Gale Austria, Janine Dizon, Justine Sia June 15, 2021

Bio 116 Lec 2

Part 1:

Slide 2

The general external features of mollusks

includes a ventral muscular foot, which is
used for locomotion and capturing prey
(Pechenik, 2015). They also have a mantle or
pallium, which is the dorsal secretory
epithelium of the shell. A mantle cavity is also
present, which is the space between the
mantle and foot where waste products are
emptied out. As for their shell, these are
almost always present, and are secreted by
the mantle to form a dorsal protective shield
for the underlying visceral mass or internal
organs. These shells also serve for defense
and protection.

Mollusks have a respiratory system that is

composed of ctenidia or gills, which project
from the anterior wall of the mantle cavity with
afferent and efferent blood vessels, muscles,
and nerves. It also consists of a long flattened
axis with lateral and alternating filaments on
either side. Moreover, the ctenidia divides the
mantle cavity into dorsal and ventral
chambers.

In its digestive system, it is composed of an

anterior mouth, a chitin-lined buccal cavity
(consisting of a radula and radula sac), a
tubular esophagus, a stomach, an intestine,
and an anus that is open midorsally at the
end of the mantle cavity.

Mollusks also have an open type of

circulatory system and a coelom. Lastly, they
have one pair of metanephridia as kidneys
(Pechenik, 2015).

Pechenik, Jan A. (2015). Biology of the

Invertebrates (7th ed). Boston:McGraw-Hill.
Slide 3

The circulatory system of mollusks consists of

a heart to move blood throughout the body,
while aquatic mollusks' respiratory system
revolves on their gills (Pechenik, 2015).
Ctenidia are the gills of mollusks, and they
are made up of a series of thin filaments of
tissue that resemble comb teeth. These
filaments take oxygen from the air and
transport it to the bloodstream. They also
take in carbon dioxide from the blood and
expel it into the environment. The lungs of
terrestrial mollusks are basic, absorbing
oxygen directly from the air they breathe. All
mollusks except those in class Cephalopoda
have an open circulatory system.

Mollusks' excretory system consists of

nephridia, tubular organs that filter waste
from internal body fluids. The waste produced
by the organism's cells is deposited into the
coelom, a fluid-filled interior body cavity.
Small tubes that enter into the coelom are
known as nephridia. They feature hair-like
cilia around the tube apertures that allow fluid
to pass from the coelom into the nephridia
tubules. Non-waste molecules like
carbohydrates and water can be reabsorbed
into the animal's body after the fluid enters
the nephridia. What's left in the tubes is
concentrated waste, which is eventually
expelled through the mantle cavity's nephridia
exit pores.

Pechenik, Jan A. (2015). Biology of the

Invertebrates (7th ed). Boston:McGraw-Hill.
Slide 4

The gills of molluscs function for both

respiration and feeding. The lateral cilia serve
in generating the water currents that will allow
gas exchange and the regulation of food
intake and waste removal (Carroll &
Catapane, 2007). By beating, the cilia on the
gills generate a flow of oxygenated water
through the mantle cavity, also carrying off
the carbon dioxide and nitrogenous wastes.
From each of the ctenidium, cilia beat in order
to facilitate the movement of water.

Carroll, M. & Catapane, E. (2007). The

nervous system control of lateral ciliary
activity of the gill of the bivalve mollusc,
Crassostrea virginica. Comp Biochem Physiol
A Mol Integr Physiol 148(2): 445-450.
http://doi.org/10.1016/j.cbpa.2007.06.003

Slide 5

The toothed chitinous ribbon in gastropods'

mouths is known scientifically as radula
(Academic Kids, n.d.). It is used to graze and
scrape diatoms and other small algae from
rock surfaces and other substrates with it.
Except for bivalves, which eat by siphoning
and filtering minute particles from water, all
mollusks have radulas.

The gastropod's mouth is placed underneath

the mollusk's anterior section. The radula sac
is contained in a pocket-like buccal cavity.

Moreover, radula apparatus consists of two

parts:

First is the odontophore, also known as the

cartilaginous base, with the odontophore
protractor muscle, the radula protractor
muscle and the radula retractor muscle. The
second is the radula itself, which consists of
longitudinal rows of chitinous and recurved
teeth that are referred to as the cuticula
(Academic Kids, n.d.).

The odontophore can move and protrude,

and the radula can move across the
odontophore. The radula teeth are erected as
a result of this movement. The odontophore's
tip scrapes the surface, while the teeth cut
and scoop up the food, transporting it to the
digestive tract through the oesophagus.

However, these acts wear down the front

teeth over time. In the radula sac, new teeth
are continually generated at the posterior end
of the buccal cavity. A gradual forward
movement of the ribbon brings them forward
to the tip, where they will be replaced as they
wear out. Teeth are produced quickly, and the
number of teeth depends on the species. The
mucus of the salivary gland, directly above
the radula, lubricates the radula's teeth. Food
particles become stuck in the sticky mucus,
smoothing the passage of food down the
oesophagus.

Academic Kids. (n.d.). Radula. Retrieved

from
https://academickids.com/encyclopedia/index.
php/Radula

Slide 6

For the molluscs, there are different types of

radula (Steneck & Watling, 1982). The first is
the rhipidoglossan radula, which is a large,
central, and symmetrical tooth. On the sides,
there are several lateral teeth, as well as
numerous closely packed flabellate marginals
that are called uncini.

As for the taenioglossan radula, there are

seven teeth in each row, with one middle
tooth that is placed in between one lateral
and two marginal teeth. These function like
rakes by scraping algae and gathering
detritus.

The docoglossan radula consists of a small

central tooth in each row, which has one to
three laterals and a few hooked marginals on
its sides. All the teeth are placed in a fixed
position. This type is said to be the most
primitive radular type, as the radula functions
like a chain of shovels. The rigid structure,
hardened by biomineralization, allows it to act
like a rasp in scraping hardened macroalgae.

Next, the stenoglossan or rachiglossan radula

has a central tooth at each row, with a lateral
tooth on each side.

Lastly, the toxoglassan radula has middle

teeth that are either very small or not present
at all. For each row, there are two teeth that
can only be at use one at a time. These teeth
are described to be grooved, very long, and
pointex. They also have neurotoxins or
venom channels, though the teeth are not
fixed firmly to the basal plate. Because of
this, they can be individually moved to the
proboscis and ejected into the prey.

The figure on the right shows a microscopic

image of a stenoglossan or rachiglossan
radula.

Steneck, R. & Watling, L. (1982). Feeding

capabilities and limitation of herbivorous
molluscs: A functional group approach.
Marine Biology, 68.
https://doi.org/10.1007/BF00409596

Slide 7

The Phylum Mollusca can be first divided into

two, the Aculifera and Conchifera (Pechenik,
2015). They are grouped based on the
presence of a conch shell, as the Aculifera do
not have one, while the Conchifera do.

The Aculifera contains two classes, which are

the Aplacophora and Polyplacophora. The
Aplacophora are known as the worm-like
solenogastres with poorly developed heads.
Two general forms of these exist: the
creepers and the burrowers. The
Polyplacophora are commonly called the
chitons, benthic marine mollusks with ovoid
bodies that are flattened dorsoventrally.

As for the Conchifera, there are five classes

found under it. Here, the mantle secretes one
or more calcareous shells but no spicules.
First, there are the Bivalvia, which have shells
composed of two valves. This includes
scallops, clams and muscles. The
Monoplacophora are called limpet-like living
fossils and have 31 described species. The
Gastropoda is a large class, composed of
snails, slugs, limpets and sea hares. The
Cephalopoda consist of the squids,
octopuses, nautilus, and ammonites. Lastly,
the Scaphopoda are known for their
tusk-shaped shells.

Pechenik, Jan A. (2015). Biology of the

Invertebrates (7th ed). Boston:McGraw-Hill.

Slide 8

This is a cladogram showing all the seven

major classes of the mollusks, as described
in the previous slide, with the characteristics
that distinguish them from one another in
lineage.

The smaller diagram shows the body forms of

sample specimens from each of the classes.
The one in the middle is the ancestral form,
and all seven are modified from it. All have
radula, muscular foot, mantle, and reduced
coelo,.

Slide 9

The molluscan shell can be divided into three

layers: the outer periostracum, middle
prismatic layer, and the inner nacreous layer
(Hechenberger, 2014). The periostracum is
not calcified, and incorporates conchiolinum,
which grows peripherally. It is secreted by a
band of cells found at the mollusc’s outer
edge. Moreover, it protects the chalk layer
from degradation and drilling organisms and
other predators.

Next, the middle prismatic layer is

calcareous, and is also known as the calcium
carbonate layer. It is built upon calcit,
prismatic chalk. The prisms are positioned
vertically to the surface, coming from a fluid
layer, and are located between the
periostracum and the mantle.
Lastly, the inner nacreous layer is made of
aragonite and known as the “mother of pearl.”
It is created by the epithelial cells of the
mantle tissue.

