymorpha is considered a harmful
exotic invader. They were carried from Europe
in ship ballast water to the Great Lakes and caused
commercial damage by clogging the water pipes
of power plants as well as cooling systems. They
are driving many native freshwater bivalve spe-
cies to extinction. Teredo worms, commonly
called shipworms, burrow into the wooden struc-
tures immersed in the sea and cause severe dam-
ages to ships, piers, etc. (Avila et al. 2004).
Mantle and Mantle Cavity
One of the unique features of mollusks is the
presence of the mantle and mantle cavity. A thin
fleshy double fold of tissue that lies beneath the
shell and the part of the epidermis/skin is known
Mollusk
as the mantle. This mantle secretes calcium car-
bonate layers, which form the mollusk’s shell
(Canti 2017) (Fig. 1). The mantle is rigidly
attached to the shell only at the outer edges and
covers the visceral organs. The mantle also
encloses a cavity called the mantle cavity between
the mantle and the body. The mantle cavity func-
tions as a respiratory chamber in most of the
mollusks. The mantle’s own exposed surface
serves for gaseous exchange. In addition to that,
the mantle cavity also possesses anus,
osphradium, nephridia, and gonophores. So prod-
ucts from the digestive, excretory, and reproduc-
tive systems are emptied into the mantle cavity. In
aquatic mollusk, there is a continuous water cur-
rent, which kept moving by the surface of cilia or
by muscular pumping, brings in oxygen and in
some forms flushes out the food waste, and carries
reproductive products (egg/sperm) out to the envi-
ronment. In aquatic mollusks, the mantle is usu-
ally equipped with sensory receptors to detect the
quality of environmental water current. In cepha-
lopods, the muscular mantle and mantle cavity
create a jet propulsion, which helps in locomotion.
In cephalopods (squids and octopuses), the mus-
cular mantle and its cavity create a jet propulsion
used in locomotion. Many mollusks can withdraw
their head or foot into the mantle cavity,
surrounded by the shell, for protection.
Mollusk, Fig. 1 Mantle cavity. http://www.mesa.edu.au/molluscs/molluscs_01.asp
3
Shell
Mollusks possess a calcareous exoskeleton
known as a shell, which encloses, supports, and
protects their visceral organs. There is great vari-
ation in shell structure. The first shell appears
during the larval period and grows continuously
throughout life. Most of the members have an
external shell, whereas the members, like a slug,
cuttlefish, squid, etc., have an internal shell, but it
is absent in octopus (Linstädter and Kehl 2012).
The mantle secretes the shell made up of several
layers consisting mainly of chitin and a protein
hardened with calcium carbonate called conchiolin
(Barnes et al. 2009).
Typically the shell consists of three layers
(Fig. 2):
a) The outer layer periosteum – It is the horny
layer composed of an organic substance called
conchiolin consist of quinone-tanned protein,
which helps protect underlying calcareous
layers from erosion by boring organisms.
A fold of mantle edge secretes this, and growth
occurs only at the shell’s margin, and at the
older parts of the shell, this periostracum often
becomes worn out.
b) The middle layer prismatic layer, composed of
densely packed prisms of calcium carbonate, is
laid down in a protein matrix that is
4 Mollusk

Shell

Shell Mantle

Periostracum

Prismatic
layer

Nacre

Mantle Pearl

Mantle
Outer folds
mantle
epdithelium
New
a periostracum
b

Mollusk, Fig. 2 (a) Diagrammatic vertical section and mantle of a bivalve. (b) Formation of a pearl. https://
biocyclopedia.com/index/general_zoology/form_and_function02.php

