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Bivalvia

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Bivalvia
Temporal range: Early Cambrian – recent[1][2]
Ernst Haeckel's "Acephala"
"Acephala", from Ernst Haeckel's Kunstformen der Natur (1904)
Scientific classification e
Kingdom: Animalia
Phylum: Mollusca
Class: Bivalvia
Linnaeus, 1758
Subclasses

And see text

Empty shell of the giant clam  (Tridacna gigas)
Empty shell of the giant clam
(Tridacna gigas)
Empty shells of the sword razor  (Ensis ensis)
Empty shells of the sword razor
(Ensis ensis)

Bivalvia (/bˈvælviə/), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bivalves have no head and they lack some usual molluscan organs, like the radula and the odontophore. The class includes the clams, oysters, cockles, mussels, scallops, and numerous other families that live in saltwater, as well as a number of families that live in freshwater. The majority are filter feeders. The gills have evolved into ctenidia, specialised organs for feeding and breathing. Most bivalves bury themselves in sediment, where they are relatively safe from predation. Others lie on the sea floor or attach themselves to rocks or other hard surfaces. Some bivalves, such as the scallops and file shells, can swim. The shipworms bore into wood, clay, or stone and live inside these substances.

The shell of a bivalve is composed of calcium carbonate, and consists of two, usually similar, parts called valves. These are joined together along one edge (the hinge line) by a flexible ligament that, usually in conjunction with interlocking "teeth" on each of the valves, forms the hinge. This arrangement allows the shell to be opened and closed without the two halves detaching. The shell is typically bilaterally symmetrical, with the hinge lying in the sagittal plane. 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 human populations. Oysters were cultured in ponds by the Romans, and mariculture 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 of eating raw or undercooked shellfish has led to improved storage and processing. Pearl oysters (the common name of two very different families in salt water and fresh water) are the most common source of natural pearls. 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 first in the early Cambrian more than 500 million years ago. The total number of known living species is about 9,200. These species are placed within 1,260 genera and 106 families. Marine bivalves (including brackish water and estuarine species) represent about 8,000 species, combined in four subclasses and 99 families with 1,100 genera. The largest recent marine families are the Veneridae, with more than 680 species and the Tellinidae and Lucinidae, each with over 500 species. The freshwater bivalves include seven families, the largest of which are the Unionidae, with about 700 species.

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Class (biology)

Class (biology)

In biological classification, class is a taxonomic rank, as well as a taxonomic unit, a taxon, in that rank. It is a group of related taxonomic orders. Other well-known ranks in descending order of size are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order.

Head

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 not have a head, but many bilaterally symmetric forms do, regardless of size.

Clam

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 shells of equal size connected by two adductor muscles and have a powerful burrowing foot. They live in both freshwater and marine environments; in salt water they prefer to burrow down into the mud and the turbidity of the water required varies with species and location; the greatest diversity of these is in North America.

Cockle (bivalve)

Cockle (bivalve)

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.

Family (biology)

Family (biology)

Family is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between order and genus. A family may be divided into subfamilies, which are intermediate ranks between the ranks of family and genus. The official family names are Latin in origin; however, popular names are often used: for example, walnut trees and hickory trees belong to the family Juglandaceae, but that family is commonly referred to as the "walnut family".

Filter feeder

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 feeding are clams, krill, sponges, baleen whales, and many fish. Some birds, such as flamingos and certain species of duck, are also filter feeders. Filter feeders can play an important role in clarifying water, and are therefore considered ecosystem engineers. They are also important in bioaccumulation and, as a result, as indicator organisms.

Gill

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 kept moist. The microscopic structure of a gill presents a large surface area to the external environment. Branchia is the zoologists' name for gills.

Ctenidium (mollusc)

Ctenidium (mollusc)

A ctenidium is a respiratory organ or gill which is found in many molluscs. This structure exists in bivalves, cephalopods, Polyplacophorans (chitons), and in aquatic gastropods such as freshwater snails and marine snails. Some aquatic gastropods possess one ctenidium known as monopectinate and others have a pair of ctenidia known as bipectinate.

Calcium carbonate

Calcium carbonate

Calcium carbonate is a chemical compound with the chemical formula CaCO3. It is a common substance found in rocks as the minerals calcite and aragonite and is the main component of eggshells, gastropod shells, shellfish skeletons and pearls. Things containing much calcium carbonate or resembling it are described as calcareous. Calcium carbonate is the active ingredient in agricultural lime and is created when calcium ions in hard water react with carbonate ions to create limescale. It has medical use as a calcium supplement or as an antacid, but excessive consumption can be hazardous and cause hypercalcemia and digestive issues.

Hinge

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 translations or rotations being prevented, and thus a hinge has one degree of freedom. Hinges may be made of flexible material or of moving components. In biology, many joints function as hinges, like the elbow joint.

Cambrian

Cambrian

The Cambrian Period is 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 years ago (mya) to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, and its base, are somewhat in flux. The period was established as "Cambrian series" by Adam Sedgwick, who named it after Cambria, the Latin name for 'Cymru' (Wales), where Britain's Cambrian rocks are best exposed. Sedgwick identified the layer as part of his task, along with Roderick Murchison, to subdivide the large "Transition Series", although the two geologists disagreed for a while on the appropriate categorization. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells. As a result, our understanding of the Cambrian biology surpasses that of some later periods.

Brackish water

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 and fresh water together, as in estuaries, or it may occur in brackish fossil aquifers. The word comes from the Middle Dutch root brak. Certain human activities can produce brackish water, in particular civil engineering projects such as dikes and the flooding of coastal marshland to produce brackish water pools for freshwater prawn farming. Brackish water is also the primary waste product of the salinity gradient power process. Because brackish water is hostile to the growth of most terrestrial plant species, without appropriate management it is damaging to the environment.

Etymology

The taxonomic term Bivalvia was first used by Linnaeus in the 10th edition of his Systema Naturae in 1758 to refer to animals having shells composed of two valves.[3] More recently, the class was known as Pelecypoda, meaning "axe-foot" (based on the shape of the foot of the animal when extended).

The name "bivalve" is derived from the Latin bis, meaning "two", and valvae, meaning "leaves of a door".[4] ("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 gastropods (small sea snails in the family Juliidae),[5] members of the phylum Brachiopoda[6] and the minute crustaceans known as ostracods[7] and conchostracans.[8]

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Linnaean taxonomy

Linnaean taxonomy

Linnaean taxonomy can mean either of two related concepts:The particular form of biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his Systema Naturae (1735) and subsequent works. In the taxonomy of Linnaeus there are three kingdoms, divided into classes, and they, in turn, into lower ranks in a hierarchical order. A term for rank-based classification of organisms, in general. That is, taxonomy in the traditional sense of the word: rank-based scientific classification. This term is especially used as opposed to cladistic systematics, which groups organisms into clades. It is attributed to Linnaeus, although he neither invented the concept of ranked classification nor gave it its present form. In fact, it does not have an exact present form, as "Linnaean taxonomy" as such does not really exist: it is a collective (abstracting) term for what actually are several separate fields, which use similar approaches.

Carl Linnaeus

Carl Linnaeus

Carl Linnaeus, also known after his ennoblement in 1761 as Carl von Linné, was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the modern system of naming organisms. He is known as the "father of modern taxonomy". Many of his writings were in Latin; his name is rendered in Latin as Carolus Linnæus and, after his 1761 ennoblement, as Carolus a Linné.

10th edition of Systema Naturae

10th edition of Systema Naturae

The 10th edition of Systema Naturae is a book written by Swedish naturalist Carl Linnaeus and published in two volumes in 1758 and 1759, which marks the starting point of zoological nomenclature. In it, Linnaeus introduced binomial nomenclature for animals, something he had already done for plants in his 1753 publication of Species Plantarum.

Systema Naturae

Systema Naturae

Systema Naturae 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 nomenclature, was partially developed by the Bauhin brothers, Gaspard and Johann, Linnaeus was first to use it consistently throughout his book. The first edition was published in 1735. The full title of the 10th edition (1758), which was the most important one, was Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis or translated: "System of nature through the three kingdoms of nature, according to classes, orders, genera and species, with characters, differences, synonyms, places".

Valve (mollusc)

Valve (mollusc)

A valve is each articulating part of the shell of a mollusc or another multi-shelled animal such as brachiopods and some crustaceans. Each part is known as a valve or in the case of chitons, a "plate". Members of two classes of molluscs, the Bivalvia (clams) and the Polyplacophora (chitons), have valves.

Axe

Axe

An axe 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 many forms and specialised uses but generally consists of an axe head with a handle, or helve.

Latin

Latin

Latin 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 around present-day Rome, but through the power of the Roman Republic it became the dominant language in the Italian region and subsequently throughout the Roman Empire. Even after the fall of Western Rome, Latin remained the common language of international communication, science, scholarship and academia in Europe until well into the 18th century, when other regional vernaculars supplanted it in common academic and political usage, and it eventually became a dead language in the modern linguistic definition.

Sea snail

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 absence of a visible shell.

Juliidae

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.