Hechenberger, C. (2014, June 26). Structure,

mechanics, and function of the mollusc shell.
Retrieved from
https://www.uibk.ac.at/zoology/marinbio/semi
narvortraege/seminar_calvi2014/mollusc-shel
l.pdf

Slide 10

To further go into the layers of the shell, the

periostracum has keratin-like conchiolin that
provide rigidity and shape to the cell
(Hechenberger, 2014). The prismatic layer
contains aragonitic or calcitic crystals, as well
as some proteoid material. Lastly, the
nacreous layer has stacks of aragonite that
alternate with sheets of conchiolin. Both the
nacreous and prismatic layers have their
respective regions of secretion. The
periostracum is also secreted from a groove
in the mantle, which is called the periostracal
groove.

In the inside of the shell, there is a pallial line

with different lobes. The lobes indicate how
large the siphon was, and the larger it is, the
deeper the bivalve dug into the bottom. There
are three types of pallial lobes: inner
(muscular), middle (sensory), and outer
(secretory).

Hechenberger, C. (2014, June 26). Structure,

mechanics, and function of the mollusc shell.
Retrieved from
https://www.uibk.ac.at/zoology/marinbio/semi
narvortraege/seminar_calvi2014/mollusc-shel
l.pdf
Slide 12

The radula is of the docoglossan type, and

there is a crawling foot, a primitive trait in
mollusks (Kazlev, 2002). The radula is
calcified with magnetite. The head is
underdeveloped and can only be seen from
the ventral side. The girdle is a belt of
muscular tissue that extends along the
animal's dorsal periphery. Small calcareous
spines, scales, or spicules, similar to those of
the very primitive and wormlike Caudofoveata
and Solenogastre Aplacophora, are typically
embedded in the girdle.

Kazlev, A. (2002, September 29).

Polyplacophora. Retrieved from
http://palaeos.com/metazoa/mollusca/polypla
cophora/polyplacophora.html

Slide 13

The image shows the internal anatomy of a

chiton, where the valves are overlying the
visceral mass. In between the plates or
valves, there are openings which have
photosensory organs called aesthetes.

There is a circulatory and nervous system

present, as well as a digestive system. The
haemocoel serves as the body cavity
containing the circulatory fluid. Atria are also
present. A nerve ring is used in the nervous
system, while nephridiopores are used in the
excretory system.

Slide 14

Those in Subclass Neomeniomorpha are also

called the Solenogastres (Pechenik, 2015).
All members have a thin foot that sits in a
small groove running down the ventral
surface from just beyond the mouth to the
posterior mantle chamber. Many animals lack
a radula, whereas the others have a radula
that is extremely simple. None of the animals
have real ctenidia. All the species are
hermaphroditic. Members of this subclass
may either crawl over the sediment or reside
among the Cnidarians.
Pechenik, Jan A. (2015). Biology of the
Invertebrates (7th ed). Boston:McGraw-Hill.

Slide 15

The figures show the sagittal section, with the

anterior on the left and the posterior on the
right. All the different components of the
internal anatomy of a Solenogastres have
been labeled. There is the cuticle,
esophagus, foot, gonad, mantle cavity,
midgut, oral cavity, radula, rectum, and
ventricle, to name a few.

Slide 17

The defining characteristics of the members

of the Class Monoplacophora are its three to
six pairs of ctenidia, six to seven pairs of
nephridia, and multiple pairs of foot or pedal
retractor muscles (Pechenik, 2015). They are
known for their oval body, and produce a
single, unhinged, cap-shaped shell. The
shells of adults are flat rather than spirally
wound, though for larvae they are spiral.
Aside from the gills, there are a number of
foot retractor muscles, heart auricles and
ventricles, gonads, and nephridia. They also
have a radula present and a linear gut. The
nervous system includes lateral and pedal
nerve cords.

Pechenik, Jan A. (2015). Biology of the

Invertebrates (7th ed). Boston:McGraw-Hill.
Slide 18

The figure shows the internal anatomy of the

Monoplacophora. The parts are labeled as
follows:

Ve - velum
Ao - aorta
Mr - retractor muscle
Ct - ctenidium
Gd - gonoduct or genital duct
Au - auricle of the heart
A - anus
Ne - nephridium
V - ventricle
Np - nephiridopore
G - gonad
Nv - nerve cord
B - buccal cavity

Slide 19

There are 300 to 400 species in the Class

Scaphopoda, which all live marine and
sedentary lives in sand and mud substrates
or in deep water (Pechenik, 2015). Their
defining characteristics include having a
tusk-shaped conical shell that is open at both
ends, and the development of anterior,
threadlike, and adhesive feeding tentacles.
They possess the usual molluscan
characteristics, such as a foot, mantle tissue,
mantle cavity, radula, and shell. As
mentioned, the shell is shaped like a tusk,
and is thus not spirally wound, instead
growing linearly like a hollow curved tube.
Water enters through the nower end, and is
expelled through the same opening.

Moreover, their head is projected from the

anterior side, and is reduced so that it is just
a conical proboscis that bears the mouth.
There are two lobes on either side of the
head which bear a large number of
thread-like tentacles that are called
captacula. The foot is also conical in shape,
extending from the anterior as an elongated
structure. Its top can be expanded as an
anchor, and there are no ctenidia. As for its
reproduction, the gonads are unpaired and
shed gametes through the right nephridium
where external fertilization occurs. The larvae
are often called veliger larvae.

Pechenik, Jan A. (2015). Biology of the

Invertebrates (7th ed). Boston:McGraw-Hill.

Slide 20

Veligers are planktonic larvae of a variety of

bivalve and gastropod mollusks with a shell,
foot, and velum (Grove, 2011). The
prototroch, a pre-oral ciliated ring in the
trochophore larva, is where the velum is
derived from. The veliger's shell is secreted
by a dorsal shell gland. A veliger's shell is
bivalved, whereas a gastropod veliger's shell
resembles a spiraling snail shell.

For spiral shells, the bi-lobed ciliated velum

extends from the shell. The metatroch is a
supplementary (post-oral) ciliated band seen
underneath the velum's primary ciliated band.
The cilia are used for locomotion and feeding.

Some larva have a bivalved shell. One valve

is in focus while the other valve is located on
the opposite side of the larva. The velum of
this larva is pulled into the shell, however its
cilia can be just barely seen protruding
beyond the edge of the shell.

Grove, M. (2011, May 28). Veliger larvae of

molluscs. Retrieved from
http://invert-embryo.blogspot.com/2011/05/vel
iger-larvae-of-molluscs.html
Slide 21

This is a typical adult. The tusk shells are

burrowing animals, burrowed into the
substrate with an opening at the bottom
where the mouth is. The foot can also be
found at the end. On the opposite side is the
shell opening where water exits after passing
through the body of the animal. The
captacula can also be seen, which surrounds
the mouth, and it also has tentacles for filter
feeding or prey capture.

Slide 22

The Class Gastropoda is composed of the

snails and slugs. It is the largest molluscan
class and contains over at least 60,000
species, which is more than half of all the
molluscs (Pechenik, 2015). These organisms
are distributed among marine, freshwater,
and terrestrial environments, occupying
diverse habitats such as rivers, lakes, trees,
and the deep sea.

They also have a distinct head that can be

retracted into the shell when in danger
(University of Texas, n.d.). Their heads may
also have scraping radula, and some species
are noted to have eyes.

A typical snail consists of a visceral mass (all

the internal organs) that sits on top of a
muscular foot. This mass is usually protected
by a univalved coiled shell. It also contains
well-developed ganglionic bodies in its
nervous system.

In its excretory system, they have one pair of

metanephridia (kidneys), where one end is
connected to the nephrostome, while the
other leads to the nephridiopore located at
the posterior of the mantle cavity. In some
species, the right nephridium is absent,
leaving a single excretory organ.

In their reproductive system, they have a

single gonad. Most snails are also oviparous,
releasing the egg immediately after formation
(Tompa, 2013).

Pechenik, Jan A. (2015). Biology of the

Invertebrates (7th ed). Boston:McGraw-Hill.

Tompa, A. (2013). Studies on the

reproductive biology of gastropods: Part 1.
The systematic distribution of egg retention in
the subclass Pulmonata (Gastropoda).
Journal of the Malacological Society of
Australia.
https://doi.org/10.1080/00852988.1979.10673
920

University of Texas. (n.d.). Gastropods.

Retrieved from
https://www.utsa.edu/fieldscience/gastropod_i
nfo.htm

Slide 23

The nervous system of molluscs can be

found at the ventral side of the animal
(Nordsieck & Eleveld, n.d.). Most ganglia are
found in pairs as part of the rope ladder
nervous system. Commissurae, or lateral
connections, join ganglia of the same kind,
while gangli of different types are connected
through longitudinal connections called
connectives. The cerebral ganglia of the
brain, the pedal ganglia of the foot, the
pleural and parietal ganglia of the pallial
cavity, and the visceral ganglia of the inner
organs are the most significant ganglion
types. The buccal ganglia of the pharynx are
located in front of the cerebral ganglia.

