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Peter Shaw
BIOH20.043S - Invertebrate
Phylum Mollusca
■ One of the most important phyla in
terms of species richness, and
nearly ubiquitous in most
■ They invariably have a
eponymously soft, mucus-covered
body. Many forms have a hard,
calcareous shell protecting most of
the body.
The mollusc
■Is unsegmented
■ Coelomate, but coelom is reduced to pericardium +
gonadial cavity.
■ The main body cavity is a haemocoel.
■ Primitively: shelled, with 2 broad body regions
◆ a muscular foot, with mouth + sense organs at
anterior end which protrudes from the shell
◆ a non-muscular visceral hump staying protected
inside the shell, wherein most action is ciliary.
The ancestral
■ A favourite parlour game for invert.
Zoologists is to create a
hypothetical ancestral mollusc.
■ In 1957 a monoplacophoran
Neopilina galatheae was dredged
up from 5000m off Mexico, and
was found to fit the bill - except
that, rather embarrassingly, it
showed signs of metamerism.
■ Are very much disputed, but current best
guess is close to the Ectoprocta (=
Bryozoa = Kamptozoa, depending on your
book!), whose larval forms can be similar
in general form to an ancestral mollusc.
■ This scenario assumes a ciliary/gliding
animal which in one radiation acquires a
shell, the other evolves lophophorate filter
■ This seems to have changed over
the years, due to increased
‘splitting‘ of obscure minor forms in
recent years
■ To a good first approximation there
are 5 main classes, but you need
to know of another three oddball
■ Most molluscs are aquatic and bear
paired gills or ctenidia, lying in the
mantle cavity. These bear many
triangular leaflets, and water is carried
over these by ciliary action.
■ Water is “tasted” before flowing over the
gill by a sense organ the osphradium.
■ The anus and renal pores lie in the
exhalant stream.
■ Neopilina has no ctenidia, but 5
pairs of muscular gills along its
body. Primitive or derived?
■ Terrestrial gastropods often lose
ctenidia, and have “lungs” in their
■ Usually consist of a unique molluscan
organ the radula. This is a muscular
cord bearing teeth, which rasps slowly
away at the surface of food grating
fragments into the mouth.
■ The radula is born on a supporting
structure the odontophore.
■ The radula has undergone much
modification - filter feeders + some odd
deep water forms lack a radula,
cephalopods have a parrot-like beak.
■ Primitively this involved food particles being
scraped off a surface by the radula/odontophore
and carried in a mucus rope into the stomach,
where it is size-sorted by cilia.
■ Filter-feeders retain the mucus rope, but collect
particles off the ctenidia.
■ Cephalopods bite lumps of flesh with a chitinous
beak prior to radula.
The shell
■ This is invariably calcium carbonate
(aragonite, not calcite - the crystal structure
differs slightly) reinforced with chitin.
■ The superficial appearance differs hugely
between groups, from a tight spiral through
loose tubes to nearly flat.
■ In fact one growth equation with 3
biologically realistic parameters can fit the
whole spectrum
Principal mollusc
C e p h a l o p o d s
O c t o p i , s q u i d s , c u t t

G a s t r o p o d s
S l u g s , s n a i l s w i n k l e

Mollusca P o l y p l a c o p h o r a
C h i t o n s , c o a t - o f - m a

S c a p h o p o d s
e l e p h a n t t u s k s h e

L a m m e l l i b r a n c h i a
b i v a l v e s : m u s s e l s , o
Minor classes
M o l lu s c s