Anatomy

Drawing of freshwater pearl mussel (Margaritifera margaritifera) anatomy: .mw-parser-output .thumb .image-key>ol{margin-left:1.3em;margin-top:0}.mw-parser-output .thumb .image-key>ul{margin-top:0}.mw-parser-output .thumb .image-key li{page-break-inside:avoid;break-inside:avoid-column}@media(min-width:300px){.mw-parser-output .thumb .image-key,.mw-parser-output .thumb .image-key-wide{column-count:2}.mw-parser-output .thumb .image-key-narrow{column-count:1}}@media(min-width:450px){.mw-parser-output .thumb .image-key-wide{column-count:3}}posterior adductoranterior adductorouter left gill demibranchinner left gill demibranchexcurrent siphonincurrent siphonfootteethhingemantleumbo
Drawing of freshwater pearl mussel (Margaritifera margaritifera) anatomy:
  1. posterior adductor
  2. anterior adductor
  3. outer left gill demibranch
  4. inner left gill demibranch
  5. excurrent siphon
  6. incurrent siphon
  7. foot
  8. teeth
  9. hinge
  10. mantle
  11. umbo
Interior of the left valve of a venerid
Interior of the left valve of a venerid
Main parts of a bivalve shell: sagittal planegrowth linesligamentumbo
Main parts of a bivalve shell:
  1. sagittal plane
  2. growth lines
  3. ligament
  4. umbo

Bivalves have bilaterally symmetrical and laterally flattened bodies, with a blade-shaped foot, vestigial head and no radula.[9][10] At the dorsal or back region of the shell is the hinge point or line, which contain the umbo and beak and the lower, curved margin is the ventral or underside region. The anterior or front of the shell is where the byssus (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.[11][12] Many bivalves such as clams, which appear upright, are evolutionarily lying on their side.

Mantle and shell

The shell is composed of two calcareous valves held together by a ligament. The valves are made of either calcite, as is the case in oysters, or both calcite and aragonite. Sometimes, the aragonite forms an inner, nacreous layer, as is the case in the order Pteriida. In other taxa, alternate layers of calcite and aragonite are laid down.[13] The ligament and byssus, if calcified, are composed of aragonite.[13] The outermost layer of the shell is the periostracum, a thin layer composed of horny conchiolin. The periostracum is secreted by the outer mantle and is easily abraded.[14] 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.[15]

In all molluscs, the mantle forms a thin membrane 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, byssus threads (where present), and teeth.[16] The posterior mantle edge may have two elongated extensions known as siphons, through one of which water is inhaled, and the other expelled.[17] The siphons retract into a cavity, known as the pallial sinus.[18]

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.[19]

The shipworms, in the family Teredinidae 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.[20]

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, 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.[21] Retractor muscles connect the mantle to the edge of the shell, along a line known as the pallial line.[22][16] These muscles pull the mantle though the valves.[16]

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.[23] 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.[24] Paired pedal protractor and retractor muscles operate the animal's foot.[11][25][26]

Nervous system

The sedentary habits of the bivalves have meant that in general the nervous system is less complex than in most other molluscs. The animals have no brain; the nervous system consists of a nerve network and a series of paired ganglia. In all but the most primitive bivalves, two cerebropleural ganglia are on either side of the oesophagus. 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. Bivalves with long siphons may also have siphonal ganglia to control them.[27][28]

Senses

The sensory organs of bivalves are largely located on the posterior mantle margins. The organs are usually mechanoreceptors or chemoreceptors, in some cases located on short tentacles. The osphradium is a patch of sensory cells located below the posterior adductor muscle that may serve to taste the water or measure its turbidity. Statocysts within the organism help the bivalve to sense and correct its orientation.[28] In the order Anomalodesmata, the inhalant siphon is surrounded by vibration-sensitive tentacles for detecting prey.[29] 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.[30] Scallops have more complex eyes with a lens, a two-layered retina, and a concave mirror.[31] All bivalves have light-sensitive cells that can detect a shadow falling over the animal.[27]

Circulation and respiration

Four filaments of the gills of the blue mussel (Mytilus edulis) a) part of four filaments showing ciliated interfilamentar junctions (cj) b) diagram of a single filament showing the two lamellae connected at intervals by interlamellar junctions (ilj) and the position of the ciliated interfilamentar junctions (cp)
Four filaments of the gills of the blue mussel (Mytilus edulis) a) part of four filaments showing ciliated interfilamentar junctions (cj) b) diagram of a single filament showing the two lamellae connected at intervals by interlamellar junctions (ilj) and the position of the ciliated interfilamentar junctions (cp)

Bivalves have an open circulatory system that bathes the organs in blood (hemolymph). The heart has three chambers: two auricles receiving blood from the gills, and a single ventricle. The ventricle is muscular and pumps hemolymph into the aorta, 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.[32] The hemolymph usually lacks any respiratory pigment.[33] In the carnivorous genus Poromya, the hemolymph has red amoebocytes containing a haemoglobin pigment.[34]

The paired gills are located posteriorly and consist of hollow tube-like filaments with thin walls for gas exchange. 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.[35][36] Oysters, including the Pacific oyster (Magallana gigas), are recognized as having varying metabolic responses to environmental stress, with changes in respiration rate being frequently observed.[37]

Digestive system

Modes of feeding

Most bivalves are filter feeders, using their gills to capture particulate food such as phytoplankton 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, or flap. The tentacles are covered in mucus, 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.[38]

In more advanced bivalves, water is drawn into the shell from the posterior ventral 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 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.[38][39]

A few bivalves, such as the granular poromya (Poromya granulata), are carnivorous, eating much larger prey 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.[34]

The unusual genus, Entovalva, is endosymbiotic, being found only in the oesophagus of sea cucumbers. 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's throat. The sea cucumber is unharmed.[40]

Digestive tract

The digestive tract of typical bivalves consists of an oesophagus, stomach, and intestine. 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" 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.[41] 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.[42]

Carnivorous bivalves generally have reduced crystalline styles and the stomach has thick, muscular walls, extensive cuticular linings and diminished sorting areas and gastric chamber sections.[43]

Excretory system

The excretory organs of bivalves are a pair of nephridia. Each of these consists of a long, looped, glandular tube, which opens into the pericardium, and a bladder 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 near the front of the upper part of the mantle cavity and excreted.[44][45]

Reproduction and development

The sexes are usually separate in bivalves but some hermaphroditism is known. The gonads and either open into the nephridia, or through a separate pore into a chamber over the gills.[46][47] The ripe gonads of males and females release sperm and eggs into the water column. Spawning 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.[48]

Fertilization is usually external. Typically, a short stage lasts a few hours or days before the eggs hatch into trochophore larvae. These later develop into veliger larvae which settle on the seabed and undergo metamorphosis into adults.[46][49] In some species, such as those in the genus Lasaea, 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.[50]

Most of the bivalve larvae that hatch from eggs in the water column feed on diatoms or other phytoplankton. In temperate regions, about 25% of species are lecithotrophic, depending on nutrients stored in the yolk of the egg where the main energy source is lipids. 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) 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) 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.[51]

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 larvae that develop within the female's shell. Later they are released and attach themselves parasitically to the gills or fins of a fish host. After several weeks they drop off their host, undergo metamorphosis and develop into adults on the substrate.[46]

Some of the species in the freshwater mussel family, Unionidae, 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. 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 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.[52]

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Freshwater pearl mussel

Freshwater pearl mussel

The freshwater pearl mussel is an endangered species of freshwater mussel, an aquatic bivalve mollusc in the family Margaritiferidae.

Ligament

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 ligament: a fold of peritoneum or other membranes. Fetal remnant ligament: the remnants of a fetal tubular structure. Periodontal ligament: a group of fibers that attach the cementum of teeth to the surrounding alveolar bone.

Beak (bivalve)

Beak (bivalve)

The beak is part of the shell of a bivalve mollusk, i.e. part of the shell of a saltwater or freshwater clam. The beak is the basal projection of the oldest part of the valve of the adult animal. The beak usually, but not always, coincides with the umbo, the highest and most prominent point on the valve. Because by definition, all bivalves have two valves, the shell of a bivalve has two umbones, and two beaks.

Byssus

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 (Mytilidae), and Dreissenidae.

Bivalve shell

Bivalve shell

A bivalve shell is part of the body, the exoskeleton or shell, of a bivalve mollusk. In life, the shell of this class of mollusks is composed of two hinged parts or valves. Bivalves are very common in essentially all aquatic locales, including saltwater, brackish water, and freshwater. The shells of bivalves commonly wash up on beaches and along the edges of lakes, rivers, and streams. Bivalves by definition possess two shells or valves, a "right valve" and a "left valve", that are joined by a ligament. The two valves usually articulate with one another using structures known as "teeth" which are situated along the hinge line. In many bivalve shells, the two valves are symmetrical along the hinge line—when truly symmetrical, such an animal is said to be equivalved; if the valves vary from each other in size or shape, inequivalved. If symmetrical front-to-back, the valves are said to be equilateral, and are otherwise considered inequilateral.

Calcareous

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.

Calcite

Calcite

Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratch hardness comparison. Large calcite crystals are used in optical equipment, and limestone composed mostly of calcite has numerous uses.

Aragonite

Aragonite

Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, CaCO3. It is formed by biological and physical processes, including precipitation from marine and freshwater environments.

Nacre

Nacre

Nacre, also known as mother of pearl, is an organic–inorganic 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.

Conchiolin

Conchiolin

Conchiolins are complex proteins which are secreted by a mollusc's outer epithelium.