The cerebral, pedal, and pleural ganglia are

concentrated and organized together in a
buccal ring enclosing the oesophagus near
the head in most modern gastropod groups
(Nordsieck & Eleveld, n.d.). Above the
oesophagus are the cerebral and pleural
ganglia, and underneath it are the pedal
ganglia. The visceral and parietal ganglia of
marine gastropods remain still in their original
positions in the body, with lengthy
connectives connecting them to the buccal
ring. Because there are four primary neural
strands in the mollusk nervous system, it is
called a tetraneural nervous system. On the
ventral side, two pairs of connectives connect
the cerebral ganglia to the pedal ganglia,
another to the visceral and parietal ganglia on
the dorsal side, traveling through the pleural
ganglia.

These allow the molluscs to have a quicker

response to things, as a result of the
cephalization with clusters of ganglia that
create a 6-lobed brain.

Nordieck, R. & Eleveld, M. (n.d.). The

nervous system of snails. Retrieved from
http://www.molluscs.at/gastropoda/index.html
?/gastropoda/morphology/nervous_system.ht
ml

Slide 24

In a shell, there is an apex at the tip, which is

the extremity of a gastropod shell opposite to
the anterior region (Leal, n.d.). The spire is a
series of successive whorls in a gastropod
shell. Between two adjacent whorls, there is a
line or region of junction called the suture. At
the center of the shell, the columella acts as a
column or pillar. The opening is called the
aperture, with the edges called the lip. On the
opposite end of the axis is the coiling axis.
There is also a varix, which is an axial
sculptural element more prominent than a
costa, but is more widely spaced. Its growth
stops when a thickened lip is developed.
Lastly, the siphonal canal is a projection of
the anterior part of the shell, shaped like a
tube, and it is used to protect the anterior
siphon.

The surface of the shell may be varied. These

textural ornamentations of the surface are
called sculptures, and they may be either
raised or incised (Royal Museum, n.d.). One
type are malleations, which are shallow,
irregular depressions. There are also striae,
which are fine lines or ridges that may be
either regular in size and spacing or irregular
and wrinkle-like. These are also usually
incised but sometimes raised. Lamellar ribs
are rib-like elements with narrow, bladelike
edges. Lastly, ribs are much larger, raised,
angular or rounded ridges that can either be
regular or irregular in size.

Leal, J. (n.d.). Gastropods. Retrieved from

http://www.fao.org/3/y4160e/y4160e08.pdf

Royal Museum. (n.d.). Identification of

species. Retrieved from
https://royalbcmuseum.bc.ca/exhibits/living-la
ndscapes/cbasin/molluscs/introduction4.html

Slide 25

The shape of protoconches may come in

different types (Terrestrial Mollusc Tool, n.d.).
First, the conical shell shape has an
elongated spire that tapers to a point. It is
slightly broader at the base. The mamimilate
is a pointed extension of the spire. The
obtuse conical is short and slightly larger. As
the name suggests, the dome-shaped shell
takes the same shape as a dome. Lastly, the
deviated pausicpiral has an axis that is
slightly deviated from the main axis. These
types are used as a basis for taxonomic
classification.

Moreover, there are two sculptures present.

The axial sculpture runs parallel to the axis of
coiling, which is an imaginary line that runs
through the apex of the shell where the
whorls are coiled. The spiral sculpture, on the
other hand, follows the spiral growth of the
shell.

Terrestrial Mollusc Tool. (n.d.). Glossary of

terms. Retrieved from
https://idtools.org/id/mollusc/glossary.php
Slide 26

Here are examples of gastropods which

develop from a trochophore larva. The middle
image in the first show shows a veliger, with a
ciliated velum on top. The rightmost image in
the first row and the images on the second
row are juveniles. The right image on the
second row is the specimen Haliotis,
commonly known as abalone, and can be
identified by the several openings on its shell.

Slide 27

The general form of coiled shells can be

described using different terms (Royal
Museum, n.d.). First, heliciform resembles the
genus Helix, as the width and height are
approximately equal and the whole shell is
round in shape. The depressed-heliciform is
similar to the heliciform, but the shell is short
and wide. The conic-heliciform is also like the
heliciform but the spire is more cone-shaped.
The cylindric shape has all sides nearly
parallel and ends bluntly. From this, the
subcylindric form is close in appearance to
the cylindrical form, though the sides are not
as parallel. The elongate-tapering form has
the sides of the spire distinctly tapered to a
point and a pointed apex. The subovate is
shaped like an egg, with the sides of the spire
like a convex. The conic-ovate is like an
ovate, and is apically conic. Lastly, the
elongate-ovate is elliptical.

As shown in the figure on the right, the

columellar muscle passes below the mantle,
thus thickening the body wall, and through
the foot. At the end, it is attached to the inner
face of the operculum.

Royal Museum. (n.d.). Identification of

species. Retrieved from
https://royalbcmuseum.bc.ca/exhibits/living-la
ndscapes/cbasin/molluscs/introduction4.html
Slide 28

During larval development, visceral organs

rotate in an anticlockwise orientation across
an angle of 180° on the remainder of the
body (Encyclopedia.com,2016). The
phenomena occurs in free-swimming (veliger)
snail larvae, transforming the symmetrical
larva into an asymmetrical adult.
Rotation may be caused by the contraction of
the larval retractor muscles and unequal
development.

Asymmetry is seen in Veliger larvae at an

early stage when the mesodermal bands
develop asymmetrically. On the right side, the
mesodermal band is larger than on the left.
Five mesoderm cells comprise the right band,
which extend to form muscle cells. The
visceral hump is shifted to the left side due to
muscle cell transformation.

The mantle chamber was initially located

behind the head, but following torsion, it now
opens just behind the skull, and its related
parts have migrated forward. The anus and
ctenidia are oriented backwards before
torsion, and the auricles are located behind
the ventricle. The anus and ctenidia come
forward after torsion, and the auricles lay in
front of the ventricle. The alimentary canal,
which was formerly straight, is twisted into a
loop, and the mouth and anus are brought
closer together. The pleuro-visceral
connectives must have crossed because the
pallial complex must have shifted from the
posterior to the anterior half of the body and
became twisted in the form of 8.

The right connective, which has a parietal

ganglion, goes above the gut and the left
connective, which runs below the intestine, is
known as the supraintestinal. After torsion,
the coil of the visceral sac that was largely
dorsal or exogastric becomes ventral or
endogastric. The coiling of the shell is
unrelated to torsion and was a separate
evolutionary process, with the shell remaining
symmetrical. It's caused by anus moving to
the right side of the mantle cavity and the loss
or reduction of paired sections of the
primitively left or topographi­cally right side.

Encyclopedia.com . (2016, May 27). Torsion

and Detorsion in Gastropoda. Biology
Discussion.
https://www.biologydiscussion.com/invertebra
te-zoology/phylum-mollusca/torsion-and-detor
sion-in-gastropoda/33044.

Slide 29

The mantle chamber was initially located

behind the head, but following torsion, it now
opens just behind the skull, and its related
parts have migrated forward. The anus and
ctenidia are oriented backwards before
torsion, and the auricles are located behind
the ventricle. The anus and ctenidia come
forward after torsion, and the auricles lay in
front of the ventricle. The alimentary canal,
which was formerly straight, is twisted into a
loop, and the mouth and anus are brought
closer together. The pleuro-visceral
connectives must have crossed because the
pallial complex must have shifted from the
posterior to the anterior half of the body and
became twisted in the form of 8.

The right connective, which has a parietal

ganglion, goes above the gut and the left
connective, which runs below the intestine, is
known as the supraintestinal. After torsion,
the coil of the visceral sac that was largely
dorsal or exogastric becomes ventral or
endogastric. The coiling of the shell is
unrelated to torsion and was a separate
evolutionary process, with the shell remaining
symmetrical. It's caused by anus moving to
the right side of the mantle cavity and the loss
or reduction of paired sections of the
primitively left or topographi­cally right side.

Jain, K. (2016, May 27). Torsion and

Detorsion in Gastropoda. Biology Discussion.
https://www.biologydiscussion.com/invertebra
te-zoology/phylum-mollusca/torsion-and-detor
sion-in-gastropoda/33044.
Slide 30

There are three subclasses under

Gastropoda. The Prosobranchia subclass
contains the majority of gastropods; the
Pulmonata subclass contains land snails; and
the Opisthobranchia subclass contains sea
hares and sea slugs (Infoplease,n.d.).
Animals having reduced or no shells belong
to the latter subclass. Although the majority of
gastropods are mobile, others, such as the
slipper shell (Crepidula), are sedentary. Sea
butter, for example, is one of them.

Infoplease. (n.d.). Mollusca: Class

Gastropoda. Infoplease.
https://www.infoplease.com/encyclopedia/eco
logy/animals/invertebrates/mollusca/class-ga
stropoda.