strengthened due to more calcium carbonate the foreign body, and it grows over time, and a
crystals made of columnar calcite. The man- pearl is formed.
tle’s glandular margin secretes this layer, and
the increase in shell size occurs at the shell
margin as the animal grows.
Foot
c) The inner layer called the nacreous layer, made
of laminated calcite, is responsible for pearl
The ventral body wall of mollusks is modified as a
formation. This layer lies next to the mantle
muscular foot. It varies in shape and function in
and is secreted continuously, so it increases in
different classes of mollusks. In gastropods, the
thickness during the life of the animal. The
foot is broad and expanded with a creeping sole.
calcareous nacre is laid down in thin layers.
Rhythmic contractions of the muscular foot per-
Fragile and wavy layers produce the iridescent
form the locomotion. Moreover, the mucus-
mother of pearl. A freshwater mollusk usually
secreting glands present in the foot secrete
has a thick periostracum, whereas it is very
mucus on which the animal slides. In bivalves
thin, or in some members, it is absent in marine
(Pelecypoda), the foot is a large plow or wedge-
forms.
shaped adapted for burrowing. A cephalopod’s
foot is used for jet propulsion, and the tentacles
When a foreign body or microbe/stone enters
and arms are derived from the foot.
and touches the mantle, the mantle’s shell secretes
Inpolyplacophora, the foot is the broad flat mus-
cells that build up layers of the nacreous layer
cular, glandular creeping sole and ventral in posi-
around it, and pearls are formed. To protect itself,
tion. In scaphopods, the foot is long, narrow
the mollusk secretes the substance, a mineral
protrusible spade-like, and is used for digging.
called aragonite and conchiolin, a protein that is
For mollusks such as limpets, the foot acts as a
the same substance it secretes to form its shell.
sucker attaching the animal to a hard surface
The composite of all these is called nacre or
(Kotpal 2009).
mother of pearl. The layers are deposited around
Mollusk 5

Radula The pattern and number of teeth in a row are

teeth species-specific. The movement of the radula is
back and forth over the odontophore cartilage.
During feeding, the animal opens the mouth; the
Odontophore odontophore is thrust forward, and the radula
gives a strong scrape backward, brings food into
the pharynx, and closes the mouth. This process is
repeated rhythmically. The radula is generally
used for feeding and is present in many species
and serves to shred or scrape food like bacteria
and algae of rocks. The mouth of mollusks is
provided with slimy mucus-secreting glands to
which the food particles are trapped. The trapped
Esophagus food enters the stomach by ciliary action in such a
Radula way that the mucus forms a long string called food
sac Odontophore string (Alison et al. 2020). The stomach has some
acidic nature, which makes the mucus less sticky
and frees the food particles from it, and reaches
the structure called pyrostyle at the region of the
hindgut. The pyrostyle is rotated by another set of
cilia, which act as a bobbin and send the smaller
particles, mainly minerals, to the pyrostyle to get
excreted. Compared to the smaller particles, the
Radula large food particles are sent to the stomach’s
Mouth
retractor cecum for digestion and absorption. Excretory
Odontophore organs consist of paired nephridia, which filter
retractor Radula the waste products and open to the mantle cavity
Radula and push them out with the help of water current
protractor Odontophore produced by ctenidia (Jordan and Verma 2001).
A protractors