Mantle (mollusc)

Mantle (mollusc)

The mantle is a significant part of the anatomy of molluscs: it is the dorsal body wall which covers the visceral mass and usually protrudes in the form of flaps well beyond the visceral mass itself.

Biological membrane

Biological membrane

A biological membrane, biomembrane or cell membrane is a selectively permeable membrane that separates the interior of a cell from the external environment or creates intracellular compartments by serving as a boundary between one part of the cell and another. Biological membranes, in the form of eukaryotic cell membranes, consist of a phospholipid bilayer with embedded, integral and peripheral proteins used in communication and transportation of chemicals and ions. The bulk of lipids in a cell membrane provides a fluid matrix for proteins to rotate and laterally diffuse for physiological functioning. Proteins are adapted to high membrane fluidity environment of the lipid bilayer with the presence of an annular lipid shell, consisting of lipid molecules bound tightly to the surface of integral membrane proteins. The cell membranes are different from the isolating tissues formed by layers of cells, such as mucous membranes, basement membranes, and serous membranes.

Comparison with brachiopods

Anadara, a bivalve with taxodont dentition from the Pliocene of Cyprus
Anadara, a bivalve with taxodont dentition from the Pliocene of Cyprus
A fossil Jurassic brachiopod with the lophophore support intact
A fossil Jurassic brachiopod with the lophophore support intact

Brachiopods 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. 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. In modern times, brachiopods are not as common as bivalves.[53]

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, a coiled, rigid cartilaginous internal apparatus adapted for filter feeding, a feature shared with two other major groups of marine invertebrates, the bryozoans and the phoronids. Some brachiopod shells are made of calcium phosphate but most are calcium carbonate in the form of the biomineral calcite, whereas bivalve shells are always composed entirely of calcium carbonate, often in the form of the biomineral aragonite.[54]

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Anadara

Anadara

Anadara is a genus of saltwater bivalves, ark clams, in the family Arcidae. It is also called Scapharca.

Ark clam

Ark clam

Ark clam is the common name for a family of small to large-sized saltwater clams or marine bivalve molluscs in the family Arcidae. Ark clams vary both in shape and size. They number about 200 species worldwide.

Cyprus

Cyprus

Cyprus, officially the Republic of Cyprus, is an island country located south of the Anatolian Peninsula in the eastern Mediterranean Sea. It is geographically in Western Asia, but its cultural ties and geopolitics are overwhelmingly Southeastern European. Cyprus is the third-largest and third-most populous island in the Mediterranean. It is located north of Egypt, east of Greece, south of Turkey, and west of Lebanon and Syria. Its capital and largest city is Nicosia. The northeast portion of the island is de facto governed by the self-declared Turkish Republic of Northern Cyprus.

Lophophore

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. All lophophores are found in aquatic organisms.

Brachiopod

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, while the front can be opened for feeding or closed for protection. Two major categories are traditionally recognized, articulate and inarticulate brachiopods. The word "articulate" is used to describe the tooth-and-groove structures of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. This is the leading diagnostic skeletal feature, by which the two main groups can be readily distinguished as fossils. Articulate brachiopods have toothed hinges and simple, vertically-oriented opening and closing muscles. Conversely, inarticulate brachiopods have weak, untoothed hinges and a more complex system of vertical and oblique (diagonal) muscles used to keep the two valves aligned. In many brachiopods, a stalk-like pedicle projects from an opening near the hinge of one of the valves, known as the pedicle or ventral valve. The pedicle, when present, keeps the animal anchored to the seabed but clear of sediment which would obstruct the opening.

Ecological niche

Ecological niche

In ecology, a niche is the match of a species to a specific environmental condition. It describes how an organism or population responds to the distribution of resources and competitors and how it in turn alters those same factors. "The type and number of variables comprising the dimensions of an environmental niche vary from one species to another [and] the relative importance of particular environmental variables for a species may vary according to the geographic and biotic contexts".

Convergent evolution

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 common ancestor of those groups. The cladistic term for the same phenomenon is homoplasy. The recurrent evolution of flight is a classic example, as flying insects, birds, pterosaurs, and bats have independently evolved the useful capacity of flight. Functionally similar features that have arisen through convergent evolution are analogous, whereas homologous structures or traits have a common origin but can have dissimilar functions. Bird, bat, and pterosaur wings are analogous structures, but their forelimbs are homologous, sharing an ancestral state despite serving different functions.

Bryozoa

Bryozoa

Bryozoa are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about 0.5 millimetres long, they have a special feeding structure called a lophophore, a "crown" of tentacles used for filter feeding. Most marine bryozoans live in tropical waters, but a few are found in oceanic trenches and polar waters. The bryozoans are classified as the marine bryozoans (Stenolaemata), freshwater bryozoans (Phylactolaemata), and mostly-marine bryozoans (Gymnolaemata), a few members of which prefer brackish water. 5,869 living species are known. At least two genera are solitary ; the rest are colonial.

Calcium phosphate

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 solids of nutritious value and are found in many living organisms, e.g., bone mineral and tooth enamel. In milk, it exists in a colloidal form in micelles bound to casein protein with magnesium, zinc, and citrate–collectively referred to as colloidal calcium phosphate (CCP). Various calcium phosphate minerals are used in the production of phosphoric acid and fertilizers. Overuse of certain forms of calcium phosphate can lead to nutrient-containing surface runoff and subsequent adverse effects upon receiving waters such as algal blooms and eutrophication (over-enrichment with nutrients and minerals).

Calcium carbonate

Calcium carbonate

Calcium carbonate is a chemical compound with the chemical formula CaCO3. It is a common substance found in rocks as the minerals calcite and aragonite and is the main component of eggshells, gastropod shells, shellfish skeletons and pearls. Things containing much calcium carbonate or resembling it are described as calcareous. Calcium carbonate is the active ingredient in agricultural lime and is created when calcium ions in hard water react with carbonate ions to create limescale. It has medical use as a calcium supplement or as an antacid, but excessive consumption can be hazardous and cause hypercalcemia and digestive issues.

Calcite

Calcite

Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratch hardness comparison. Large calcite crystals are used in optical equipment, and limestone composed mostly of calcite has numerous uses.

Aragonite

Aragonite

Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, CaCO3. It is formed by biological and physical processes, including precipitation from marine and freshwater environments.

Evolutionary history

The Cambrian explosion 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.[55]

Possible early bivalves include Pojetaia and Fordilla; these probably lie in the stem rather than crown group. Watsonella and Anabarella are perceived to be (earlier) close relatives of these taxa.[56] Only five genera of supposed Cambrian "bivalves" exist, the others being Tuarangia, Camya and Arhouriella and potentially Buluniella.[57]

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, 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 and Carboniferous periods, siphons first appeared, which, with the newly developed muscular foot, allowed the animals to bury themselves deep in the sediment.[58]

By the middle of the Paleozoic, around 400 Mya, the brachiopods were among the most abundant filter feeders in the ocean, and over 12,000 fossil species are recognized.[59] By the Permian–Triassic extinction event 250 Mya, bivalves were undergoing a huge radiation of diversity. The bivalves were hard hit by this event, but re-established themselves and thrived during the Triassic period that followed. In contrast, the brachiopods lost 95% of their species diversity.[54] 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 in soft sediments by developing spines on the shell, gaining the ability to swim, and in a few cases, adopting predatory habits.[58]

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 events.[60]

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Cambrian explosion

Cambrian explosion

The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately 538.8 million years ago in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. It lasted for about 13 – 25 million years and resulted in the divergence of most modern metazoan phyla. The event was accompanied by major diversification in other groups of organisms as well.

Fordilla

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. The genus currently contains three described species, Fordilla germanica, Fordilla sibirica, and the type species Fordilla troyensis.

Camya

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, Camya asy.

Arhouriella

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, Arhouriella opheodontoides, was named and described by Gerd Geyer & Michael Streng in 1998.

Devonian

Devonian

The Devonian is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, 419.2 million years ago (Mya), to the beginning of the Carboniferous, 358.9 Mya. It is named after Devon, England, where rocks from this period were first studied.

Carboniferous

Carboniferous

The Carboniferous is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period 358.9 million years ago (Mya), to the beginning of the Permian Period, 298.9 million years ago. The name Carboniferous means "coal-bearing", from the Latin carbō ("coal") and ferō, and refers to the many coal beds formed globally during that time.

Paleozoic

Paleozoic

The Paleozoic 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 and zōḗ, "life", meaning "ancient life").

Permian–Triassic extinction event

Permian–Triassic extinction event

The Permian–Triassic extinction event, also known as the End-Permian extinction event and colloquially as the Great Dying, forms the boundary between the Permian and Triassic geologic periods, and with them the Paleozoic and Mesozoic eras respectively, approximately 251.9 million years ago. As the largest of the "Big Five" mass extinctions of the Phanerozoic, it is the Earth's most severe known extinction event, with the extinction of 57% of biological families, 83% of genera, 81% of marine species and 70% of terrestrial vertebrate species. It is the largest known mass extinction of insects. There is evidence for one to three distinct pulses, or phases, of extinction.

Evolutionary radiation

Evolutionary radiation

An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rapid or gradual; where they are rapid, and driven by a single lineage's adaptation to their environment, they are termed adaptive radiations.

