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87-515: And see text Bivalvia ( / b aɪ ˈ v æ l v i ə / ) or bivalves , in previous centuries referred to as the Lamellibranchiata and Pelecypoda , is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consisting of a hinged pair of half- shells known as valves . As a group, bivalves have no head and lack some typical molluscan organs such as

174-466: A convenient "artificial key" according to his Systema Sexuale , largely based on the arrangement of flowers. In botany, classes are now rarely discussed. Since the first publication of the APG system in 1998, which proposed a taxonomy of the flowering plants up to the level of orders, many sources have preferred to treat ranks higher than orders as informal clades . Where formal ranks have been assigned,

261-420: 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

348-502: A distinct head, and most of them without feet. They are principally distinguished by their tentacles (or feelers). By the Ancients they were not improperly called imperfect animals, as being destitute of ears, nose, head, eyes and legs; and are therefore totally distinct from Insects." Linnaeus divided the "Vermes" based upon the structure of the body, into the following orders and genera: The second volume, published in 1759, detailed

435-527: 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 . 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

522-403: 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

609-459: 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

696-457: 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 . Class (biology) The class as a distinct rank of biological classification having its own distinctive name – and not just called a top-level genus (genus summum) – was first introduced by French botanist Joseph Pitton de Tournefort in

783-464: A long time without food, are tenacious of life, and have the power of reproducing parts which have been destroyed or lost; some undergo a metamorphosis; some cast (shed) their skin; some appear to live promiscuously on land or in the water, and some are torpid during the winter." Linnaeus divided the amphibians based upon the limb structures and the way they breathed, into the following orders and genera: Linnaeus described fish as: "Always inhabiting

870-497: 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. The sexes are usually separate in bivalves but some hermaphroditism

957-483: 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

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1044-643: A resting state, even when they are permanently submerged. In oysters, for example, their behaviour follows very strict circatidal and circadian rhythms according to the relative positions of the moon and sun. During neap tides, they exhibit much longer closing periods than during spring tides. Although many non-sessile bivalves use their muscular foot to move around, or to dig, members of the freshwater family Sphaeriidae 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

1131-423: A shadow falling over the animal. 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

1218-436: 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,

1305-430: 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 ligament and byssus, if calcified, are composed of aragonite. The outermost layer of

1392-554: 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

1479-408: 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

1566-521: Is to say a particular layout of organ systems. This said, the composition of each class is ultimately determined by the subjective judgment of taxonomists . In the first edition of his Systema Naturae (1735), Carl Linnaeus divided all three of his kingdoms of nature ( minerals , plants , and animals ) into classes. Only in the animal kingdom are Linnaeus's classes similar to the classes used today; his classes and orders of plants were never intended to represent natural groups, but rather to provide

1653-430: 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

1740-610: 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 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. By

1827-725: 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 . 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

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1914-586: 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. 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,

2001-408: 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

2088-596: 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. 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

2175-466: The radula and the odontophore . Their gills have evolved into ctenidia , specialised organs for feeding and breathing. Common bivalves include clams , oysters , cockles , mussels , scallops , and numerous other families that live in saltwater, as well as a number of families that live in freshwater. Majority of the class are benthic filter feeders that bury themselves in sediment, where they are relatively safe from predation . Others lie on

2262-400: 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

2349-456: The Arctic, about 140 species being known from that zone. 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

2436-498: The English title is A General System of Nature ) 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 . Before 1758, most biological catalogues had used polynomial names for

2523-462: 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

2610-469: The air, can gape the shell slightly and gas exchange can take place. 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. Most bivalves are filter feeders , using their gills to capture particulate food such as phytoplankton from the water. Protobranchs feed in

2697-424: The animal's foot. 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

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2784-479: The application of their different perceptions. They all originate from an egg. Their external and internal structure; their comparative anatomy, habits, instincts, and various relations to each other, are detailed in authors who professedly treat on their subjects." The list has been broken down into the original six classes Linnaeus described for animals; Mammalia , Aves , Amphibia , Pisces , Insecta , and Vermes . These classes were ultimately created by studying

