118-553: See text. The Donacidae , the "bean clams" or "wedge shells", are a family of bivalve molluscs of the superfamily Tellinoidea . The family is related to the Tellina . The Donacidae are prolific filter feeders and are an important part of coastal food chains where they occur. The family is sensitive to coastal industry such as dam-building and dredging . Members of this family have asymmetric, elongated, compressed shells. The two siphons are short but are completely divided, and
236-410: 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 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
354-443: 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 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
472-467: A constant rate. These " molecular clocks ", however, are fallible, and provide only approximate timing: for example, they are not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first evolved, and estimates produced by different techniques may vary by a factor of two. Organisms are only rarely preserved as fossils in the best of circumstances, and only
590-420: 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
708-528: 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
826-426: A fraction of such fossils have been discovered. This is illustrated by the fact that the number of species known through the fossil record is less than 5% of the number of known living species, suggesting that the number of species known through fossils must be far less than 1% of all the species that have ever lived. Because of the specialized and rare circumstances required for a biological structure to fossilize, only
944-495: A hierarchical classification system still in use today. Darwin and his contemporaries first linked the hierarchical structure of the tree of life with the then very sparse fossil record. Darwin eloquently described a process of descent with modification, or evolution, whereby organisms either adapt to natural and changing environmental pressures, or they perish. When Darwin wrote On the Origin of Species by Means of Natural Selection, or
1062-460: 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
1180-402: 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 . Fossil record A fossil (from Classical Latin fossilis , lit. ' obtained by digging ' ) is any preserved remains, impression, or trace of any once-living thing from
1298-418: A mid-Ordovician age. Such index fossils must be distinctive, be globally distributed and occupy a short time range to be useful. Misleading results are produced if the index fossils are incorrectly dated. Stratigraphy and biostratigraphy can in general provide only relative dating ( A was before B ), which is often sufficient for studying evolution. However, this is difficult for some time periods, because of
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#17328768042381416-526: A mutation first appeared. Phylogenetics and paleontology work together in the clarification of science's still dim view of the appearance of life and its evolution. Niles Eldredge 's study of the Phacops trilobite genus supported the hypothesis that modifications to the arrangement of the trilobite's eye lenses proceeded by fits and starts over millions of years during the Devonian . Eldredge's interpretation of
1534-420: 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 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
1652-499: 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
1770-399: 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 . Though the fossil record is incomplete, numerous studies have demonstrated that there is enough information available to give a good understanding of
1888-570: A portion of the deceased organism, usually that portion that was partially mineralized during life, such as the bones and teeth of vertebrates , or the chitinous or calcareous exoskeletons of invertebrates . Fossils may also consist of the marks left behind by the organism while it was alive, such as animal tracks or feces ( coprolites ). These types of fossil are called trace fossils or ichnofossils , as opposed to body fossils . Some fossils are biochemical and are called chemofossils or biosignatures . Gathering fossils dates at least to
2006-484: 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
2124-644: 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
2242-400: A richly diverse assembly of early multicellular eukaryotes . The fossil record and faunal succession form the basis of the science of biostratigraphy or determining the age of rocks based on embedded fossils. For the first 150 years of geology , biostratigraphy and superposition were the only means for determining the relative age of rocks. The geologic time scale was developed based on
2360-424: 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
2478-436: 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,
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#17328768042382596-448: A small percentage of life-forms can be expected to be represented in discoveries, and each discovery represents only a snapshot of the process of evolution. The transition itself can only be illustrated and corroborated by transitional fossils, which will never demonstrate an exact half-way point. The fossil record is strongly biased toward organisms with hard-parts, leaving most groups of soft-bodied organisms with little to no role. It
2714-424: A steady pull. Paired pedal protractor and retractor muscles operate 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