Mollusk, Fig. 3 Lateral section of a gastropod head

showing the radula and radula sac. https://biocyclopedia.
com/index/general_zoology/form_and_function02.php
Feeding and Digestion
Most members of this family enjoy intracellular
digestion. The buccal cavity’s interior region has a
tongue-like organ called a radula (Fig. 3). The
radula bears a ribbon of chitinous tooth-like orna-
mentation, supported by a muscular structure
known as odontophore. The teeth are pointed
backward. There are complex muscles that move
the radula and its supporting cartilages. There may
be a few or as many as 250,000 teeth. When the
radula wears away, new rows of teeth are contin-
uously replaced by secretion at its posterior end.
Respiration
In mollusk, the process of gaseous exchange
occurs through the body surface, particularly
through the mantle and in specialized respiratory
organs such as ctenidia, secondary gills, and
lungs. Respiration in aquatic members of mol-
lusks is performed by gills or ctenidia. The mem-
bers of the terrestrial habitat perform pulmonary
respiration. The feather-like gills are filamentous
and provided with cilia. The ctenidia/gills are
located on opposite sides of the mantle cavity
and are arranged so that the cavity is functionally
divided into an incurrent chamber and an excur-
rent chamber where the water enters through the
bottom exit near the top (Fig. 4). There are three
cilia sets, one set makes the water current into the
6 Mollusk
Mollusk, Fig. 4 Respiratory system. https://www.daviddarling.info/images3/bivalve_section.jpg
mantle cavity, and two other sets keep the ctenidia
clean. If the osphridia detects any unfavorable
entry, the ciliary beating stops and prevents cur-
rent water entry. The gills are supplied with blood
vessels connected to the hemocoel (Jordan and
Verma 2001). Cilia between gill filaments propel
many leaf-like gill filaments project from the cen-
tral axis, and water and blood diffuse from an
afferent vessel in the central axis through the
filament to an efferent vessel. The direction of
blood flow is opposite to the water current,
thereby establishing a counter current exchange
mechanism.
Nervous System and Organs of
Sensation
In general, their nervous system has paired cere-
bral pleural, pedal, and visceral ganglia and sim-
pler than annelids and arthropods. Typically a
mollusk consists of a pair of cerebral ganglia
(mass of nerve cells bodies) that innervate the
head, mouth, and associated sense organs. Arising
from the dorsal ganglia, two pairs of longitudinal
nerve cords arise: a pair of lateral (pleural) nerve
cords often forming pleural ganglia which inner-
vate the mantle and a pair of pedal nerve cords that
often include pedal ganglia which innervate the
foot. In primitive forms, both nerve cords are
interconnected by lateral branches of nerve fibers.
A buccal nerve loop with paired ganglia generally
supplies the radular apparatus in the head. Poste-
rior paired visceral ganglia innervate the viscera.
Because of torsion (twisting of the body during
development), special nerve configurations are
found in gastropods. In cephalopods, a cartilagi-
nous capsule encloses the concentrated mass of
ganglia. The cephalopods have two pairs of nerve
cords with four paired ganglia. The visceral nerve
cord serves the internal organs, whereas the pedal
nerve cord serves the foot. The ganglia located
above the esophagus are cerebral, pleural, and
visceral, and the ganglia situated below the esoph-
agus are the pedal (Jordan and Verma 2001).
Monoplacophora and Aplacophora have a very
primitive ladder-like nervous system (Fig. 5).
The cephalopod members possess well-developed
vertebrate-like eyes, and there are photoreceptors
on the mantle margins of some bivalves.
Circulatory System
Most mollusks’ circulatory systems are mainly
open type, where the blood or hemolymph
pumps into the open space around the visceral
organs and nourishes the visceral organs. As mol-
lusks are true coelomates, the main body cavity is
called hemocoel, through which the coelomic
fluid and blood circulate (Pechenik 2015). The
hemocoelic spaces act as a hydrostatic skeleton.
Mollusk
Mollusk, Fig. 5 Nervous system of gastropods. https://commons.wikimedia.org/wiki/File:Gastropod_nervous_system.
gif
The heart possesses one or two atria/auricle pairs
that receive the gills’ oxygenated blood and
pumps it into ventricles. From there, it enters
into the main blood vessel called the aorta that
opens into the hemocoel. The auricle/atria per-
form the excretory system’s function partly by
filtering waste products out of the blood and
dumping them into the coelom as urine (Ruppert
et al. 2004). The body fluids are transported
largely within sinuses devoid of distinct epithelial
walls; the heart is posterior-dorsal, enclosed in a
pericardium that typically consists of a ventricle
and two posterior auricles. The blood or hemo-
lymph is drained from ctenidia, gills, or the spe-
cialized respiratory epithelia into respective
chambers. The ventricle pumps the hemolymph
through the middorsal sinuses (in solenogasters
and scaphopods) or vessel (aorta) into the body
tissues. Hemocyanin is the respiratory pigment
that contains copper rather than iron: in more
advanced forms, hemoglobin is bound to blood
cells. In chiton and some members of mono-
placophorans, the heart is also the site of purifica-
tion discharged waste into the pericardium and via
a pair of pericardial outlets (modified kidney/
nephridia) (Fig. 6).
7
Excretory System
The anatomical structure of the renal gland varies
from species to species. Still, a common plan is
evident. The atria/auricle of the heart also func-
tions as part of the excretory system by filtering
waste products known as the organs of Bojanus.
The renal gland is a relatively wide tube opening
from a sac (pericardium) that surrounds the heart
at one end and to the mantle cavity at the other
end. Some mollusks possess Keber’s organ, oth-
erwise called the pericardial gland. It is a large
reddish-brown glandular mass situated ante-
rolateral to the pericardium into which the waste
materials are excreted. A majority of mollusks are
aquatic, so they excrete nitrogen in the form of
ammonia (Baeumler et al. 2012). In octopus the
excretion is in the form of ammonium chloride.
Terrestrial snails and slugs excrete uric acid and
excrete ammonia when the animal is in moist
surroundings. In all mollusks, the primary urine
production appears to be filtration of blood. This
may take place through the heart walls into the
pericardium or from blood vessels that supply the
glandular part of the renal gland. These are located
at the rear of the coelom, extract any reusable
8 Mollusk

