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Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
of marine and freshwater
molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant taxon, extant species of molluscs are recognized. The number of fossil sp ...
that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bivalves have no
head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may ...
and they lack some usual molluscan organs, like the
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
and the
odontophore The odontophore is part of the feeding mechanism in molluscs. It is the cartilage which underlies and supports the radula, a ribbon of teeth. The radula is found in every class of molluscs except for the bivalves. The feeding apparatus can be ...
. They include the
clam Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two she ...
s,
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not al ...
s, cockles,
mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and Freshwater bivalve, freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other ...
s,
scallop Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families ...
s, and numerous other
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
that live in saltwater, as well as a number of families that live in freshwater. The majority are
filter feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s. The
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
s have evolved into ctenidia, specialised organs for feeding and breathing. Most bivalves bury themselves in sediment, where they are relatively safe from
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
. Others lie on the sea floor or attach themselves to rocks or other hard surfaces. Some bivalves, such as the scallops and
file shell The Limidae or file shells are members of the only family of bivalve molluscs in the order Limida. The family includes 130 living species, assigned to 10 genera. Widely distributed in all seas from shallow to deep waters, the species are usually ...
s, can swim. The
shipworms The shipworms are marine bivalve molluscs in the family Teredinidae: a group of saltwater clams with long, soft, naked bodies. They are notorious for boring into (and commonly eventually destroying) wood that is immersed in sea water, including ...
bore into wood, clay, or stone and live inside these substances. The
shell Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses ** Thin-shell structure Science Biology * Seashell, a hard ou ...
of a bivalve is composed of calcium carbonate, and consists of two, usually similar, parts called
valves A valve is a device or natural object that regulates, directs or controls the flow of a fluid (gases, liquids, fluidized solids, or slurries) by opening, closing, or partially obstructing various passageways. Valves are technically fittings ...
. These are joined together along one edge (the
hinge line A hinge line is an imaginary longitudinal line along the dorsal edge of the shell of a bivalve mollusk where the two valves hinge or articulate. The hinge line can easily be perceived in these images of a mussel shell and an ark shell.Invertebrat ...
) by a flexible
ligament A ligament is the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the: * Peritoneal li ...
that, usually in conjunction with interlocking "teeth" on each of the valves, forms the
hinge A hinge is a mechanical bearing that connects two solid objects, typically allowing only a limited angle of rotation between them. Two objects connected by an ideal hinge rotate relative to each other about a fixed axis of rotation: all other ...
. This arrangement allows the shell to be opened and closed without the two halves detaching. The shell is typically
bilaterally symmetrical Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
, with the hinge lying in the
sagittal plane The sagittal plane (; also known as the longitudinal plane) is an anatomical plane that divides the body into right and left sections. It is perpendicular to the transverse and coronal planes. The plane may be in the center of the body and divid ...
. Adult shell sizes of bivalves vary from fractions of a millimetre to over a metre in length, but the majority of species do not exceed 10 cm (4 in). Bivalves have long been a part of the diet of coastal and
riparian A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the terrestrial biomes of the Earth. Plant habitats and communities along the river margins and banks ar ...
human populations. Oysters were
cultured Culture () is an umbrella term which encompasses the social behavior, institutions, and norms found in human societies, as well as the knowledge, beliefs, arts, laws, customs, capabilities, and habits of the individuals in these groups.Tylor ...
in ponds by the Romans, and
mariculture Mariculture or marine farming is a specialized branch of aquaculture (which includes freshwater aquaculture) involving the cultivation of marine organisms for food and other animal products, in enclosed sections of the open ocean ( offshore mari ...
has more recently become an important source of bivalves for food. Modern knowledge of molluscan reproductive cycles has led to the development of hatcheries and new culture techniques. A better understanding of the potential
hazards A hazard is a potential source of harm. Substances, events, or circumstances can constitute hazards when their nature would allow them, even just theoretically, to cause damage to health, life, property, or any other interest of value. The probabi ...
of eating raw or undercooked
shellfish Shellfish is a colloquial and fisheries term for exoskeleton-bearing aquatic invertebrates used as food, including various species of molluscs, crustaceans, and echinoderms. Although most kinds of shellfish are harvested from saltwater envir ...
has led to improved storage and processing.
Pearl oysters A pearl is a hard, glistening object produced within the soft tissue (specifically the mantle) of a living shelled mollusk or another animal, such as fossil conulariids. Just like the shell of a mollusk, a pearl is composed of calcium carb ...
(the common name of two very different families in salt water and fresh water) are the most common source of natural
pearls A pearl is a hard, glistening object produced within the soft tissue (specifically the mantle (mollusc), mantle) of a living animal shell, shelled mollusk or another animal, such as fossil conulariids. Just like the shell of a mollusk, a pea ...
. The shells of bivalves are used in craftwork, and the manufacture of jewellery and buttons. Bivalves have also been used in the biocontrol of pollution. Bivalves appear in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
first in the early
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
more than 500 million years ago. The total number of known living
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
is about 9,200. These species are placed within 1,260 genera and 106 families. Marine bivalves (including
brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...
and
estuarine An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environment ...
species) represent about 8,000 species, combined in four subclasses and 99 families with 1,100 genera. The largest
recent The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene together ...
marine families are the
Veneridae The Veneridae or venerids, common name: Venus clams, are a very large family of minute to large, saltwater clams, marine bivalve molluscs. Over 500 living species of venerid bivalves are known, most of which are edible, and many of which are e ...
, with more than 680 species and the
Tellinidae The Tellinidae are a family of marine bivalve molluscs of the order Cardiida. Commonly known as tellins or tellens, they live fairly deep in soft sediments in shallow seas and respire using long siphons that reach up to the surface of the sedime ...
and
Lucinidae Lucinidae, common name hatchet shells, is a family of saltwater clams, marine bivalve molluscs. These bivalves are remarkable for their endosymbiosis with sulphide-oxidizing bacteria. Characteristics The members of this family have a worldwi ...
, each with over 500 species. The freshwater bivalves include seven families, the largest of which are the
Unionidae The Unionidae are a family of freshwater mussels, the largest in the order Unionida, the bivalve molluscs sometimes known as river mussels, or simply as unionids. The range of distribution for this family is world-wide. It is at its most diverse ...
, with about 700 species.


Etymology

The
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
term Bivalvia was first used by
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
in the 10th edition of his ''
Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial nomen ...
'' in 1758 to refer to animals having shells composed of two
valves A valve is a device or natural object that regulates, directs or controls the flow of a fluid (gases, liquids, fluidized solids, or slurries) by opening, closing, or partially obstructing various passageways. Valves are technically fittings ...
. More recently, the class was known as Pelecypoda, meaning "
axe An axe ( sometimes ax in American English; see spelling differences) is an implement that has been used for millennia to shape, split and cut wood, to harvest timber, as a weapon, and as a ceremonial or heraldic symbol. The axe has ma ...
-foot" (based on the shape of the foot of the animal when extended). The name "bivalve" is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''bis'', meaning "two", and ''valvae'', meaning "leaves of a door". ("Leaf" is an older word for the main, movable part of a door. We normally consider this the door itself.) Paired shells have evolved independently several times among animals that are not bivalves; other animals with paired valves include certain
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s (small
sea snail Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the ...
s in the family
Juliidae Juliidae, common name the bivalved gastropods, is a family of minute sea snails, marine gastropod mollusks or micromollusks in the superfamily Oxynooidea, an opisthobranch group. MolluscaBase eds. (2021). MolluscaBase. Juliidae E. A. Smith, 188 ...
), members of the phylum Brachiopoda and the minute crustaceans known as
ostracods Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typical ...
and conchostrachans.


Anatomy

Bivalves have
bilaterally symmetrical Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
and laterally flattened bodies, with a blade-shaped foot, vestigial head and no
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
. At the dorsal or back region of the shell is the hinge point or line, which contain the umbo and
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for food ...
and the lower, curved margin is the ventral or back region. The anterior or front of the shell is where the
byssus A byssus () is a bundle of filaments secreted by many species of bivalve mollusc that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells (Pinnidae), true mussels (Mytilid ...
(when present) and foot are located, and the posterior of the shell is where the siphons are located. With the hinge uppermost and with the anterior edge of the animal towards the viewer's left, the valve facing the viewer is the left valve and the opposing valve the right.


Mantle and shell

The shell is composed of two
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
valves held together by a ligament. The valves are made of either
calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
, as is the case in oysters, or both calcite and
aragonite Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including prec ...
. Sometimes, the aragonite forms an inner,
nacre Nacre ( , ), also known as mother of pearl, is an organicinorganic composite material produced by some molluscs as an inner shell layer; it is also the material of which pearls are composed. It is strong, resilient, and iridescent. Nacre is f ...
ous layer, as is the case in the order
Pteriida The Pteriida are an order of large and medium-sized marine bivalve mollusks. It includes five families, among them the Pteriidae (pearl oysters and winged oysters). 2010 taxonomy In 2010, a new proposed classification system for the Bivalvia ...
. In other
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, alternate layers of calcite and aragonite are laid down. The ligament and byssus, if calcified, are composed of aragonite. The outermost layer of the shell is the
periostracum The periostracum ( ) is a thin, organic coating (or "skin") that is the outermost layer of the shell of many shelled animals, including molluscs and brachiopods. Among molluscs, it is primarily seen in snails and clams, i.e. in gastropods and ...
, a thin layer composed of horny
conchiolin Conchiolins (sometimes referred to as conchins) are complex proteins which are secreted by a mollusc's outer epithelium (the mantle). These proteins are part of a matrix of organic macromolecules, mainly proteins and polysaccharides, that assem ...
. The periostracum is secreted by the outer mantle and is easily abraded. The outer surface of the valves is often sculpted, with clams often having concentric striations, scallops having radial ribs and oysters a latticework of irregular markings. In all molluscs, the mantle forms a thin
membrane A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. B ...
that covers the animal's body and extends 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 A ligament is the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the: * Peritoneal li ...
, byssus threads (where present), and
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
. The posterior mantle edge may have two elongated extensions known as siphons, through one of which water is inhaled, and the other expelled. The siphons retract into a cavity, known as the
pallial sinus The pallial sinus is an indentation or inward bending in the pallial line on the interior of a bivalve mollusk shell's valves that corresponds to the position of the siphons in those types of clams which have siphons (i.e. siphonate). The posit ...
. The shell grows larger when more material is secreted by the mantle edge, and the valves themselves thicken as more material is secreted from the general mantle surface. Calcareous matter comes from both its diet and the surrounding seawater. Concentric rings on the exterior of a valve are commonly used to age bivalves. For some groups, a more precise method for determining the age of a shell is by cutting a cross section through it and examining the incremental growth bands. The
shipworm The shipworms are marine bivalve molluscs in the family Teredinidae: a group of saltwater clams with long, soft, naked bodies. They are notorious for boring into (and commonly eventually destroying) wood that is immersed in sea water, including ...
s, in the family
Teredinidae The shipworms are marine bivalve molluscs in the family Teredinidae: a group of saltwater clams with long, soft, naked bodies. They are notorious for boring into (and commonly eventually destroying) wood that is immersed in sea water, including ...
have greatly elongated bodies, but their shell valves are much reduced and restricted to the anterior end of the body, where they function as scraping organs that permit the animal to dig tunnels through wood.


