Phylum Mollusca

• Second-largest phylum in numbers of

species- over 100,000 described.
• Ecologically widespread- marine,
freshwater, terrestrial (gastropods very
successful on land)
• Great variety of body plan (therefore,
many classes within the phylum)
• Great variety in body size- from ~1 mm to
~18 m (60 feet). 80% are under 5 cm, but
many are large and therefore significant as
food for man.

Extant Molluscan classes

Gastropoda Cephalopoda Bivalvia
(snails) (octopus, squid, (clams, mussels)
nautilus)

Aplacophora
Polyplacophora
Monoplacophora (chitons)

Scaphopoda
(tusk shells)

1
 Some basic molluscan characteristics
• Ciliated body surface
• Calcareous shell- composed of three primary layers- outer
periostracum, middle prismatic layer (columnar crystals of
calcite) and inner nacre (flat crystals of calcite)
• Mantle- dorsal surface of body wall, modified to secrete
shell
• Radula- a rasping “tongue” with chitin teeth, sometimes also
chitinous jaws
• Ctenidia- ciliated gills for respiratory gas exchange, usually
located in a mantle cavity
• Open circulatory system (hemocoel)- coelom is reduced

Shell
Mantle

mouth

Mantle cavity

Heart: ventricle
atrium
pericardium

ctenidium

2
 digestive gland

radula

stomach & style sac

Digestive system:
Radula on protrusible
rasping tongue

Radula in action- a prosobranch gastropod on glass

3
 Gastropods and cephalopods
also have chitin cutting plates
or “jaws”
(Beak and radula of a squid)

Molluscan digestive tract, continued

• Style sac and crystalline style in monoplacophorans, some

gastropods, most bivalves.

• Digestive gland- also called “hepatopancreas”

• produces digestive enzymes for extracellular digestion
(pancreas-like function)
• Stores glycogen (liver-like)
• Intracellular digestion

• Intestine – more absorption- anus into mantle cavity

Bivalve digestive system

Digestive gland
(hepatopancreas)
Food
string
Style

Style sac

Stomach

Gut

4
 Juvenile freshwater bivalve- note the rotation of food in
the stomach, and movement of cilia on the ctenidia

Molluscan circulatory system

• Multichambered heart (1-2 atria, 1 ventricle)

• Arterial vessels to organs and hemocoel
• Blood from ctenidia, enters atrium of heart, then
ventricle pumps it out.
• Open circulatory system-
no separation of blood and extracellular fluid-
no separate lymphatic system
• Body cavity is hemocoel
• Heart lies within pericardium (coelomic remnant)

Mollusc blood is hemolymph

(because there is no
separation of blood and ECF)
Hemolymph contains an
oxygen transport pigment
called hemocyanin- blue
when oxygenated
Some mollusca also use
myoglobin in the muscles

5
 Excretory system
Heart acts as filter- filtrate enters pericardium-
from there enters nephridia
Nephridia are analogous to kidney tubules
they modify the filtrate by selective
reabsorption and secretion – becomes urine
Gonoducts
Primitively, the gonads also empty into the
excretory ducts, (e.g. archaegastropods) but in
most molluscs there are separate gonoducts

gonad
ventricle excretory
atrium system

pericardium
artery
vein

ctenidium

Reproduction
Most molluscs are
dioecious, and most have
internal fertilization
Primitive larval stage of
marine molluscs is a
trochophore.
Annelids have a similar
larva.

6
Class Polyplacophora (chitons)

• ~800 species, all marine, many intertidal

• Shell is distinctive- 8 overlapping plates

imbedded partly or entirely in tough “girdle”.
• Mantle space extends around perimeter of
animal (not just posterior).
• Ctenidia are lateral and multiple.
• Very conservative class.
Fossils date to mid/late Cambrian (500 my).

7
8
9
 A collection of chitons

Class Monoplacophora (“single shell carriers”)

• Originally described from fossils- thought

to be extinct since Devonian (350 mya)
• first live specimens recovered in 1952
from 3,600 m depth
• Primitive class, most similar to
gastropods, chitons
• Organ systems show metamerism, similar
to annelids and arthropods
• (Most molluscs are not metameric)

Monoplacophorans are chiton-like in having multiple,

lateral ctenidia. However, the excretory organs , muscles
and gonads are also multiple.
Unlike chitons, but similar to gastropods, they have a single
cap-shaped shell

10
 Class Aplacophora (“no-shell carriers”)
• Wormlike, 2-25 mm long. Body surface covered
with calcareous spicules
• Primitive or derived?
• Two subclasses- solenogasters and chaetogasters

Solenogasters have a
ventral groove and are
errant.
They feed on cnidarians

Chaetogasters have no ventral groove.

