Mollusca (Molluscs)
Gerhard Haszprunar, Zoologische Staatssammlung München, Munich, Germany
Mollusca are bilateral (or secondarily asymmetrical) coelomate Protostomia with spiral
cleavage and mostly a (often modified) trochophore-like larva. Most species are provided
with spicule or shell (plates), a mantle cavity with gills, a ciliary or muscular gliding sole
(foot), a visceral portion with heart and excretory organs and alimentary tract with rasping
tongue (radula).
Diversity
The Mollusca is one of the best known and (after the
Arthropoda) the second biggest phylum of the animal
kingdom, having about 100 000 extant species and about
70 000 described fossil species. Molluscs have an unambig-
uous body plan, but show at the same time tremendous
variation. Molluscan diversity concerns all aspects of
biology and also includes a long and extremely rich and
informative fossil record. The most familiar groups are
limpets, whelks, snails and slugs (class Gastropoda: 70 000
species), clams, mussels and scallops (class Bivalvia, also
known as Lamellibranchia or Pelecypoda: 20 000 species),
and squids and octopuses (class Cephalopoda or Sipho-
nopoda: 900 extant species, about 40 000 fossil species).
There are five more main taxa, which are usually referred as
classes: two worm-like, aplacophoran (shell-less) groups,
the classes Solenogastres (or Neomeniomorpha or Ven-
troplicita: 200 species) and Caudofoveata (or Chaetoder-
momorpha: 120 species), chitons (class Polyplacophora:
1000 species), tusk shells (class Scaphopoda: 500 species),
and the limpet-like Tryblidia (class Monoplacophora: 25
species) (Figure 1). The size of extant molluscs ranges from
0.4 mm (omalogyrid gastropods) to 20 m (giant squids of
the genus Architeuthis).
The many relationships between humans and molluscs
include food, money and jewels, art and holy nimbus, but
molluscs are also vectors of severe diseases and serious
pests. In the science arena, molluscs play an important role
as model organisms, particularly in neurobiology and
evolutionary biology. No other group of animals has so
many species under threat of extinction.
Basic Design
One must clearly distinguish between the concepts of a
hypothetical stem species, which is a reconstruction of a
probable historical reality, and that of a body plan
(bauplan), which is an idealistic design containing all
features shared by the majority of species of a phylum.
Molluscs are characterized by a very distinct body plan, yet
there is only a single, cytological, character (rhogocytes,
ENCYCLOPEDIA OF LIFE SCIENCES © 2002, John Wiley & Sons, Ltd. www.els.net
Introductory article
Article Contents
. Diversity
. Basic Design
. Habitats and Abundance
. Habits and Lifestyles
. Life Histories
. Fossil Record
. Phylogeny
see below) that is diagnostic for all molluscan species. In
what follows each major molluscan feature is described
from an evolutionary point of view.
External morphology
External morphology of molluscs varies enormously.
Although chitons, snails, slugs, mussels, scallops or squids
are well known to the public, nearly every major taxon
contains certain species or subgroups that are amazingly
different from the usual design. Particularly odd examples
are shipworms (teredinid bivalves), certain entoparasitic
eulimoid caenogastropods, heteropods like Carinaria or
Pterotrachea, and various pelagic and interstitial opistho-
branchs. Other molluscan classes, however, show very
uniform external morphology, e.g. the Caudofoveata,
Solenogastres, Polyplacophora, Tryblidia (extant Mono-
placophora), Scaphopoda, and the (largely extinct) Nau-
tiloidea (Cephalopoda).
By heredity from their ancestors among the spiralian
protostomes, the earliest molluscs were bilaterally symme-
trical worm-like creatures and probably were small
(millimetre range). The limpet-like monoplacophoran
shape probably gave rise to the other conchiferan (molluscs
with a single shell, including bivalves) groups.
External covering and integument
The dorsal epidermis of the ancestral mollusc was probably
covered by a chitinous cuticle with aragonitic spicules and
scales. The ventral (pedal) sole was ciliated with a
microvillous border. This level is still represented by the
Solenogastres.