Muscles and ligaments

The main muscular system in bivalves is the posterior and anterior adductor muscles. These muscles connect the two valves and contract to close the shell. The valves are also joined dorsally by the hinge
ligament A ligament is the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the: * Peritoneal li ...
, which is an extension of the periostracum. The ligament is responsible for opening the shell, and works against the adductor muscles when the animal opens and closes. Retractor muscles connect the mantle to the edge of the shell, along a line known as the
pallial line The pallial line is a mark (a line) on the interior of each valve of the shell of a bivalve mollusk. This line shows where all of the mantle A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle m ...
. These muscles pull the mantle though the valves. In sedentary or recumbent bivalves that lie on one valve, such as the oysters and scallops, the anterior adductor muscle has been lost and the posterior muscle is positioned centrally. In species that can swim by flapping their valves, a single, central adductor muscle occurs. These muscles are composed of two types of muscle fibres, striated muscle bundles for fast actions and smooth muscle bundles for maintaining a steady pull. Paired pedal protractor and retractor muscles operate the animal's foot.


Nervous system

The sedentary habits of the bivalves have meant that in general the
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...
is less complex than in most other molluscs. The animals have no
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
; the nervous system consists of a
nerve net A nerve net consists of interconnected neurons lacking a brain or any form of cephalization. While organisms with bilateral body symmetry are normally associated with a condensation of neurons or, in more advanced forms, a central nervous syst ...
work and a series of paired
ganglia A ganglion is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system there are both sympatheti ...
. In all but the most primitive bivalves, two cerebropleural ganglia are on either side of the
oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
. The cerebral ganglia control the sensory organs, while the pleural ganglia supply nerves to the mantle cavity. The pedal ganglia, which control the foot, are at its base, and the visceral ganglia, which can be quite large in swimming bivalves, are under the posterior adductor muscle. These ganglia are both connected to the cerebropleural ganglia by
nerve fibres An axon (from Greek ἄξων ''áxōn'', axis), or nerve fiber (or nerve fibre: see spelling differences), is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action po ...
. Bivalves with long siphons may also have siphonal ganglia to control them.


Senses

The sensory organs of bivalves are largely located on the posterior mantle margins. The organs are usually
mechanoreceptor A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, ...
s or
chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
s, in some cases located on short
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
s. The
osphradium The osphradium is a pigmented chemosensory epithelium patch in the mantle cavity present in six of the eight extant classes of molluscs (it is absent in the scaphopoda and monoplacophora; among cephalopoda, only the nautilus has what appears to be ...
is a patch of sensory cells located below the posterior adductor muscle that may serve to taste the water or measure its
turbidity Turbidity is the cloudiness or haziness of a fluid caused by large numbers of individual particles that are generally invisible to the naked eye, similar to smoke in air. The measurement of turbidity is a key test of water quality. Fluids can ...
.
Statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in ''Xenoturbella''. The statocyst cons ...
s within the organism help the bivalve to sense and correct its orientation. In the order
Anomalodesmata Anomalodesmata is an superorder of saltwater clams, marine bivalve molluscs. This grouping was formerly recognised as a taxonomic subclass. It is called a superorder in the current World Register of Marine Species, despite having no orders, to ...
, the inhalant siphon is surrounded by vibration-sensitive tentacles for detecting prey. Many bivalves have no eyes, but a few members of the Arcoidea, Limopsoidea, Mytiloidea, Anomioidea, Ostreoidea, and Limoidea have simple eyes on the margin of the mantle. These consist of a pit of photosensory cells and a
lens A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements''), ...
.
Scallop Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families ...
s have more complex eyes with a lens, a two-layered
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
, and a concave mirror. All bivalves have light-sensitive cells that can detect a shadow falling over the animal.


Circulation and respiration

Bivalves have an open
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
that bathes the organs in blood (
hemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
). The
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
has three chambers: two auricles receiving blood from the gills, and a single ventricle. The ventricle is muscular and pumps hemolymph into the
aorta The aorta ( ) is the main and largest artery in the human body, originating from the left ventricle of the heart and extending down to the abdomen, where it splits into two smaller arteries (the common iliac arteries). The aorta distributes ...
, and then to the rest of the body. Some bivalves have a single aorta, but most also have a second, usually smaller, aorta serving the hind parts of the animal. The hemolymph usually lacks any respiratory pigment. In the carnivorous genus '' Poromya'', the hemolymph has red
amoebocyte An amebocyte or amoebocyte () is a mobile cell (moving like an amoeba) in the body of invertebrates including cnidaria, echinoderms, molluscs, tunicates, sponges and some chelicerates. They move by pseudopodia. Similarly to some of the white blood c ...
s containing a haemoglobin pigment. The paired gills are located posteriorly and consist of hollow tube-like filaments with thin walls for
gas exchange Gas exchange is the physical process by which gases move passively by Diffusion#Diffusion vs. bulk flow, diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liqui ...
. The respiratory demands of bivalves are low, due to their relative inactivity. Some freshwater species, when exposed to the air, can gape the shell slightly and gas exchange can take place. Oysters, including the
Pacific oyster The Pacific oyster, Japanese oyster, or Miyagi oyster (''Magallana gigas''), is an oyster native to the Pacific coast of Asia. It has become an introduced species in North America, Australia, Europe, and New Zealand. Etymology The genus ''Mag ...
(''Magallana gigas''), are recognized as having varying metabolic responses to environmental stress, with changes in respiration rate being frequently observed.


Digestive system


Modes of feeding

Most bivalves are
filter feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s, using their gills to capture particulate food such as
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
from the water. Protobranchs feed in a different way, scraping detritus from the seabed, and this may be the original mode of feeding used by all bivalves before the gills became adapted for filter feeding. These primitive bivalves hold on to the bottom with a pair of tentacles at the edge of the mouth, each of which has a single
palp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") ...
, or flap. The tentacles are covered in
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It is ...
, which traps the food, and cilia, which transport the particles back to the palps. These then sort the particles, rejecting those that are unsuitable or too large to digest, and conveying others to the mouth. In more advanced bivalves, water is drawn into the shell from the posterior
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...
surface of the animal, passes upwards through the gills, and doubles back to be expelled just above the intake. There may be two elongated, retractable siphons reaching up to the seabed, one each for the inhalant and exhalant streams of water. The gills of filter-feeding bivalves are known as ctenidia and have become highly modified to increase their ability to capture food. For example, the
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
on the gills, which originally served to remove unwanted sediment, have become adapted to capture food particles, and transport them in a steady stream of mucus to the mouth. The filaments of the gills are also much longer than those in more primitive bivalves, and are folded over to create a groove through which food can be transported. The structure of the gills varies considerably, and can serve as a useful means for classifying bivalves into groups. A few bivalves, such as the granular poromya (''Poromya granulata''), are
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
, eating much larger
prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
than the tiny microalgae consumed by other bivalves. Muscles draw water in through the inhalant siphon which is modified into a cowl-shaped organ, sucking in prey. The siphon can be retracted quickly and inverted, bringing the prey within reach of the mouth. The gut is modified so that large food particles can be digested. The unusual genus, '' Entovalva'', is
endosymbiotic An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
, being found only in the oesophagus of
sea cucumbers Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothuria ...
. It has mantle folds that completely surround its small valves. When the sea cucumber sucks in sediment, the bivalve allows the water to pass over its gills and extracts fine organic particles. To prevent itself from being swept away, it attaches itself with byssal threads to the
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman * Michel Host ...
's throat. The sea cucumber is unharmed.


Digestive tract

The digestive tract of typical bivalves consists of an
oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
,
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
, and
intestine The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
. Protobranch stomachs have a mere sac attached to them while filter-feeding bivalves have elongated rod of solidified mucus referred to as the "
crystalline style A style, sometimes referred to as a crystalline style (though there are no other biological kinds), is a rod made of glycoprotein located in the midgut of most bivalves and some gastropods The gastropods (), commonly known as snails and slugs, ...
" projected into the stomach from an associated sac. Cilia in the sac cause the style to rotate, winding in a stream of food-containing mucus from the mouth, and churning the stomach contents. This constant motion propels food particles into a sorting region at the rear of the stomach, which distributes smaller particles into the digestive glands, and heavier particles into the intestine. Waste material is consolidated in the rectum and voided as pellets into the exhalent water stream through an anal pore. Feeding and digestion are synchronized with diurnal and tidal cycles. Carnivorous bivalves generally have reduced crystalline styles and the stomach has thick, muscular walls, extensive
cuticular A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
linings and diminished sorting areas and gastric chamber sections.


Excretory system

The excretory organs of bivalves are a pair of
nephridia The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Neph ...
. Each of these consists of a long, looped, glandular tube, which opens into the
pericardium The pericardium, also called pericardial sac, is a double-walled sac containing the heart and the roots of the great vessels. It has two layers, an outer layer made of strong connective tissue (fibrous pericardium), and an inner layer made of ...
, and a
bladder The urinary bladder, or simply bladder, is a hollow organ in humans and other vertebrates that stores urine from the kidneys before disposal by urination. In humans the bladder is a distensible organ that sits on the pelvic floor. Urine enters ...
to store urine. They also have pericardial glands either line the auricles of the heart or attach to the pericardium, and serve as extra filtration organs. Metabolic waste is voided from the bladders through a
nephridiopore A nephridiopore is part of the nephridium, an excretory organ found in many organisms, such as flatworms and annelids. Polychaetes typically release their gametes into the water column using nephridiopores. Nephridia are homologous to nephrons or ...
near the front of the upper part of the mantle cavity and excreted.