Buoyancy

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 pressure at the bottom of a column of fluid is greater than at the top of the column. Similarly, the pressure at the bottom of an object submerged in a fluid is greater than at the top of the object. The pressure difference results in a net upward force on the object. The magnitude of the force is proportional to the pressure difference, and is equivalent to the weight of the fluid that would otherwise occupy the submerged volume of the object, i.e. the displaced fluid.

Competition (biology)

Competition (biology)

Competition is an interaction between organisms or species in which both require a resource that is in limited supply. Competition lowers the fitness of both organisms involved since the presence of one of the organisms always reduces the amount of the resource available to the other.

Ecological niche

Ecological niche

In ecology, a niche is the match of a species to a specific environmental condition. It describes how an organism or population responds to the distribution of resources and competitors and how it in turn alters those same factors. "The type and number of variables comprising the dimensions of an environmental niche vary from one species to another [and] the relative importance of particular environmental variables for a species may vary according to the geographic and biotic contexts".

Diversity of extant bivalves

The adult maximum size of living species of bivalve ranges from 0.52 mm (0.02 in) in Condylonucula maya,[61] a nut clam, to a length of 1,532 millimetres (60.3 in) in Kuphus polythalamia, an elongated, burrowing shipworm.[62] However, the species generally regarded as the largest living bivalve is the giant clam Tridacna gigas, which can grow to a length of 1,200 mm (47 in) and a weight of more than 200 kg (441 lb).[63] The largest known extinct bivalve is a species of Platyceramus whose fossils measure up to 3,000 mm (118 in) in length.[64]

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.[65] 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:

Subclass Superfamilies Families Genera Species
Heterodonta 64 (incl. 1 freshwater) 800 (16 freshwater) 5600 (270 freshwater)
Arcticoidea 2 6 13
Cardioidea 2 38 260
Chamoidea 1 6 70
Clavagelloidea 1 2 20
Crassatelloidea 5 65 420
Cuspidarioidea 2 20 320
Cyamioidea 3 22 140
Cyrenoidea 1 6 (3 freshwater) 60 (30 freshwater)
Cyrenoidoidea 1 1 6
Dreissenoidea 1 3 (2 freshwater) 20 (12 freshwater)
Galeommatoidea ca. 4 about 100 about 500
Gastrochaenoidea 1 7 30
Glossoidea 2 20 110
Hemidonacoidea 1 1 6
Hiatelloidea 1 5 25
Limoidea 1 8 250
Lucinoidea 2 about 85 about 500
Mactroidea 4 46 220
Myoidea 3 15 (1 freshwater) 130 (1 freshwater)
Pandoroidea 7 30 250
Pholadoidea 2 34 (1 freshwater) 200 (3 freshwater)
Pholadomyoidea 2 3 20
Solenoidea 2 17 (2 freshwater) 130 (4 freshwater)
Sphaerioidea (1 freshwater) (5 freshwater) (200 freshwater)
Tellinoidea 5 110 (2 freshwater) 900 (15 freshwater)
Thyasiroidea 1 about 12 about 100
Ungulinoidea 1 16 100
Veneroidea 4 104 750
Verticordioidea 2 16 160
Palaeoheterodonta 7 (incl. 6 freshwater) 171 (170 freshwater) 908 (900 freshwater)
Trigonioidea 1 1 8
Unionoidea (6 freshwater) (170 freshwater) (900 freshwater)
Protobranchia 10 49 700
Manzanelloidea 1 2 20
Nuculanoidea 6 32 460
Nuculoidea 1 8 170
Sareptoidea 1 about 5 10
Solemyoidea 1 2 30
Pteriomorphia 25 240 (2 freshwater) 2000 (11 freshwater)
Anomioidea 2 9 30
Arcoidea 7 60 (1 freshwater) 570 (6 freshwater)
Dimyoidea 1 3 15
Limoidea 1 8 250
Mytiloidea 1 50 (1 freshwater) 400 (5 freshwater)
Ostreoidea 2 23 80
Pectinoidea 4 68 500
Pinnoidea 1 3 (+) 50
Plicatuloidea 1 1 20
Pterioidea 5 9 80

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Condylonucula maya

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 500 μm (0.020 in) and is believed to be the smallest living bivalve. It is found in shallow waters in the Caribbean Sea off the coast of Mexico.

Kuphus

Kuphus

Kuphus is a genus of shipworms, marine bivalve molluscs in the family Teredinidae. While there are four extinct species in the genus, the only extant species is Kuphus polythalamius. It is the longest bivalve mollusc in the world, where the only known permanent natural habitat is Kalamansig, Sultan Kudarat in the Philippines.

Giant clam

Giant clam

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, the most commonly intended species referred to as "the giant clam".

Extinction

Extinction

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 recover may have been lost before this point. Because a species' potential range may be very large, determining this moment is difficult, and is usually done retrospectively. This difficulty leads to phenomena such as Lazarus taxa, where a species presumed extinct abruptly "reappears" after a period of apparent absence.

Platyceramus

Platyceramus

Platyceramus was a genus of Cretaceous bivalve molluscs belonging to the extinct inoceramid lineage. It is sometimes classified as a subgenus of Inoceramus.

Cardioidea

Cardioidea

Cardioidea is a taxonomic superfamily of saltwater clams, marine bivalve molluscs consisting of the extant Cardiidae (cockles) and the extinct Pterocardiidae.

Crassatelloidea

Crassatelloidea

Crassatelloidea is a superfamily of bivalves in the order Carditida. In the World Register of Marine Species (WoRMS), Astartoidea is considered a junior synonym of Crassatelloidea, whereas in ITIS Astartoidea is a separate family containing Astartidae and Cardiniidae – Cardiniidae itself being classified instead in Carditoidea by WoRMS.

Dreissenoidea

Dreissenoidea

Dreissenoidea is a superfamily of brackish water and freshwater false mussels, aquatic bivalve molluscs in the order Myida.

Tellinoidea

Tellinoidea

Tellinoidea is a taxonomic superfamily of saltwater clams, marine bivalve molluscs in the order Cardiida.

Distribution

Zebra mussels encrusting a water velocity meter in Lake Michigan
Zebra mussels encrusting a water velocity meter in Lake Michigan

The bivalves are a highly successful class of invertebrates found in aquatic habitats throughout the world. Most are infaunal 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 and sublittoral 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, careful sampling produced an estimate of 1.44 million cockles (Cerastoderma edule) per acre of beach.[66]

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.[67] The Antarctic scallop, Adamussium colbecki, lives under the sea ice at the other end of the globe, where the subzero temperatures mean that growth rates are very slow.[68] The giant mussel, Bathymodiolus thermophilus, and the giant white clam, Calyptogena magnifica, both live clustered around hydrothermal vents at abyssal depths in the Pacific Ocean. They have chemosymbiotic bacteria in their gills that oxidise hydrogen sulphide, and the molluscs absorb nutrients synthesized by these bacteria.[69] The saddle oyster, Enigmonia aenigmatica, is a marine species that could be considered amphibious. It lives above the high tide mark in the tropical Indo-Pacific on the underside of mangrove leaves, on mangrove branches, and on sea walls in the splash zone.[70]

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 in Arkansas and Oklahoma, and like several other freshwater mussel species from the southeastern US, it is in danger of extinction.[71] In contrast, a few species of freshwater bivalves, including the golden mussel (Limnoperna fortunei), are dramatically increasing their ranges. The golden mussel has spread from Southeast Asia to Argentina, where it has become an invasive species.[72] Another well-travelled freshwater bivalve, the zebra mussel (Dreissena polymorpha) 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 ecosystems.[73]

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Intertidal zone

Intertidal zone

The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide. This area can include several types of habitats with various species of life, such as seastars, sea urchins, and many species of coral with regional differences in biodiversity. Sometimes it is referred to as the littoral zone or seashore, although those can be defined as a wider region.

South Wales

South Wales

South Wales 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 to include Carmarthenshire and Pembrokeshire. In the western extent, from Swansea westwards, local people would probably recognise that they lived in both south Wales and west Wales. The Brecon Beacons National Park covers about a third of south Wales, containing Pen y Fan, the highest British mountain south of Cadair Idris in Snowdonia.

Bathymodiolus thermophilus

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 exploring the deep ocean. It occurs on the sea bed, often in great numbers, close to hydrothermal vents where hot, sulphur-rich water wells up through the floor of the Pacific Ocean.

Calyptogena magnifica

Calyptogena magnifica

Calyptogena magnifica is a species of giant white clam found clustered around hydrothermal vents at abyssal depths in the Pacific Ocean.

Hydrothermal vent

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 hotspots. Hydrothermal deposits are rocks and mineral ore deposits formed by the action of hydrothermal vents.

Abyssal zone

Abyssal zone

The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. "Abyss" derives from the Greek word ἄβυσσος, meaning bottomless. At depths of 4,000 to 6,000 metres, this zone remains in perpetual darkness. It covers 83% of the total area of the ocean and 60% of Earth's surface. The abyssal zone has temperatures around 2 to 3 °C through the large majority of its mass. Due to there being no light, there are no plants producing oxygen, which instead primarily comes from ice that had melted long ago from the polar regions. The water along the seafloor of this zone is actually devoid of oxygen, resulting in a death trap for organisms unable to quickly return to the oxygen-enriched water above. This region also contains a much higher concentration of nutrient salts, like nitrogen, phosphorus, and silica, due to the large amount of dead organic material that drifts down from the above ocean zones and decomposes. The water pressure can reach up to 76 megapascal.