Mollusk, Fig. 6
Circulatory system

material from the urine, dump additional waste
products into it, and then eject it via tubes that
discharge into the mantle cavity (Brusca et al.
2014). In most freshwater mollusks, the
reabsorption process takes place by glandular
tube and in the wide tubule. So the final urine is
more dilute than the blood.
Reproduction
Most mollusks are dioecious, and fertilization
occurs externally, except terrestrial mollusks
such as snails, slugs, and cephalopods (Furuhashi
et al. 2009). In adult cephalopods and other rep-
resentatives, the paired dorsal gonad retains the
developmental connection with the pericardium.
Some solenogasters egg and sperms are
discharged into the pericardial cavity and through
pericardial outlets discharge to an environment
where the fertilization takes place. In more
advanced mollusks, separate gonoducts
(oviducts for female gametes vas deferens for
male gametes) produce the egg and sperm respec-
tively to the mantle cavity. In some mollusks, the
zygote hatches and undergoes two larval stages –
trochophore and veliger – before becoming a
young adult; bivalves may exhibit a third larval
stage, glochidia. Eventually, the larva sinks to the
seafloor and metamorphoses into an adult form
(Buchsbaum et al. 2013).
Classification of Phylum Mollusca
There are seven living classes and two extinct
groups. First, we will see the living classes:
1. Gastropoda (Gr. gaster-belly+podios-foot):
This is the largest class of mollusks and
includes snails, conchs, abalones, and whelks
sea hares, garden slugs, etc. Because their
stomach is situated on foot, they are named
gastropods. The member’s habitat includes
marine, freshwater, or terrestrial. Their body
is asymmetrical with a distinct head, which is
well developed with tentacles and eyes. The
foot is broad and expanded with a creeping
sole. Rhythmic contractions of the muscular
foot perform the locomotion. Some mucus-
secreting glands in the foot secrete mucus on
which the animal slides (Ruppert et al. 2004).
The body is asymmetrical due to the “rotation
of visceral mass in its axis” (torsion). There is a
lid at the opening of the shell called the oper-
culum. Respiration is by gills or pulmonary
sac. The mantle encloses the visceral mass.
The buccal cavity contains the radula with
odontophore, which works as a file to scrape
and tear food materials. The nervous system is
well developed with paired ganglia and their
connectives and commissures, with distinct
cerebral and pleural ganglia besides pedal and
visceral ganglia. Sexes are separate, and the
development is indirect through a veliger
larva or trochophore larva. The animal is very
active during the night or dim light. As the
slugs have no shell, the mucus layer gives
Mollusk
protection to the body, enabling the animal to
live in wet and moist areas. Slugs and land
snails cause severe damage to plants and agri-
cultural cultivations as they eat leaves and
stems (Jordan and Verma 2001; Louise 2006).
Examples include Pila (Fig. 7), Xancus,
Aplysia, Doris, and Dolabella.
2. Pelecypoda (Lamellibranchiate or Bivalvia):
Pelecypodes (Gr pelekus-hatchet+podos-foot)
are bilaterally symmetrical, laterally com-
pressed mollusks with a two-lobed mantle.
The body is enclosed within two valved shells,
hence the name bivalve. The head is rudimen-
tary without eyes and tentacles but with a pair of
labial palps. The foot is large plow or wedge-
shaped, adapted for burrowing. The respiratory
organs are plate-like gills called ctenidia. The
intestine is highly coiled and devoid of the
radula. Excretory organs consist of paired
nephridia. The nervous system includes three
pairs of nerve ganglia – cerebropleural, pedal,
and visceral (Ruppert et al. 2004). Sexes are
separate, gonads are single, and development
includes a parasitic glochidium larva
(freshwater mussel) or free-living veliger larva
(Pinctada). This group includes clams, oysters,
and scallops. They are the food sources for other
living beings, including humans. This group of
animals is well adapted for living in the water.
For protection, clams burrow deep into the sand
Mollusk, Fig. 7 Pila. https://pixabay.com/photos/snail-garden-snail-shell-1634852/
9