Reproduction and development

The sexes are usually separate in bivalves but some
hermaphroditism In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have separ ...
is known. The
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
s and either open into the nephridia, or through a separate pore into a chamber over the gills. The ripe gonads of males and females release sperm and eggs into the
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
.
Spawning Spawn is the eggs and sperm released or deposited into water by aquatic animals. As a verb, ''to spawn'' refers to the process of releasing the eggs and sperm, and the act of both sexes is called spawning. Most aquatic animals, except for aquati ...
may take place continually or be triggered by environmental factors such as day length, water temperature, or the presence of sperm in the water. Some species are "dribble spawners", releasing gametes during protracted period that can extend for weeks. Others are mass spawners and release their gametes in batches or all at once. Fertilization is usually external. Typically, a short stage lasts a few hours or days before the eggs hatch into
trochophore A trochophore (; also spelled trocophore) is a type of free-swimming planktonic marine larva with several bands of cilia. By moving their cilia rapidly, they make a water eddy, to control their movement, and to bring their food closer, to captur ...
larvae. These later develop into
veliger A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells. Description The veliger is the characteristic larva of the gastropod, bivalve and scaphopod ...
larvae which settle on the seabed and undergo
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
into adults. In some species, such as those in the genus ''
Lasaea Lasaea or Lasaia ( grc, Λασαία) was a city on the south coast of ancient Crete, near the roadstead of the "Fair Havens" where apostle Paul landed. This place is not mentioned by any other writer, under this name but is probably the same as ...
'', females draw water containing sperm in through their inhalant siphons and fertilization takes place inside the female. These species then brood the young inside their mantle cavity, eventually releasing them into the water column as veliger larvae or as crawl-away juveniles. Most of the bivalve larvae that hatch from eggs in the water column feed on
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s or other phytoplankton. In
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions, about 25% of species are
lecithotrophic Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and mo ...
, depending on nutrients stored in the yolk of the egg where the main energy source is
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
s. The longer the period is before the larva first feeds, the larger the egg and yolk need to be. The reproductive cost of producing these energy-rich eggs is high and they are usually smaller in number. For example, the Baltic tellin (''
Macoma balthica ''Limecola balthica'', commonly called the Baltic macoma, Baltic clam or Baltic tellin,Sartori, André F. (2016)''Limecola balthica'' (Linnaeus, 1758).In: Sartori, André F. (2016). Limecola balthica (Linnaeus, 1758). In: MolluscaBase (2016). W ...
'') produces few, high-energy eggs. The larvae hatching out of these rely on the energy reserves and do not feed. After about four days, they become D-stage larvae, when they first develop hinged, D-shaped valves. These larvae have a relatively small dispersal potential before settling out. The common mussel (''
Mytilus edulis The blue mussel (''Mytilus edulis''), also known as the common mussel, is a medium-sized edible marine (ocean), marine bivalve mollusc in the family (biology), family Mytilidae, the mussels. Blue mussels are subject to commercial use and intensiv ...
'') produces 10 times as many eggs that hatch into larvae and soon need to feed to survive and grow. They can disperse more widely as they remain planktonic for a much longer time. Freshwater bivalves have different lifecycle. Sperm is drawn into a female's gills with the inhalant water and internal fertilization takes place. The eggs hatch into
glochidia The glochidium (plural glochidia) is a microscopic larval stage of some freshwater mussels, aquatic bivalve mollusks in the families Unionidae and Margaritiferidae, the river mussels and European freshwater pearl mussels. These larvae are ...
larvae that develop within the female's shell. Later they are released and attach themselves parasitically to the
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
or fins of a fish host. After several weeks they drop off their host, undergo metamorphosis and develop into adults on the substrate. Some of the species in the freshwater mussel family,
Unionidae The Unionidae are a family of freshwater mussels, the largest in the order Unionida, the bivalve molluscs sometimes known as river mussels, or simply as unionids. The range of distribution for this family is world-wide. It is at its most diverse ...
, commonly known as pocketbook mussels, have evolved an unusual reproductive strategy. The female's mantle protrudes from the shell and develops into an imitation small fish, complete with fish-like markings and false eyes. This decoy moves in the current and attracts the attention of real fish. Some fish see the decoy as prey, while others see a
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organ ...
. They approach for a closer look and the mussel releases huge numbers of larvae from its gills, dousing the inquisitive fish with its tiny, parasitic young. These glochidia larvae are drawn into the fish's gills, where they attach and trigger a tissue response that forms a small
cyst A cyst is a closed sac, having a distinct envelope and cell division, division compared with the nearby Biological tissue, tissue. Hence, it is a cluster of Cell (biology), cells that have grouped together to form a sac (like the manner in which ...
around each larva. The larvae then feed by breaking down and digesting the tissue of the fish within the cysts. After a few weeks they release themselves from the cysts and fall to the stream bed as juvenile molluscs.


Comparison with brachiopods

Brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s are shelled marine organisms that superficially resembled bivalves in that they are of similar size and have a hinged shell in two parts. However, brachiopods evolved from a very different ancestral line, and the resemblance to bivalves only arose because they occupy similar
ecological niches In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
. The differences between the two groups are due to their separate ancestral origins. Different initial structures have been adapted to solve the same problems, a case of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. In modern times, brachiopods are not as common as bivalves. Both groups have a shell consisting of two valves, but the organization of the shell is quite different in the two groups. In brachiopods, the two valves are positioned on the dorsal and ventral surfaces of the body, while in bivalves, the valves are on the left and right sides of the body, and are, in most cases, mirror images of one other. Brachiopods have a
lophophore The lophophore () is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Hyolitha, and Phoronida, which collectively constitute the protostome group Lophophorata.bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns and the
phoronid Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that filter-feed with a lophophore (a "crown" of tentacles), and build upright tubes of chitin to support and protect their soft bodies. ...
s. Some brachiopod shells are made of
calcium phosphate The term calcium phosphate refers to a family of materials and minerals containing calcium ions (Ca2+) together with inorganic phosphate anions. Some so-called calcium phosphates contain oxide and hydroxide as well. Calcium phosphates are white ...
but most are calcium carbonate in the form of the biomineral
calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
, whereas bivalve shells are always composed entirely of calcium carbonate, often in the form of the biomineral
aragonite Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including prec ...
.


Evolutionary history

The
Cambrian explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil recor ...
took place around 540 to 520 million years ago (Mya). In this geologically brief period, all the major animal phyla diverged and these included the first creatures with mineralized skeletons. Brachiopods and bivalves made their appearance at this time, and left their fossilized remains behind in the rocks. Possible early bivalves include ''
Pojetaia ''Pojetaia'' is an extinct genus of early bivalves, one of two genera in the extinct family Fordillidae. The genus is known solely from Early to Middle Cambrian fossils found in North America, Greenland, Europe, North Africa, Asia, and Australia ...
'' and ''
Fordilla ''Fordilla'' is an extinct genus of early bivalves, one of two genera in the extinct family Fordillidae. The genus is known solely from Early Cambrian fossils found in North America, Greenland, Europe, the Middle East, and Asia.Tuarangia ''Tuarangia'' is a Cambrian shelly fossil interpreted as an early bivalve, though alternative classifications have been proposed and its systematic position remains controversial.Elicki, O., & Gürsu, S. (2009). First record of ~Pojetaia runnegar ...
'', ''
Camya ''Camya'' is an extinct genus of early bivalve and is the only genus in the extinct family Camyidae. The genus is known solely from early Middle Cambrian fossils found in Europe. The genus currently contains a solitary accepted species, ''C ...
'' and ''
Arhouriella ''Arhouriella'' is arguably the oldest example of a bivalve mollusc in the fossil record. Arguably because there are older contenders to this crown, and because there is not a watertight case that it ''is'' a bivalve. The type and only species, ' ...
'' and potentially ''
Buluniella ''Pojetaia'' is an extinct genus of early bivalves, one of two genera in the extinct family Fordillidae. The genus is known solely from Early to Middle Cambrian fossils found in North America, Greenland, Europe, North Africa, Asia, and Austra ...
''. Bivalve fossils can be formed when the sediment in which the shells are buried hardens into rock. Often, the impression made by the valves remains as the fossil rather than the valves. During the
Early Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. ...
, a great increase in the diversity of bivalve species occurred, and the dysodont, heterodont, and taxodont dentitions evolved. By the Early Silurian, the gills were becoming adapted for filter feeding, and during the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
and
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
periods, siphons first appeared, which, with the newly developed muscular foot, allowed the animals to bury themselves deep in the sediment. By the middle of the
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
, around 400 Mya, the brachiopods were among the most abundant filter feeders in the ocean, and over 12,000 fossil species are recognized. By the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
250 Mya, bivalves were undergoing a huge
radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes: * ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...
of diversity. The bivalves were hard hit by this event, but re-established themselves and thrived during the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
period that followed. In contrast, the brachiopods lost 95% of their
species diversity Species diversity is the number of different species that are represented in a given community (a dataset). The effective number of species refers to the number of equally abundant species needed to obtain the same mean proportional species abundan ...
. The ability of some bivalves to burrow and thus avoid predators may have been a major factor in their success. Other new adaptations within various families allowed species to occupy previously unused evolutionary niches. These included increasing relative
buoyancy Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the p ...
in soft sediments by developing spines on the shell, gaining the ability to swim, and in a few cases, adopting predatory habits. For a long time, bivalves were thought to be better adapted to aquatic life than brachiopods were, outcompeting and relegating them to minor niches in later ages. These two taxa appeared in textbooks as an example of replacement by competition. Evidence given for this included the fact that bivalves needed less food to subsist because of their energetically efficient ligament-muscle system for opening and closing valves. All this has been broadly disproven, though; rather, the prominence of modern bivalves over brachiopods seems due to chance disparities in their response to
extinction event An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. I ...
s.


Diversity of extant bivalves

The adult maximum size of
living Living or The Living may refer to: Common meanings *Life, a condition that distinguishes organisms from inorganic objects and dead organisms ** Living species, one that is not extinct *Personal life, the course of an individual human's life * Hu ...
species of bivalve ranges from in ''
Condylonucula maya ''Condylonucula maya'' is a tiny species of saltwater clam, a marine bivalve mollusk or micromollusk in the family Nuculidae, the nut clams. This species grows to a length of about and is believed to be the smallest living bivalve.
'', a nut clam, to a length of in ''
Kuphus polythalamia ''Kuphus polythalamius'' is a species of shipworm, a marine bivalve mollusc in the family Teredinidae. Description The tube of ''Kuphus polythalamius'' is known as a crypt and is a calcareous secretion designed to enable the animal to live in it ...
'', an elongated, burrowing shipworm. However, the species generally regarded as the largest living bivalve is the giant clam ''
Tridacna gigas The giant clams are the members of the clam genus ''Tridacna'' that are the largest living bivalve mollusks. There are actually several species of "giant clams" in the genus ''Tridacna'', which are often misidentified for ''Tridacna gigas'', th ...
'', which can grow to a length of and a weight of more than 200 kg (441 lb). The largest known
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
bivalve is a species of ''
Platyceramus ''Platyceramus'' was a genus of Cretaceous bivalve molluscs belonging to the extinct inoceramid lineage. It is sometimes classified as a subgenus of ''Inoceramus''. Size The largest and best known species is ''P. platinus''. Individuals of th ...
'' whose fossils measure up to in length. In his 2010 treatise, ''Compendium of Bivalves'', Markus Huber gives the total number of living bivalve species as about 9,200 combined in 106 families. Huber states that the number of 20,000 living species, often encountered in literature, could not be verified and presents the following table to illustrate the known diversity:


Distribution

The bivalves are a highly successful class of invertebrates found in aquatic habitats throughout the world. Most are
infauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. Zoo ...
l and live buried in sediment on the seabed, or in the sediment in freshwater habitats. A large number of bivalve species are found in the
intertidal The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of Marine habitat, habitats with var ...
and
sublittoral The littoral zone or nearshore is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely inundated), to coastal areas ...
zones of the oceans. A sandy sea beach may superficially appear to be devoid of life, but often a very large number of bivalves and other invertebrates are living beneath the surface of the sand. On a large beach in
South Wales South Wales ( cy, De Cymru) is a loosely defined region of Wales bordered by England to the east and mid Wales to the north. Generally considered to include the historic counties of Glamorgan and Monmouthshire, south Wales extends westwards ...
, careful sampling produced an estimate of 1.44 million cockles (''
Cerastoderma edule The common cockle (''Cerastoderma edule'') is a species of edible saltwater clam, a marine bivalve mollusc in the family Cardiidae, the cockles. It is found in waters off Europe, from Iceland in the north, south into waters off western Africa ...
'') per acre of beach. Bivalves inhabit the tropics, as well as temperate and boreal waters. A number of species can survive and even flourish in extreme conditions. They are abundant in the Arctic, about 140 species being known from that zone. The Antarctic scallop, ''
Adamussium colbecki The Antarctic scallop (''Adamussium colbecki'') is a species of bivalve mollusc in the large family of scallops, the Pectinidae. It was thought to be the only species in the genus ''Adamussium'' until an extinct Pliocene species was described ...
'', lives under the sea ice at the other end of the globe, where the subzero temperatures mean that growth rates are very slow. The giant mussel, ''
Bathymodiolus thermophilus ''Bathymodiolus thermophilus'' is a species of large, deep water mussel, a marine bivalve mollusc in the family Mytilidae, the true mussels. The species was discovered at abyssal depths when submersible vehicles such as DSV Alvin began expl ...
'', and the giant white clam, ''
Calyptogena magnifica ''Calyptogena magnifica'' is a species of giant white clam found clustered around hydrothermal vents at abyssal depths in the Pacific Ocean. Description The systematics of the family Vesicomyidae is unclear because of the small number of spec ...
'', both live clustered around
hydrothermal vent A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspot ...
s at
abyssal The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. "Abyss" derives from the Greek word , meaning bottomless. At depths of , this zone remains in perpetual darkness. It covers 83% of the total area of the ocean an ...
depths in the Pacific Ocean. They have chemosymbiotic bacteria in their gills that oxidise
hydrogen sulphide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...
, and the molluscs absorb nutrients synthesized by these bacteria. The saddle oyster, ''
Enigmonia aenigmatica ''Enigmonia'' is a genus of saltwater clams, a marine bivalve mollusc in the family Anomiidae, the jingle shells. ''Enigmonia aenigmatica'', the mangrove jingle shell clam, is the only species in this monotypic genus. It is found living on mangr ...
'', is a marine species that could be considered
amphibious Amphibious means able to use either land or water. In particular it may refer to: Animals * Amphibian, a vertebrate animal of the class Amphibia (many of which live on land and breed in water) * Amphibious caterpillar * Amphibious fish, a fish ...
. It lives above the high tide mark in the tropical Indo-Pacific on the underside of
mangrove A mangrove is a shrub or tree that grows in coastal saline water, saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evoluti ...
leaves, on mangrove branches, and on sea walls in the splash zone. Some freshwater bivalves have very restricted ranges. For example, the Ouachita creekshell mussel, '' Villosa arkansasensis'', is known only from the streams of the
Ouachita Mountains The Ouachita Mountains (), simply referred to as the Ouachitas, are a mountain range in western Arkansas and southeastern Oklahoma. They are formed by a thick succession of highly deformed Paleozoic strata constituting the Ouachita Fold and Thru ...
in Arkansas and Oklahoma, and like several other freshwater mussel species from the southeastern US, it is in danger of extinction. In contrast, a few species of freshwater bivalves, including the golden mussel (''
Limnoperna fortunei ''Limnoperna fortunei'', the golden mussel, is a medium-sized freshwater bivalve mollusc of the family Mytilidae. The native range of the species is China, but it has accidentally been introduced to South America and several Asian countries w ...
''), are dramatically increasing their ranges. The golden mussel has spread from Southeast Asia to Argentina, where it has become an
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
. Another well-travelled freshwater bivalve, the zebra mussel (''
Dreissena polymorpha The zebra mussel (''Dreissena polymorpha'') is a small freshwater mussel. The species originates from the lakes of southern Russia and Ukraine, but has been accidentally Introduced species, introduced to numerous other areas and has become an inv ...
'') originated in southeastern Russia, and has been accidentally introduced to inland waterways in North America and Europe, where the species damages water installations and disrupts local
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s.


Behaviour

Most bivalves adopt a sedentary or even
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
lifestyle, often spending their whole lives in the area in which they first settled as juveniles. The majority of bivalves are infaunal, living under the seabed, buried in soft substrates such as sand, silt, mud, gravel, or coral fragments. Many of these live in the
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species o ...
where the sediment remains damp even when the tide is out. When buried in the sediment, burrowing bivalves are protected from the pounding of waves, desiccation, and overheating during low tide, and variations in salinity caused by rainwater. They are also out of the reach of many predators. Their general strategy is to extend their siphons to the surface for feeding and respiration during high tide, but to descend to greater depths or keep their shell tightly shut when the tide goes out. They use their muscular foot to dig into the substrate. To do this, the animal relaxes its adductor muscles and opens its shell wide to anchor itself in position while it extends its foot downwards into the substrate. Then it dilates the tip of its foot, retracts the adductor muscles to close the shell, shortens its foot and draws itself downwards. This series of actions is repeated to dig deeper. Other bivalves, such as
mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and Freshwater bivalve, freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other ...
s, attach themselves to hard surfaces using tough
byssus A byssus () is a bundle of filaments secreted by many species of bivalve mollusc that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells (Pinnidae), true mussels (Mytilid ...
threads made of
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
and
elastin Elastin is a protein that in humans is encoded by the ''ELN'' gene. Elastin is a key component of the extracellular matrix in gnathostomes (jawed vertebrates). It is highly elastic and present in connective tissue allowing many tissues in the bod ...
proteins. Some species, including the true oysters, the jewel boxes, the jingle shells, the thorny oysters and the kitten's paws, cement themselves to stones, rock or larger dead shells. In oysters the lower valve may be almost flat while the upper valve develops layer upon layer of thin horny material reinforced with calcium carbonate. Oysters sometimes occur in dense beds in the
neritic zone The neritic zone (or sublittoral zone) is the relatively shallow part of the ocean above the drop-off of the continental shelf, approximately in depth. From the point of view of marine biology it forms a relatively stable and well-illuminated ...
and, like most bivalves, are filter feeders. Bivalves filter large amounts of water to feed and breathe but they are not permanently open. They regularly shut their valves to enter a resting state, even when they are permanently submerged. In oysters, for example, their behaviour follows very strict circatidal and circadian rhythms according to the relative positions of the moon and sun. During neap tides, they exhibit much longer closing periods than during spring tides. Although many non-sessile bivalves use their muscular foot to move around, or to dig, members of the freshwater family
Sphaeriidae Sphaeriidae is a family of small to minute freshwater bivalve molluscs in the order Sphaeriida. In the US, they are commonly known as pea clams or fingernail clams. Heard, William H. 1977. Reproduction of fingernail clams (Sphaeriidae: ''Sphaeri ...
are exceptional in that these small clams climb about quite nimbly on weeds using their long and flexible foot. The European fingernail clam (''
Sphaerium corneum ''Sphaerium corneum'', also known as the European fingernailclam, is a very small freshwater clam, an aquatic bivalve mollusk in the family Sphaeriidae, the fingernail clams. Description The shell is fairly globular and can grow up to 9–13.5& ...
''), for example, climbs around on
water weed ''Elodea'' is a genus of 6 species of aquatic plants often called the waterweeds described as a genus in 1803. Classified in the frog’s-bit family (Hydrocharitaceae), ''Elodea'' is native to the Americas and is also widely used as aquarium vege ...
s at the edges of lakes and ponds; this enables the clam to find the best position for filter feeding.


Predators and defence

The thick shell and rounded shape of bivalves make them awkward for potential predators to tackle. Nevertheless, a number of different creatures include them in their diet. Many species of
demersal fish Demersal fish, also known as groundfish, live and feed on or near the bottom of seas or lakes (the demersal zone).Walrond Carl . "Coastal fish - Fish of the open sea floor"Te Ara - the Encyclopedia of New Zealand. Updated 2 March 2009 They occ ...
feed on them including the
common carp The Eurasian carp or European carp (''Cyprinus carpio''), widely known as the common carp, is a widespread freshwater fish of eutrophic waters in lakes and large rivers in Europe and Asia.Fishbase''Cyprinus carpio'' Linnaeus, 1758/ref>Arkive The ...
(''Cyprinus carpio''), which is being used in the upper Mississippi River to try to control the invasive zebra mussel (''Dreissena polymorpha''). Birds such as the
Eurasian oystercatcher The Eurasian oystercatcher (''Haematopus ostralegus'') also known as the common pied oystercatcher, or palaearctic oystercatcher, or (in Europe) just oystercatcher, is a wader in the oystercatcher bird family Haematopodidae. It is the most widesp ...
(''Haematopus ostralegus'') have specially adapted beaks which can pry open their shells. The
herring gull Herring gull is a common name for several birds in the genus ''Larus'', all formerly treated as a single species. Three species are still combined in some taxonomies: * American herring gull (''Larus smithsonianus'') - North America * European he ...
(''Larus argentatus'') sometimes drops heavy shells onto rocks in order to crack them open.
Sea otter The sea otter (''Enhydra lutris'') is a marine mammal native to the coasts of the northern and eastern North Pacific Ocean. Adult sea otters typically weigh between , making them the heaviest members of the weasel family, but among the small ...
s feed on a variety of bivalve species and have been observed to use stones balanced on their chests as anvils on which to crack open the shells. The
Pacific walrus The walrus (''Odobenus rosmarus'') is a large flippered marine mammal with a discontinuous distribution about the North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere. The walrus is the only living species in the fa ...
(''Odobenus rosmarus divergens'') is one of the main predators feeding on bivalves in Arctic waters. Shellfish have formed part of the human diet since prehistoric times, a fact evidenced by the remains of mollusc shells found in ancient middens. Examinations of these deposits in Peru has provided a means of dating long past El Niño events because of the disruption these caused to bivalve shell growth. Further changes in shell development due to environmental stress has also been suggested to cause increased mortality in oysters due to reduced shell strength. Invertebrate predators include crustaceans, starfish and octopuses. Crustaceans crack the shells with their pincers and starfish use their
water vascular system The water vascular system is a hydraulic system used by echinoderms, such as sea stars and sea urchins, for locomotion, food and waste transportation, and respiration. The system is composed of canals connecting numerous tube feet. Echinoderms mov ...
to force the valves apart and then insert part of their stomach between the valves to digest the bivalve's body. It has been found experimentally that both crabs and starfish preferred molluscs that are attached by byssus threads to ones that are cemented to the substrate. This was probably because they could manipulate the shells and open them more easily when they could tackle them from different angles. Octopuses either pull bivalves apart by force, or they bore a hole into the shell and insert a digestive fluid before sucking out the liquified contents. Certain carnivorous gastropod snails such as whelks (
Buccinidae The Buccinidae are a very large and diverse taxonomic family of large sea snails, often known as whelks or true whelks.Bouchet, P.; Gofas, S. (2010). Buccinidae. In: Bouchet, P.; Gofas, S.; Rosenberg, G. (2010) World Marine Mollusca database. A ...
) and murex snails (
Muricidae Muricidae is a large and varied taxonomic family of small to large predatory sea snails, marine gastropod mollusks, commonly known as murex snails or rock snails. With about 1,600 living species, the Muricidae represent almost 10% of the Neogas ...
) feed on bivalves by boring into their shells. A dog whelk (''
Nucella ''Nucella'', common name dog whelks or dog winkles, is a genus of small to medium-sized predatory sea snails, marine gastropod mollusks in the subfamily Ocenebrinae which is part of the large family Muricidae, the murex snails or rock snails ...
'') drills a hole with its
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
assisted by a shell-dissolving secretion. The dog whelk then inserts its extendible proboscis and sucks out the body contents of the victim, which is typically a
blue mussel The blue mussel (''Mytilus edulis''), also known as the common mussel, is a medium-sized edible marine (ocean), marine bivalve mollusc in the family (biology), family Mytilidae, the mussels. Blue mussels are subject to commercial use and intensiv ...
.
Razor shell A razor is a bladed tool primarily used in the removal of body hair through the act of shaving. Kinds of razors include straight razors, safety razors, disposable razors, and electric razors. While the razor has been in existence since before ...
s can dig themselves into the sand with great speed to escape predation. When a
Pacific razor clam The Pacific razor clam, ''Siliqua patula'', is a species of large Marine (ocean), marine bivalve mollusc in the family (biology), family Pharidae. Range Pacific razor clams can be found along the West Coast of the United States, Pacific West Co ...
(''Siliqua patula'') is laid on the surface of the beach it can bury itself completely in seven seconds and the
Atlantic jackknife clam The Atlantic jackknife clam, ''Ensis leei'', also known as the bamboo clam, American jackknife clam or razor clam, is a large edible marine bivalve mollusc found on the North American Atlantic coast, from Canada to South Carolina. The species ...
, ''Ensis directus'', can do the same within fifteen seconds. Scallops and file clams can swim by opening and closing their valves rapidly; water is ejected on either side of the hinge area and they move with the flapping valves in front. Scallops have simple eyes around the margin of the mantle and can clap their valves shut to move sharply, hinge first, to escape from danger. Cockles can use their foot to move across the seabed or leap away from threats. The foot is first extended before being contracted suddenly when it acts like a spring, projecting the animal forwards. In many bivalves that have
siphon A siphon (from grc, σίφων, síphōn, "pipe, tube", also spelled nonetymologically syphon) is any of a wide variety of devices that involve the flow of liquids through tubes. In a narrower sense, the word refers particularly to a tube in a ...
s, they can be retracted back into the safety of the shell. If the siphons inadvertently get attacked by a predator, in some cases they snap off. The animal can regenerate them later, a process that starts when the cells close to the damaged site become activated and remodel the tissue back to its pre-existing form and size. On the other hand in some cases it does not snap off. If the siphon is exposed it is the key for a predatory fish to obtain the entire body. This tactic has been observed against bivalves with an
infaunal Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. Zoo ...
lifestyle. File shells such as ''
Limaria fragilis ''Limaria fragilis'', the fragile file clam, is a species of bivalve mollusc in the family Limidae. It is found in shallow waters in the Indian and Pacific Oceans and has the ability to swim. Description The fragile file clam has a pair of hinge ...
'' can produce a noxious secretion when stressed. It has numerous tentacles which fringe its mantle and protrude some distance from the shell when it is feeding. If attacked, it sheds tentacles in a process known as
autotomy Autotomy (from the Greek language, Greek ''auto-'', "self-" and ''tome'', "severing", wikt:αὐτοτομία, αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usual ...
. The toxin released by this is distasteful and the detached tentacles continue to writhe which may also serve to distract potential predators.