Probably by fusion of the spicules, the dorsal integument
became equipped with a series of shell plates, as still seen in
the Polyplacophora. It is uncertain whether the final,
single, molluscan shell evolved by fusion of these shell
plates or as an evolutionary novelty. The presence of a true
shell characterizes the conchiferan level of molluscan
organization, although the shell has been tremendously
modified, reduced, or even lost in various lines. In many
conchiferan taxa the shell has additional functions apart
1
 Mollusca (Molluscs)

Figure 1 Phylogenetic relationships and schematic organizations of the extant classes of the Mollusca, including a diagram of the hypothetical molluscan
stem species. Note that concrete exemplary anatomies are schematized rather than general bauplans. Modified from Salvini-Plawen and Steiner (1996)
and Haszprunar (2000).

2
 Mollusca (Molluscs)

from, or additional to, protection of the soft body: The adductor (shell-closing) muscle system of the
examples are the shells of teredinid shipworms (boring), Bivalvia is diagnostic of the group. Numerous parallel
the shells of scallops (swimming), and the chambered shell lines lead from original ‘dimyarian’ (both adductors of
of the Nautilus (buoyancy). Shell structure varies signifi- equal size), through heteromyarian (the posterior adductor
cantly between taxa, but usually an outer organic layer is larger than the anterior one) to the monomyarian (the
(periostracum) is underlain by one to several calcareous anterior retractor is lost) condition.
layers stabilized by an organic matrix. Old and new data on bivalves and recent studies on
The coleoid cephalopods (sepiids, squids and octopuses) gastropods provide clear evidence that the larval muscu-
have changed their external covering by successive reduc- lature of both groups is entirely independent from the adult
tion of the shell and are famous for their powers of condition whereas scaphopods do not have specific larval
camouflage and colour change by means of chromophores muscles.
and iridocytes.
Molluscs produce many glandular secretion such as
mucus for gliding or to prevent desiccation, and acid Mantle cavity
secretion against potential predators. Certain glandular
The molluscan mantle cavity contains the excretory,
products, like the gastropod operculum, effectively protect
genital (or urinogenital) and anal openings, but also
the retracted body, and byssus threads fix larval clams or
certain typical organs like the chemoreceptive osphradium
adult mussels on the hard bottom. Various glands produce
(see below under sensory organs), ctenidia or gills, and
pheromone-like substances used during courtship.
various distinct (‘hypobranchial’ or ‘pallial’) glands.
Originally situated in the posterior part of the animal’s
body (still in the aplacophoran groups), the mantle cavity is
Muscle systems circumpedal (i.e. surrounds the whole pedal sole) in Poly-
and Monoplacophora, Bivalvia and Scaphopoda. In
The worm-like molluscan stem species had a more or less
contrast, a restricted mantle cavity is present in Cephalo-
complete layer of body wall musculature. Clearly second-
poda and Gastropoda. In Gastropoda the unique torsion
ary systems of body wall musculature have been indepen-
process results in an anterior position of the mantle cavity.
dently established in most pulmonate gastropods
Many opisthobranch gastropods (e.g. shell-less sacoglo-
(particularly slugs), worm-like opisthobranchs and in
ssans, all nudibranchs) have entirely lost the mantle cavity.
coleoid cephalopods. Whereas extension of the body or
Since the Solenogastres (as one of the most primitive
of body parts is usually facilitated by haemolymph
taxa) show none or secondary gills, it is unclear whether or
pressure, prosobranch gastropods and cephalopods have
not the typical molluscan ctenidium was a feature of the
a muscular hydrostatic system where, similar to the
molluscan stem species. Because of the small size of early
vertebrate tongue muscle, contractions cause the extension
molluscs, the primary function of the heavily ciliated
of the animals as a whole or of any appendages, e.g.
ctenidia was ventilation rather than respiration; the latter
tentacles or arms.