2832-449: Is a notable example of how knowledge encoded by the fossil record continues to contribute otherwise unattainable information on the emergence and development of life on Earth. For example, the research suggests Markuelia has closest affinity to priapulid worms, and is adjacent to the evolutionary branching of Priapulida , Nematoda and Arthropoda . Despite significant advances in uncovering and identifying paleontological specimens, it
2950-432: 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
3068-479: Is generally accepted that the fossil record is vastly incomplete. Approaches for measuring the completeness of the fossil record have been developed for numerous subsets of species, including those grouped taxonomically, temporally, environmentally/geographically, or in sum. This encompasses the subfield of taphonomy and the study of biases in the paleontological record. Paleontology seeks to map out how life evolved across geologic time. A substantial hurdle
3186-557: 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
3304-438: Is replete with the mollusks , the vertebrates , the echinoderms , the brachiopods and some groups of arthropods . Fossil sites with exceptional preservation—sometimes including preserved soft tissues—are known as Lagerstätten —German for "storage places". These formations may have resulted from carcass burial in an anoxic environment with minimal bacteria, thus slowing decomposition. Lagerstätten span geological time from
3422-408: 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
3540-470: Is the difficulty of working out fossil ages. Beds that preserve fossils typically lack the radioactive elements needed for radiometric dating . This technique is our only means of giving rocks greater than about 50 million years old an absolute age, and can be accurate to within 0.5% or better. Although radiometric dating requires careful laboratory work, its basic principle is simple: the rates at which various radioactive elements decay are known, and so
3658-413: Is the science of deciphering the "layer-cake" that is the sedimentary record. Rocks normally form relatively horizontal layers, with each layer younger than the one underneath it. If a fossil is found between two layers whose ages are known, the fossil's age is claimed to lie between the two known ages. Because rock sequences are not continuous, but may be broken up by faults or periods of erosion , it
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3776-472: 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. The shell
3894-432: 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
4012-459: Is very difficult to match up rock beds that are not directly adjacent. However, fossils of species that survived for a relatively short time can be used to match isolated rocks: this technique is called biostratigraphy . For instance, the conodont Eoplacognathus pseudoplanus has a short range in the Middle Ordovician period. If rocks of unknown age have traces of E. pseudoplanus , they have
4130-757: The Cambrian period to the present . Worldwide, some of the best examples of near-perfect fossilization are the Cambrian Maotianshan Shales and Burgess Shale , the Devonian Hunsrück Slates , the Jurassic Solnhofen Limestone , and the Carboniferous Mazon Creek localities. A fossil is said to be recrystallized when the original skeletal compounds are still present but in a different crystal form, such as from aragonite to calcite . Replacement occurs when
4248-545: 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
4366-721: 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
4484-581: The Phacops fossil record was that the aftermaths of the lens changes, but not the rapidly occurring evolutionary process, were fossilized. This and other data led Stephen Jay Gould and Niles Eldredge to publish their seminal paper on punctuated equilibrium in 1971. Synchrotron X-ray tomographic analysis of early Cambrian bilaterian embryonic microfossils yielded new insights of metazoan evolution at its earliest stages. The tomography technique provides previously unattainable three-dimensional resolution at
4602-559: The Renaissance . Leonardo da Vinci concurred with Aristotle's view that fossils were the remains of ancient life. For example, Leonardo noticed discrepancies with the biblical flood narrative as an explanation for fossil origins: If the Deluge had carried the shells for distances of three and four hundred miles from the sea it would have carried them mixed with various other natural objects all heaped up together; but even at such distances from
4720-591: The Song dynasty during the 11th century, who kept a specific seashell fossil with his own poem engraved on it. In his Dream Pool Essays published in 1088, Song dynasty Chinese scholar-official Shen Kuo hypothesized that marine fossils found in a geological stratum of mountains located hundreds of miles from the Pacific Ocean was evidence that a prehistoric seashore had once existed there and shifted over centuries of time . His observation of petrified bamboos in
4838-571: 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,
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4956-591: 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
5074-402: The law of superposition ) preserved different assemblages of fossils, and that these assemblages succeeded one another in a regular and determinable order. He observed that rocks from distant locations could be correlated based on the fossils they contained. He termed this the principle of faunal succession . This principle became one of Darwin's chief pieces of evidence that biological evolution
5192-517: 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. The sensory organs of bivalves are largely located on
