Lophotrochozoa

The Mollusca
Sea slugs, squid, snails, and scallops

An introduction
Mollusca is one of the
most diverse groups of
animals on the planet,
with at least 50,000
living species (and
more likely around
200,000). It includes
such familiar organisms
as snails, octopuses,
squid, clams, scallops,
oysters, and chitons.
Mollusca also includes
some lesser known
groups like the A cuttlefish, a coleoid cephalopod, moves primarily
monoplacophorans, a by undulating its body fins.
group once thought to
be extinct for millions of years until one was found in 1952 in the deep ocean off
the coast of Costa Rica.

Molluscs are a clade of organisms that all have soft bodies which typically have
a "head" and a "foot" region. Often their bodies are covered by a hard
exoskeleton, as in the shells of snails and clams or the plates of chitons.

A part of almost every ecosystem in the world, molluscs are extremely important
members of many ecological communities. They range in distribution from
terrestrial mountain tops to the hot vents and cold seeps of the deep sea, and
range in size from 20-meter-long giant squid to microscopic aplacophorans, a
millimeter or less in length, that live between sand grains.

These creatures have been important to humans throughout history as a source of

food, jewelry, tools, and even pets. For example, on the Pacific coast of
California, Native Americans consumed large quantities of abalone and
especially owl limpets. However, the impact of Native Americans on these
molluscan communities pales by comparison to the overharvesting of some
molluscan taxa by the United States in the 1960s and 1970s. Species whose
members once numbered in the millions, now teeter on the verge of extinction.
For example, fewer than 100 white abalone remain after several million
individuals were captured and sold as meat in the 1970s. Besides having yummy
soft parts, molluscs often have desirable hard parts. The shells of some molluscs
are considered quite beautiful and valuable. Molluscs can also be nuisances,
such as the common garden snail; and molluscs make up a major component of
fouling communities both on docks and on the hulls of ships.

On the left is a marine snail, the California Trivia (Trivia californiana). Here the mantle
covers much of the shell. Note how a portion of the mantle is rolled into a tube shape to
form the siphon just above the head. At the right is a restoration of one of the largest of all
molluscs, the Giant Squid (Architeuthis).

They also have a very long and rich fossil record going back more than 550
million years, making them one of the most common types of organism used by
paleontologists to study the history of life.

Systematics
Molluscan systematics are still in flux. As you can see from the cladogram
below, there is still no agreement on some of the major relationships. The
polytomies shown indicate that the question of which molluscs are the most
closely related is still a matter of debate.

However, new types of data and much larger and more sophisticated analyses
continue to be performed. The resolved relationships shown (such as
cephalopods, scaphopods, and gastropods) are recent discoveries.

Visit the mollusca pages on the Tree of Life for more on molluscan systematics.

Morphology
Despite their amazing diversity, all molluscs share some unique characteristics
that define their body plan. The body has a head, a foot and a visceral mass. This
is all covered with a mantle (also known as a pallium) that typically secretes the
shell. In some groups, like slugs and octopuses, the mantle is secondarily lost,
while in others, it is used for other activities, such as respiration.

The buccal cavity, at the anterior of the

mollusc, contains a radula (lost in
bivalves) — a ribbon of teeth
supported by an odontophore, a
muscular structure. The radula is
generally used for feeding. The ventral
foot is used in locomotion. This foot
propels the mollusc by utilizing
muscular waves and/or cilia in
combination with mucus. The freshwater Sinistral Pond Snail
Typically, at least in the more primitive (Physella sp.) scrapes algae from the
glass with its radula, the two "toothy"
members of each group, there are one
arcs you can see lining the mouth.
or more pairs of gills (called ctenidia)
Click on the photo for a closer look.
which lie in a posterior cavity (the
pallial cavity) or in a posterolateral groove surrounding the foot. The pallial
cavity typically contains a pair of sensory osphradia (for smelling) and is the
space into which the kidneys, gonads, and anus open.

Molluscs are coelomate, although the coelom is reduced and represented by the
kidneys, gonads, and pericardium, the main body cavity which surrounds the
heart.

Life history and ecology

Molluscs occur in almost every habitat found on Earth, where they are often the
most conspicuous organisms. While most are found in the marine environment,
extending from the intertidal to the deepest oceans, several major gastropod
clades live predominantly in freshwater or terrestrial habitats. Remarkably, one
study found around 3000 species within a single locality at a coral reef in New
Caledonia. In terrestrial communities, gastropods can achieve reasonably high
diversity and abundance: as many as 60-70 species may coexist in a single
habitat and abundance in leaf litter can exceed more than 500 individuals in four
liters of litter.