function was (and in most cases still is) provided by the
A system of specific dorsoventral muscles (shell muscles or
mantle epithelium or (in heterobranch gastropods) also by
pedal retractor muscles) is present in most molluscs, but
the pallially situated excretory organ. Only cephalopods
shows great modification and reduction. Originally, they
have truly respiratory ctenidia. Often equipped with
were numerous and serially arranged. In chitons they are
chitinous skeletal rods, ctenidia are additionally used for
organized into eight distinct bundle pairs inserting at the
filter feeding in the majority of bivalves and several
shell plates, and this condition remains in the monoplaco-
gastropod groups. Inhabitants of reduced habitats such
phorans despite the single shell. During conchiferan evolu-
as micro-oxygenated mud or hydrothermal vents and seeps
tion the number of these bundles was repeatedly reduced.
often house symbiotic bacteria in the ctenidial leaflets or
Primitive gastropods have a single pair, but most gastropods
filaments.
retain only the single (posttorsional) left shell muscle, known
Because of their overall similarity, it is extremely difficult
as the columellar muscle. A similar trend occurred in
to provide clear evidence for homology of the various
bivalves, where primitive species show seven to eight pairs,
pallial glands with each other. This is true for pallial glands
whereas advanced taxa have only three pairs. Scaphopods
between classes, as well as for mantle glands within
and cephalopods show two pairs of shell muscles.
gastropods in particular.
The detailed anatomy of the buccal musculature has
been widely modified during molluscan evolution, as has
the buccal apparatus as a whole. Bivalves show total loss of Heart and circulatory system
this system.
A mantle retractor system is found in all shelled With few exceptions (e.g. all scaphopods, certain Mono-
conchiferans; the respective insertion area at the shell is placophora, certain opisthobranchs), all molluscs have a
known as the mantle line. heart consisting of one to four pairs of atria and a paired

3
 Mollusca (Molluscs)
but usually fused ventricle. Most gastropods show a
monotocard (a single, left atrium) rather than a diotocard
(one pair of atria) condition. Coleoid cephalopods show
additional branchial hearts to increase the volume and
speed of the blood flow through the ctenidia.
The molluscan circulatory system is ‘sinusoid’, which
means that the blood (haemolymph) runs within well-
defined spaces (sinuses and lacunae) that lack an endothe-
lium. Only stylommatophoran pulmonate gastropods and
Cephalopoda show true ‘vessels’ with endothelia; other-
wise this condition is restricted to the aorta.
Usually the haemolymph transports dissolved oxygen,
but true erythrocytes are known from all Solenogastres,
several bivalvian and certain gastropod taxa. The respira-
tory pigment is haemocyanin in most cases, although many
species have haemoglobin. The buccal muscle cells are
probably always equipped with a myoglobin system.
Aside from erythrocytes, there are many cell types that
have immunogenic or phagocygotic function. Depending
on their shape or content, these haemocytes are referred as
granulocytes, hyalinocytes or amoebocytes. A specific
function concerns the rhogocytes (often referred as pore
cells), which are diagnostic for molluscs: they show
ultrafiltration sites at their surface and play a major role
in metal ion metabolism.
Coelomic conditions, excretory system
There is a long-running discussion as to whether or not
molluscs are coelomates. Additional confusion occurs
because the term ‘coelom’ is differently defined by different
authors. Recent comparative ultrastructural studies have
revealed that the Mollusca are coelomates in the broad
histological sense (i.e. presence of an epithelial secondary
body cavity, namely the gonopericardium), but are not
eucoelomates (i.e. there is no coelomatic cavity homo-
logous to that of, for example, the Sipunculida or
Annelida, where the coelom is intimately associated with
the body wall musculature).
Ultrafiltration in molluscs takes place through podo-
cytes in the atrial wall into the pericardial cavity.