M o n o p l a c o p A h p o l ar a c o p h o r a
N e o p i li n a

n e o m e n i o C r hp ah es t o d e r
■ Chitons, or coat of mail shells
■ Primitive marine molluscs with 8 small
dorsal shells, partly or wholly buried in
■ This is not segmentation but an
adaptation to curling in tight crevices
■ Feeding is by grazing on algae.
■ Mantle cavity runs along side, almost up
to head, with paired gills along its length
■ “Tusk shells”: 350 spp
■ mantle secretes a tubular shell,
open at both ends
■ Burrow in soft marine sediments,
collecting food particles with sticky
clubbed captactula
Bivalvia =
■ 20,000 spp
■ Sedentary/ sessile molluscs encased in
paired shells
■ Lacking eyes, radula tentacles, though
these may re-evolve around mantle
■ Foot retained, often used to burrow into
■ Live by filter feeding, mainly using
ctenidia as filtering surfaces
■ The most numerous group of molluscs,
with c. 76,000 species (75%)
■ Mainly shelled, except for the
■ The shell is always single, usually spiral
■ Locomotion is by muscular waves
passing along the foot, except for a few
tiny spp that use cilia
Gastropods, 2
■ A typical gastropod is a squat mollusc
crawling with its muscular foot.
■ The cephalic sensory organs and radula
are well developed but can retract fully
into the shell
■ Eg common limpet Patella
■ Many gastropods have a
calcareous/chitinous plug to the shell -
the operculum
Gastropods 3
■ All have undergone a curious modification
of body form known as torsion
■ The visceral hump is rotated 180o so that
the mantle cavity, gills and anus all point
■ Perhaps to allow the head to retract into the
generous space of the mantle cavity +
allow osphradium to sense oncoming water
■ But having your anus discharging over your head
is poor design, even for a mollusc
■ The solution is to modify the exhalent stream:
◆ evolve a special slit in the shell
◆ lose one ctenidium (R side) giving a
unidirectional flow
◆ Loss of R gill => loss of R kidney, osphradium
+ R side heart
◆ L (inhalent) mantle protrudes as a siphon
■ Many spp have undergone 90o detorsion!

G a s t r o p o d s

P r o s o b r a n Oc h p s i s t h o b r a n p c u h l ms o n a
a n c e s t r a l g a Ms t r a o i np lo y d m s a mr i n a e i n l y t e
■ Mainly marine “snails”, with
traditional body/shell design
■ Most graze algae, sessile animals,
or sediment
■ Many have notable shells, ie
■ One order Stenoglossa have
effective predators: Conus spp
hunt+kill fish!
■ Marine, shell-less: sea slugs, sea
hares, pteropods etc.
■ Some are pelagic, hunting cnidaria
■ one group are pelagic filter feeders
using mucus nets
■ Ctenidia are reduced or absent:
the body surface is often adequate,
or 2ndry gills have evolved
■ Mainly terrestrial radiations
■ Mantle cavity becomes an air-
breathing lung with a contractile
opening the pneumatostome
■ Most have a shell, but
stylommatophora have lost it: land
■ A few pulmonates have returned to
■ The most intelligent, fastest moving
and highly modified molluscs
■ all are marine predators with good/
excellent vision
■ include the largest invertebrate (giant
■ foot is modified into multiple
tentacles with suckers
■ swim by jet propulsion
Cephalopoda N
a u t i l i d
a u t i l u
A m m o n
e x t i n c t
O c t o p o

T e u t h i d
s q u i d s

V a m p y r o
'V a m p i r e

S e p i i d a
c u t t l e f i s
■ We are lucky to have alive an ancestral
form, Nautilus (6 spp). This has a spiral
shell used for protection and buoyancy.
■ The eye is a pinhole camera with no
lens but a contractile iris
■ It lives as a deep water scavenger,
probably explaining its survival of the
K/T boundary event
■ Its fossil ancestors had straight shells
(<= 4m long)
■ Were large open-water Nautiloids that went
extinct at the end of the Cretaceous
■ Up to 2m diameter shells which differed from
Nautilus by heavy sculpturing of sutures
■ We can only guess at their ecology - ?pelagic
■ Some big bulky forms ?plankton feeders
■ Evolutionary trends of their shells are complex if
not haphazard!
■ Is the sub-class containing the remaining
◆ octopoda
◆ teuthida
◆ sepiida
◆ Vampyromorpha
◆ These have a closed blood circulation, non-
ciliated gills, a good compound eye and well
developed CNS. Are oddly semelparous
Cephalopod sex

These animals have some remarkable reproductive behaviour.

Sexes are separate and fertilisation is internal, but there is no

penis. Instead one of the male’s arms is differentiated (usually
with smaller suckers at the tip), is used to pull the torpedo-
shaped chitinous sperm packet out of his genital opening, and
insert it into the female’s body cavity.

In some deep sea squids the sperm packet may be jammed into
non-standard regions - the mantle, or a tentacle base. How
this effects sperm transfer is still unclear - these animals are
effectively impossible to study alive.
Note the reduced suckers on the hectocotylus arm

arm on
South African