Slide 31

For the exhalant mantle cavity currents, these

primitive archeogastropods feature a linear
series of perforations (Burch,2001). Excurrent
water exiting the mantle cavity is aided by
slits in the mantle and shell. Because of the
mantle cavity with the anus, nephridiopores,
and gonopores placed over the head, this is a
primitive solution to the sanitation problems
produced by torsion.

Eogastropoda was a historical taxonomic

classification for snails and gastropods
(Eogastropoda,n.d.). It was one of the two
major divisions (subclasses) of the snail class
Gastropoda. The Ortho Gastropoda was the
other gastropod subclass. Eogastropoda is a
significantly older gastropod family. All real
limpets belonged to this category.

Burch, J. B. (2001). Archaeogastropoda.

Animal Diversity Web.
https://animaldiversity.org/accounts/Archaeog
astropoda/.

Eogastropoda. The Free Encyclopedia. (n.d.).

https://en.jejakjabar.com/wiki/Eogastropoda.
Slide 32

Orthogastropoda was a prominent taxonomic

grouping of snails and slugs, a massive
subclass within the Gastropoda class
(Orthogastropoda,n.d.). The Orthogastropoda
looked to form a clade that was backed up by
clear synapomorphies. The clade's defining
characteristics were synapomorphies i.e a set
of characteristics are present in its members
but not in the other forms from which it
diverged.

Eyes with a vitreous body on eyestalks and

paired jaws free of the buccal bulk are some
of the characteristics. A solitary kidney is
found on the right side of the pericardium, as
well as a flexoglossate radula with a flexible
radular membrane. The radula is a rasping
instrument made from the snail's tongue.

Orthogastropoda. DBPedia. (n.d.).

https://dbpedia.org/page/Orthogastropoda.

Slide 33

Superorder From the intertidal zone to the

deep sea, Vetigastropoda is a huge and
diverse group of marine gastropods
(Encyclopedia.com,2021). Most organ
systems have some bilateral asymmetry, and
many have shells with slits or other
secondary apertures.

Although certain deep-sea members of the

group are hermaphrodites, most
vetigastropods are dioecious. Typically,
vetigastropods release their gametes directly
into the sea for fertilization and development.
In response to other predatory snails as well
as sea stars, certain vetigastropods have
been shown to make escape responses.
Swaying and tilting the shell to avoid the tube
feet of the sea star, as well as short spurts of
rapid movement away from potential
predators after tissue contact, are examples
of escape responses.
Encyclopedia.com. (2021, June 23).
Vetigastropoda (Slit And Top Shells).
Encyclopedia.com.
https://www.encyclopedia.com/environment/e
ncyclopedias-almanacs-transcripts-and-maps
/vetigastropoda-slit-and-top-shells.

Slide 34

Vetigastropod shells come in a variety of

shapes and sizes, ranging from squat and
globe-shaped to elongate turreted
constructions (Encyclopedia.com,2021).
Limpet morphology in the vetigastropod has
changed at least six times. Simple concentric
growth lines, which may be barely visible on
the shell surface, to substantial radial and
axial ribbing, as well as everything in
between, are all examples of shell sculpture.
The shell aperture, also known as the
opening, is usually elliptical and runs parallel
to the coiling axis. After the head and foot
have been brought back into the shell, most
species have an operculum that covers the
opening. The animals feature a unique snout
with a mouth and a single pair of cephalic
tentacles. Sensory epipodial tentacles are
usually seen on the animal's lateral surfaces.

Encyclopedia.com. (2021, June 23).

Vetigastropoda (Slit And Top Shells).
Encyclopedia.com.
https://www.encyclopedia.com/environment/e
ncyclopedias-almanacs-transcripts-and-maps
/vetigastropoda-slit-and-top-shells.
Slide 35

As seen in the images presented in the slide,

the internal organs are located under the
shell and around the foot (Facts About
Abalone,n.d.). The crescent-shaped gonad,
the most visible organ, is gray or green in
females and cream in men. It wraps around
the abalone's back and around the side
opposite the pores. The abalone features two
eyes, a mouth, and a pair of elongated
tentacles. The radula is a long, file-like
tongue that sits inside the mouth.

The gill chamber is located between the

mouth and the respiratory pores. Water is
sucked in under the shell's edge, then passes
through the gills and out the pores. The
abalone is classified as a primitive animal
since it lacks a visible brain component. It
does, however, have a heart on its left side,
and blood is circulated through the arteries,
sinuses, and veins with the help of the
surrounding tissues and muscles.

Facts About Abalone. (n.d.). Facts About

Abalone. FISHTECH .
http://www.fishtech.com/facts.html.

Slide 36

The Neritopsina are the first gastropod clade

to go through the massive evolutionary
radiations seen throughout the Gastropoda
(Encyclopedia.com,2021). Because
Neritopsina's shells are coiled differently than
those of other coiled gastropods, they lack a
central shell axis, the columella, and most
species absorb the shell's internal partitions
as they grow.

Neritopsina shell morphology encompasses

the majority of Gastropoda forms, from coiled
to limpet, and even shellless slugs. The
juvenile shell, or protoconch, of the
Neritopsina is unusual, and most species
absorb the internal partitions of the shell as
they grow, allowing the snail's body to stay
limpet-like rather than coiled.
Neritopsis radula does not absorb the internal
partitions of its shell and is regarded to be the
most basic taxon based on this and other
anatomical characteristics. The Helicinidae
and Hydrocenidae terrestrial groups have the
most common coiled morphologies, which are
similar to those seen in coiled vetigastropod
groups like the Turbinidae.

Encyclopedia.com. (2021, June 24).

Neritopsina (Nerites And Relatives).
Encyclopedia.com.
https://www.encyclopedia.com/environment/e
ncyclopedias-almanacs-transcripts-and-maps
/neritopsina-nerites-and-relatives.

Slide 37

Mesogastropoda is a subclass of
Mesogastropoda. Individuals in the
Prosobranchia order of prosobranch
gastropods have one pectinibranch gill and
one kidney (Encyclopedia.com,2021). The
shell is helical in shape, with an entire
opening. The radula has a variety of
morphologies, and the sexes are distinct.

Previously classified as "lower

mesogastropods," members of the
Architaenoglossa are now recognized as the
Sorbeoconcha's sibling taxa (which also
includes the Neogastropoda) (Sadler,2021).
The Architaenoglossa and Sorbeoconcha are
thus classified as belonging to the
Caenogastropoda. The freshwater
Am-pullarioidea and the terrestrial
Cyclophoroidea are both found in the
Architaenoglossa.

Encyclopedia.com. (2021, June 24).

Mesogastropoda. Encyclopedia.com.
https://www.encyclopedia.com/science/diction
aries-thesauruses-pictures-and-press-release
s/mesogastropoda.

Sadler, P. (2021, June 22). Evolution and

systematics - Reproductive Biology. GUWS
Medical.
https://www.guwsmedical.info/reproductive-bi
ology-2/evolution-and-systematics-erl.html.

Slide 38

Caenogastropod shells are usually coiled,

and they come in a variety of shapes and
sizes, from flat squat shells to globose shells
and long, narrow, densely coiled ones
(Encyclopedia.com,2021). The massive,
ciliated osphradium, as well as the ctenidium,
propel water flow through the pallial cavity of
the Sorbeoconcha. In several taxa, the
creation of an anterior notch, or siphon, in the
shell and mantle is also observed.

Encyclopedia.com. (2021, June 24).

Caenogastropoda (Caenogastropods).
Encyclopedia.com.
https://www.encyclopedia.com/environment/e
ncyclopedias-almanacs-transcripts-and-maps
/caenogastropoda-caenogastropods.

Slide 39

Caenogastropods have figured more

significantly in human culture than most other
gastropod groupings due to their remarkable
diversity in form and color
(Encyclopedia.com,2021). The Phoenicians
were known throughout the ancient
Mediterranean world for their royal purple
dye, which was created from organs found in
Muricidae family members. Circa China,
cowry shells were first utilized as currency in
1200 B.C.

Because of their vibrant colors and forms,

caenogastropoda shells are frequently used
to embellish still-life paintings. Cone shells
are harmful to humans in about 20 species,
with three species causing fatalities as a
result of stings. Cone shell venom, on the
other hand, is being exploited to develop
painkillers. Finally, shell collecting has been
popular since the early 1800s, with certain
caenogastropoda groups including cone
shells, cowries, and Murex among the most
valuable and sought-after species.

Encyclopedia.com. (2021, June 24).

Caenogastropoda (Caenogastropods).
Encyclopedia.com.
https://www.encyclopedia.com/environment/e
ncyclopedias-almanacs-transcripts-and-maps
/caenogastropoda-caenogastropods.

Slide 40

Cone snails (Conidae), tulip shells

(Fasciolariidae), and whelks (Buccinidae) are
all members of the Neogastropoda family
(Modica & Holford,2010). Neogastropods are
often prominent members of the benthic
community at the top of the food chain due to
their incredible predatory specializations.