2871-620: The birds based upon the characters of the bill and feet, into the following 6 orders and 63 genera: Linnaeus described his "Amphibia" (comprising reptiles and amphibians ) as: "Animals that are distinguished by a body cold and generally naked; stern and expressive countenance; harsh voice; mostly lurid color; filthy odor; a few are furnished with a horrid poison; all have cartilaginous bones, slow circulation, exquisite sight and hearing, large pulmonary vessels, lobate liver, oblong thick stomach, and cystic, hepatic, and pancreatic ducts: they are deficient in diaphragm, do not transpire (sweat), can live

2958-571: 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'. ("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

3045-420: The classification of plants that appeared in his Eléments de botanique of 1694. Insofar as a general definition of a class is available, it has historically been conceived as embracing taxa that combine a distinct grade of organization—i.e. a 'level of complexity', measured in terms of how differentiated their organ systems are into distinct regions or sub-organs—with a distinct type of construction, which

3132-435: The edges of lakes and ponds; this enables the clam to find the best position for filter feeding. 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

3219-399: The family Juliidae ), members of the phylum Brachiopoda and the minute crustaceans known as ostracods and conchostracans . 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

3306-420: 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 . 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

3393-432: 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,

3480-433: 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. Brachiopods are shelled marine organisms that superficially resemble bivalves in that they are of similar size and have

3567-475: The fishes based upon the position of the ventral and pectoral fins, into the following orders and genera: Linnaeus described his "Insecta" (comprising all arthropods , including insects , crustaceans , arachnids and others) as: "A very numerous and various class consisting of small animals, breathing through lateral spiracles, armed on all sides with a bony skin, or covered with hair; furnished with many feet, and moveable antennae (or horns), which project from

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3654-465: The gills are also much longer than those in more primitive bivalves, and are folded over to create a groove through which food can be transported. The structure of the gills varies considerably, and can serve as a useful means for classifying bivalves into groups. A few bivalves, such as the granular poromya ( Poromya granulata ), are carnivorous , eating much larger prey than the tiny microalgae consumed by other bivalves. Muscles draw water in through

3741-405: The head, and are the probable instruments of sensation." Linnaeus divided the insects based upon the form of the wings, into the following orders and genera: Linnaeus described his "Vermes" as: "Animals of slow motion, soft substance, able to increase their bulk and restore parts which have been destroyed, extremely tenacious of life, and the inhabitants of moist places. Many of them are without

3828-594: 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

3915-431: The hind parts of the animal. The hemolymph usually lacks any respiratory pigment. In the carnivorous genus Poromya , the hemolymph has red amoebocytes containing a haemoglobin pigment. 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

4002-434: 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. The main muscular system in bivalves is the posterior and anterior adductor muscles. These muscles connect the two valves and contract to close

4089-523: 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 . Scallops have more complex eyes with a lens, a two-layered retina , and a concave mirror. All bivalves have light-sensitive cells that can detect

4176-420: The inhalant siphon which is modified into a cowl-shaped organ, sucking in prey. The siphon can be retracted quickly and inverted, bringing the prey within reach of the mouth. The gut is modified so that large food particles can be digested. The unusual genus, Entovalva , is endosymbiotic , being found only in the oesophagus of sea cucumbers . It has mantle folds that completely surround its small valves. When

4263-461: The internal anatomy, as seen in his key: By current standards Pisces and Vermes are informal groupings, Insecta also contained arachnids and crustaceans , and one order of Amphibia comprised sharks , lampreys , and sturgeons . Linnaeus described mammals as: "Animals that suckle their young by means of lactiferous teats. In external and internal structure they resemble man: most of them are quadrupeds; and with man, their natural enemy, inhabit

4350-440: 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. The excretory organs of bivalves are