They are burrowers.
They have a pair of ctenidia (gills) at
the posterior end.

11
 Class Scaphopoda (“spade-foots”)
• All marine, ~350 species
• More similar to Bivalves than to the
other mollusc classes
• Tubular shell, burrowing habit, tip
of shell protrudes from the substrate
• Feeding tentacles probe for
meiofauna, e.g. forams.
• Tidal ventilation of mantle cavity
via posterior end- ctenidia are lost
• Scaphopods not large (2 mm-15 cm)
but have external fertilization.

Scaphopod shells were used

as ornamental beads since
ancient times and were used
for trade money by some NA
Indian tribes

Class Gastropoda (“belly-foot”)

• snails, slugs, nudibranchs, limpets, cowrys,
abalones, etc.
• By far the largest and most diverse molluscan
class- over 80% of mollusc species are gastropods
• Marine benthic, pelagic, freshwater benthic,
terrestrial (mesic & xeric)
• Grazers or predators
• Prominent head, with well-developed sensory
structures (second only to cephalopods)

12
 Veliger larva of a
marine gastropod

The veliger stage occurs in

most marine and some
freshwater gastropods,
bivalves, and scaphopods.

The veliger follows the trochophore stage.

The trochophore’s equatorial band of cilia is expanded
into velar lobes on either side of the head.
The velar lobes are used for swimming and food capture

Torsion- rotation of visceral hump, bringing mantle

cavity anterior.
Unusual developmental process that is peculiar to
gastropods and apparently a primitive feature in this
class.

13
14
15
 Gastropod shell
Basically a cone, straight or coiled, with new shell
deposited at the margin of the open end during growth

New growth

limpets snails

Operculum is present in many prosobranch gastropods.

“Door” of shell on the dorsal foot that closes aperture of shell
when the snail retracts

Class Gastropoda >40K species (largest class)

Subclass Prosobranchia (“gills before”) >20K
mainly marine, some freshwater and few terrestrial
generally operculate, with ctenidium (many exceptions)
Subclass Opisthobranchia (“gills behind”) ~2K
mainly marine, few elsewhere
generally shell reduced or absent, detorted
ctenidia often replaced by secondary respiratory surfaces
Subclass Pulmonata (“having lungs”) ~20K
mainly freshwater and terrestrial
no operculum, no ctenidium, mantle cavity is lung

16
Subclass Prosobranchia*
Order Archaeogastropoda primitive snails, often
with excurrent openings in shell. Abalones, slit shells
Order Mesogastropoda many marine families and
most freshwater & land prosobranchs. Conchs,
cowries, periwinkles, cerithiaceans, helmets, slipper
shells, moon shells, et al.
Order Neogastropoda nearly all marine- Long
incurrent siphon and associated shell modification.
Muricids, whelks, volutes, olives, cones, et al.
Order Neritacea small order- marine and freshwater,
operculum has a “peg” attachment to foot

* You don’t have to know the orders for examination

Gastropoda, Prosobranchia, Archeogastropoda

Abalone are large, valuable snails that are harvested and
cultured for food and the iridescent shells

17
Archaeogastropoda- Slitsnails- dibranchiate, with
excurrent opening at posterior of mantle cavity

Andanson's slitsnail
(Entemnotrochus adansonianus).

King Midas slitsnail

(Perotrochus midas)
feeding on a sponge.

Gastropoda,
Prosobranchia,
Mesogastropoda

Example- conchs (Strombus ~50

species)

The queen conch (Strombus gigas)

herbivorous- it is common in the
Caribbean and is harvested for its
meat and shell.

This species is economically

important and of conservation
concern due to overharvesting and
pollution

Conch pearls

18
cowries
Some other mesogastropod
prosobranchs

tritons

Gastropoda,
Prosobranchia,
Neogastropoda
Example- cone snails-
Conus
500-1000 species.
All cone snails inject venom
and several of the fish
eaters can be fatal to
humans

19
 Cone snails
are able to engulf
their prey whole
after paralyzing it
with venom

Cone snails have a unique venom strategy.