Mariculture

Oysters Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not al ...
,
mussels Mussel () is the common name used for members of several families of bivalve mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,0 ...
, clams, scallops and other bivalve species are grown with food materials that occur naturally in their culture environment in the sea and lagoons.FAO State of Fisheries and Aquaculture 2012
/ref> One-third of the world's farmed food fish harvested in 2010 was achieved without the use of feed, through the production of bivalves and filter-feeding
carp Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
s. European flat oysters (''Ostrea edulis'') were first farmed by the Romans in shallow ponds and similar techniques are still in use. Seed oysters are either raised in a hatchery or harvested from the wild. Hatchery production provides some control of the broodstock but remains problematic because disease-resistant strains of this oyster have not yet been developed. Wild spats are harvested either by broadcasting empty mussel shells on the seabed or by the use of long, small-mesh nets filled with mussel shells supported on steel frames. The oyster larvae preferentially settle out on the mussel shells. Juvenile oysters are then grown on in nursery trays and are transferred to open waters when they reach in length. Many juveniles are further reared off the seabed in suspended rafts, on floating trays or cemented to ropes. Here they are largely free from bottom-dwelling predators such as starfish and crabs but more labour is required to tend them. They can be harvested by hand when they reach a suitable size. Other juveniles are laid directly on the seabed at the rate of per hectare. They grow on for about two years before being harvested by
dredging Dredging is the excavation of material from a water environment. Possible reasons for dredging include improving existing water features; reshaping land and water features to alter drainage, navigability, and commercial use; constructing da ...
. Survival rates are low at about 5%. The
Pacific oyster The Pacific oyster, Japanese oyster, or Miyagi oyster (''Magallana gigas''), is an oyster native to the Pacific coast of Asia. It has become an introduced species in North America, Australia, Europe, and New Zealand. Etymology The genus ''Mag ...
(''Crassostrea gigas'') is cultivated by similar methods but in larger volumes and in many more regions of the world. This oyster originated in Japan where it has been cultivated for many centuries. It is an estuarine species and prefers
salinities Salinity () is the saltiness or amount of salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensionless and equal ...
of 20 to 25
parts per thousand In science and engineering, the parts-per notation is a set of pseudo-units to describe small values of miscellaneous dimensionless quantities, e.g. mole fraction or mass fraction. Since these fractions are quantity-per-quantity measures, they ...
. Breeding programmes have produced improved stock that is available from hatcheries. A single female oyster can produce 50–80 million eggs in a batch so the selection of broodstock is of great importance. The larvae are grown on in tanks of static or moving water. They are fed high quality
microalgae Microalgae or microphytes are microscopic algae invisible to the naked eye. They are phytoplankton typically found in freshwater and marine systems, living in both the water column and sediment. They are unicellular species which exist indiv ...
and diatoms and grow fast. At metamorphosis the juveniles may be allowed to settle on PVC sheets or pipes, or crushed shell. In some cases, they continue their development in "upwelling culture" in large tanks of moving water rather than being allowed to settle on the bottom. They then may be transferred to transitional, nursery beds before being moved to their final rearing quarters. Culture there takes place on the bottom, in plastic trays, in mesh bags, on rafts or on long lines, either in shallow water or in the intertidal zone. The oysters are ready for harvesting in 18 to 30 months depending on the size required. Similar techniques are used in different parts of the world to cultivate other species including the
Sydney rock oyster ''Saccostrea glomerata'', is an oyster species belonging to the family Ostreidae.MolluscaBase eds. (2022). MolluscaBase. Saccostrea Dollfus & Dautzenberg, 1920. Accessed through: World Register of Marine Species at: https://www.marinespecies.org/ ...
(''Saccostrea commercialis''), the northern quahog (''Mercenaria mercenaria''), the blue mussel (''Mytilus edulis''), the
Mediterranean mussel The Mediterranean mussel (''Mytilus galloprovincialis'') is a species of bivalve, a marine (ocean), marine mollusc in the family (biology), family Mytilidae. It is an invasive species in many parts of the world, and also an object of aquaculture ...
(''Mytilus galloprovincialis''), the
New Zealand green-lipped mussel ''Perna canaliculus'', the New Zealand green-lipped mussel, also known as the New Zealand mussel, the greenshell mussel, ''kuku'', and ''kutai'', is a bivalve mollusc in the family Mytilidae (the true mussels). ''P. canaliculus'' has economic im ...
(''Perna canaliculus''), the
grooved carpet shell The grooved carpet shell, or Palourde clam, ''Ruditapes decussatus'', or ''Venerupis decussatus'', is a clam (bivalve mollusc) in the family Veneridae. It is distributed worldwide and due to its ecological and economic interest has been proposed ...
(''Ruditapes decussatus''), the Japanese carpet shell (''Venerupis philippinarum''), the pullet carpet shell (''Venerupis pullastra'') and the Yesso scallop (''Patinopecten yessoensis''). Production of bivalve molluscs by mariculture in 2010 was 12,913,199 tons, up from 8,320,724 tons in 2000. Culture of clams, cockles and ark shells more than doubled over this time period from 2,354,730 to 4,885,179 tons. Culture of mussels over the same period grew from 1,307,243 to 1,812,371 tons, of oysters from 3,610,867 to 4,488,544 tons and of scallops from 1,047,884 to 1,727,105 tons.


Use as food

Bivalves have been an important source of food for humans at least since Roman times and empty shells found in middens at archaeological sites are evidence of earlier consumption.
Oysters Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not al ...
,
scallops Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related famil ...
,
clams Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shel ...
, ark clams,
mussels Mussel () is the common name used for members of several families of bivalve mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,0 ...
and cockles are the most commonly consumed kinds of bivalve, and are eaten cooked or raw. In 1950, the year in which the
Food and Agriculture Organization The Food and Agriculture Organization of the United Nations (FAO)french: link=no, Organisation des Nations unies pour l'alimentation et l'agriculture; it, Organizzazione delle Nazioni Unite per l'Alimentazione e l'Agricoltura is an intern ...
(FAO) started making such information available, world trade in bivalve molluscs was 1,007,419 tons. By 2010, world trade in bivalves had risen to 14,616,172 tons, up from 10,293,607 tons a decade earlier. The figures included 5,554,348 (3,152,826) tons of clams, cockles and ark shells, 1,901,314 (1,568,417) tons of mussels, 4,592,529 (3,858,911) tons of oysters and 2,567,981 (1,713,453) tons of scallops. China increased its consumption 400-fold during the period 1970 to 1997. It has been known for more than a century that consumption of raw or insufficiently cooked shellfish can be associated with infectious diseases. These are caused either by bacteria naturally present in the sea such as ''
Vibrio ''Vibrio'' is a genus of Gram-negative bacteria, possessing a curved-rod (comma) shape, several species of which can cause foodborne infection, usually associated with eating undercooked seafood. Being highly salt tolerant and unable to survive ...
spp.'' or by viruses and bacteria from sewage
effluent Effluent is wastewater from sewers or industrial outfalls that flows directly into surface waters either untreated or after being treated at a facility. The term has slightly different meanings in certain contexts, and may contain various pollut ...
that sometimes contaminates coastal waters. As filter feeders, bivalves pass large quantities of water through their gills, filtering out the organic particles, including the microbial pathogens. These are retained in the animals' tissues and become concentrated in their liver-like digestive glands. Another possible source of contamination occurs when bivalves contain marine
biotoxin A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849– ...
s as a result of ingesting numerous
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s. These microalgae are not associated with sewage but occur unpredictably as
algal bloom An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term ''algae'' encompas ...
s. Large areas of a sea or lake may change colour as a result of the proliferation of millions of single-cell algae, and this condition is known as a
red tide A harmful algal bloom (HAB) (or excessive algae growth) is an algal bloom that causes negative impacts to other organisms by production of natural phycotoxin, algae-produced toxins, mechanical damage to other organisms, or by other means. HABs are ...
.