5310-403: The thunderbird . There is no such direct mythological connection known from prehistoric Africa, but there is considerable evidence of tribes there excavating and moving fossils to ceremonial sites, apparently treating them with some reverence. In Japan, fossil shark teeth were associated with the mythical tengu , thought to be the razor-sharp claws of the creature, documented some time after
5428-400: 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
5546-449: The 16th century. Roman naturalist Pliny the Elder wrote of " tongue stones ", which he called glossopetra . These were fossil shark teeth, thought by some classical cultures to look like the tongues of people or snakes. He also wrote about the horns of Ammon , which are fossil ammonites , whence the group of shelled octopus-cousins ultimately draws its modern name. Pliny also makes one of
5664-739: The 19th century that certain fossils were associated with certain rock strata led to the recognition of a geological timescale and the relative ages of different fossils. The development of radiometric dating techniques in the early 20th century allowed scientists to quantitatively measure the absolute ages of rocks and the fossils they host. There are many processes that lead to fossilization , including permineralization , casts and molds, authigenic mineralization , replacement and recrystallization, adpression, carbonization , and bioimmuration. Fossils vary in size from one- micrometre (1 μm) bacteria to dinosaurs and trees, many meters long and weighing many tons. A fossil normally preserves only
5782-413: The 8th century AD. In medieval China, the fossil bones of ancient mammals including Homo erectus were often mistaken for " dragon bones" and used as medicine and aphrodisiacs . In addition, some of these fossil bones are collected as "art" by scholars, who left scripts on various artifacts, indicating the time they were added to a collection. One good example is the famous scholar Huang Tingjian of
5900-454: 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
6018-457: 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
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#17328768042386136-597: The Preservation of Favoured Races in the Struggle for Life , the oldest animal fossils were those from the Cambrian Period, now known to be about 540 million years old. He worried about the absence of older fossils because of the implications on the validity of his theories, but he expressed hope that such fossils would be found, noting that: "only a small portion of the world is known with accuracy." Darwin also pondered
6254-555: The Proterozoic and deeper still in the Archean is only "recounted by microscopic fossils and subtle chemical signals." Molecular biologists, using phylogenetics , can compare protein amino acid or nucleotide sequence homology (i.e., similarity) to evaluate taxonomy and evolutionary distances among organisms, with limited statistical confidence. The study of fossils, on the other hand, can more specifically pinpoint when and in what organism
6372-498: 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
6490-467: 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
6608-413: The beach, indicating the fossils were once living animals. He had previously explained them in terms of vaporous exhalations , which Persian polymath Avicenna modified into the theory of petrifying fluids ( succus lapidificatus ). Recognition of fossil seashells as originating in the sea was built upon in the 14th century by Albert of Saxony , and accepted in some form by most naturalists by
6726-710: The beginning of recorded history. The fossils themselves are referred to as the fossil record. The fossil record was one of the early sources of data underlying the study of evolution and continues to be relevant to the history of life on Earth . Paleontologists examine the fossil record to understand the process of evolution and the way particular species have evolved. Fossils have been visible and common throughout most of natural history, and so documented human interaction with them goes back as far as recorded history, or earlier. There are many examples of paleolithic stone knives in Europe, with fossil echinoderms set precisely at
6844-568: 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
6962-660: The deity Sopdu , the Morning Star, equivalent of Venus in Roman mythology. Fossils appear to have directly contributed to the mythology of many civilizations, including the ancient Greeks. Classical Greek historian Herodotos wrote of an area near Hyperborea where gryphons protected golden treasure. There was indeed gold mining in that approximate region , where beaked Protoceratops skulls were common as fossils. A later Greek scholar, Aristotle , eventually realized that fossil seashells from rocks were similar to those found on
7080-466: The dry northern climate zone of what is now Yan'an , Shaanxi province, China, led him to advance early ideas of gradual climate change due to bamboo naturally growing in wetter climate areas. In medieval Christendom , fossilized sea creatures on mountainsides were seen as proof of the biblical deluge of Noah's Ark . After observing the existence of seashells in mountains, the ancient Greek philosopher Xenophanes (c. 570 – 478 BC) speculated that
7198-415: The earlier known references to toadstones , thought until the 18th century to be a magical cure for poison originating in the heads of toads, but which are fossil teeth from Lepidotes , a Cretaceous ray-finned fish. The Plains tribes of North America are thought to have similarly associated fossils, such as the many intact pterosaur fossils naturally exposed in the region, with their own mythology of