by contracting and relaxing their muscular foot.
Mussels and oysters attach themselves with a
strong thread or cement to solid surfaces, called
byssus threads. Examples include Lamellidens,
Milieus/Perna, Pinctada (Fig. 8), and Solen.
3. Polyplacophora (Loricata or Amphineura)
(Gr. Amphi-both+nervous-nerve): This group
is considered as most primitive of existing
mollusks. The members of this group include
chitons. They are exclusively marine, bilater-
ally symmetrical, and dorsoventrally flattened
animals. They have an articulated shell com-
posed of eight hinged and overlapping longi-
tudinal/transverse plates or valves (Budd and
Jensen 2000). A thick leathery mantle covers
the body. They have an indistinct head without
eyes and tentacles. They have a well-
developed radula mounted on an odontophore.
The foot is the broad, flat muscular, glandular
creeping sole and ventral in position. The man-
tle cavity is restricted to mantle grooves on
either side of the ventral side. Numerous
bipectinate gills occur on either side of the
body in the mantle groove. The nervous system
is primitive and ladder-like, which resembles
the Platyhelminthes (turbellarians). They have
a cosmopolitan distribution, from cold waters
through to the tropics (Jordan and Verma
2001). They live on hard surfaces, such as on
or under rocks or in rock crevices, inhabit
10
Mollusk, Fig. 8 Pinctada. https://en.wikipedia.org/wiki/Pinctada#/media/File:Pinctada_margaritifera_MHNT.CON.
2002.893.jpg
intertidal or subtidal zones, and are exposed to
the air and light for long periods. They feed on
algae and seaweeds that grow on rocks by their
radula. Examples include chitons (Fig. 9),
Leptochiton, and Chaetopleura.
4. Cephalopoda (Gr kephale-head+podos foot):
The word cephalopod means “head-footed.”
All are exclusively marine, most specialized,
and complex animals such as squid, octopuses,
and cuttlefish. The members possess a large
head, a crown of oral arms, a well-developed
nervous system with centralized brains, and
large eyes similar to the human eyes. They
are the only mollusks having an advanced cir-
culatory system, an accessory bronchial heart,
and closed blood vessels. Their foot is divided
into many tentacles with strong suction cups or
hooks, capturing the prey. They feed on crus-
taceans, worms, and other small mollusks
(Canti 2017). They have thick muscular, and
leathery mantle. There is an external or internal
shell, which can be absent in some species.
They have a cartilaginous endoskeleton for
supporting the body, attachment of muscles,
and protection of nerve centers. They possess
Mollusk
ink glands as a defensive adaptation. These
glands secrete melanin-containing dark color
ink to discolor the surrounding water and
escape enemies (Stoger et al. 2013). Examples
include Sepia, Loligo, octopus (Fig. 10), and
Nautilus.
5. Scaphopoda (Solenoconcha): Scaphopoda
(Gr. Shapha-boat+podos-foot) includes ele-
phant tusk shells. A group of aberrant marine
mollusks of very low grade of organization.
According to the fossil record, it is believed
that scaphopods from the youngest mollusks
and appeared from Mississippian onwards.
The members of this class are exclusively
marine. The shells’ size ranges from about 0.5
to 15 cm in length, and they like to live in soft
substrates offshore. Long cylindrical and bilat-
erally symmetrical body without eyes, sensory
tentacles, and gills (David and Winston 2001).
Tubular and tusk-like shell open at both ends.
Mantle edge fuses to form a mantle tube open
anterior and posterior. The foot is long, narrow
protrusible spade-like and is used for digging.
The mouth is surrounded by lobular out-
growths, cephalic filaments, or protrusible
Mollusk
Mollusk, Fig. 9 Chiton. https://www.flickr.com/photos/jsjgeology/15947947127
tentacular processes called captacula, which
serve as tactile organs to capture food particles.
As gills are absent, the respiration is by the
exchange of gases through the mantle surface.
The circulatory system is simple, and the heart
is rudimentary or absent. The nervous system
has a pair of cerebral and pleural ganglia. Sexes
are separate, and development includes a veli-
ger larva. According to molecular data, it is
suggested that the scaphopods are considered