Amphibious fish

Amphibious fish

Amphibious fish are fish that are able to leave water for extended periods of time. About 11 distantly related genera of fish are considered amphibious. This suggests that many fish genera independently evolved amphibious traits, a process known as convergent evolution. These fish use a range of terrestrial locomotory modes, such as lateral undulation, tripod-like walking, and jumping. Many of these locomotory modes incorporate multiple combinations of pectoral-, pelvic-, and tail-fin movement.

Mangrove

Mangrove

A mangrove is a shrub or tree that grows in coastal 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 evolution in several plant families. They occur worldwide in the tropics and subtropics and even some temperate coastal areas, mainly between latitudes 30° N and 30° S, with the greatest mangrove area within 5° of the equator. Mangrove plant families first appeared during the Late Cretaceous to Paleocene epochs, and became widely distributed in part due to the movement of tectonic plates. The oldest known fossils of mangrove palm date to 75 million years ago.

Splash zone

Splash zone

In offshore construction, the splash zone is the transition from air to water when lowering heavy burdens into the sea. The overall efforts applied on the crane change dramatically when the load starts touching water, up to the point where it is completely submerged. Its buoyancy reduces the static mass that the crane has to support, but contact with the waves creates widely fluctuating dynamic forces.

Ouachita Mountains

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 Thrust Belt, one of the important orogenic belts of North America. The Ouachitas continue in the subsurface to the northeast, where they make a poorly understood connection with the Appalachians and to the southwest, where they join with the Marathon uplift area of West Texas. Together with the Ozark Plateaus, the Ouachitas form the U.S. Interior Highlands. The highest natural point is Mount Magazine at 2,753 feet (839 m).

Limnoperna fortunei

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 where it has become an invasive species. It is considered to be an ecosystem engineer because it alters the nature of the water and the bottom habitats of lakes and rivers and modifies the associated invertebrate communities. It also has strong effects on the properties of the water column, modifying nutrient proportions and concentrations, increasing water transparency, decreasing phytoplankton and zooplankton densities, on which it feeds, and enhancing the growth of aquatic macrophytes. Because mussels attach to hard substrata, including the components of industrial, water-treatment and power plants, they have become a major biofouling problem in the areas invaded.

Invasive species

Invasive species

An invasive or alien species is an introduced species to an environment that becomes overpopulated and harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native species that become harmful to their native environment after human alterations to its food web – for example the purple sea urchin which has decimated kelp forests along the northern California coast due to overharvesting of its natural predator, the California sea otter. Since the 20th century, invasive species have become a serious economic, social, and environmental threat.

Behaviour

A large number of live venerid bivalves underwater with their siphons visible
A large number of live venerid bivalves underwater with their siphons visible
Pacific oyster equipped with activity electrodes to follow its daily behaviour
Pacific oyster equipped with activity electrodes to follow its daily behaviour

Most bivalves adopt a sedentary or even sessile 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 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.[74] 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.[74] 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.[75]

Other bivalves, such as mussels, attach themselves to hard surfaces using tough byssus threads made of collagen and elastin proteins.[76] 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.[77] 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 and, like most bivalves, are filter feeders.[78]

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.[79]

Although many non-sessile bivalves use their muscular foot to move around, or to dig, members of the freshwater family Sphaeriidae 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), for example, climbs around on water weeds at the edges of lakes and ponds; this enables the clam to find the best position for filter feeding.[80]

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 feed on them including the common carp (Cyprinus carpio), which is being used in the upper Mississippi River to try to control the invasive zebra mussel (Dreissena polymorpha).[81] Birds such as the Eurasian oystercatcher (Haematopus ostralegus) have specially adapted beaks which can pry open their shells.[82] The herring gull (Larus argentatus) sometimes drops heavy shells onto rocks in order to crack them open.[83] Sea otters 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.[84] The Pacific walrus (Odobenus rosmarus divergens) is one of the main predators feeding on bivalves in Arctic waters.[85] 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.[86] Further changes in shell development due to environmental stress has also been suggested to cause increased mortality in oysters due to reduced shell strength. [37]

Invertebrate predators include crustaceans, starfish and octopuses. Crustaceans crack the shells with their pincers and starfish use their water vascular system 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.[77] 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.[87] Certain carnivorous gastropod snails such as whelks (Buccinidae) and murex snails (Muricidae) feed on bivalves by boring into their shells. A dog whelk (Nucella) drills a hole with its radula 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.[88]

Razor shells can dig themselves into the sand with great speed to escape predation. When a Pacific razor clam (Siliqua patula) is laid on the surface of the beach it can bury itself completely in seven seconds [89] and the Atlantic jackknife clam, Ensis directus, can do the same within fifteen seconds.[90] 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.[91] 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.[91] 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.[92]

In many bivalves that have siphons, 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.[93] 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 lifestyle.[94][95]

File shells such as Limaria fragilis 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. The toxin released by this is distasteful and the detached tentacles continue to writhe which may also serve to distract potential predators.[96]

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Pacific oyster

Pacific oyster

The Pacific oyster, Japanese oyster, or Miyagi oyster, is an oyster native to the Pacific coast of Asia. It has become an introduced species in North America, Australia, Europe, and New Zealand.

Intertidal zone

Intertidal zone

The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide. This area can include several types of habitats with various species of life, such as seastars, sea urchins, and many species of coral with regional differences in biodiversity. Sometimes it is referred to as the littoral zone or seashore, although those can be defined as a wider region.

Mussel

Mussel

Mussel is the common name used for members of several families of bivalve molluscs, from saltwater and freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other edible clams, which are often more or less rounded or oval.

Byssus

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 (Mytilidae), and Dreissenidae.

Collagen

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-body protein content. Collagen consists of amino acids bound together to form a triple helix of elongated fibril known as a collagen helix. It is mostly found in connective tissue such as cartilage, bones, tendons, ligaments, and skin.

Elastin

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. It is highly elastic and present in connective tissue allowing many tissues in the body to resume their shape after stretching or contracting. Elastin helps skin to return to its original position when it is poked or pinched. Elastin is also an important load-bearing tissue in the bodies of vertebrates and used in places where mechanical energy is required to be stored.

Chamidae

Chamidae

Chamidae, common name the jewel boxes or jewel box clams, is a taxonomic family of saltwater clams, a group of marine bivalve mollusks in the order Venerida.

Anomiidae

Anomiidae

Anomiidae is a family of saltwater clams, marine bivalve molluscs related to scallops and oysters, and known as anomiids. It contains seven genera.

Neritic zone

Neritic zone

The neritic zone is the relatively shallow part of the ocean above the drop-off of the continental shelf, approximately 200 meters (660 ft) in depth. From the point of view of marine biology it forms a relatively stable and well-illuminated environment for marine life, from plankton up to large fish and corals, while physical oceanography sees it as where the oceanic system interacts with the coast.

Demersal fish

Demersal fish

Demersal fish, also known as groundfish, live and feed on or near the bottom of seas or lakes. They occupy the sea floors and lake beds, which usually consist of mud, sand, gravel or rocks. In coastal waters they are found on or near the continental shelf, and in deep waters they are found on or near the continental slope or along the continental rise. They are not generally found in the deepest waters, such as abyssal depths or on the abyssal plain, but they can be found around seamounts and islands. The word demersal comes from the Latin demergere, which means to sink.

Eurasian oystercatcher

Eurasian oystercatcher

The Eurasian oystercatcher also known as the common pied oystercatcher, or palaearctic oystercatcher, or just oystercatcher, is a wader in the oystercatcher bird family Haematopodidae. It is the most widespread of the oystercatchers, with three races breeding in western Europe, central Eurosiberia, Kamchatka, China, and the western coast of Korea. No other oystercatcher occurs within this area. The extinct Canary Islands oystercatcher, formerly considered a distinct species, may have actually been an isolated subspecies or distinct population of the Eurasian oystercatcher.

European herring gull

European herring gull

The European herring gull is a large gull, up to 66 cm (26 in) long. One of the best-known of all gulls along the shores of Western Europe, it was once abundant. It breeds across Northern Europe, Western Europe, Central Europe, Eastern Europe, Scandinavia, and the Baltic states. Some European herring gulls, especially those resident in colder areas, migrate further south in winter, but many are permanent residents, such as in Ireland, Britain, Iceland, or on the North Sea shores. They have a varied diet, including fish, crustaceans, as well as some plants, and are also scavengers, consuming carrion and food left by or stolen from humans.