4437-491: 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

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4524-810: The kingdom Plantae, in which Linnaeus included true plants , as well as fungi , algae and lichens . In addition to repeating the species he had previously listed in his Species Plantarum (1753), and those published in the intervening period, Linnaeus described several hundred new plant species. The species from Species Plantarum were numbered sequentially, while the new species were labelled with letters. Many were sent to Linnaeus by his correspondents overseas, including Johannes Burman and David de Gorter in South Africa , Patrick Browne , Philip Miller and John Ellis in America, Jean-François Séguier , Carlo Allioni and Casimir Christoph Schmidel in

4611-482: 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). The largest known extinct bivalve is a species of Platyceramus whose fossils measure up to 3,000 mm (118 in) 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

4698-482: 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

4785-415: 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

4872-422: 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: 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

4959-506: 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

5046-421: The oysters and scallops, the anterior adductor muscle has been lost and the posterior muscle is positioned centrally. In species that can swim by flapping their valves, a single, central adductor muscle occurs. These muscles are composed of two types of muscle fibres, striated muscle bundles for fast actions and smooth muscle bundles for maintaining a steady pull. Paired pedal protractor and retractor muscles operate

5133-399: The palps. These then sort the particles, rejecting those that are unsuitable or too large to digest, and conveying others to the mouth. In more advanced bivalves, water is drawn into the shell from the posterior ventral 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

5220-523: 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

5307-442: 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. The sensory organs of bivalves are largely located on

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5394-408: 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. In the order Anomalodesmata ,

5481-425: 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

5568-524: The prominence of modern bivalves over brachiopods seems due to chance disparities in their response to extinction events . The adult maximum size of living species of bivalve ranges from 0.52 mm (0.02 in) in Condylonucula maya , a nut clam, to a length of 1,532 millimetres (60.3 in) in Kuphus polythalamia , an elongated, burrowing shipworm. However, the species generally regarded as

5655-480: The ranks have been reduced to a very much lower level, e.g. class Equisitopsida for the land plants, with the major divisions within the class assigned to subclasses and superorders. The class was considered the highest level of the taxonomic hierarchy until George Cuvier 's embranchements , first called Phyla by Ernst Haeckel , were introduced in the early nineteenth century. 10th edition of Systema Naturae The 10th edition of Systema Naturae (Latin;

5742-503: The rules. The only work which takes priority over the 10th edition is Carl Alexander Clerck 's Svenska Spindlar or Aranei Suecici , which was published in 1757, but is also to be treated as if published on January 1, 1758. During Linnaeus' lifetime, Systema Naturae was under continuous revision. Progress was incorporated into new and ever-expanding editions; for example, in his 1st edition (1735), whales and manatees were originally classified as species of fish (as

5829-495: 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. 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

5916-450: The sea floor or attach themselves to rocks or other hard surfaces. Some bivalves, such as scallops and file shells , can swim . 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 valves are for feeding and for disposal of waste. These are joined together along one edge (the hinge line ) by

6003-419: 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

6090-468: The sediment remains damp even when the tide is out. When buried in the sediment, burrowing bivalves are protected from the pounding of waves, desiccation, and overheating during low tide, and variations in salinity caused by rainwater. They are also out of the reach of many predators. Their general strategy is to extend their siphons to the surface for feeding and respiration during high tide, but to descend to greater depths or keep their shell tightly shut when

6177-435: 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

6264-466: The shell is the periostracum , a thin layer composed of horny conchiolin . The periostracum is secreted by the outer mantle and is easily abraded. The outer surface of the valves is often sculpted, with clams often having concentric striations, scallops having radial ribs and oysters a latticework of irregular markings. In all molluscs, the mantle forms a thin membrane that covers the animal's body and extends out from it in flaps or lobes. In bivalves,