Neogastropods have created various prey

hunting inventions using specific ingredients
pharmaceutical corporations could only
dream about, ranging from digging holes to
darting harpoons (Burch, n.d.). The lengthy
incurrent siphon and accompanying siphonal
structure on the base of the shell
distinguishes neogastropods. Almost of
neogastropods live in the sea.

Modica M.V., Holford M. (2010). The

Neogastropoda: Evolutionary Innovations of
Predatory Marine Snails with Remarkable
Pharmacological Potential. In: Pontarotti P.
(eds) Evolutionary Biology – Concepts,
Molecular and Morphological Evolution.
Springer, Berlin, Heidelberg.
https://doi.org/10.1007/978-3-642-12340-5_1
5

Burch, J. B. (n.d.). Neogastropoda. Animal

Diversity Web.
https://animaldiversity.org/accounts/Neogastr
opoda/.
Slide 41

The visceral hump, mantle, head, and foot

are the four primary sections of a gastropod's
body (Encyclopædia Britannica, inc, n.d.). A
single columellar muscle or a group of
muscles attaches the body to the shell. Snails
can retract their heads and feet into their
shells, but many species have shells that are
too small to hold the body; slugs, on the other
hand, have either an internal shell remnant or
no shell at all.

The digestive, reproductive, excretory, and

respiratory systems are all housed in the
visceral hump, or visceral mass, of
gastropods, which is always contained within
the shell. The mantle, or pallial, hollow makes
up a large portion of the visceral hump. The
mantle is the fleshy covering that covers the
shell's outer wall and covers the mantle
chamber. The mouth is in the middle of the
ventral border, and the head is bilaterally
symmetrical, with one or two pairs of
tentacles, typically with supplementary palps.
The mouth is frequently extended into a
proboscis that reaches well forward of the
tentacles. A propodium and a metapodium
are frequently divided anteriorly and
posteriorly, with the former being capable of
reflexing over the shell.

Encyclopædia Britannica, inc. (n.d.).

Gastropod-The foot. Encyclopædia
Britannica.
https://www.britannica.com/animal/gastropod/
The-foot.
Slide 42

Heterobranch mollusks (molluscs) are one of

the most successful and diverse radiations of
the class Gastropoda, and even of the
phylum Mollusca, in terms of species counts
and ecological diversity (Malaquias, 2020).
They have evolved adaptations to all possible
conditions in the sea and on land

About 25 families and 2000 species of bubble

shells and seaslugs, as well as the and
Aplysiidae, make up the opisthobranchs
(Heterobranchia,n.d). Almost all
opisthobranchs are marine, with the majority
of shell-bearing taxa having no operculum as
adults and only a few of the "primitive"
shell-bearing taxa possessing an operculum.

Malaquias, M. A. E. (2020). Heterobranchia.

AccessScience.
https://doi.org/10.1036/1097-8542.315650

Heterobranchia. Mollusca: Gastropoda:

Heterobranchia. (n.d.).
https://ucmp.berkeley.edu/mollusca/mollusca/
gastropoda/heterobranchia/heterobranchia.ht
ml.

Slide 43

The cephalaspids are the opisthobranch

orders' most basic members
(Cephalaspidea,n.d.). They have some
similarities to the Prosobranchia, but they are
classified with the Opisthobranchia since the
transition from the Cephalaspidea to the other
opisthobranch groups is less abrupt. A huge
head shield distinguishes cephalaspids,
preventing sand and debris from entering the
mantle chamber as the animals plow through
the ground. Two "eyes," pigmented
light-sensitive organs, are located in the
center of the skull shield. These and other
opisthobranchs have translucent, colorless
shells.

The color and design are on the mantle and

are visible through the clear, transparent
shell. When an animal dies, its shells
frequently wash up on beaches, where they
are difficult to distinguish from the bubbles
created by waves splashing on the shore,
hence the term "bubble shells."

Cephalaspidea. Animal Diversity Web. (n.d.).

https://animaldiversity.org/accounts/Cephalas
pidea/.

Slide 44

The classic spiralled gastropod shell is

absent in cephalaspids; most species have a
bubble-shaped shell with a short or involute
spire (because only one whorl is totally
visible) (Malaquias,2014). Some groups don't
have a shell at all, or have a shell with a
smaller inside shell. Although the shell might
be heavily calcified, it is usually thin, delicate,
and translucent.

Although the majority of species are less than

1 cm in length, certain genera include larger
species on average. These animals' bodies
range in color from pale to brownish, although
some families, especially those found in
tropical areas, include species with brilliant
colors.

Malaquias, M. A. E. (2014, June 30). Bubble

shells& Cephalaspidea. Artsdatabanken -
Kunnskapsbank for naturmangfold.
https://www.biodiversity.no/Pages/149434/Bu
bble_shells__small_Cephalaspidea__small_.

Slide 45

Parapodia of Anaspidea have a tendency to

grow and, along with the mantle, to surround
the fragile shell (with increasingly reduced
and involute spire) (Cephalaspidea,2000)..
The mantle cavity is also shrinking and being
pushed to the right, while the visceropallium
(visceral mass and mantle) is being
dominated by the head-foot.

The Aplysidae have tentacles (rhinophores)

that resemble rabbit ears, earning them the
nickname "sea hares." (Anaspidea,2000). A
neck-like area commonly exists behind the
head, which combines into a fat trunk. Warts
and other protrusion commonly cover the
entire body.

Cephalaspidea. Animal Diversity Web.

(2000).
https://animaldiversity.org/accounts/Cephalas
pidea/.

Anaspidea. Animal Diversity Web. (2000).

https://animaldiversity.org/accounts/Anaspide
a/.

Slide 46

The Aplysioidea family includes small species

that grow to less than 2 cm in length and
giant species that grow to over 70 cm in
length (Rudman,2004). They normally have a
pair of enrolled rhinophores on their heads,
as well as enormous flattened, enrolled oral
tentacles on each side of the mouth. When
present, the shell is reduced to a weakly
calcified interior plate that protects the gills
and heart.

A pair of enormous parapodial lobes surround

this severely diminished mantle cavity, which
in some taxa are partially merged to form a
secondary parapodial chamber. The
presence of an opaline gland on the floor and
a purple gland on the roof in the mantle cavity
are two characteristics shared by both the
Akeroidea and the Aplysioidea.

The oesophageal gizzard, which has an

anterior chamber with huge chitinous plates
and a posterior chamber with numerous fine
spines, is a third common feature.
Anaspideans are all 'herbivores,' meaning
they eat algae or cyanobacteria.

Rudman, W. B. (2004, December 22).

Anaspidea. The Sea Slug Forum.
http://www.seaslugforum.net/find/anaspidea.
Slide 47

Nudibranchs are a group of marine

gastropods that includes over 3,000 species
(Laidlaw,2020). They are slug-like soft-bodied
animals with a variety of vivid colors and
elaborate body configurations. Nudibranchs
breathe through a 'bare gill' that is fashioned
into branchial plumes and is external to their
body, similar to the alveoli of a human lung.

The sexual apertures are on the right side of

the body in all nudibranchs that have both
female and male sex organs. They do not
have the ability to self-fertilize, therefore they
still need a mate to reproduce. Two
nudibranchs will cross-fertilize after a
dance-like courtship, with one laying eggs
fertilized by the other.

Because of their slug-like appearance,

nudibranchs are commonly referred to as sea
slugs. They were originally classified as part
of the Opisthobranchia family, which
comprised side-gill slugs, bubble snails, sea
slugs, sea hares, and others.

Laidlaw, S. (2020, October 5). Nudibranch -

Facts and Beyond. Biology Dictionary.
https://biologydictionary.net/nudibranch/.

Slide 48

This phyllidiid on the slide is elongate-oval in

shape, with three longitudinal ridges made up
of fused or discontinuous simple tubercles
(Pittman & Fience,n.d.). The tubercles are
topped with dark yellow and ringed with
blue-gray basally on a black background.
Around the mantle margin, crescent-shaped
blue-gray patches with black dots and
yellow-capped tubercles are common. The
rhinophores are yellow in color and the foot is
a uniform gray color.

Hypselodoris is a genus of soft-bodied,

bilaterally symmetrical slugs without shells
(Jones,2019). Their bright colors and patterns
are well-known. The east Pacific and Atlantic
clades are predominantly blue, with dots or
stripes of a variety of hues excluding red. The
Indo-Pacific clade has a wider spectrum of
colour, ranging from pink/purple to
orange/yellow.

Pittman , C., & Fience, P. (n.d.). Phyllidia sp.

#1: main page.
http://seaslugsofhawaii.com/species/Phyllidia-
sp1-a.html.

Jones, M. (2019). Hypselodoris. Animal

Diversity Web.
https://animaldiversity.org/accounts/Hypselod
oris/.

