Bivalvia

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. 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 , meaning 'two', and , meaning 'leaves of a door' members of the phylum Brachiopoda and the minute crustaceans known as ostracods and conchostracans. == Anatomy ==

(Margaritifera margaritifera) anatomy: Bivalves have bilaterally symmetrical and laterally flattened bodies, with a blade-shaped foot, vestigial head and no radula. 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. 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. The outer surface of the valves is often sculpted, with clams often having concentric striations, scallops having radial ribs and oysters a latticework of irregular markings. In all molluscs, the mantle forms a thin membrane 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. The posterior mantle edge may have two elongated extensions known as siphons, through one of which water is inhaled, and the other expelled. The siphons retract into a cavity, known as the pallial sinus. The shell grows larger when more material is secreted by the mantle edge, and the valves themselves thicken as more material is secreted from the general mantle surface. Calcareous matter comes from both its diet and the surrounding seawater. Concentric rings on the exterior of a valve are commonly used to age bivalves. For some groups, a more precise method for determining the age of a shell is by cutting a cross section through it and examining the incremental growth bands. The 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. 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. Retractor muscles connect the mantle to the edge of the shell, along a line known as the pallial line. In species that can swim by flapping their valves, a single, central adductor muscle occurs. These muscles are composed of two types of muscle fibres, striated muscle bundles for fast actions and smooth muscle bundles for maintaining a steady pull. Paired pedal protractor and retractor muscles operate the animal's foot. Nervous system The sedentary habits of the bivalves have meant that in general the nervous system 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. In the order Anomalodesmata, the inhalant siphon is surrounded by vibration-sensitive tentacles for detecting prey. Many bivalves have no eyes, but a few members of the Arcoidea, Limopsoidea, Mytiloidea, Anomioidea, Ostreoidea, and Limoidea have simple eyes on the margin of the mantle. These consist of a pit of photosensory cells and a lens. All bivalves have light-sensitive cells that can detect a shadow falling over the animal. The hemolymph usually lacks any respiratory pigment. In the carnivorous genus Poromya, the hemolymph has red amoebocytes containing a haemoglobin pigment. 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. 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. 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. 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. Waste material is consolidated in the rectum and voided as pellets into the exhalent water stream through an anal pore. Feeding and digestion are synchronized with diurnal and tidal cycles. Carnivorous bivalves generally have reduced crystalline styles and the stomach has thick, muscular walls, extensive cuticular linings and diminished sorting areas and gastric chamber sections. 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. Reproduction and development The sexes are usually separate in bivalves but some hermaphroditism is known. The gonads either open into the nephridia or through a separate pore into a chamber over the gills. The ripe gonads of males and females release sperm and eggs into the water column. 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. 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. 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. 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. 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. == Comparison with brachiopods ==

'', a bivalve with taxodont dentition from the Pliocene of Cyprus support intact Brachiopods are shelled marine organisms that superficially resemble 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. 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. == Evolutionary history ==

The Cambrian explosion took place around 540 to 520 million years ago (Mya). In this geologically brief period, most major animal phyla diverged including some of the first creatures with mineralized skeletons. Brachiopods and bivalves made their appearance at this time, and left their fossilized remains behind in the rocks. Possible early bivalves include Pojetaia 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. Only five genera of supposed Cambrian "bivalves" exist, the others being Tuarangia, Camya and Arhouriella and potentially Buluniella. 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. 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. == Diversity of extant bivalves ==

The adult maximum size of living species of bivalve ranges from in Condylonucula maya, a nut clam, to a length of in Kuphus polythalamia, an elongated, burrowing shipworm. However, the species generally regarded as the largest living bivalve is the giant clam Tridacna gigas, which can grow to a length of and a weight of more than 200 kg (441 lb). The largest known extinct bivalve is a species of Platyceramus whose fossils measure up to in length. In his 2010 treatise, Compendium of Bivalves, Markus Huber gives the total number of living bivalve species as about 9,200 combined in 106 families. Huber states that the number of 20,000 living species, often encountered in literature, could not be verified and presents the following table to illustrate the known diversity: == Distribution ==

The bivalves are a highly successful class of invertebrates found in aquatic habitats throughout the world. Most are 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. 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. 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. Some species are found in the hadal zone, like Vesicomya sergeevi, which occurs at depths of 7600–9530 meters. 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. 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. 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. 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. == Behaviour ==

visible 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. Other bivalves, such as mussels, attach themselves to hard surfaces using tough byssus threads made of collagen and elastin proteins. Some species, including the true oysters, the jewel boxes, the jingle shells, the thorny oysters and the kitten's paws, cement themselves to stones, rock or larger dead shells. 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. 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). Birds such as the Eurasian oystercatcher (Haematopus ostralegus) have specially adapted beaks which can pry open their shells. The herring gull (Larus argentatus) sometimes drops heavy shells onto rocks in order to crack them open. 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. The Pacific walrus (Odobenus rosmarus divergens) is one of the main predators feeding on bivalves in Arctic waters. Shellfish have formed part of the human diet since prehistoric times, a fact evidenced by the remains of mollusc shells found in ancient middens. Examinations of these deposits in Peru has provided a means of dating long past El Niño events because of the disruption these caused to bivalve shell growth. 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. 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 and the Atlantic jackknife clam, Ensis directus, can do the same within fifteen seconds. Scallops and file clams can swim by opening and closing their valves rapidly; water is ejected on either side of the hinge area and they move with the flapping valves in front. 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. In some other 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. 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. == Mariculture ==