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#17328768042387316-550: The earliest known stromatolites are over 3.4 billion years old. The fossil record is life's evolutionary epic that unfolded over four billion years as environmental conditions and genetic potential interacted in accordance with natural selection. The Virtual Fossil Museum Paleontology has joined with evolutionary biology to share the interdisciplinary task of outlining the tree of life, which inevitably leads backwards in time to Precambrian microscopic life when cell structure and functions evolved. Earth's deep time in
7434-460: The earth during earthquake and subsidences, and petrifies whatever comes into contact with it. As a matter of fact, the petrifaction of the bodies of plants and animals is not more extraordinary than the transformation of waters. From the 13th century to the present day, scholars pointed out that the fossil skulls of Deinotherium giganteum , found in Crete and Greece, might have been interpreted as being
7552-436: 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
7670-478: The existence of a world previous to ours, destroyed by some kind of catastrophe. Interest in fossils, and geology more generally, expanded during the early nineteenth century. In Britain, Mary Anning 's discoveries of fossils, including the first complete ichthyosaur and a complete plesiosaurus skeleton, sparked both public and scholarly interest. Early naturalists well understood the similarities and differences of living species leading Linnaeus to develop
7788-402: 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
7906-421: 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
8024-434: 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,
8142-434: 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
8260-497: The foot is large. They are vigorous burrowers. This bivalve -related article is a stub . You can help Misplaced Pages by expanding it . Bivalve 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
8378-467: 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
8496-519: The hand grip, dating back to Homo heidelbergensis and Neanderthals . These ancient peoples also drilled holes through the center of those round fossil shells, apparently using them as beads for necklaces. The ancient Egyptians gathered fossils of species that resembled the bones of modern species they worshipped. The god Set was associated with the hippopotamus , therefore fossilized bones of hippo-like species were kept in that deity's temples. Five-rayed fossil sea urchin shells were associated with
8614-577: 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
8732-434: 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
8850-399: 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
8968-521: 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
9086-421: 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
9204-443: 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
9322-489: 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
9440-485: 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
9558-535: The limits of fossilization. Fossils of two enigmatic bilaterians, the worm-like Markuelia and a putative, primitive protostome , Pseudooides , provide a peek at germ layer embryonic development. These 543-million-year-old embryos support the emergence of some aspects of arthropod development earlier than previously thought in the late Proterozoic. The preserved embryos from China and Siberia underwent rapid diagenetic phosphatization resulting in exquisite preservation, including cell structures. This research
9676-418: 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
9794-452: The mantle though the valves. In sedentary or recumbent bivalves that lie on one valve, such as the oysters and scallops, the anterior adductor muscle has been lost and the posterior muscle is positioned centrally. In species that can swim by flapping their valves, a single, central adductor muscle occurs. These muscles are composed of two types of muscle fibres, striated muscle bundles for fast actions and smooth muscle bundles for maintaining
9912-626: 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
10030-422: 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
10148-499: 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
10266-400: 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
10384-474: The pattern of diversification of life on Earth. In addition, the record can predict and fill gaps such as the discovery of Tiktaalik in the arctic of Canada . Paleontology includes the study of fossils: their age, method of formation, and evolutionary significance. Specimens are usually considered to be fossils if they are over 10,000 years old. The oldest fossils are around 3.48 billion years to 4.1 billion years old. The observation in
10502-519: 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
10620-409: 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 ,
10738-545: The problems involved in matching rocks of the same age across continents . Family-tree relationships also help to narrow down the date when lineages first appeared. For instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an ancestor of B and C, then A must have evolved earlier. It is also possible to estimate how long ago two living clades diverged, in other words approximately how long ago their last common ancestor must have lived, by assuming that DNA mutations accumulate at