Marine molluscs occur on a large

variety of substrates including rocky
shores, coral reefs, mud flats, and
sandy beaches. Gastropods and chitons
are characteristic of these hard
substrates, and bivalves are commonly
associated with softer substrates where
they burrow into the sediment.
However, there are many exceptions:
the largest living bivalve, Tridacna
Many marine molluscs emerge from
gigas, lives on coral reefs, and many
their eggs as planktonic trochophore
bivalves (e.g., mussels and oysters)
larvae, however, Sinistral Pond Snails
attach themselves to hard substrates.
(Physella sp.) emerge from their eggs
Some microscopic gastropods even
as young snails. The whitish,
live interstitially between sand grains.
jellybean-shaped organisms are
ostracodes (crustaceans). Large concentrations of gastropods
and bivalves are found at hydrothermal
vents in the deep sea. Living in these or other dysoxic habitats appears to be a
plesiomorphic condition for the Mollusca and several outgroups. For example,
the fauna of Palaeozoic hydrothermal vent communities includes the molluscan
groups Bivalvia, Monoplacophora and Gastropoda as well as the outgroups
Brachiopoda and Annelida.

The adoption of different feeding habits appears to have had a profound

influence on molluscan evolution. The change from grazing to other forms of
food acquisition is one of the major features in the radiation of the group. Based
on our current understanding of relationships, the earliest molluscs grazed on
encrusting animals and detritus. Such feeding may have been selective or
indiscriminate and will have encompassed algal, diatom, or cyanobacterial films
and mats, or encrusting colonial animals. Truly herbivorous grazers are
relatively rare and are limited to some polyplacophorans and a few gastropod
groups. Most chaetodermomorph aplacophorans, monoplacophorans and
scaphopods feed on protists and/or bacteria while neomeniomorph
aplacophorans graze on cnidarians. Cephalopods are mainly active predators as
are some gastropods, while a few chitons and septibranch bivalves capture
microcrustaceans. Most bivalves are either suspension or deposit feeders that
indiscriminately take in particles, but then elaborately sort them based on size
and weight, typically assimilating bacteria, protists, and diatoms.

The fossil record

The Mollusca include some of the oldest metazoans known. Late Precambrian
rocks of southern Australia and the White Sea region in northern Russia contain
bilaterally symmetrical, benthic animals with a univalved shell (Kimberella) that
resembles those of molluscs. The earliest unequivocal molluscs are helcionelloid
molluscs that date from Late Ediacaran (Vendian) rocks. In the Early Cambrian
the Coeloscleritophora are also present. Most of the familiar groups, including
gastropods, bivalves, monoplacophorans, and rostroconchs, all date from the
Early Cambrian, whereas cephalopods are first found in the Middle Cambrian,
polyplacophorans in the Late Cambrian, and the Scaphopoda in the Middle
Ordovician. Most of these early taxa tend to be small (‹10 mm in length). The
Late Vendian-Early Cambrian taxa bear little resemblance to the Cambrian-
Ordovician taxa (most of which remain extant today).

On the left is Inoceramus sp., a bivalve from the Cretaceous of Alameda County, CA. At right
is Turritella andersoni, a gastropod from the Eocene of Ventura County, CA.

After their initial appearance, molluscan taxonomic diversity tended to remain

low until the Ordovician, when gastropods, bivalves, and cephalopods show
large increases in diversity. For bivalves and gastropods this diversification
increases throughout the Phanerozoic, with relatively small losses at the end-
Permian and end-Cretaceous extinction events. Cephalopod diversity is much
more variable through the Phanerozoic, whereas the remaining groups
(monoplacophorans, rostroconchs, polyplacophorans, and scaphopods) maintain
low diversity over the entire Phanerozoic or became extinct.

Original text by Paul Bunje, 2003. Photos of cuttlefish, California Trivia, Giant
Squid, Sinistral Pond Snail radula and eggs, all © Larry Jon Friesen. Photos of
Inoceramus and Turritella andersoni by Sarah Rieboldt, © UCMP. Mollusca
phylogeny based on Sigwart, J.D., and M.D. Sutton. 2007. Deep molluscan
phylogeny: synthesis of palaeontological and neontological data. Proceedings of
The Royal Society B 274(1624):2413-2419; and suggestions from Gonzalo
Giribet, Harvard University.