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. 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. Production of bivalve molluscs by mariculture in 2010 was 12,913,199 tons, up from 8,320,724 tons in 2000. Culture of clams, cockles and ark shells more than doubled over this time period from 2,354,730 to 4,885,179 tons. Culture of mussels over the same period grew from 1,307,243 to 1,812,371 tons, of oysters from 3,610,867 to 4,488,544 tons and of scallops from 1,047,884 to 1,727,105 tons. == Use as food ==

'') from France Bivalves have been an important source of food for humans at least since Roman times and empty shells found in middens at archaeological sites are evidence of earlier consumption. 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. 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. ==Ecosystem services==

s are used as examples but other bivalves like oysters can also provide these nutrient extraction services. 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. , the Atlantic pearl-oyster (Pinctada radiata) is considered to be a useful bioindicator of heavy metals. Crushed shells, available as a by-product of the seafood canning industry, can be used to remove pollutants from water. It has been found that, as long as the water is maintained at an alkaline pH, crushed shells will remove cadmium, lead and other heavy metals from contaminated waters by swapping the calcium in their constituent aragonite for the heavy metal, and retaining these pollutants in a solid form. The rock oyster (Saccostrea cucullata) 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. == Other uses ==

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. Shells are used decoratively in many ways. They can be pressed into concrete or plaster to make decorative paths, steps or walls and can be used to embellish picture frames, mirrors or other craft items. They can be stacked up and glued together to make ornaments. They can be pierced and threaded onto necklaces or made into other forms of jewellery. Shells have had various uses in the past as body decorations, utensils, scrapers and cutting implements. Carefully cut and shaped shell tools dating back 32,000 years have been found in a cave in Indonesia. In this region, shell technology may have been developed in preference to the use of stone or bone implements, perhaps because of the scarcity of suitable rock materials. The indigenous peoples of the Americas 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. The Ho-Chunk people from Wisconsin had numerous uses for freshwater mussels including using them as spoons, cups, ladles and utensils. They notched them to provide knives, graters and saws. They carved them into fish hooks and lures. They incorporated powdered shell into clay to temper 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. Buttons have traditionally been made from a variety of freshwater and marine shells. 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. Crushed shells are added as a calcareous supplement to the diet of laying poultry. Oyster shell and cockle shell are often used for this purpose and are obtained as a by-product from other industries. Pearls and mother-of-pearl Mother-of-pearl or nacre is the naturally occurring lustrous layer that lines some mollusc shells. It is used to make pearl buttons and in artisan craftwork to make organic jewellery. It has traditionally been inlaid into furniture and boxes, particularly in China. It has been used to decorate musical instruments, watches, pistols, fans and other products. The import and export of goods made with nacre are controlled in many countries under the International Convention of Trade in Endangered Species of Wild Fauna and Flora. A pearl is created in the mantle of a mollusc 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. 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. 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. From this, the scallop and other bivalve shells came to be used as a symbol for fertility. Its depiction is used in architecture, furniture and fabric design and it is the logo of Royal Dutch Shell, the global oil and gas company. == Bivalvian taxonomies ==

, England of Wisconsin. and attached mytilid bivalves in a Jurassic limestone (Matmor Formation) in southern Israel ''; 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, which employed a classification system based on general shell shape, microstructures and hinge configuration. 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. 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. Proposed classification of Class Bivalvia (under the redaction of Rüdiger Bieler, Joseph G. Carter and Eugene V. Coan) (all taxa marked † are extinct) : Grade Euprotobranchia • Order Fordillida :: 2 families (2†) • Order Tuarangiida :: 1 family (1†) Subclass Heterodonta Infraclass Archiheterodonta • Order Carditida :: 4 families Infraclass Euheterodonta • Unassigned Euheterodonta :: 4 families • Order Pholadomyida (=Anomalodesmata) :: 16 families • Order Myida :: 4 families • Order Lucinida :: 2 families • Order Venerida :: 30 families Subclass Palaeoheterodonta • Order Trigoniida :: 16 families (15†) • Order Unionida :: 15 families (8†) Subclass Protobranchia • Order Nuculanida :: 8 families • Order Nuculida :: 3 families (1†) • Order Solemyida :: 2 families Subclass Pteriomorphia • Order Arcida :: 7 families Infraclass Eupteriomorphia • Order Ostreida :: 2 families • Suborder Pectinida :: 7 families • Suborder Limida :: 1 family • Suborder Mytilida :: 1 family • Suborder Pteriida :: 4 families == References ==