Additional ultrafiltration sites occur in many bivalves
(called ‘pericardial glands’ and in all cephalopods (called
‘appendages of branchial hearts’). In the heart-less
scaphopods, ultrafiltration takes place from a circumrectal
blood sinus directly into the excretory cavity, and similar
conditions are found in pulmonate gastropods. As it passes
towards the external environment the primary urine is
more or less concentrated and modified, particularly in
freshwater and terrestrial species (the latter are usually
provided with a distinct ureter). Starting with the
Polyplacophora the pericardial duct is developed towards
a distinct excretory organ also called nephridium, emunc-
torium or kidney. Owing to the different mode of
ontogenetic development, the molluscan excretory organ
4
is not homologous with nephridia of other animal phyla,
whereas the type of ultrafiltration (from a large primary
into a restricted secondary body cavity) may be function-
ally called ‘metanephridial’. The excretory organ of higher
gastropods (Heterobranchia including opisthobranchs
and pulmonates) is located in the mantle roof and,
additionally, plays an important role in respiration.
In molluscs there is always an intimate association
between the gonad and the pericardial–nephridial system,
so that a gonopericardial system is formed. Common
anlage and heterochronic processes have caused highly
variable conditions throughout molluscan evolution.
Thus, aplacophoran taxa release their gametes through
the pericardium and its ducts, monoplacophorans and
several primitive gastropod groups have urinogenital ducts
and openings, and even advanced gastropods show
‘gonopericardial’ ducts. There is, however, a general trend
to separate the excretory and genital system resulting in
true nephro-and gonopores.
Genital system
The genital system of molluscs is extremely diverse and
ranges from very simple sac-like gonads with a simple
gonoduct to highly differentiated genital systems that are
among the most complicated in the animal kingdom.
Originally a paired genital system was present in molluscs,
but Scaphopoda, many cephalopods and all Gastropoda
show pronounced asymmetry. Whereas Caudofoveata,
most Polyplacophora, most Monoplacophora, most Bi-
valvia, Cephalopoda, and most primitive (prosobranch)
Gastropoda are gonochoristic, all Solenogastres, hetero-
branch Gastropoda and certain taxa from the remaining
groups show hermaphroditic conditions. As mentioned
above, many primitive groups show urinogenital condi-
tions in that the gametes are released through the excretory
organ (duct) or via a urinogenital opening into the free
environment.
Entaquatic (i.e. within the mantle cavity) or internal (i.e.
within the soft body) fertilization have been repeatedly
evolved among the molluscan taxa. Spermatophores are
well known from many cephalopods and gastropods, and
copulatory organs have been formed by various organs
(e.g. cephalopod arms called ‘hectocotylus’ or cephalic
tentacles in many gastropods); the latter are often provided
with penial stylets.
Alimentary tract
The one-way gut of the Mollusca also shows all levels of
evolution, the only common structure (except for the
Bivalvia) being the buccal apparatus with the radula. The
latter organ is broadly known as a rasping tongue with
chitinous teeth, but may be modified into a series of (or a
single) forceps (Caudofoveata), a crusher organ (Scapho-

poda), a hollow harpoon combined with a poison gland
(conid neogastropods), or a filtering fan (certain trochid
vetigastropods). In addition, conchiferans are generally
provided with a pair of jaws.
Starting with a simple, straight, mucoid gut (Soleno-
gastres), most molluscs have specialized the alimentary
canal into an oesophagus (transport, first enzymatic
activity by secretions from the salivary and/or oesophageal
glands), a stomach (sorting area), midgut or digestive
gland(s) (major part of true digestion), intestine and
rectum (transport, further digestion). Relative size and
shape as well as detailed structure vary greatly between the
molluscan taxa.
Nervous system
Like many other organs, the molluscan nervous system is
highly diverse. Primitive groups show simple neural cords
throughout the whole nervous system, whereas advanced
taxa have independently evolved true ganglia that are
interconnected by commissures (between symmetrical
parts) and connectives (between different parts).
The primitive condition of the molluscan nervous system
is characterized by: (1) a cerebral commissure that
innervates the head and oral region; (2) a buccal system
that supplies the buccal apparatus; (3) a pedal system
supplying the foot sole; and (4) a pleurovisceral system,
which innervates the mantle cavity and its organs as well as
the viscera. The larvae are additionally equipped with an
apical ganglion that supplies the apical sensory organ with
its ciliary tuft.