Mariculture

Oyster culture in Brittany, France
Oyster culture in Brittany, France

Oysters, mussels, clams, scallops and other bivalve species are grown with food materials that occur naturally in their culture environment in the sea and lagoons.[97] 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 carps.[97] European flat oysters (Ostrea edulis) were first farmed by the Romans in shallow ponds and similar techniques are still in use.[98] 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 5 to 6 millimetres (0.20 to 0.24 in) in length.[98]

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 50 to 100 kilograms (110 to 220 lb) per hectare. They grow on for about two years before being harvested by dredging. Survival rates are low at about 5%.[98]

The Pacific oyster (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.[99] It is an estuarine species and prefers salinities of 20 to 25 parts per thousand. 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 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.[99]

Similar techniques are used in different parts of the world to cultivate other species including the Sydney rock oyster (Saccostrea commercialis), the northern quahog (Mercenaria mercenaria), the blue mussel (Mytilus edulis), the Mediterranean mussel (Mytilus galloprovincialis), the New Zealand green-lipped mussel (Perna canaliculus), the grooved carpet shell (Ruditapes decussatus), the Japanese carpet shell (Venerupis philippinarum), the pullet carpet shell (Venerupis pullastra) and the Yesso scallop (Patinopecten yessoensis).[100]

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.[101]

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Oyster farming

Oyster farming

Oyster farming is an aquaculture practice in which oysters are bred and raised mainly for their pearls, shells and inner organ tissue, which is eaten. Oyster farming was practiced by the ancient Romans as early as the 1st century BC on the Italian peninsula and later in Britain for export to Rome. The French oyster industry has relied on aquacultured oysters since the late 18th century.

Scallop aquaculture

Scallop aquaculture

Scallop aquaculture is the commercial activity of cultivating (farming) scallops until they reach a marketable size and can be sold as a consumer product. Wild juvenile scallops, or spat, were collected for growing in Japan as early as 1934. The first attempts to fully cultivate scallops in farm environments were not recorded until the 1950s and 1960s. Traditionally, fishing for wild scallops has been the preferred practice, since farming can be expensive. However worldwide declines in wild scallop populations have resulted in the growth of aquaculture. Globally the scallop aquaculture industry is now well established, with a reported annual production totalling over 1,200,000 metric tonnes from about 12 species. China and Japan account for about 90% of the reported production.

Carp

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 Africa, Australia and most of the United States.

Ostrea edulis

Ostrea edulis

Ostrea edulis, commonly known as the European flat oyster, is a species of oyster native to Europe. In Britain and Ireland, regional names include Colchester native oyster, mud oyster, or edible oyster. In France, Ostrea edulis are known as huîtres plates except for those that come from the Belon River estuary in Brittany, France, which are known as Belons.

Dredging

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 dams, dikes, and other controls for streams and shorelines; and recovering valuable mineral deposits or marine life having commercial value. In all but a few situations the excavation is undertaken by a specialist floating plant, known as a dredger.

Pacific oyster

Pacific oyster

The Pacific oyster, Japanese oyster, or Miyagi oyster, is an oyster native to the Pacific coast of Asia. It has become an introduced species in North America, Australia, Europe, and New Zealand.

Salinity

Salinity

Salinity is the saltiness or amount of salt dissolved in a body of water, called saline water. It is usually measured in g/L or g/kg.

Microalgae

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 individually, or in chains or groups. Depending on the species, their sizes can range from a few micrometers (μm) to a few hundred micrometers. Unlike higher plants, microalgae do not have roots, stems, or leaves. They are specially adapted to an environment dominated by viscous forces.

Mediterranean mussel

Mediterranean mussel

The Mediterranean mussel is a species of bivalve, a marine mollusc in the family Mytilidae. It is an invasive species in many parts of the world, and also an object of aquaculture.

Grooved carpet shell

Grooved carpet shell

The grooved carpet shell, or Palourde clam, Ruditapes decussatus, or Venerupis decussatus, is a clam in the family Veneridae. It is distributed worldwide and due to its ecological and economic interest has been proposed as a bioindicator.

Use as food

Flat oysters (Ostrea edulis) from France
Flat oysters (Ostrea edulis) from France

Bivalves have been an important source of food for humans at least since Roman times[102] and empty shells found in middens at archaeological sites are evidence of earlier consumption.[86] Oysters, scallops, clams, ark clams, mussels 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 (FAO) started making such information available, world trade in bivalve molluscs was 1,007,419 tons.[103] 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.[103] China increased its consumption 400-fold during the period 1970 to 1997.[104]

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 spp. or by viruses and bacteria from sewage effluent 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.[104][105] Another possible source of contamination occurs when bivalves contain marine biotoxins as a result of ingesting numerous dinoflagellates. These microalgae are not associated with sewage but occur unpredictably as algal blooms. 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.[104]

Viral and bacterial infections

In 1816 in France, a physician, J. P. A. Pasquier, described an outbreak of typhoid linked to the consumption of raw oysters. The first report of this kind in the United States was in Connecticut 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 have been killed by the treatment of sewage.[104]

Since the 1970s, outbreaks of oyster-vectored diseases have occurs throughout the world. The mortality rate of one disease causing bacteria Vibrio vulnificus, was high at 50%.[104] 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.[106] In 1988, an outbreak of hepatitis A associated with the consumption of inadequately cooked clams (Anadara subcrenata) took place in the Shanghai area of China. An estimated 290,000 people were infected and there were 47 deaths.[107] 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.[104]

Paralytic shellfish poisoning

Paralytic shellfish poisoning (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 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 established the connection between blooms of these phytoplankton and PSP.[108] The biotoxin remains potent even when the shellfish are well-cooked.[108] In the United States, there is a regulatory limit of 80 µg/g of saxitoxin equivalent in shellfish meat.[108]

Amnesic shellfish poisoning

Amnesic shellfish poisoning (ASP) was first reported in eastern Canada in 1987. It is caused by the substance domoic acid found in certain diatoms of the genus Pseudo-nitzschia. Bivalves can become toxic when they filter these microalgae out of the water. Domoic acid is a low-molecular weight amino acid that is able to destroy brain cells causing memory loss, gastroenteritis, 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.[108] In the United States and Canada, a regulatory limit of 20 µg/g of domoic acid in shellfish meat is set.[109]

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Ostrea edulis

Ostrea edulis

Ostrea edulis, commonly known as the European flat oyster, is a species of oyster native to Europe. In Britain and Ireland, regional names include Colchester native oyster, mud oyster, or edible oyster. In France, Ostrea edulis are known as huîtres plates except for those that come from the Belon River estuary in Brittany, France, which are known as Belons.

Oyster

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 all oysters are in the superfamily Ostreoidea.

Clam

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 shells of equal size connected by two adductor muscles and have a powerful burrowing foot. They live in both freshwater and marine environments; in salt water they prefer to burrow down into the mud and the turbidity of the water required varies with species and location; the greatest diversity of these is in North America.

Ark clam

Ark clam

Ark clam is the common name for a family of small to large-sized saltwater clams or marine bivalve molluscs in the family Arcidae. Ark clams vary both in shape and size. They number about 200 species worldwide.

Mussel

Mussel

Mussel is the common name used for members of several families of bivalve molluscs, from saltwater and freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other edible clams, which are often more or less rounded or oval.

Cockle (bivalve)

Cockle (bivalve)

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.

Food and Agriculture Organization

Food and Agriculture Organization

The Food and Agriculture Organization of the United Nations (FAO) is an international organization that leads international efforts to defeat hunger and improve nutrition and food security. Its Latin motto, fiat panis, translates to "let there be bread." It was founded on 16 October 1945.

Effluent

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 pollutants depending on the source. Treating wastewater efficiently is challenging, but improved technology allows for enhanced removal of specific materials, increased re-use of water, and energy production from waste.

Dinoflagellate

Dinoflagellate

The dinoflagellates are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are mostly marine plankton, but they also are common in freshwater habitats. Their populations vary with sea surface temperature, salinity, and depth. Many dinoflagellates are photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey.

Algal bloom

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 encompasses many types of aquatic photosynthetic organisms, both macroscopic multicellular organisms like seaweed and microscopic unicellular organisms like cyanobacteria. Algal bloom commonly refers to the rapid growth of microscopic unicellular algae, not macroscopic algae. An example of a macroscopic algal bloom is a kelp forest.

Connecticut

Connecticut

Connecticut is the southernmost state in the New England region of the Northeastern United States. As of the 2020 United States census, Connecticut was home to over 3.6 million residents, its highest decennial count count ever, growing every decade since 1790. The state is bordered by Rhode Island to its east, Massachusetts to its north, New York to its west, and Long Island Sound to its south. Its capital is Hartford, and its most populous city is Bridgeport. Historically, the state is part of New England as well as the tri-state area with New York and New Jersey. The state is named for the Connecticut River which approximately bisects the state. The word "Connecticut" is derived from various anglicized spellings of "Quinnetuket”, a Mohegan-Pequot word for "long tidal river".

Coliform bacteria

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 aerobes, and are a commonly used indicator of low sanitary quality of foods, milk, and water. Coliforms can be found in the aquatic environment, in soil and on vegetation; they are universally present in large numbers in the feces of warm-blooded animals as they are known to inhabit the gastrointestinal system. While coliform bacteria are not normally causes of serious illness, they are easy to culture, and their presence is used to infer that other pathogenic organisms of fecal origin may be present in a sample, or that said sample is not safe to consume. Such pathogens include disease-causing bacteria, viruses, or protozoa and many multicellular parasites.