The venom contains many different small peptides
that target many different receptors and enzymes.
One species may have over 100 different peptides,
with great variety among species as well.
Conotoxins have a very useful characteristic: they
have exceptionally high affinity for receptors and a
very high target specificity.
Great interest in these peptides as
pharmacological agents for research and for drug
use

20
 Some other neogastropod prosobranchs

Volutes

Muricids whelks

Subclass Opisthobranchia (gills-behind)

some major groups by common name
• Bubble snails (cephalaspids)
• Sea slugs
• Nudibranchs: (dorids, aeolids) most are
carnivorous, feed on cnidarians
• Sacoglossans: most are herbivorous- many
adopt chloroplasts
• Pteropods: (thecosomes, gymnosomes)
pelagic, foot modified into winglike lobes
• Sea Hares: (anaspids) important in neurobiology

Bubble shells (Order Cephalaspida)

Most are aeolian carnivores on other gastropods and
annelids

21
Nudibranchs (Order Nudibranchia)
Shell, mantle cavity and ctenida are gone
Possess cerata (singular = ceras) dorsal projections of
the body wall and hemocoel that act as secondary gills

Most are carnivorous and feed largely on cnidarian

polyps.
There are two suborders-
Doridina (dorids) and Aeolidina (aeolids).
Nudibranchs are some of the most incredibly colorful
animals on earth

Dorid nudibranchs

More dorid nudibranchs

(suborder Doridina))

22
 Aeolid nudibranchs
(suborder Aeolidina)

http://www.mbayaq.org/video/sea_slug_reproduction320x240.mov

Borrowed weapons
Aeolids feed on cnidarians and store the functional
nematocysts at the tips of their cerata in cnidosacs
Each ceras contains a branch of the digestive gland. A
duct connects the cnidosac to the digestive gland.

Digestive gland
cnidosac Stained section of cnidosac showing
nematocysts at tip

Saccoglossan sea slugs-

Shell reduced or absent
most saccoglossans are herbivores
that suck plant cytoplasm- some can
hijack chloroplasts

23
 Solar-powered Opisthobranchs

Left: sacoglossan Placida showing network of ducts containing

green chloroplasts from its algal food.
Right: aeolid nudibranch Pteraeolidia "farms" colonies of
brown single-celled algae (zooxanthellae) in its cerata (stolen
from cnidarian prey).

Julia and Berthelina are remarkable

sacoglossan 'snails' with bivalve shells.
Convergent evolution!

Mimicry among dorid nudibranchs and polyclad flatworms-

can you tell which is which?

24
 PHOTO: Three common
marine polyclad flatworms
from south-eastern
Australia.
A: Paraplanocera
oligoglena.
B: Pseudoceros sp.
C: Eurylepta
fuscopunctatus.

Opisthobranch gastropods, continued

Pteropods (“winged feet”)

• Two groups (orders) of planktonic sea slugs with either an

external lightly calcified shell or a leathery pseudoconch.

• Thecosome pteropods are small suspension feeders,

catching plankton by producing mucous nets, some of
which can be up to 2 meters in diameter.

• Gymnosome pteropods are larger predators and capture

thecosomes and other plankton

A thecosome pteropod

Creseis has an elongate

external shell, and a pair of
wing-like flaps which are
used in swimming.

It feeds by trapping
plankton in a transparent
mucous web suspended
above it in the water.

The web is drawn into the

mouth at intervals and
ingested

25
Gymnosome pteropod
Cliopsis is a highly specialised
carnivore.

It feeds on thecosome pteropods,

which it locates by encountering
their nets.

The wing-like lobes are used for

swimming,

Pterotrachea coronata
“sea elephant”
A pelagic prosobranch
(Mesogastropoda:
Heteropoda) that is
convergent on pteropods
The foot is a sculling fin

26
 Sea hares- (O. Cephalaspida)
internal shell, ink defense,
neurobiology subjects

Review:
Subclass Opisthobranchia (gills-behind)
some major groups by common name
• Bubble snails (cephalaspids)
• Sea slugs
• Nudibranchs: (dorids, aeolids) most are
carnivorous, feed on cnidarians
• Sacoglossans: most are herbivorous- many
adopt chloroplasts
• Pteropods: (thecosomes, gymnosomes)
pelagic, foot modified into winglike lobes
• Sea Hares: (anaspids) important in neurobiology

Class Gastropoda
Subclass Prosobranchia (“gills before”) >20K
mainly marine, many freshwater and few terrestrial
generally operculate, with ctenidia (many exceptions)

Subclass Opisthobranchia (“gills behind”) ~2K

mainly marine, few elsewhere
generally shell reduced or absent, detorted
ctenidia often replaced by secondary respiratory surfaces

Subclass Pulmonata (“having lungs”) ~20K

mainly freshwater and terrestrial
no operculum, no ctenidium, mantle cavity is lung

27
Subclass Pulmonata- see next file

28