Viral and bacterial infections

In 1816 in France, a physician, J. P. A. Pasquier, described an outbreak of
typhoid Typhoid fever, also known as typhoid, is a disease caused by '' Salmonella'' serotype Typhi bacteria. Symptoms vary from mild to severe, and usually begin six to 30 days after exposure. Often there is a gradual onset of a high fever over several ...
linked to the consumption of raw oysters. The first report of this kind in the United States was in
Connecticut Connecticut () is the southernmost state in the New England region of the Northeastern United States. It is bordered by Rhode Island to the east, Massachusetts to the north, New York to the west, and Long Island Sound to the south. Its cap ...
in 1894. As sewage treatment programmes became more prevalent in the late 19th century, more outbreaks took place. This may have been because sewage was released through outlets into the sea providing more food for bivalves in estuaries and coastal habitats. A causal link between the bivalves and the illness was not easy to demonstrate because the illness might come on days or even weeks after the ingestion of the contaminated shellfish. One viral pathogen is the '' Norwalk'' virus. This is resistant to treatment with chlorine-containing chemicals and may be present in the marine environment even when
coliform bacteria Coliform bacteria are defined as either motile or non-motile Gram-negative non- spore forming Bacilli that possess β-galactosidase to produce acids and gases under their optimal growth temperature of 35-37°C. They can be aerobes or facultative ...
have been killed by the treatment of sewage. Since the 1970s, outbreaks of oyster- vectored diseases have occurs throughout the world. The mortality rate of one disease causing bacteria ''
Vibrio vulnificus ''Vibrio vulnificus'' is a species of Gram-negative, motile, curved rod-shaped (bacillus), pathogenic bacteria of the genus ''Vibrio''. Present in marine environments such as estuaries, brackish ponds, or coastal areas, ''V. vulnificus'' is relat ...
'', was high at 50%. In 1978, an oyster-associated gastrointestinal infection affecting more than 2,000 people occurred in Australia. The causative agent was found to be the ''Norwalk'' virus and the epidemic caused major economic difficulties to the oyster farming industry in the country. In 1988, an outbreak of
hepatitis A Hepatitis A is an infectious disease of the liver caused by ''Hepatovirus A'' (HAV); it is a type of viral hepatitis. Many cases have few or no symptoms, especially in the young. The time between infection and symptoms, in those who develop them ...
associated with the consumption of inadequately cooked clams (''
Anadara subcrenata ''Anadara kagoshimensis'' is an ark clam in the family Arcidae. It can be found in shallow water in temperate parts of the west Pacific Ocean and is cultivated in China, Japan, and Korea for human consumption. It is known as maohan in China and ...
'') took place in the
Shanghai Shanghai (; , , Standard Mandarin pronunciation: ) is one of the four direct-administered municipalities of the People's Republic of China (PRC). The city is located on the southern estuary of the Yangtze River, with the Huangpu River flow ...
area of China. An estimated 290,000 people were infected and there were 47 deaths. In the United States and the European Union, since the early 1990s regulations have been in place that are designed to prevent shellfish from contaminated waters entering restaurants.


Paralytic shellfish poisoning

Paralytic shellfish poisoning Paralytic shellfish poisoning (PSP) is one of the four recognized syndromes of shellfish poisoning, which share some common features and are primarily associated with bivalve mollusks (such as mussels, clams, oysters and scallops). These shellfis ...
(PSP) is primarily caused by the consumption of bivalves that have accumulated toxins by feeding on toxic dinoflagellates, single-celled protists found naturally in the sea and inland waters.
Saxitoxin Saxitoxin (STX) is a potent neurotoxin and the best-known paralytic shellfish toxin (PST). Ingestion of saxitoxin by humans, usually by consumption of shellfish contaminated by toxic algal blooms, is responsible for the illness known as paralytic ...
is the most virulent of these. In mild cases, PSP causes tingling, numbness, sickness and diarrhoea. In more severe cases, the muscles of the chest wall may be affected leading to paralysis and even death. In 1937, researchers in
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
established the connection between blooms of these phytoplankton and PSP. The biotoxin remains potent even when the shellfish are well-cooked. In the United States, there is a regulatory limit of 80 
µg In the metric system, a microgram or microgramme is a unit of mass equal to one millionth () of a gram. The unit symbol is μg according to the International System of Units (SI); the recommended symbol in the United States and United Kingdom whe ...
/g of saxitoxin equivalent in shellfish meat.


Amnesic shellfish poisoning

Amnesic shellfish poisoning Amnesic shellfish poisoning (ASP) is an illness caused by consumption of shellfish that contain the marine biotoxin called domoic acid. In mammals, including humans, domoic acid acts as a neurotoxin, causing permanent short-term memory loss, brain ...
(ASP) was first reported in eastern Canada in 1987. It is caused by the substance
domoic acid Domoic acid (DA) is a kainic acid-type neurotoxin that causes amnesic shellfish poisoning (ASP). It is produced by algae and accumulates in shellfish, sardines, and anchovies. When sea lions, otters, cetaceans, humans, and other predators eat cont ...
found in certain diatoms of the genus ''
Pseudo-nitzschia ''Pseudo-nitzschia'' is a marine planktonic diatom genus that accounts for 4.4% of pennate diatoms found worldwide. Some species are capable of producing the neurotoxin domoic acid (DA), which is responsible for the neurological disorder in huma ...
''. Bivalves can become toxic when they filter these microalgae out of the water. Domoic acid is a low-molecular weight
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
that is able to destroy brain cells causing memory loss,
gastroenteritis Gastroenteritis, also known as infectious diarrhea and gastro, is an inflammation of the gastrointestinal tract including the stomach and intestine. Symptoms may include diarrhea, vomiting, and abdominal pain. Fever, lack of energy, and dehydra ...
, long-term neurological problems or death. In an outbreak in the western United States in 1993, finfish were also implicated as vectors, and seabirds and mammals suffered neurological symptoms. In the United States and Canada, a regulatory limit of 20 µg/g of domoic acid in shellfish meat is set.


Ecosystem services

Ecosystem services Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. Th ...
provided by marine bivalves in relation to nutrient extraction from the coastal environment have gained increased attention to mitigate adverse effects of excess nutrient loading from human activities, such as agriculture and sewage discharge. These activities damage coastal ecosystems and require action from local, regional, and national environmental management. Marine bivalves filter particles like
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. Ph ...
, thereby transforming
particulate organic matter Particulate organic matter (POM) is a fraction of total organic matter operationally defined as that which does not pass through a filter pore size that typically ranges in size from 0.053 to 2 millimeters. Particulate organic carbon (POC) is ...
into bivalve tissue or larger faecal pellets that are transferred to the
benthos Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.Nutrient extraction from the coastal environment takes place through two different pathways: (i) harvest/removal of the bivalves – thereby returning nutrients back to land; or (ii) through increased
denitrification Denitrification is a microbially facilitated process where nitrate (NO3−) is reduced and ultimately produces molecular nitrogen (N2) through a series of intermediate gaseous nitrogen oxide products. Facultative anaerobic bacteria perform denitr ...
in proximity to dense bivalve aggregations, leading to loss of nitrogen to the atmosphere. Active use of marine bivalves for nutrient extraction may include a number of secondary effects on the ecosystem, such as filtration of
particulate Particulates – also known as atmospheric aerosol particles, atmospheric particulate matter, particulate matter (PM) or suspended particulate matter (SPM) – are microscopic particles of solid or liquid matter suspended in the air. The ter ...
material. This leads to partial transformation of particulate-bound nutrients into dissolved nutrients via bivalve excretion or enhanced mineralization of faecal material. When they live in polluted waters, bivalve molluscs have a tendency to accumulate substances such as
heavy metals upright=1.2, Crystals of osmium, a heavy metal nearly twice as dense as lead">lead.html" ;"title="osmium, a heavy metal nearly twice as dense as lead">osmium, a heavy metal nearly twice as dense as lead Heavy metals are generally defined as ...
and
persistent organic pollutants Persistent organic pollutants (POPs), sometimes known as "forever chemicals", are organic compounds that are resistant to environmental degradation through chemical, biological, and photolytic processes. They are toxic chemicals that adversely ...
in their tissues. This is because they ingest the chemicals as they feed but their enzyme systems are not capable of metabolising them and as a result, the levels build up. This may be a health hazard for the molluscs themselves, and is one for humans who eat them. It also has certain advantages in that bivalves can be used in
monitoring Monitoring may refer to: Science and technology Biology and healthcare * Monitoring (medicine), the observation of a disease, condition or one or several medical parameters over time * Baby monitoring * Biomonitoring, of toxic chemical compounds, ...
the presence and quantity of pollutants in their environment. There are limitations to the use of bivalves as
bioindicator A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
s. The level of pollutants found in the tissues varies with species, age, size, time of year and other factors. The quantities of pollutants in the water may vary and the molluscs may reflect past rather than present values. In a study near
Vladivostok Vladivostok ( rus, Владивосто́к, a=Владивосток.ogg, p=vɫədʲɪvɐˈstok) is the largest city and the administrative center of Primorsky Krai, Russia. The city is located around the Zolotoy Rog, Golden Horn Bay on the Sea ...
it was found that the level of pollutants in the bivalve tissues did not always reflect the high levels in the surrounding sediment in such places as harbours. The reason for this was thought to be that the bivalves in these locations did not need to filter so much water as elsewhere because of the water's high nutritional content. A study of nine different bivalves with widespread distributions in tropical marine waters concluded that the mussel, ''
Trichomya hirsuta ''Trichomya'' is a monotypic genus of marine bivalve molluscs in the family Mytilidae, the mussels. The only species is ''Trichomya hirsuta'' which is endemic to southern and eastern Australia. Its common names include the hairy mussel, the gree ...
'', most nearly reflected in its tissues the level of heavy metals (Pb, Cd, Cu, Zn, Co, Ni, and Ag) in its environment. In this species there was a linear relationship between the sedimentary levels and the tissue concentration of all the metals except zinc. In the
Persian Gulf The Persian Gulf ( fa, خلیج فارس, translit=xalij-e fârs, lit=Gulf of Persis, Fars, ), sometimes called the ( ar, اَلْخَلِيْجُ ٱلْعَرَبِيُّ, Al-Khalīj al-ˁArabī), is a Mediterranean sea (oceanography), me ...
, the Atlantic pearl-oyster ('' Pinctada radiata'') is considered to be a useful bioindicator of heavy metals. Crushed shells, available as a by-product of the seafood canning industry, can be used to remove pollutants from water. It has been found that, as long as the water is maintained at an alkaline pH, crushed shells will remove cadmium, lead and other heavy metals from contaminated waters by swapping the calcium in their constituent aragonite for the heavy metal, and retaining these pollutants in a solid form. The rock oyster (''
Saccostrea cucullata ''Saccostrea cucullata'', the hooded oyster or Natal rock oyster, is a species of rock oyster found mainly in the Indo-Pacific Ocean. It was first described by the Czech mineralogist, metallurgist, and malacologist Ignaz von Born in 1778. De ...
'') has been shown to reduce the levels of
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
and cadmium in contaminated waters in the Persian Gulf. The live animals acted as biofilters, selectively removing these metals, and the dead shells also had the ability to reduce their concentration.