Ecosystem services

Nutrient extraction services provided by bivalves. Blue mussels are used as examples but other bivalves like oysters can also provide these nutrient extraction services.[110]
Nutrient extraction services provided by bivalves. Blue mussels are used as examples but other bivalves like oysters can also provide these nutrient extraction services.[110]

Ecosystem services 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, thereby transforming particulate organic matter into bivalve tissue or larger faecal pellets that are transferred to the benthos. 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 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 material. This leads to partial transformation of particulate-bound nutrients into dissolved nutrients via bivalve excretion or enhanced mineralization of faecal material.[110]

When they live in polluted waters, bivalve molluscs have a tendency to accumulate substances such as heavy metals and persistent organic pollutants 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 the presence and quantity of pollutants in their environment.[111]

Economic value of bivalve nutrient extraction, linking processes to services to economic values.[110]
Economic value of bivalve nutrient extraction, linking processes to services to economic values.[110]

There are limitations to the use of bivalves as bioindicators. 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 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.[112]

A study of nine different bivalves with widespread distributions in tropical marine waters concluded that the mussel, Trichomya hirsuta, 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.[113] In the Persian Gulf, the Atlantic pearl-oyster (Pinctada radiata) is considered to be a useful bioindicator of heavy metals.[114]

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.[115] The rock oyster (Saccostrea cucullata) has been shown to reduce the levels of copper 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.[116]

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Blue mussel

Blue mussel

The blue mussel, also known as the common mussel, is a medium-sized edible marine bivalve mollusc in the family Mytilidae, the mussels. Blue mussels are subject to commercial use and intensive aquaculture. A species with a large range, empty shells are commonly found on beaches around the world.

Oyster

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 all oysters are in the superfamily Ostreoidea.

Phytoplankton

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 φυτόν, meaning 'plant', and πλαγκτός, meaning 'wanderer' or 'drifter'.

Particulate organic matter

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.

Benthos

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. This community lives in or near marine or freshwater sedimentary environments, from tidal pools along the foreshore, out to the continental shelf, and then down to the abyssal depths.

Nutrient cycle

Nutrient cycle

A nutrient cycle is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cyclic. Mineral cycles include the carbon cycle, sulfur cycle, nitrogen cycle, water cycle, phosphorus cycle, oxygen cycle, among others that continually recycle along with other mineral nutrients into productive ecological nutrition.

Denitrification

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 denitrification as a type of respiration that reduces oxidized forms of nitrogen in response to the oxidation of an electron donor such as organic matter. The preferred nitrogen electron acceptors in order of most to least thermodynamically favorable include nitrate (NO3−), nitrite (NO2−), nitric oxide (NO), nitrous oxide (N2O) finally resulting in the production of dinitrogen (N2) completing the nitrogen cycle. Denitrifying microbes require a very low oxygen concentration of less than 10%, as well as organic C for energy. Since denitrification can remove NO3−, reducing its leaching to groundwater, it can be strategically used to treat sewage or animal residues of high nitrogen content. Denitrification can leak N2O, which is an ozone-depleting substance and a greenhouse gas that can have a considerable influence on global warming.

Biomonitoring

Biomonitoring

In analytical chemistry, biomonitoring is the measurement of the body burden of toxic chemical compounds, elements, or their metabolites, in biological substances. Often, these measurements are done in blood and urine. Biomonitoring is performed in both environmental health, and in occupational safety and health as a means of exposure assessment and workplace health surveillance.

Bioindicator

Bioindicator

A bioindicator is any 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 small water crustaceans that are present in many water bodies can be monitored for changes that may indicate a problem within their ecosystem. Bioindicators can tell us about the cumulative effects of different pollutants in the ecosystem and about how long a problem may have been present, which physical and chemical testing cannot.

Persian Gulf

Persian Gulf

The Persian Gulf, sometimes called the Arabian Gulf, is a mediterranean sea in Western Asia. The body of water is an extension of the Indian Ocean located between Iran and the Arabian Peninsula. It is connected to the Gulf of Oman in the east by the Strait of Hormuz. The Shatt al-Arab river delta forms the northwest shoreline.

Pinctada radiata

Pinctada radiata

Pinctada radiata, commonly known as the Atlantic pearl-oyster or the Gulf pearl oyster is a species of pearl oyster distributed throughout the Indo-Pacific and in the Mediterranean. Its range extends as far north as Japan and as far south as the Australian state of Victoria.

PH

PH

In chemistry, pH, also referred to as acidity, historically denoting "potential of hydrogen", is a scale used to specify the acidity or basicity of an aqueous solution. Acidic solutions are measured to have lower pH values than basic or alkaline solutions.

Other uses

Carved shell miniatures
Carved shell miniatures

Conchology 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, which houses in excess of 20 million specimens.[117]

1885 wampum belt
1885 wampum belt
Freshwater mussel shell used for making buttons
Freshwater mussel shell used for making buttons
Carved nacre in a 16th-century altarpiece
Carved nacre in a 16th-century altarpiece

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.[118]

The indigenous peoples of the Americas living near the east coast used pieces of shell as wampum. The channeled whelk (Busycotypus canaliculatus) and the quahog (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.[119] The Winnebago Tribe 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 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.[120]

Buttons have traditionally been made from a variety of freshwater and marine shells.[121] 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 in the Indus Valley.[122]

Sea silk is a fine fabric woven from the byssus threads of bivalves, particularly the pen shell (Pinna nobilis). It used to be produced in the Mediterranean region where these shells are endemic. It was an expensive fabric and overfishing has much reduced populations of the pen shell.[123]

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.[124]

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.[125]

A pearl 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 in the family Pteriidae and freshwater mussels in the families Unionidae and Margaritiferidae 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.[126]

Symbolism

The scallop is the symbol of St James and is called Coquille Saint-Jacques in French. It is an emblem carried by pilgrims on their way to the shrine of Santiago de Compostela 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.[127]

Roman myth has it that Venus, the goddess of love, was born in the sea and emerged accompanied by fish and dolphins, with Botticelli 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.[128] From this, the scallop and other bivalve shells came to be used as a symbol for fertility.[129] Its depiction is used in architecture, furniture and fabric design and it is the logo of Royal Dutch Shell, the global oil and gas company.[130]

Discover more about Other uses related topics

Conchology

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 includes the study of land and freshwater mollusc shells as well as seashells and extends to the study of a gastropod's operculum.

Altarpiece

Altarpiece

An altarpiece is an artwork such as a painting, sculpture or relief representing a religious subject made for placing at the back of or behind the altar of a Christian church. Though most commonly used for a single work of art such as a painting or sculpture, or a set of them, the word can also be used of the whole ensemble behind an altar, otherwise known as a reredos, including what is often an elaborate frame for the central image or images. Altarpieces were one of the most important products of Christian art especially from the late Middle Ages to the era of the Counter-Reformation.

Indigenous peoples of the Americas

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.

Ho-Chunk

Ho-Chunk

The Ho-Chunk, also known as Hoocągra or Winnebago, are a Siouan-speaking Native American people whose historic territory includes parts of Wisconsin, Minnesota, Iowa, and Illinois. Today, Ho-Chunk people are enrolled in two federally recognized tribes, the Ho-Chunk Nation of Wisconsin and the Winnebago Tribe of Nebraska.

Mohenjo-daro

Mohenjo-daro

Mohenjo-daro is an archaeological site in the province of Sindh, Pakistan. Built around 2500 BCE, it was the largest settlement of the ancient Indus Valley Civilisation, and one of the world's earliest major cities, contemporaneous with the civilizations of ancient Egypt, Mesopotamia, Minoan Crete, and Norte Chico. With an estimated population of at least 40,000 people, Mohenjo-daro prospered until around 1700 BCE.

Sea silk

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. The byssus is used by the clam to attach itself to the sea bed.

Byssus

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 (Mytilidae), and Dreissenidae.

Pinna nobilis

Pinna nobilis

Pinna nobilis, whose common name is 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.

Endemism

Endemism

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 elsewhere. For example, the Cape sugarbird is found exclusively in southwestern South Africa and is therefore said to be endemic to that particular part of the world.

Overfishing

Overfishing

Overfishing is the removal of a species of fish from a body of water at a rate greater than that the species can replenish its population naturally, resulting in the species becoming increasingly underpopulated in that area. Overfishing can occur in water bodies of any sizes, such as ponds, wetlands, rivers, lakes or oceans, and can result in resource depletion, reduced biological growth rates and low biomass levels. Sustained overfishing can lead to critical depensation, where the fish population is no longer able to sustain itself. Some forms of overfishing, such as the overfishing of sharks, has led to the upset of entire marine ecosystems. Types of overfishing include: growth overfishing, recruitment overfishing, ecosystem overfishing.

Marquetry

Marquetry

Marquetry is the art and craft of applying pieces of veneer to a structure to form decorative patterns, designs. The technique may be applied to case furniture or even seat furniture, to decorative small objects with smooth, veneerable surfaces or to freestanding pictorial panels appreciated in their own right.

CITES

CITES

CITES is a multilateral treaty to protect endangered plants and animals from the threats of international trade. It was drafted as a result of a resolution adopted in 1963 at a meeting of members of the International Union for Conservation of Nature (IUCN). The convention was opened for signature in 1973 and CITES entered into force on 1 July 1975.