Aside from ganglionization, there is a general trend
towards concentration of the nervous system. This is
particularly obvious in the cephalopods and gastropods,
where the pleurovisceral loop runs within the shell muscles.
In both taxa true brains have been evolved, while
cephalopods additionally have developed a cartilaginous
‘skull’ around the brain.
Primitive gastropods are famous for their crossover of
their visceral nerve loop due to the torsion process. This
‘streptoneuran’ condition is modified towards an euthy-
neuran (no crossover) one by (1) concentration of the
visceral loop (several higher prosobranch taxa); (2)
detorsion of the visceral portion (many opisthobranchs);
or (3) both phenomena (pulmonates). Euthyneuran
gastropods are additionally characterized by giant nerve
cells (up to 2 mm diameter) and specific neurosecretory
centres (e.g. dorsal bodies, rhinophoral ganglion or
procerebrum, pulmonate cerebral gland), which make
them ideal model systems for neurobiological studies.
Sensory organs
Molluscs have many sensory organs, some of which are
more or less generally present, whereas others are restricted
to distinct taxa. Only the most important are listed here.
Mollusca (Molluscs)
So-called osphradia (chemoreceptors) are present in
most molluscs (except Monoplacophora and Scaphopoda)
near the ctenidia or outside the mantle cavity (Solenogas-
tres and Caudofoveata). Despite high variability in shape
(simple epithelial patches or grooves to gill-like organs)
and even fine structure, these organs are homologous
throughout the phylum.
Sensory tentacles are typical for most conchiferan
groups and occur at the head (cephalic), at the mantle
(pallial) or along the foot (epipodial), where they usually
combine chemoreceptive and mechanoreceptive functions.
True cerebral eyes are restricted to Cephalopoda and
Gastropoda and show all levels of eye evolution from the
simple pigment cups to highly evolved complex eyes with
lens and cornea. Similar variability is found in the mantle
eyes of Bivalvia and the shell-plate eyes (aesthetes) of the
Polyplacophora.
True (cerebropleurally innervated) statocysts occur only
in conchiferan molluscs and are equipped with many small
statoconia, or with a single, concentrically structured
statolith.
Habitats and Abundance
Whereas most molluscan classes are restricted to marine
habitats, bivalves have repeatedly colonized freshwater
ecosystems. Here they often represent a major part of the
benthic biomass, as they do in certain soft-bottom marine
ecosystems such as muddy grounds. There are many
independent gastropod lines that have invaded brackish
and freshwater as well as terrestrial ecosystems. Indeed,
gastropods are found in practically all ecosystems on Earth
and are certainly one of the most successful colonizers in
the animal kingdom. Ecologically dominating species are
rare – by contrast, very many terrestrial species show
extremely restricted distribution (e.g. single islands or
valleys) and consequently are often endangered.
Habits and Lifestyles
Other than flying, molluscs have evolved every type of
locomotion. Most of them crawl by the foot sole, and many
use head or foot to burrow into the soft bottom, or to
hobble or jump. Some are able to swim actively by means of
various organs: well-known swimmers are heteropod
caenogastropods, certain opisthobranchs such as the
‘Spanish Dancer’ and the sea hare Aplysia, and squids.
On the other hand, there are many (semi-)sessile molluscs
that move rarely or are cemented to the substrate.
All types of feeding strategies can be found in the
Mollusca: predators, grazers, detritivores, filter feeders
and parasites are all found in the phylum. Several
gastropods and bivalves house symbiotic bacteria or
5
 Mollusca (Molluscs)
zooxanthellae to become at least in part chemo- or
photoautotrophic or to use certain biochemically inert
food (e.g. wood) for nourishment.
Life Histories
The original molluscan life cycle starts with spiral quartet
cleavage and protostomy. Embryonic development is
followed by a planktic, but lecithotrophic trochophore-
like larva, which is equipped with a prototroch, an apical
sense organ with a ciliary tuft and protonephridia. The
benthic adult condition is reached via a metamorphosis.