Other uses

Conchology Conchology () is the study of mollusc shells. Conchology is one aspect of malacology, the study of molluscs; however, malacology is the study of molluscs as whole organisms, whereas conchology is confined to the study of their shells. It includ ...
is the scientific study of mollusc shells, but the term conchologist is also sometimes used to describe a collector of shells. Many people pick up shells on the beach or purchase them and display them in their homes. There are many private and public collections of mollusc shells, but the largest one in the world is at the
Smithsonian Institution The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums and education and research centers, the largest such complex in the world, created by the U.S. government "for the increase and diffusion of knowledge". Founded ...
, which houses in excess of 20 million specimens. Shells are used decoratively in many ways. They can be pressed into concrete or plaster to make decorative paths, steps or walls and can be used to embellish picture frames, mirrors or other craft items. They can be stacked up and glued together to make ornaments. They can be pierced and threaded onto necklaces or made into other forms of jewellery. Shells have had various uses in the past as body decorations, utensils, scrapers and cutting implements. Carefully cut and shaped shell tools dating back 32,000 years have been found in a cave in Indonesia. In this region, shell technology may have been developed in preference to the use of stone or bone implements, perhaps because of the scarcity of suitable rock materials. The
indigenous peoples of the Americas The Indigenous peoples of the Americas are the inhabitants of the Americas before the arrival of the European settlers in the 15th century, and the ethnic groups who now identify themselves with those peoples. Many Indigenous peoples of the A ...
living near the east coast used pieces of shell as
wampum Wampum is a traditional shell bead of the Eastern Woodlands tribes of Native Americans. It includes white shell beads hand-fashioned from the North Atlantic channeled whelk shell and white and purple beads made from the quahog or Western Nort ...
. The
channeled whelk ''Busycotypus canaliculatus'', commonly known as the channeled whelk, is a very large predatory sea snail, a marine prosobranch gastropod, a busycon whelk, belonging to the family Busyconidae.Fraussen, K.; Rosenberg, G. (2012). Busycotypus canal ...
(''Busycotypus canaliculatus'') and the
quahog The hard clam (''Mercenaria mercenaria''), also known as the round clam, hard-shell (or hard-shelled) clam, or the quahog, is an edible marine bivalve mollusk that is native to the eastern shores of North America and Central America from Prince ...
(''Mercenaria mercenaria'') were used to make white and purple traditional patterns. The shells were cut, rolled, polished and drilled before being strung together and woven into belts. These were used for personal, social and ceremonial purposes and also, at a later date, for currency. The
Winnebago Tribe The Winnebago Tribe of Nebraska ( win, Nįįšoc Hoocąk) is one of two federally recognized tribes of Ho-Chunk Native Americans. The other is the Ho-Chunk Nation of Wisconsin. Tribe members often refer to themselves as ''Hochungra'' – "People ...
from Wisconsin had numerous uses for freshwater mussels including using them as spoons, cups, ladles and utensils. They notched them to provide knives, graters and saws. They carved them into fish hooks and lures. They incorporated powdered shell into clay to
temper Temper, tempered or tempering may refer to: Heat treatment * Tempering (metallurgy), a heat treatment technique to increase the toughness of iron-based alloys ** Temper mill, a steel processing line * Tempering (spices), a cooking technique where ...
their pottery vessels. They used them as scrapers for removing flesh from hides and for separating the scalps of their victims. They used shells as scoops for gouging out fired logs when building canoes and they drilled holes in them and fitted wooden handles for tilling the ground.
Button A button is a fastener that joins two pieces of fabric together by slipping through a loop or by sliding through a buttonhole. In modern clothing and fashion design, buttons are commonly made of plastic but also may be made of metal, wood, ...
s have traditionally been made from a variety of freshwater and marine shells. At first they were used decoratively rather than as fasteners and the earliest known example dates back five thousand years and was found at
Mohenjo-daro Mohenjo-daro (; sd, موئن جو دڙو'', ''meaning 'Mound of the Dead Men';Indus Valley The Indus ( ) is a transboundary river of Asia and a trans-Himalayan river of South and Central Asia. The river rises in mountain springs northeast of Mount Kailash in Western Tibet, flows northwest through the disputed region of Kashmir, ...
.
Sea silk Sea silk is an extremely fine, rare, and valuable fabric that is made from the long silky filaments or byssus secreted by a gland in the foot of pen shells (in particular ''Pinna nobilis''). The byssus is used by the clam to attach itself to t ...
is a fine fabric woven from the
byssus A byssus () is a bundle of filaments secreted by many species of bivalve mollusc that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells (Pinnidae), true mussels (Mytilid ...
threads of bivalves, particularly the pen shell (''
Pinna nobilis ''Pinna nobilis'', common name the noble pen shell or fan mussel, is a large species of Mediterranean clam, a marine bivalve mollusc in the family Pinnidae, the pen shells. *''Pinna gigas'' Chemnitz It reaches up to of shell length.Zavodnik, D ...
''). It used to be produced in the Mediterranean region where these shells are
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
. It was an expensive fabric and
overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in th ...
has much reduced populations of the pen shell. Crushed shells are added as a calcareous supplement to the diet of laying poultry. Oyster shell and cockle shell are often used for this purpose and are obtained as a by-product from other industries.


Pearls and mother-of-pearl

Mother-of-pearl or nacre is the naturally occurring lustrous layer that lines some mollusc shells. It is used to make pearl buttons and in artisan craftwork to make organic jewellery. It has traditionally been inlaid into furniture and boxes, particularly in China. It has been used to decorate musical instruments, watches, pistols, fans and other products. The import and export of goods made with nacre are controlled in many countries under the International Convention of Trade in Endangered Species of Wild Fauna and Flora. A
pearl A pearl is a hard, glistening object produced within the soft tissue (specifically the mantle) of a living shelled mollusk or another animal, such as fossil conulariids. Just like the shell of a mollusk, a pearl is composed of calcium carb ...
is created in the mantle of a mollusk when an irritant particle is surrounded by layers of nacre. Although most bivalves can create pearls,
oysters Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not al ...
in the family
Pteriidae Pteriidae, also called the feather oysters, is a family of medium-sized to large saltwater clams. They are pearl oysters, marine bivalve molluscs in the order Pteriida. Some of the species in this family are important economically as the sou ...
and freshwater mussels in the families
Unionidae The Unionidae are a family of freshwater mussels, the largest in the order Unionida, the bivalve molluscs sometimes known as river mussels, or simply as unionids. The range of distribution for this family is world-wide. It is at its most diverse ...
and
Margaritiferidae Margaritiferidae is a family of medium-sized freshwater mussels, aquatic bivalve molluscs in the order Unionida. Text may have been copied from this source, which is available under Attribution 4.0 International (CC BY 4.0)licence. It is the m ...
are the main source of commercially available pearls because the calcareous concretions produced by most other species have no lustre. Finding pearls inside oysters is a very chancy business as hundreds of shells may need to be pried open before a single pearl can be found. Most pearls are now obtained from cultured shells where an irritant substance has been purposefully introduced to induce the formation of a pearl. A "mabe" (irregular) pearl can be grown by the insertion of an implant, usually made of plastic, under a flap of the mantle and next to the mother-of-pearl interior of the shell. A more difficult procedure is the grafting of a piece of oyster mantle into the gonad of an adult specimen together with the insertion of a shell bead nucleus. This produces a superior, spherical pearl. The animal can be opened to extract the pearl after about two years and reseeded so that it produces another pearl. Pearl oyster farming and pearl culture is an important industry in Japan and many other countries bordering the Indian and Pacific Oceans.


Symbolism

The scallop is the symbol of
St James Saint James or St. James may refer to: People Saints *James, brother of Jesus (died 62 or 69), also known as James the Just *James the Great (died 44), Apostle, also known as James, son of Zebedee, or Saint James the Greater **Saint James Matamoro ...
and is called ''
Coquille Saint-Jacques Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related famil ...
'' in French. It is an emblem carried by pilgrims on their
way Way may refer to: Paths * a road, route, path or pathway, including long-distance paths. * a straight rail or track on a machine tool, (such as that on the bed of a lathe) on which part of the machine slides * Ways, large slipway in shipbuildin ...
to the shrine of
Santiago de Compostela Santiago de Compostela is the capital of the autonomous community of Galicia, in northwestern Spain. The city has its origin in the shrine of Saint James the Great, now the Cathedral of Santiago de Compostela, as the destination of the Way of St ...
in Galicia. The shell became associated with the pilgrimage and came to be used as a symbol showing hostelries along the route and later as a sign of hospitality, food and lodging elsewhere.
Roman Roman or Romans most often refers to: *Rome, the capital city of Italy *Ancient Rome, Roman civilization from 8th century BC to 5th century AD *Roman people, the people of ancient Rome *''Epistle to the Romans'', shortened to ''Romans'', a letter ...
myth has it that
Venus Venus is the second planet from the Sun. It is sometimes called Earth's "sister" or "twin" planet as it is almost as large and has a similar composition. As an interior planet to Earth, Venus (like Mercury) appears in Earth's sky never fa ...
, the goddess of love, was born in the sea and emerged accompanied by fish and dolphins, with
Botticelli Alessandro di Mariano di Vanni Filipepi ( – May 17, 1510), known as Sandro Botticelli (, ), was an Italian painter of the Early Renaissance. Botticelli's posthumous reputation suffered until the late 19th century, when he was rediscovered ...
depicting her as arriving in a scallop shell. The Romans revered her and erected shrines in her honour in their gardens, praying to her to provide water and verdant growth. From this, the scallop and other bivalve shells came to be used as a symbol for fertility. Its depiction is used in architecture, furniture and fabric design and it is the logo of
Royal Dutch Shell Shell plc is a British multinational oil and gas company headquartered in London, England. Shell is a public limited company with a primary listing on the London Stock Exchange (LSE) and secondary listings on Euronext Amsterdam and the New Yo ...
, the global oil and gas company.


Bivalvian taxonomies

For the past two centuries no consensus has existed on bivalve
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
from the many classifications developed. In earlier taxonomic systems, experts used a single characteristic feature for their classifications, choosing among shell morphology, hinge type or gill type. Conflicting naming schemes proliferated due to these taxonomies based on single organ systems. One of the most widely accepted systems was that put forward by
Norman D. Newell Norman Dennis Newell (January 27, 1909 – April 18, 2005) was professor of geology at Columbia University, and chairman and curator of invertebrate paleontology at the American Museum of Natural History in New York City. Personal life Newell was ...
in Part N of the ''
Treatise on Invertebrate Paleontology The ''Treatise on Invertebrate Paleontology'' (or ''TIP'') published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and co ...
'', which employed a classification system based on general shell shape, microstructures and hinge configuration. Because features such as hinge morphology, dentition, mineralogy, shell morphology and shell composition change slowly over time, these characteristics can be used to define major taxonomic groups. Since the year 2000, taxonomic studies using
cladistical Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
analyses of multiple organ systems, shell morphology (including fossil species) and modern
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
have resulted in the drawing up of what experts believe is a more accurate phylogeny of the Bivalvia. Based upon these studies, a new proposed classification system for the Bivalvia was published in 2010 by Bieler, Carter & Coan. In 2012, this new system was adopted by the
World Register of Marine Species The World Register of Marine Species (WoRMS) is a taxonomic database that aims to provide an authoritative and comprehensive list of names of marine organisms. Content The content of the registry is edited and maintained by scientific specialist ...
(WoRMS) for the classification of the Bivalvia. Some experts still maintain that Anomalodesmacea should be considered a separate subclass, whereas the new system treats it as the order Anomalodesmata, within the subclass
Heterodonta Heteroconchia is a taxonomic infraclass of saltwater clams, marine bivalve molluscs, belonging to the subclass Autobranchia This infraclass includes the edible clams, the cockles and the Venus clams. Description These bivalves are distinguis ...
. Molecular phylogenetic work continues, further clarifying which Bivalvia are most closely related and thus refining the classification.