Bivalvian taxonomies

Mussels in the intertidal zone in Cornwall, England
Mussels in the intertidal zone in Cornwall, England
Mytilarca is a distant relative of the mussels; from the Middle Devonian of Wisconsin.
Mytilarca is a distant relative of the mussels; from the Middle Devonian of Wisconsin.
Fossil gastropod and attached mytilid bivalves in a Jurassic limestone (Matmor Formation) in southern Israel
Fossil gastropod and attached mytilid bivalves in a Jurassic limestone (Matmor Formation) in southern Israel
Aviculopecten subcardiformis; a fossil of an extinct scallop from the Logan Formation of Wooster, Ohio (external mold)
Aviculopecten subcardiformis; a fossil of an extinct scallop from the Logan Formation of Wooster, Ohio (external mold)

For the past two centuries no consensus has existed on bivalve phylogeny 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 in Part N of the Treatise on Invertebrate Paleontology,[131] which employed a classification system based on general shell shape, microstructures and hinge configuration.[132] 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 analyses of multiple organ systems, shell morphology (including fossil species) and modern molecular phylogenetics have resulted in the drawing up of what experts believe is a more accurate phylogeny of the Bivalvia.[133][134][135][136][137] Based upon these studies, a new proposed classification system for the Bivalvia was published in 2010 by Bieler, Carter & Coan.[138] In 2012, this new system was adopted by the World Register of Marine Species (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. Molecular phylogenetic work continues, further clarifying which Bivalvia are most closely related and thus refining the classification.[139][140]

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)
Schizodonta: Big bifurcating teeth (e.g. Trigonia spp.)
Isodonta: Equal teeth (e.g. Spondylus spp.)
Dysodonta: Absent teeth and ligaments joins the valves.
Subclass:Teleodesmacea
Order
Heterodonta: Different teeth (e.g. family Cardiidae). [ Lower Ordovician – Recent]
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.[141][142]

Taxonomy based upon hinge tooth morphology

The systematic layout presented here follows Newell's 1965 classification based on hinge tooth morphology (all taxa marked † are extinct) :[132]

Subclass Order
Palaeotaxodonta Nuculoida (nut shells)
Cryptodonta Praecardioida

Solemyoida

Pteriomorphia Arcoida (ark shells)

Cyrtodontoida

Limoida (file shells)

Mytiloida (true mussels)

Ostreoida (oysters, formerly included in Pterioida)

Praecardioida

Pterioida (pearl oysters, pen shells)

Palaeoheterodonta Trigonioida (Neotrigonia is the only extant genus)

Unionoida (freshwater mussels)

Modiomorpha

Heterodonta Cycloconchidae

Hippuritoida

Lyrodesmatidae

Myoida (soft-shell clams, geoducks, shipworms)

Redoniidae

Veneroida (hard-shell clams, cockles, razor shells)

Anomalodesmata Pholadomyoida

The monophyly of the subclass Anomalodesmata is disputed. The standard view now is that it resides within the subclass Heterodonta.[133][136][143]

Taxonomy based upon gill morphology

An alternative systematic scheme exists using gill morphology.[144] 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 and are carnivorous, having a muscular septum instead of filamentous gills.[145]

2010 taxonomy

In May 2010, a new taxonomy of the Bivalvia was published in the journal Malacologia. 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 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.[138] This classification has since been adopted by WoRMS.[146]

Proposed classification of Class Bivalvia (under the redaction of Rüdiger Bieler, Joseph G. Carter and Eugene V. Coan) (all taxa marked † are extinct) :[147]

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Cornwall

Cornwall

Cornwall is a historic county and ceremonial county in South West England. It is recognised as one of the Celtic nations, and is the homeland of the Cornish people. Cornwall is bordered to the north and west by the Atlantic Ocean, to the south by the English Channel, and to the east by the county of Devon, with the River Tamar forming the border between them. Cornwall forms the westernmost part of the South West Peninsula of the island of Great Britain. The southwesternmost point is Land's End and the southernmost Lizard Point. Cornwall has a population of 568,210 and an area of 3,563 km2 (1,376 sq mi). The county has been administered since 2009 by the unitary authority, Cornwall Council. The ceremonial county of Cornwall also includes the Isles of Scilly, which are administered separately. The administrative centre of Cornwall is Truro, its only city.

Devonian

Devonian

The Devonian is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, 419.2 million years ago (Mya), to the beginning of the Carboniferous, 358.9 Mya. It is named after Devon, England, where rocks from this period were first studied.

Mytilidae

Mytilidae

Mytilidae are a family of small to large marine and brackish-water bivalve molluscs in the order Mytilida. One of the genera, Limnoperna, even inhabits freshwater environments. The order has only this one family which contains some 52 genera.

Limestone

Limestone

Limestone is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of CaCO3. Limestone forms when these minerals precipitate out of water containing dissolved calcium. This can take place through both biological and nonbiological processes, though biological processes, such as the accumulation of corals and shells in the sea, have likely been more important for the last 540 million years. Limestone often contains fossils which provide scientists with information on ancient environments and on the evolution of life.

Matmor Formation

Matmor Formation

The Matmor Formation is a geologic formation of up to 100 metres (330 ft) thick, that is exposed in Hamakhtesh Hagadol in southern Israel. The Matmor Formation contains fossils from a Jurassic equatorial shallow marine environment. Bivalves, gastropods, sponges, corals, echinoderms, and sclerobionts are present in the Matmor Formation to various degrees. The stratigraphy of the Matmor Formation consists of alternating layers of limestone and marl.

Israel

Israel

Israel, officially the State of Israel, is a country in Western Asia. Situated in the Southern Levant, it is bordered by Lebanon to the north, by Syria to the northeast, by Jordan to the east, by the Red Sea to the south, by Egypt to the southwest, by the Mediterranean Sea to the west, and by the Palestinian territories — the West Bank along the east and the Gaza Strip along the southwest. Tel Aviv is the economic and technological center of the country, while its seat of government is in its proclaimed capital of Jerusalem, although Israeli sovereignty over East Jerusalem is unrecognized internationally.

Fossil

Fossil

A fossil 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 amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Scallop

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 within the superfamily Pectinoidea, which also includes the thorny oysters.

Logan Formation

Logan Formation

The Logan Formation is the name given to a Lower Carboniferous siltstone, sandstone and conglomeratic unit exposed in east-central Ohio and parts of western West Virginia, USA.

Norman D. Newell

Norman D. Newell

Norman Dennis Newell 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.

Cladistics

Cladistics

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 characteristics (synapomorphies) that are not present in more distant groups and ancestors. However, from an empirical perspective, common ancestors are inferences based on a cladistic hypothesis of relationships of taxa whose character states can be observed. Theoretically, a last common ancestor and all its descendants constitute a (minimal) clade. Importantly, all descendants stay in their overarching ancestral clade. For example, if the terms worms or fishes were used within a strict cladistic framework, these terms would include humans. Many of these terms are normally used paraphyletically, outside of cladistics, e.g. as a 'grade', which are fruitless to precisely delineate, especially when including extinct species. Radiation results in the generation of new subclades by bifurcation, but in practice sexual hybridization may blur very closely related groupings.

Molecular phylogenetics

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 determine the processes by which diversity among species has been achieved. The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree. Molecular phylogenetics is one aspect of molecular systematics, a broader term that also includes the use of molecular data in taxonomy and biogeography.

Source: "Bivalvia", Wikipedia, Wikimedia Foundation, (2023, March 4th), https://en.wikipedia.org/wiki/Bivalvia.

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References
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  147. ^ Carter, J. G.; Altaba, C. R.; Anderson, L. C.; Araujo, R.; Biakov, A. S.; Bogan, A. E.; Campbell, D. C.; Campbell, M.; Chen, J.; Cope, J. C. W.; Delvene. G.; Dijkstra, H. H.; Fang, Z.; Gardner, R. N.; Gavrilova, V. A.; Goncharova, I. A.; Harries, P. J.; Hartman, J. H.; Hautmann, M.; Hoeh, W. R.; Hylleberg, J.; Jiang, B.; Johnston, P.; Kirkendale, L.; Kleemann, K.; Koppka, J.; Kříž, J.; Machado, D.; Malchus, N.; Márquez-Aliaga, A.; Masse, J-P.; McRoberts, C. A.; Middelfart, P. U.; Mitchell, S.; Nevesskaja, L. A.; Özer, S.; Pojeta, J. Jr.; Polubotko, I. V.; Pons, J. M.; Popov, S.; Sánchez, T.; Sartori, A. F.; Scott, R. W.; Sey, I. I.; Signorelli, J. H.; Silantiev, V. V.; Skelton, P. W.; Steuber, T.; Waterhouse, J. B.; Wingard, G. L.; Yancey, T. (2011). "A synoptical classification of the Bivalvia (Mollusca)" (PDF). Paleontological Contributions. 4: 1–47.
Further reading
  • Schneider, Jay A. (2001). "Bivalve systematics during the 20th century". Journal of Paleontology. 75 (6): 1119–1127. doi:10.1666/0022-3360(2001)0752.0.CO;2. ISSN 0022-3360.
  • Poutiers, J.-M.; Bernard, F. R. (1995). "Carnivorous bivalve molluscs (Anomalodesmata) from the tropical western Pacific Ocean, with a proposed classification and a catalogue of recent species". In Bouchet, P. (ed.). Résultats des Campagnes Musorstom. Mémoires Muséum National d'Histoire Naturelle. Vol. 167. pp. 107–188.
  • Vaught, K. C. (1989). A Classification of the Living Mollusca. American Malacologists. ISBN 978-0-915826-22-3.

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