10856-516: 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
10974-443: The ratio of the radioactive element to its decay products shows how long ago the radioactive element was incorporated into the rock. Radioactive elements are common only in rocks with a volcanic origin, and so the only fossil-bearing rocks that can be dated radiometrically are volcanic ash layers, which may provide termini for the intervening sediments. Consequently, palaeontologists rely on stratigraphy to date fossils. Stratigraphy
11092-462: The relative ages of rock strata as determined by the early paleontologists and stratigraphers . Since the early years of the twentieth century, absolute dating methods, such as radiometric dating (including potassium/argon , argon/argon , uranium series , and, for very recent fossils, radiocarbon dating ) have been used to verify the relative ages obtained by fossils and to provide absolute ages for many fossils. Radiometric dating has shown that
11210-429: The sea and that they were still living when the strait of Gibraltar was cut through. In the mountains of Parma and Piacenza multitudes of shells and corals with holes may be seen still sticking to the rocks.... In 1666, Nicholas Steno examined a shark, and made the association of its teeth with the "tongue stones" of ancient Greco-Roman mythology, concluding that those were not in fact the tongues of venomous snakes, but
11328-496: 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
11446-423: The sea we see the oysters all together and also the shellfish and the cuttlefish and all the other shells which congregate together, found all together dead; and the solitary shells are found apart from one another as we see them every day on the sea-shores. And we find oysters together in very large families, among which some may be seen with their shells still joined together, indicating that they were left there by
11564-419: 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
11682-469: 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
11800-435: 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
11918-468: 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,
12036-571: 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,
12154-420: 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
12272-400: 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
12390-736: 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
12508-581: The skulls of the Cyclopes of Greek mythology , and are possibly the origin of that Greek myth. Their skulls appear to have a single eye-hole in the front, just like their modern elephant cousins, though in fact it's actually the opening for their trunk. In Norse mythology , echinoderm shells (the round five-part button left over from a sea urchin) were associated with the god Thor , not only being incorporated in thunderstones , representations of Thor's hammer and subsequent hammer-shaped crosses as Christianity
12626-577: 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
12744-416: 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
12862-467: The sudden appearance of many groups (i.e. phyla ) in the oldest known Cambrian fossiliferous strata. Since Darwin's time, the fossil record has been extended to between 2.3 and 3.5 billion years. Most of these Precambrian fossils are microscopic bacteria or microfossils . However, macroscopic fossils are now known from the late Proterozoic. The Ediacara biota (also called Vendian biota) dating from 575 million years ago collectively constitutes
12980-451: The teeth of some long-extinct species of shark. Robert Hooke (1635–1703) included micrographs of fossils in his Micrographia and was among the first to observe fossil forams . His observations on fossils, which he stated to be the petrified remains of creatures some of which no longer existed, were published posthumously in 1705. William Smith (1769–1839) , an English canal engineer, observed that rocks of different ages (based on
13098-590: 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
13216-558: 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
13334-406: 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
13452-402: 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 . These muscles pull
13570-511: The world was once inundated in a great flood that buried living creatures in drying mud. In 1027, the Persian Avicenna explained fossils' stoniness in The Book of Healing : If what is said concerning the petrifaction of animals and plants is true, the cause of this (phenomenon) is a powerful mineralizing and petrifying virtue which arises in certain stony spots, or emanates suddenly from
13688-417: 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
13806-458: Was adopted, but also kept in houses to garner Thor's protection. These grew into the shepherd's crowns of English folklore, used for decoration and as good luck charms, placed by the doorway of homes and churches. In Suffolk , a different species was used as a good-luck charm by bakers, who referred to them as fairy loaves , associating them with the similarly shaped loaves of bread they baked. More scientific views of fossils emerged during
13924-408: Was real. Georges Cuvier came to believe that most if not all the animal fossils he examined were remains of extinct species. This led Cuvier to become an active proponent of the geological school of thought called catastrophism . Near the end of his 1796 paper on living and fossil elephants he said: All of these facts, consistent among themselves, and not opposed by any report, seem to me to prove
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