Slide 49

Start with the shape of the body, or external

anatomy, to identify nudibranchs (Wolters,
n.d.). Rhinophores are horn-like tentacles on
their heads that are used to detect odor
(smell) and can be retracted as a predator
approaches. The branchial plume on his back
has gills that filter oxygen from the water
(breathe). Dorid nudibranchs have this
characteristic.

More tentacles are found in some species,

with some pointing forward (oral tentacles)
and others looking to the side (propodial
tentacles). Cerata are creatures with a lot of
tentacles or finger-like structures on their
backs. This is a common feature of aeolid
nudibranchs. The overarching cover is the
mantle, which can be skirt-like and blow
upwards.

The placement of the colors and patterns is

crucial when it comes to coloring. Consider
horizontal or vertical stripes, lines
surrounding rhinophores, and so on. It's
usually a good idea to double-check and
confirm the nudibranch name you find with
someone who knows what they're talking
about.

Wolters, M. (n.d.). 100 Stunning Nudibranchs

of Anilao - Nudibranch Capital of The World.
Dive O'Clock!
https://www.diveoclock.com/destinations/Asia
/Philippines/Anilao_nudibranchs/.

Slide 50

Pulmonates are land, freshwater, and marine

snails that have lost their gills and now
breathe through a "lung," which is a highly
vascularized saclike modification of the
mantle cavity (Encyclopædia Britannica,
inc,n.d.). Although some snails have no
external shell, most pulmonates have a spiral
shell that can be flattened or attenuated.
They have more advanced anatomical
features than other snails.

As adults, all are detorted with a bilaterally

symmetrical and unlooped nerve cord, and
the heart's auricle is anterior to the ventricle.
All of them are hermaphroditic, meaning they
have both male and female reproductive
organs in one body. Pulmonates are also
employed in the delicacy escargot.

Encyclopædia Britannica, inc. (n.d.).

Pulmonate. Encyclopædia Britannica.
https://www.britannica.com/animal/pulmonate
.

Slide 51

Eupulmonata is a clade of air-breathing snails

classified as a taxonomic group (Superorder
Eupulmonata,2020). The bulk of the snails
and slugs in this category are land snails and
slugs, however some are marine and
saltmarsh snails that can withstand saline
environments.

Superorder Eupulmonata. iNaturalist.ca.

(2020, April).
https://inaturalist.ca/taxa/775812-Eupulmonat
a.
Slide 52

A eupulmonata reproductive system is

composed of ovotestis, hermaphroditic duct,
albumen gland, penis and dart shaft. As they
are hermaphrodites, they are able to produce
female and male gametes. A muscular,
brownish capsule is developed between the
free oviduct and the vagina, here named as
vaginoviducal capsule and its location is
species-specific.

The distal end of the vaginoviducal capsule

bifurcates into a free oviduct and the duct of
the gametolytic organ. The gametolytic sac of
the gametolytic organ is a whitish, tubular
structure that becomes pinkish and
club-shaped in inseminated snails. Two to
seven spermatophores are found in the
pinkish gametolytic sac.

The spermatophore is a whip-like, translucent

structure. The epiphallus possesses a coiled
epiphallic caecum to which a penial retractor
muscle is attached. The penis is a short,
muscular structure with a distinct penial
caecum. The dart apparatus is
well-developed, having four distinct parts.
The proximal part of the dart apparatus
possesses a nondisposable muscular dart.
The dart shaft includes a central core which is
encircled by many layers of circular muscle
cells.

Reference: Soumen Roy (2020): Macro- and

microscopic morphology of the
reproductivesystem of the terrestrial snail
Macrochlamys indica (Godwin-Austen, 1883)
(Eupulmonata,Stylommatophora,
Ariophantidae), Molluscan Research.
 Slide 53

Veronicella is a genus of tropical air-breathing

land slugs in the family Veronicellidae, the
leatherleaf slugs. Veronicellidae species can
only be reliably distinguished from each other
through dissections and observation of the
genitalia.

V. cubensis: It has a varying body color.

There may be multiple shades of brown with
two dark stripes running down the length of
its back. The lines may be solid or broken up
into spots. There may also be an albino form.
Another thin, pale white stripe also runs down
the midline of the animal. The body texture
also varies, where the body may appear
smooth or granular. This slug can usually be
distinguished from other species of
Veronicellidae by the presence of its
blue-gray eye tentacles. There is also a pale
brown area around the eyespots. Adults will
measure between 50-70 mm in length,
although lengths of up to 120 mm have been
recorded.
V. sloanei: Similarly to the other species of
this genus, this animal has variable body
color, ranging from albino, to tan to grey with
varying degrees of grey markings. It has the
potential to attain a maximum length of 120
mm. The tentacles of this species are
typically blue-grey with pale brown tips.
V. moreleti: This brown-colored species
usually does not have a dorsal-median stripe.
Genitalia: The basal section of the penis is
cylindrical. The apex is twisted and the entire
region is a hardened mass.

Part 2:
Slide 54

Class Bivalvia includes both marine and

freshwater organisms such as clams, oysters,
mussels, shipworms etc. they have shells
composed of 2 similar, usually oval, convex
valves which are attached dorsally by
ligament. Their first shells are called dumbo
and have the same composition and structure
as gastropods. Their mantle is in the form of
2 lobes which possess anterior and posterior
adductors. They are able to form pearls that
are formed from mollusks with nacreous
layers in it. For the foot, it is ventral but
compressed.

Moreover, their respiration pathway goes

from inhalant siphon → gills → exhalant
siphon.

They get their food by deposit feeding since

they retain primitive features of the digestive
system.

They typically have an open vascular system

where it goes from heart → tissue sinuses →
nephridia → gills then back to the heart.

They have a pair of nephridia folded into a

long U and one end of it is glandular and
opens into the pericardial cavity.

They have sense organs which include pallial

tentacles, 1 pair of statocyst, ocelli and
osphradium.

Most of them are dioecious with paired

gonads, gonoducts, nephridia and exhalant
siphon, while some are hermaphroditic with
ventral ovary and dorsal testes.

Reference: Mollusk.pdf module

Slide 55

Internal Anatomy

Mantle

The soft parts are covered by the mantle,

which is composed of two thin sheaths of
tissue, thickened at the edges. The two
halves of the mantle are attached to the shell
from the hinge ventral to the pallial line but
are free at their edges. The thickened edges
may or may not be pigmented and have three
folds. The mantle edge often has tentacles; in
clams the tentacles are at the tips of the
siphon. In species such as scallops the
mantle edge not only has tentacles but also
numerous light sensitive organs - eyes.

The main function of the mantle is to secrete

the shell but it also has other purposes. It has
a sensory function and can initiate closure of
the valves in response to unfavourable
environmental conditions. It can control inflow
of water into the body chamber and, in
addition, it has a respiratory function. In
species such as scallops, it controls water
flow into and out of the body chamber and
hence movement of the animal when it
swims.

Gills

The prominent gills or ctenidia are a major

characteristic of lamellibranchs. They are
large leaf-like organs that are used partly for
respiration and partly for filtering food from
the water. Two pairs of gills are located on
each side of the body. At the anterior end, two
pairs of flaps, termed labial palps, surround
the mouth and direct food into the mouth.

Foot

At the base of the visceral mass is the foot. In

species such as clams it is a well developed
organ that is used to burrow into the
substrate and anchor the animal in position.
In scallops and mussels it is much reduced
and may have little function in adults but in
the larval and juvenile stages it is important
and is used for locomotion. In oysters it is
vestigial. Midway along the foot is the
opening from the byssal gland through which
the animal secretes a thread-like, elastic
substance called "byssus" by which it can
attach itself to a substrate. This is important
in species such as mussels and some
scallops enabling the animal to anchor itself
in position.

Digestive system

The large gills filter food from the water and

direct it to the labial palps, which surround the
mouth. Food is sorted and passed into the
mouth. Bivalves have the ability to select food
filtered from the water. Boluses of food,
bound with mucus, that are passed to the
mouth are sometimes rejected by the palps
and discarded from the animal as what is
termed "pseudofaeces". A short oesophagus
leads from the mouth to the stomach, which
is a hollow, chambered sac with several
openings. The stomach is completely
surrounded by the digestive diverticulum
(gland), a dark mass of tissue that is
frequently called the "liver". An opening from
the stomach leads to the much-curled
intestine that extends into the foot in clams
and into the gonad in scallops, ending in the
rectum and eventually the anus. Another
opening from the stomach leads to a closed,
sac-like tube containing the crystalline style.
The style is a clear, gelatinous rod that can
be up to 8 cm in length in some species. It is
round at one end and pointed at the other.
The round end impinges on the gastric shield
in the stomach. It is believed it assists in
mixing food in the stomach and releases
enzymes that assist in digestion. The style is
composed of layers of mucoproteins, which
release digestive enzymes to convert starch
into digestible sugars. If bivalves are held out
of water for a few hours the crystalline style
becomes much reduced and may disappear
but it is reconstituted quickly when the animal
is replaced in water.
Circulatory system

Bivalves have a simple circulatory system,

which is rather difficult to trace. The heart lies
in a transparent sac, the pericardium, close to
the adductor muscle in monomyarian
species. It consists of two irregular shaped
auricles and a ventricle. Anterior and
posterior aorta lead from the ventricle and
carry blood to all parts of the body. The
venous system is a vague series of
thin-walled sinuses through which blood
returns to the heart.