Cephalopod sister group (Ruppert et al. 2004).
An example is Dentalium (Fig. 11).
6. Aplacophora: Aplacophora is the animal
group popularly known as Solenogasters or
mud moles. They are primitive degenerative
marine mollusks that inhabit marine habitat
with cosmopolitan distribution with long, nar-
row, bilaterally symmetrical, and vermiform
bodies (Scheltema 1993). There is the absence
of head, foot, mantle, shell, and nephridia
(Ruppert et al. 2004), but they have calcareous
spicules in the skin. The digestive tract is
straight with toothed radula in the mouth.
They have the presence of a pair of
coelomoducts, which serve as gonoducts. The
heart consists of a single auricle and a
11
ventricle. Hemocoel has two portions, namely,
dorsal hemocoel and pedal hemocoel separated
from each other by septum. The aplacophorans
are traditionally considered as an ancestral
mollusk (Buchsbaum et al. 2013). They are
found at all depths of the sea, usually absent
in the littoral zone. Some of them burrow in the
mud while some are live in association with
certain colonial coelenterates, and still, others
are predators. The group comprises the two
clades Solenogasters and Caudofoveata. The
aplacophorans are traditionally known as
ancestral mollusks (Budd and Jensen 2000;
Amelie 1993). Examples include Neomenia
(Fig. 12) and Chaetoderma.
7. Monoplacophora (Tryblidiida): A small
group of mollusks with primitive features.
Body bilaterally symmetrical flat and oval
body with a minute head foot, mantle, and
radula. The body is internally segmented. The
shell is univalved, conical or cup-shaped, and
symmetrical. There is the presence of segmen-
tally paired external gills, segmentally repeat-
ing nephridia, and gonads. They have a
primitive ladder-like nervous system. The
sexes are separate in almost all forms. At
12
Mollusk, Fig. 10 Octopus. https://www.flickr.com/photos/joachim_s_mueller/6775462987
Mollusk, Fig. 11 Dentalium. https://commons.wikimedia.org/wiki/File:Dentalium_octangulatum_01.JPG
present, a monoplacophoran is represented by
a single genus, namely, Neopilina. It is a very
primitive form and considered a living fossil
and thought to have become extinct over
380 million years ago. The shells of many
monoplacophorans are limpet-like in shape.
They were not recognized as such until 1952
(Ruppert et al. 2004). An example is Neopilina
(Fig. 13).
Some Extinct Groups of Mollusks
a) Rostroconchia: These are extinct mollusks
from the early Cambrian period to the late
Permian period. This group was mistaken as
bivalves because the larval stage is univalve,
but the adult typically has a pseudo-bivalve
shell that encloses the mantle and muscular
Mollusk
foot. This group consists of more than 1,000
species of members (Budd and Jensen 2000).
b) Helcionelloida: They are also an extinct group
of mollusks, consisting of the oldest
conchiferan mollusk with mineralized shells
(Budd and Jensen 2000).
The Fossil Records
Mollusks are considered some of the oldest meta-
zoans. The evolutionary history and origin of mol-
lusks from the ancestral group Lophotrochozoa
and their evolution into modern living form and
fossil form are still a debated subject. Some of the
fossil record obtained from the south region of
Australia and North Russia shows close resem-
blance with mollusks. There is good evidence
available for the appearance of gastropods and
Mollusk 13

Mollusk, Fig. 12 Neomenia. https://commons.wikimedia.org/wiki/File:Neomenia_yamamotoi.jpeg

Mollusk, Fig. 13
Neopilina. https://digit.
snm.ku.dk/www/inv/full/
MON-000001_shell-mo.
jpg

bivalves (Parker 2017). Most familiar members of ▶ Digestive System

living members of this phylum, such as the
Gastropoda, the Bivalvia Cephalopoda, the Poly-
placophora, the Scaphopoda, the Monoplacophora,
and the Aplacophora are believed to originate from
the Cambrian period (Budd and Jensen 2000).
Now itself, the debate occurs about Kimberella, a
mollusk-like organism from 555 million years ago,
and some Ediacaran and early Cambrian fossils
(Ruppert et al. 2004).
Cross-References
▶ Bilateral Symmetry
▶ Class
▶ Fossil Record
▶ Jean Baptiste Lamarck
▶ Nervous System of Invertebrates
▶ Phylum
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