6351-570: The shell, gaining the ability to swim, and in a few cases, adopting predatory habits. For a long time, bivalves were thought to be better adapted to aquatic life than brachiopods were, outcompeting and relegating them to minor niches in later ages. These two taxa appeared in textbooks as an example of replacement by competition. Evidence given for this included the fact that bivalves needed less food to subsist because of their energetically efficient ligament-muscle system for opening and closing valves. All this has been broadly disproven, though; rather,

6438-460: 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 . These muscles pull the mantle though the valves. In sedentary or recumbent bivalves that lie on one valve, such as

6525-418: The shells and open them more easily when they could tackle them from different angles. Octopuses either pull bivalves apart by force, or they bore a hole into the shell and insert a digestive fluid before sucking out the liquified contents. Certain carnivorous gastropod snails such as whelks ( Buccinidae ) and murex snails ( Muricidae ) feed on bivalves by boring into their shells. A dog whelk ( Nucella ) drills

6612-400: 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

6699-736: 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. Further changes in shell development due to environmental stress has also been suggested to cause increased mortality in oysters due to reduced shell strength. Invertebrate predators include crustaceans, starfish and octopuses. Crustaceans crack

6786-582: 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

6873-415: The species damages water installations and disrupts local ecosystems . 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

6960-690: The surface of the Earth. The largest, though fewest in number, inhabit the ocean." Linnaeus divided the mammals based upon the number, situation, and structure of their teeth, into the following orders and genera: Linnaeus described birds as: "A beautiful and cheerful portion of created nature consisting of animals having a body covered with feathers and down; protracted and naked jaws (the beak), two wings formed for flight, and two feet. They are areal, vocal, swift and light, and destitute of external ears, lips, teeth, scrotum, womb, bladder, epiglottis, corpus callosum and its arch, and diaphragm." Linnaeus divided

7047-528: The taxa included, including earlier editions of Systema Naturae . The first work to consistently apply binomial nomenclature across the animal kingdom was the 10th edition of Systema Naturae . The International Commission on Zoological Nomenclature therefore chose 1 January 1758 as the "starting point" for zoological nomenclature and asserted that the 10th edition of Systema Naturae was to be treated as if published on that date. Names published before that date are unavailable, even if they would otherwise satisfy

7134-587: The tide goes out. They use their muscular foot to dig into the substrate. To do this, the animal relaxes its adductor muscles and opens its shell wide to anchor itself in position while it extends its foot downwards into the substrate. Then it dilates the tip of its foot, retracts the adductor muscles to close the shell, shortens its foot and draws itself downwards. This series of actions is repeated to dig deeper. Other bivalves, such as mussels , attach themselves to hard surfaces using tough byssus threads made of collagen and elastin proteins. Some species, including

7221-556: The true oysters, the jewel boxes , the jingle shells , the thorny oysters and the kitten's paws , cement themselves to stones, rock or larger dead shells. In oysters, the lower valve may be almost flat while the upper valve develops layer upon layer of thin horny material reinforced with calcium carbonate. Oysters sometimes occur in dense beds in the neritic zone and, like most bivalves, are filter feeders. Bivalves filter large amounts of water to feed and breathe but they are not permanently open. They regularly shut their valves to enter

7308-467: 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. The shell

7395-502: The waters; are swift in their motion and voracious in their appetites. They breathe by means of gills, which are generally united by a bony arch; swim by means of radiate fins, and are mostly covered over with cartilaginous scales. Besides the parts they have in common with other animals, they are furnished with a nictitant membrane, and most of them with a swim-bladder, by the contraction or dilatation of which, they can raise or sink themselves in their element at pleasure." Linnaeus divided

7482-416: 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

7569-426: Was thought to be the case then). In the 10th edition, they were both moved into the mammal class. The animal kingdom (as described by Linnaeus): "Animals enjoy sensation by means of a living organization, animated by a medullary substance; perception by nerves; and motion by the exertion of the will. They have members for the different purposes of life; organs for their different senses; and faculties (or powers) for

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