Nervous system

The nervous system is difficult to observe

without special preparation. Essentially it
consists of three pairs of ganglia with
connectives (cerebral, pedal and visceral
ganglia).

Urogenital system

Sexes of bivalves can be separate

(dioecious) or hermaphroditic (monoecious).
The gonad may be a conspicuous, well
defined organ as in scallops or occupy a
major portion of the visceral mass as in
clams. The gonad is generally only evident
during the breeding season in oysters when it
may form up to 50% of the body volume. In
some species such as scallops, the sexes
can be readily distinguished by eye when the
gonad is full since the male gonad is white in
colour and the female is red, even in
hermaphroditic species. Colour of the full
gonad may distinguish the sexes in some
species such as mussels. In other species,
microscopic examination of the gonad is
required to determine the sex of the animal. A
small degree of hermaphroditism may occur
in dioecious species.

Protandry and sex reversal may occur in

bivalves. In some species there is a
preponderance of males in smaller animals
indicating that either males develop sexually
before females or that some animals develop
as males first and then change to females as
they become larger. In some species, e.g. the
European flat oyster, Ostrea edulis, the
animal may spawn originally as a male in a
season, refill the gonad with eggs and spawn
a second time during the season as a female.

The renal system is difficult to observe in

some bivalves but is evident in such species
as scallops where the two kidneys are two
small, brown, sac-like bodies that lie flattened
against the anterior part of the adductor
muscle. The kidneys empty through large slits
into the mantle chamber. In scallops, eggs
and sperm from the gonads are extruded
through ducts into the lumen of the kidney
and then into the mantle chamber.

Reference:
https://ucmp.berkeley.edu/taxa/inverts/mollus
ca/bivalvia.php

Slide 56

External Anatomy

Adductor muscle

Removal of the mantle shows the underlying

soft body parts, a prominent feature of which
are the adductor muscles clams and mussels
or the single muscle in monomyarian species
(oysters and scallops). In clams and mussels
the two adductor muscles are located near
the anterior and posterior margins of the shell
valves. The large, single muscle is centrally
located in oysters and scallops. The
muscle(s) close the valves and act in
opposition to the ligament and resilium, which
spring the valves open when the muscles
relax.

The most prominent feature of bivalves is the

two valves of the shell that may or may not be
equal and may or may not completely
enclose the inner soft parts. They have a
variety of shapes and colours depending on
species. The valves are composed mostly of
calcium carbonate and have three layers; the
inner or nacreous layer, the middle or
prismatic layer that forms most of the shell,
and the outer layer or periostacum, a brown
leathery layer which is often missing through
abrasion or weathering in older animals.

Bivalves do not have obvious head or tail

regions, but anatomical terms used to
describe these areas in other animals are
applied to them. The umbo or hinge area,
where the valves are joined together, is the
dorsal part of the animal. The region opposite
is the ventral margin. In species with obvious
siphons (clams), the foot is in the
anterior-ventral position and the siphons are
in the posterior area. In oysters the anterior
area is at the hinge and in scallops it is where
the mouth and rudimentary foot are located.

Reference:
https://ucmp.berkeley.edu/taxa/inverts/mollus
ca/bivalvia.php

Slide 57

The shell of it is a calcium carbonate layer. It

has an outer, chalk-like prismatic layer and an
inner pearly, lamellar or nacreous layer. The
layers usually incorporate a substance called
conchiolin, often in order to help bind the
calcium carbonate crystals together.
Conchiolin is composed largely of
quinone-tanned proteins.

The periostracum and prismatic layer are

secreted by a marginal band of cells, so that
the shell grows at its outer edge. Conversely,
the nacreous layer is derived from the main
surface of the mantle. Also, Nacre is secreted
by the epithelial cells of the mantle tissue of
various molluscs. The nacre is continuously
deposited onto the inner surface of the shell,
the iridescent nacreous layer, commonly
known as mother of pearl.

Reference:
https://ucmp.berkeley.edu/taxa/inverts/mollus
ca/bivalvia.php
Slide 58

Water flows into the filaments of each

demibranch outer demibranch → inner
demibranch, then moves posterior in the
epibranchial spaces. From here are blood
vessels efferent and afferent which carries
nutrients and waste products to and from the
body.

Slide 60

Most bivalves broadcast spawn Broadcast

Spawning: Releasing gametes — eggs and
sperm — into the water for external
fertilization. by releasing their gametes
directly into the water where fertilization takes
place. Fertilization results in an embryo that
passes through several distinct larval stages,
the last of which is the veliger.

Planktonic larval type characteristic of most

mollusks (including bivalves), characterized
by a ciliated locomotory organ (velum) which
is either discarded or resorbed at
metamorphosis., which swims in the
plankton, feeds (or not), grows, then settles to
grow to adulthood.

Slide 61

Palaeoheterodonta is a subclass of bivalve

molluscs. It contains the extant orders
Unionida (freshwater mussels) and
Trigoniida. They are distinguished by having
the two halves of the shell be of equal size
and shape, but by having the hinge teeth be
in a single row, rather than separated into two
groups, as they are in the clams and cockles.
They have an inner nacreous layer, but
usually without siphons.
Slide 62

Subclass Heterodonta have shells that are

highly variable. They have hinge teeth that
may be reduced or absent. Their shell can be
crossed-lamellar or prismatic layers, but it will
never contain nacreous layers. They are
primitively isomyarian but with a wide range
of adductor muscle configurations. They have
extensively fused mantle margins.

An roder below it is Veneroida. They are

characterized with typically equivalent shells
and of outer crossed-lamellar and inner
complex crossed-lamellar layers. Their hinge
comprises radiating cardinal and lateral teeth
which are often weakly developed. According
to their habit, their adductor muscles are of
varying proportions. Their mantle margins are
also extensively fused which oftenly develop
into long active siphons.

Slide 63

Order Myoida are marine deep burrower

organisms characterized with typically thin,
equivalent shells, comprising either 2 or 3
layers. They have hinge plates with cardinal
dentition and are often degenerate. Most of
them are approximately isomyarian but with
much variation. Boring forms develop
accessory shell plates. Also, their mantle
margins are extensively fused and covered in
periostracum.
Slide 64

The Pteriomorphia comprise a subclass of

saltwater clams, marine bivalve molluscs.[1] It
contains several major orders, including the
Arcida, Ostreida, Pectinida, Limida, Mytilida,
and Pteriida. It also contains some extinct and
probably basal families, such as the Evyanidae,
Colpomyidae, Bakevelliidae, Cassianellidae,
and Lithiotidae.

This subclass of molluscs has lamellibranch

gills, and is epibenthic. Some attach to the
substrate using a byssus. The foot is reduced.
The mantle margins are not fused. Gills are
usually large and assist in feeding. This group
includes the well known mussels, scallops, pen
shells, and oysters.[2] It also includes the only
members of the class bivalvia to have
rudimentary eyes.

Order Mytiloida (common mussels)

Shell equivalve, rounded, elongate or triangular

depending on habits; anisomyarian tending
toward monomyarian; hinge edentulous; shell
microstructure of outer calcitic fibrous prisms
and inner nacre; ctenidia filibranch; mantle
margin lacking fusions; foot creeping; typically
byssate; marine, estuarine, rarely freshwater;
endobyssate and epibyssate.

Slide 65

Order Arcoida have solid, elongate or

circular-oval, often heavily ribbed shells.
They have fibrous periostracum with simple
crossed-lamellar outer layer and inner
complex crossed-lamellar layer, thereby
differing from all other pteriomorphia;
dimyarian; hinge with vertical denticulations;
ctenidia filibranch; mantle margin with
uniquely divided outer fold; foot often
byssate; marine
Slide 66

Order Pterioida (pearl oysters and fan

shells)Shell equivalve, variably shaped;
anisomyarian but often monomyarian; shell
structure of outer simple calcitic prisms and
inner nacre; ctenidia pseudo lamellibranch,
often plicate (deeply folded); mantle margin
lacking fusions; foot reduced; marine;
endobyssate or epibyssate. About 100
species.

Slide 67

Don't include

Slide 68

Monoplacophora, is a polyphyletic superclass

of molluscs with a cap-like shell now living at
the bottom of the deep sea.