Main deviations from this basic life cycle concern
primarily the larval phase: Aplacophorans and proto-
branch bivalves often show so-called pericalymma larva,
where the larval body is enveloped by large ciliary cells.
Gastropods and bivalves have independently evolved
planktotrophic veliger larvae, or show intracapsular to
direct development. The latter is typical for freshwater and
terrestrial species. Several partly parasitic larval types (e.g.
the glochidium) are found in unionoid freshwater bivalves.
Cephalopoda are unique among molluscs by producing
very large and yolky eggs, which causes superficial cleavage
similar to that in birds. Yolk digestion is provided by blood
vessels reaching into an internal yolk sac, which is
connected via a thin duct with a very large external yolk
sac. There is no sign of a trochophore-or veliger-like larva
in cephalopods.
Fossil Record
The fossil record of molluscs is long (down to the early
Cambrian), extensive (about 70 000 fossil species have been
described) and highly informative. Because the formation
of the gastropod or bivalve protoconch is correlated with
the type of larval life (e.g. lecithotrophic versus plankto-
trophic), the fossil record of molluscs also provides
informations about the life cyle of extinct species and taxa.
Phylogeny
Since the 1990s there has been substantial progress
concerning our understanding of molluscan phylogeny.
The old controversies of turbellarian versus annelid
affinities are largely overcome in that Mollusca are now
considered to be trochozoan spiralians, but not eucoelo-
mates. The direct sister group is probably the Kamptozoa
(Entoprocta).
Probable affinities within the phylum are presented in
Figure 1. The aplacophoran groups probably represent
subsequent clades with the Solenogastres as the earliest
offshoot. A common origin of Polyplacophora and
Conchifera and the monophyly of Conchifera itself (all
remaining classes) are very certain. Monoplacophora
represent an early but highly specialized offshoot among
the Conchifera. It is likely that Bivalvia, Scaphopoda (plus
the fossil Rostroconcha: Diasoma) as well as Cephalopoda
and Gastropoda (including the fossil Bellerophontida:
Cyrtosoma) are subsequent offshoots. After a revolution
of the classic gastropod system since the mid-1980s there is
increasing consensus towards a new, phylogenetic system.
Also the phylogeny of scaphopods and cephalopods is
nowadays quite well established, whereas aplacophoran,
polyplacophoran and bivalve evolution is still poorly
understood.
Further Reading
Beesley PL, Ross GJB and Wells A (eds) (1998) Mollusca: The Southern
Synthesis. Fauna of Australia, vols 5A and 5B. Melbourne: CSIRO
Publishing.
Harrison FW and Kohn AJ (eds) (1995) Microscopic Anatomy of
Invertebrates, vol 5: Mollusca I: Aplacophora, Polyplacophora,
Prosobranchia, Opisthobranchia. New York: Wiley-Liss.
Harrison FW and Kohn AJ (eds) (1997) Microscopic Anatomy of
Invertebrates, vols 6A and 6B: Mollusca II: Pulmonata, Monoplaco-
phora, Bivalvia, Scaphopoda, Cephalopoda. New York: Wiley-Liss.
Haszprunar G (2000) Is the Aplacophora monophyletic? A cladisitic
point of view. American Malacological Bulletin 15: 115–130.
Rosenberg G (1992) The Encyclopedia of Seashells. New York: The Five
Mile Press.
Salvini-Plawen L and Steiner G (1996) Synapomorphies and plesiomor-
phies in higher classification of Mollusca. In: Taylor JD (ed.) Origin
and Evolutionary Radiation of the Mollusca, pp. 29–51. Oxford:
Oxford University Press.
Taylor JD (ed.) (1996) Origin and Evolutionary Radiation of the Mollusca.
Oxford: Oxford University Press.
Wilbur KM (ed.) (1983–1988) The Mollusca, vols 1–12. Orlando, FL:
Academic Press.
Conch-Net. Conchologists of America, Inc. [http://coa.acnatsci.org/
conchnet/] [a good start with many links]