Bivalvia
is
a class
of
marine and freshwater mollusks
that have a laterally compressed shell composed of to valves (left
and right) joined dorsally by an elastic ligament. This ligament in
conjunction with interlocking "teeth" on each of the
valves, forms the hinge. This arrangement allows the shell to be
opened and closed without the two valves becoming disarticulated.
Bivalves typically display bilateral symmetry both in shell and
anatomy, but there are significant departures from this theme such as
scallops and oysters.
This
class is the second most diverse group of mollusks behind gastropods.
Adult shell sizes vary from one millimeter to over a meter in length,
but the majority of species do not exceed 10 cm.
The
name "bivalve" is derived from the Latin
bis,
meaning "two", and valvae,
meaning "leaves of a door".
Not
all animals with shells with two hinged parts are classified under
Bivalvia; other animals with paired valves include certain gastropods
(small
sea snails
in
the family Juliidae) and some members
of the phylum Brachiopoda (image).
California clam shrimp, Cyzicus californicus |
Bivalves
have long been a part of the diets of coastal human populations.
Oysters were cultured
by the Romans, and mariculture
has
more recently become an important source of bivalves for food.
Besides their use as food, oysters are the most common source of
natural pearls.
The shells of bivalves are used in craftwork and the manufacture of
jewelery and buttons. Bivalves have also been used in the biocontrol
of pollution.
Bivalves appear in the fossil record first in the early Cambrian more than 500 million years ago. The total number of living species is approximately 9,200 that inhabit most aquatics habitats. The marines species range from shallow to the deep waters, from equator to the poles. Bivalves also have colonized brackish-water estuaries and fresh water rivers, streams, and lakes.
The
bivalve shell consists of two valves that are hinged dorsally,
usually with interlocking teeth (the hinge), and always with a horny
ligament that connects the two valves along their dorsal surfaces and
acts to force the valves apart. The interior of the valves contain
scars of the various muscles attached to it, in particular the
(usually two, sometimes one) adductor muscles that, on contraction,
close the valves. The shell can also be internal, reduced or even
absent as in shipworms.
Shipworm |
Near
the hinge of the shell is the umbone
or
beak, a rounded, knobbly protuberance. This represents the oldest
portion of the shell. The hinge area is the dorsal region of the
shell and the lower margin is the ventral region. The anterior or
front of the shell is where the byssus
and
foot are located, and the posterior of the shell is where the siphons
are located. When the umbone is uppermost, the valve with the
anterior end to the left is considered to be the left valve, while
the valve with the anterior end to the right is the right valve.
The valves of bivalves are made of either calcite, as is the case in oysters, or both calcite and aragonite. The ligament and byssus, if calcified, are composed of aragonite. The shell usually consists of three layers: an outer periostracum, and outer an inner shell layers.The outermost layer of the shell is the periostracum, a skin-like layer which is composed of a hard organic substance and is usually olive or brown in colour and easily abraded. The outer layer forms surface details such as scales or spines.
In
all mollusks, the mantle
forms
a thin membrane
covering
the animal's body and extending out from it in flaps or lobes. In
bivalves, the mantle lobes secrete the valves, and the mantle crest
secretes the whole hinge mechanism consisting of ligament,
byssus threads, and teeth.
Most
bivalves have a capacious mantle cavity that accommodates large
gills, but they don't have head, radula or jaws. In addition to
respiration, the gills filter food particles from the water in the
majority of bivalves. Some primitive forms feed directly on the
organic matter in fine sediments, a few specialized groups derive
nutrition from symbiotic algae or bacteria, while others capture and
consume small crustaceans and worms in the deep sea.
The
intestine is irregularly looped and opens dorsally into the exhalant
area. Also opening into this region are the paired kidneys and, when
separate from the kidneys, the gonopores of the paired gonads. The
heart typically lies below the center of the valves and consists of
two auricles and a single ventricle that supplies both anterior and
posterior aorta. The nervous system is made up of three pairs of
ganglia.
The
bivalve foot is modified as a powerful digging tool in many groups,
while in those that live a permanently attached life (e.g., oysters),
it is very reduced.
Most
bivalves burrow in sand or mud, some in wood, clay or coral. Some
attach to hard substrates with threadlike strands of protein
(byssus), others by permanently cementing one of their valves.
The
tiny larva of a bivalve produces a single, uncalcified, caplike
shell,called a pellicle. As the larva growths, it is gradually
enveloped by two mantle lobes, each developing a separate center of
calcification – the dissoconchs, the parts of the shell produced
after the larva metamorphoses, assume the proportions and features of
the adult bivalve.
Bivalves
are divided into the following subclasses:
Protobranchia
Pteriomorpha
Pteriomorpha
Anomalodesmata
RostroconchiaHeterodonta
Chama limbula (Heterodonta) |
Spondylus regius (Pteriomorphia) |