Although the shell of many

monoplacophorans is limpet-like in shape,
they are not gastropods, nor do they have
any close relation to gastropods.
Slide 69

Octopus have large eyes with which it scans

its environment actively and which allow the
organism to discriminate objects easily. On
cursory examination, the single-chambered
eyes of octopus with their spherical lenses
resemble vertebrate eyes. However there are
also apparent differences. For example, the
retina of the octopus is everted instead of
inverted, and it is equipped with primary
rhabdomeric photoreceptors rather than
secondary ciliary variety found in the retina of
the vertebrate eye. The eyes of octopus are
well adapted to the habitat and lifestyle of the
species; the pupil closes quickly as a
response to sudden light stimuli mimicking a
situation in which the octopus leaves its den
in shallow water during daytime. Although the
many general anatomical and physiological
features of octopus vision have been
described elsewhere, our review reveals that
a lot of information is still missing.
Investigations that remain to be undertaken
include a detailed examination of the dioptric
apparatus or the visual functions such as
brightness discrimination as well as a
conclusive test for a faculty analogous to, or
in lieu of, color vision. For a better
understanding of the octopus eye and the
functions mediated by it, we suggest that
future studies focus on knowledge gaps that
we outline in the present review.

Slide 70

Nautiloids are a large and diverse group of

marine cephalopods (Mollusca) belonging to
the subclass Nautiloidea that began in the
Late Cambrian and are represented today by
the living Nautilus and Allonautilus. Nautiloids
flourished during the early Paleozoic era,
where they constituted the main predatory
animals, and developed an extraordinary
diversity of shell shapes and forms. Some
2,500 species of fossil nautiloids are known,
but only a handful of species survive to the
present day.

They are distinguished by two main

characteristics—simple concave septa,
concave in the forward direction, that produce
generally simple sutures, and a siphuncle in
which the septal necks point to the rear (i.e. is
retrosiphonate, throughout the ontogeny of the
animal).

Nautili are free swimming animals that possess

a head with two simple lens-free eyes and arms
(or tentacles). They have a smooth shell over a
large body chamber, which is divided into sub
chambers filled with an inert gas (similar to the
composition of atmospheric air, but with more
nitrogen and less oxygen) making the animal
neutrally buoyant in the water. As many as 90
tentacles are arranged in two circles around the
mouth. The animal is predatory, and has jaws
which are horny and beak-like, allowing it to
feed on crustaceans.

Slide 71

The "tentacles" of the nautiluses are actually

cirri (singular: cirrus), composed of long, soft,
flexible appendages which are retractable
into corresponding hardened sheaths. Unlike
the 8–10 head appendages of coleoid
cephalopods, nautiluses have many cirri. In
the early embryonic stages of nautilus
development a single molluscan foot
differentiates into a total of 60–90 cirri,
varying even within a species. Nautilus cirri
also differ from the tentacles of some coleoids
in that they are non-elastic and lack pads or
suckers. Instead, nautilus cirri adhere to prey
by means of their ridged surface. Nautiluses
have a powerful grip, and attempts to take an
object already grasped by a nautilus may tear
away the animal's cirri, which will remain
firmly attached to the surface of the object.
The main cirri emerge from sheaths which
cohere into a single firm fleshy mass. Also,
the pair of cirri before the eye (pre-ocular)
and the pair of cirri behind the eye
(post-ocular) are separate from the others.
These are more evidently grooved, with more
pronounced ridges. They are extensively
ciliated and are believed to serve an olfactory
purpose.

The radula is wide and distinctively has nine

teeth.

The mouth consists of a parrot-like beak

made up of two interlocking jaws capable of
ripping the animal's food— mostly
crustaceans— from the rocks to which they
are attached.Males can be superficially
differentiated from females by examining the
arrangement of tentacles around the buccal
cone: males have a spadix organ (shaped
like a spike or shovel) located on the left side
of the cone making the cone look irregular,
whereas the buccal cone of the female is
bilaterally symmetrical.

The central component of the nautilus

nervous system is the oesophageal nerve
ring which is a collection of ganglia,
commissures, and connectives that together
form a ring around the animal's oesophagus.
From this ring extend all of the nerves
forward to the mouth, tentacles, and funnel;
laterally to the eyes and rhinophores; and
posteriorly to the remaining organs.

The nerve ring does not constitute what is

typically considered a cephalopod "brain": the
upper portion of the nerve ring lacks
differentiated lobes, and most of the nervous
tissue appears to focus on finding and
consuming food (i.e., it lacks a "higher
learning" center). Nautiluses also tend to
have rather short memory spans, and the
nerve ring is not protected by any form of
brain case.

The crop is the largest portion of the digestive

tract, and is highly extensible. From the crop,
food passes to the small muscular stomach
for crushing, and then goes past a digestive
caecum before entering the relatively brief
intestine.
Slide 72

Subclass Coleoidea or Dibranchiata, is the

grouping of cephalopods containing all the
various taxa popularly thought of as
"soft-bodied" or "shell-less" (i.e., octopuses,
squid and cuttlefish). Unlike its extant sister
group, Nautiloidea, whose members have a
rigid outer shell for protection, the coleoids
have at most an internal cuttlebone, gladius,
or shell that is used for buoyancy or support.
Some species have lost their cuttlebone
altogether, while in some it has been replaced
by a chitinous support structure.
The major divisions of Coleoidea are based
upon the number of arms or tentacles and
their structure. The extinct and most primitive
form, the Belemnoidea, presumably had ten
equally-sized arms in five pairs numbered
dorsal to ventral as I, II, III, IV and V. More
modern species either modified or lost a pair
of arms. The superorder Decapodiformes has
arm pair IV modified into long tentacles with
suckers generally only on the club-shaped
distal end. Superorder Octopodiformes has
modifications to arm pair II; it is significantly
reduced and used only as a sensory filament
in the Vampyromorphida, while Octopoda
species have totally lost that arm pair.

Order Sepioidea (cuttlefishes and bottle-tailed

squids)Early Cenozoic to present; worldwide
with family exceptions; shell coiled and
chambered (Spirulidae), straight with vestigial
chambering (Sepiidae), vestigial, or lacking;
eyes covered with transparent membrane; 8
sucker-bearing arms and 2 tentacles retractile
into pockets; total length 2.5–90 cm.
Slide 73

Theuthoidea have shells that has an internal

simple, rod-like or feather-like chitinous
gladius. Ten circum-oral appendages; the
tentacles are retractile (but not retractile into
pockets). The arm and tentacle suckers are
provided with chitinous rings and/or hooks.
Radular teeth commonly with primary
projection and secondary cusp(s), especially
on the median (rachidian) and first lateral
teeth. A buccal membrane is present. The
olfactory organ consists of two projecting
papillae without protective epithelium. The
eyes are either covered with a transparent
membrane with a minute pore (Suborder
Myopsida), or without membrane and
completely exposed to the water (Suborder
Oegopsida). Gills with branchial canal
between afferent and efferent branchial blood
vessels. The digestive gland is a simple
structure.

Slide 74

Squid are soft-bodied molluscs whose forms

evolved to adopt an active predatory lifestyle.
The head and foot of the squid are at one end
of a long body, and this end is functionally
anterior, leading the animal as it moves
through the water. A set of eight arms and
two distinctive tentacles surround the mouth;
each appendage takes the form of a
muscular hydrostat and is flexible and
prehensile, usually bearing disc-like suckers.

The suckers may lie directly on the arm or be

stalked. Their rims are stiffened with chitin
and may contain minute toothlike denticles.
These features, as well as strong
musculature, and a small ganglion beneath
each sucker to allow individual control,
provide a very powerful adhesion to grip prey.
Hooks are present on the arms and tentacles
in some species, but their function is unclear.
The two tentacles are much longer than the
arms and are retractile. Suckers are limited to
the spatulate tip of the tentacle, known as the
manus. They also have Proostracum which is
the anterior horny or calcareous prolongation
of the phragmocone of belemnites and
related cephalopods.

Slide 75

Octopods are the order which contains the

common octopus and some other types.
Fossils are rare, but they do have a fossil
record starting in the later Carboniferous.

Octopods are put into two suborders. One,

the Incirrina, is composed of the well-known
Octopus of rocky shores and coral reefs and
its relatives. The other suborder, the Cirrina,
contains octopods whose tentacles are linked
by an umbrella-like mantle, so their activity is
rather different from the common octopus.

Slide 76

The smallest octopus is the Octopus wolfi. It

is smaller than an inch (2.5 cm) long and
weighs less than a gram.
Slide 77

The arms/appendages, the head and the

mantle. The mantle holds all the animal’s
internal organs, the most noticeable being the
large muscular tube called the siphon and
also the digestive cecum. The head houses
the brain which provides the basis to the
nervous system. It also consists of the beak
and the opening of the siphon. The tentacles
are the parts that contain most of the nervous
system which aids in the octopus locomotion
and other hunting and feeding strategies as
well as for defense.

Its gills, hearts, digestive system and

reproductive glands are all crammed into this
one space. The strong muscles in the mantle
protect the organs and help with respiration
and contraction. The octopus also has a
funnel, sometimes called a siphon, which is a
tubular opening that serves as a pathway for
water.