Practical taxonomy of R.C. Moore

R.C. Moore, in Moore, Lalicker, and Fischer, 1952, ''Invertebrate Fossils'', gives a practical and useful classification of pelecypods (Bivalvia) even if somewhat antiquated, based on shell structure, gill type, and hinge teeth configuration. Subclasses and orders given are: : Subclass: Prionodesmacea :: Order :: Paleoconcha :: Taxodonta: Many teeth (e.g. order
Nuculida Nuculida is an order of small saltwater clams, marine bivalve mollusks. This order belongs to the subclass Protobranchia Protobranchia is a subclass of bivalve molluscs. It contains the extant orders Nuculanida, Nuculida, and Solemyida. ...
) :: Schizodonta: Big bifurcating teeth (e.g. ''
Trigonia ''Trigonia'' is an extinct genus of saltwater clams, fossil marine bivalve mollusk in the family Trigoniidae. The fossil range of the genus spans the Paleozoic, Mesozoic and Paleocene of the Cenozoic, from 298 to 56 Ma. Description The gen ...
'' spp.) :: Isodonta: Equal teeth (e.g. ''
Spondylus ''Spondylus'' is a genus of bivalve molluscs, the only genus in the family (biology), family Spondylidae.MolluscaBase (2019). MolluscaBase. Spondylus Linnaeus, 1758. Accessed through: World Register of Marine Species at: http://www.marinespecies ...
'' spp.) :: Dysodonta: Absent teeth and ligaments joins the valves. : Subclass: Teleodesmacea :: Order ::
Heterodonta Heteroconchia is a taxonomic infraclass of saltwater clams, marine bivalve molluscs, belonging to the subclass Autobranchia This infraclass includes the edible clams, the cockles and the Venus clams. Description These bivalves are distinguis ...
: Different teeth (e.g. family
Cardiidae A cockle is an edible marine bivalve mollusc. Although many small edible bivalves are loosely called cockles, true cockles are species in the family Cardiidae. MolluscaBase eds. (2022). MolluscaBase. Cardiidae Lamarck, 1809. Accessed through: W ...
). Lower_Ordovician –_Holocene.html" "title="Lower_Ordovician.html" ;"title="Lower Ordovician">Lower Ordovician – Holocene">Recent The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene together ...
] :: Pachydonta: Large, different, deformed teeth (e.g. rudist spp.). [ Late Jurassic – Upper Cretaceous] :: Desmodonta: Hinge-teeth absent or irregular with ligaments (e.g. family Anatinidae). Prionodesmacea have a prismatic and nacreous shell structure, separated mantle lobes, poorly developed siphons, and hinge teeth that are lacking or unspecialized. Gills range from protobranch to eulamellibranch. Teleodesmacea on the other hand have a porcelanous and partly nacreous shell structure; Mantle lobes that are generally connected, well developed siphons, and specialized hinge teeth. In most, gills are eulamellibranch.


1935 taxonomy

In his 1935 work ''Handbuch der systematischen Weichtierkunde'' (Handbook of Systematic Malacology), Johannes Thiele introduced a mollusc taxonomy based upon the 1909 work by Cossmann and Peyrot. Thiele's system divided the bivalves into three orders. ''Taxodonta'' consisted of forms that had taxodont dentition, with a series of small parallel teeth perpendicular to the hinge line. ''Anisomyaria'' consisted of forms that had either a single adductor muscle or one adductor muscle much smaller than the other. ''Eulamellibranchiata'' consisted of forms with ctenidial gills. The Eulamellibranchiata was further divided into four suborders: ''Schizodonta'', ''Heterodonta'', ''Adapedonta'' and ''Anomalodesmata''.


Taxonomy based upon hinge tooth morphology

The systematic layout presented here follows Newell's 1965 classification based on hinge tooth
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
(all taxa marked † are extinct) : The
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...
of the subclass Anomalodesmata is disputed. The standard view now is that it resides within the subclass Heterodonta.


Taxonomy based upon gill morphology

An alternative systematic scheme exists using gill morphology. This distinguishes between Protobranchia, Filibranchia and Eulamellibranchia. The first corresponds to Newell's Palaeotaxodonta and Cryptodonta, the second to his Pteriomorphia, with the last corresponding to all other groups. In addition, Franc separated the Septibranchia from his eulamellibranchs because of the morphological differences between them. The septibranchs belong to the superfamily
Poromyoidea Poromyoidea is a superfamily of molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are reco ...
and are carnivorous, having a muscular septum instead of filamentous gills.


2010 taxonomy

In May 2010, a new taxonomy of the Bivalvia was published in the journal ''
Malacologia ''Malacologia'' is a peer-reviewed scientific journal in the field of malacology, the study of mollusks. The journal publishes articles in the fields of molluscan systematics, ecology, population ecology, genetics, molecular genetics, evolution, an ...
''. In compiling this, the authors used a variety of phylogenetic information including molecular analysis, anatomical analysis, shell morphology and shell microstructure as well as bio-geographic, paleobiogeographic and
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithostrati ...
information. In this classification 324 families are recognized as valid, 214 of which are known exclusively from fossils and 110 of which occur in the recent past, with or without a fossil record. This classification has since been adopted by WoRMS. Proposed classification of Class Bivalvia (under the redaction of Rüdiger Bieler, Joseph G. Carter and Eugene V. Coan) (all taxa marked † are extinct) : Grade Euprotobranchia * Order
Fordillida Fordilloidea is an extinct superfamily of early bivalves containing two described families, Fordillidae and Camyidae and the only superfamily in the order Fordillida. The superfamily is known from fossils of early to middle Cambrian age found ...
:: 2 families (2†) * Order
Tuarangiida ''Tuarangia'' is a Cambrian shelly fossil interpreted as an early bivalve, though alternative classifications have been proposed and its systematic position remains controversial.Elicki, O., & Gürsu, S. (2009). First record of ~Pojetaia runnegar ...
:: 1 family (1†) Subclass
Heterodonta Heteroconchia is a taxonomic infraclass of saltwater clams, marine bivalve molluscs, belonging to the subclass Autobranchia This infraclass includes the edible clams, the cockles and the Venus clams. Description These bivalves are distinguis ...
Infraclass
Archiheterodonta Carditida is an order of marine bivalve clams. Superfamilies and families According to the World Register of Marine Species: * Family: † Archaeocardiidae *Superfamily: Carditoidea ** Family: Carditidae ** Family: Condylocardiidae *Sup ...
* Order
Carditida Carditida is an order of marine bivalve clams. Superfamilies and families According to the World Register of Marine Species: * Family: † Archaeocardiidae *Superfamily: Carditoidea ** Family: Carditidae ** Family: Condylocardiidae *Superf ...
:: 4 families Infraclass
Euheterodonta Euheterodonta is an subterclass of Mollusca in the class Bivalvia. Families Infraclass Euheterodonta *Superorder Anomalodesmata Dall, 1889 **Superfamily Clavagelloidea d'Orbigny, 1844 *** Clavagellidae d'Orbigny, 1844 *** Penicillidae Gray, 185 ...
* Unassigned Euheterodonta :: 4 families * Order
Pholadomyida Anomalodesmata is an superorder of saltwater clams, marine bivalve molluscs. This grouping was formerly recognised as a taxonomic subclass. It is called a superorder in the current World Register of Marine Species, despite having no orders, to ...
(=
Anomalodesmata Anomalodesmata is an superorder of saltwater clams, marine bivalve molluscs. This grouping was formerly recognised as a taxonomic subclass. It is called a superorder in the current World Register of Marine Species, despite having no orders, to ...
) :: 16 families * Order
Myida Myida (formerly Myoida) is an order of saltwater and freshwater clams, marine and freshwater bivalve molluscs in the subclass Heterodonta. The order includes such bivalves as soft-shell clams, geoducks and shipworms. Description They are burrowi ...
:: 4 families * Order
Lucinida Lucinida (formerly Lucinoida) is a taxonomic order of saltwater clams, marine bivalve molluscs. Families In 2010, a new proposed classification system for the Bivalvia was published in by Bieler, Carter & Coan, revising the classification o ...
:: 2 families * Order
Venerida Venerida (formerly Veneroida) is an order of mostly saltwater but also some freshwater bivalve molluscs. This order includes many familiar groups such as many clams that are valued for food and a number of freshwater bivalves. Since the 2000s, ...
:: 30 families Subclass
Palaeoheterodonta Palaeoheterodonta is a subclass of bivalve molluscs. It contains the extant orders Unionida (freshwater mussels) and Trigoniida. They are distinguished by having the two halves of the shell be of equal size and shape, but by having the hinge t ...
* Order Trigoniida :: 16 families (15†) * Order
Unionida Unionida is a monophyletic order of freshwater mussels, aquatic bivalve molluscs.Bieler R., Carter J.G. & Coan E.V. (2010). ''Classification of Bivalve families''. pp. 113–133, in: Bouchet P. & Rocroi J.-P. (2010), ''Nomenclator of Bivalve F ...
:: 15 families (8†) Subclass
Protobranchia Protobranchia is a subclass of bivalve molluscs. It contains the extant orders Nuculanida, Nuculida, and Solemyida. These are deep water clams of a small and primitive order with a taxodont hinge (composed of many similar, small teeth), gener ...
* Order
Nuculanida Nuculanida is an order of very small saltwater clams, marine bivalve molluscs in the subclass Protobranchia. Description These bivalves are distinguished by the presence of relatively primitive, "protobranchiate" gills. There are a row of s ...
:: 8 families * Order
Nuculida Nuculida is an order of small saltwater clams, marine bivalve mollusks. This order belongs to the subclass Protobranchia Protobranchia is a subclass of bivalve molluscs. It contains the extant orders Nuculanida, Nuculida, and Solemyida. ...
:: 3 families (1†) * Order
Solemyida Solemyida is an order of bivalve mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are re ...
:: 2 families Subclass
Pteriomorphia The Pteriomorphia comprise a subclass of saltwater clams, marine bivalve molluscs. It contains several major orders, including the Arcida, Ostreida, Pectinida, Limida, Mytilida, and Pteriida. It also contains some extinct and probably basal ...
* Order
Arcida The Arcida is an extant order of bivalve molluscs. This order dates back to the lower Ordovician period. They are distinguished from related groups, such as the mussels, by having a straight hinge to the shells, and the adductor muscles being ...
:: 7 families Infraclass Eupteriomorphia * Order
Ostreida The order Ostreida includes the true oysters. One superfamily (Ostreoidea) and two extant families are recognised within it. The two families are Ostreidae, the true oysters, and Gryphaeidae The Gryphaeidae, common name the foam oysters or ...
:: 2 families * Suborder
Pectinida Pectinida is a taxonomic order of large and medium-sized saltwater clams, marine bivalve molluscs, commonly known as scallops and their allies. It is believed that they began evolutionarily in the late Middle Ordovician epoch; many species, of ...
:: 7 families * Suborder
Limida The Limidae or file shells are members of the only family of bivalve molluscs in the order Limida. The family includes 130 living species, assigned to 10 genera. Widely distributed in all seas from shallow to deep waters, the species are usually ...
:: 1 family * Suborder
Mytilida Mytilida is an order of marine bivalve molluscs, commonly known as true mussels. There is one extant superfamily, the Mytiloidea, with a single extant family, the Mytilidae. Species in the order Mytilida are found worldwide, but they are more ...
:: 1 family * Suborder
Pteriida The Pteriida are an order of large and medium-sized marine bivalve mollusks. It includes five families, among them the Pteriidae (pearl oysters and winged oysters). 2010 taxonomy In 2010, a new proposed classification system for the Bivalvia ...
:: 4 families


References


Further reading

* * * {{Authority control Extant Cambrian first appearances Mollusc classes Taxa named by Carl Linnaeus