Bivalvia
Bivalvia Temporal range:
| |
---|---|
"Acephala", from Ernst Haeckel's Kunstformen der Natur (1904) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Mollusca |
Class: | Bivalvia Linnaeus, 1758 |
Subclasses | |
|
Bivalvia ( bore into wood, clay, or stone and live inside these substances.
The
Bivalves have long been a part of the diet of coastal and
Bivalves appear in the
Etymology
The taxonomic term Bivalvia was first used by Linnaeus in the 10th edition of his Systema Naturae in 1758 to refer to animals having shells composed of two valves.[3] More recently, the class was known as Pelecypoda, meaning "axe-foot" (based on the shape of the foot of the animal when extended).
The name "bivalve" is derived from the
Anatomy
Bivalves have
Mantle and shell
The shell is composed of two calcareous valves held together by a ligament. The valves are made of either calcite, as is the case in oysters, or both calcite and aragonite. Sometimes, the aragonite forms an inner, nacreous layer, as is the case in the order Pteriida. In other taxa, alternate layers of calcite and aragonite are laid down.[13] The ligament and byssus, if calcified, are composed of aragonite.[13] The outermost layer of the shell is the periostracum, a thin layer composed of horny conchiolin. The periostracum is secreted by the outer mantle and is easily abraded.[14] The outer surface of the valves is often sculpted, with clams often having concentric striations, scallops having radial ribs and oysters a latticework of irregular markings.[15]
In all molluscs, the mantle forms a thin membrane that covers the animal's body and extends out from it in flaps or lobes. In bivalves, the mantle lobes secrete the valves, and the mantle crest secretes the whole hinge mechanism consisting of ligament, byssus threads (where present), and teeth.[16] The posterior mantle edge may have two elongated extensions known as siphons, through one of which water is inhaled, and the other expelled.[17] The siphons retract into a cavity, known as the pallial sinus.[18]
The shell grows larger when more material is secreted by the mantle edge, and the valves themselves thicken as more material is secreted from the general mantle surface. Calcareous matter comes from both its diet and the surrounding seawater. Concentric rings on the exterior of a valve are commonly used to age bivalves. For some groups, a more precise method for determining the age of a shell is by cutting a cross section through it and examining the incremental growth bands.[19]
The
Muscles and ligaments
The main muscular system in bivalves is the posterior and anterior adductor muscles. These muscles connect the two valves and contract to close the shell. The valves are also joined dorsally by the hinge ligament, which is an extension of the periostracum. The ligament is responsible for opening the shell, and works against the adductor muscles when the animal opens and closes.[21] Retractor muscles connect the mantle to the edge of the shell, along a line known as the pallial line.[22][16] These muscles pull the mantle though the valves.[16]
In sedentary or recumbent bivalves that lie on one valve, such as the oysters and scallops, the anterior adductor muscle has been lost and the posterior muscle is positioned centrally.[23] In species that can swim by flapping their valves, a single, central adductor muscle occurs. These muscles are composed of two types of muscle fibres, striated muscle bundles for fast actions and smooth muscle bundles for maintaining a steady pull.[24] Paired pedal protractor and retractor muscles operate the animal's foot.[11][25][26]
Nervous system
The sedentary habits of the bivalves have meant that in general the
Senses
The sensory organs of bivalves are largely located on the posterior mantle margins. The organs are usually
Circulation and respiration
Bivalves have an open
The paired gills are located posteriorly and consist of hollow tube-like filaments with thin walls for gas exchange. The respiratory demands of bivalves are low, due to their relative inactivity. Some freshwater species, when exposed to the air, can gape the shell slightly and gas exchange can take place.[35][36] Oysters, including the Pacific oyster (Magallana gigas), are recognized as having varying metabolic responses to environmental stress, with changes in respiration rate being frequently observed.[37]
Digestive system
Modes of feeding
Most bivalves are
In more advanced bivalves, water is drawn into the shell from the posterior
A few bivalves, such as the
The unusual genus,
Digestive tract
The digestive tract of typical bivalves consists of an
Carnivorous bivalves generally have reduced crystalline styles and the stomach has thick, muscular walls, extensive cuticular linings and diminished sorting areas and gastric chamber sections.[43]
Excretory system
The excretory organs of bivalves are a pair of nephridia. Each of these consists of a long, looped, glandular tube, which opens into the pericardium, and a bladder to store urine. They also have pericardial glands either line the auricles of the heart or attach to the pericardium, and serve as extra filtration organs. Metabolic waste is voided from the bladders through a nephridiopore near the front of the upper part of the mantle cavity and excreted.[44][45]
Reproduction and development
The sexes are usually separate in bivalves but some
Fertilization is usually external. Typically, a short stage lasts a few hours or days before the eggs hatch into trochophore larvae. These later develop into veliger larvae which settle on the seabed and undergo metamorphosis into adults.[46][49] In some species, such as those in the genus Lasaea, females draw water containing sperm in through their inhalant siphons and fertilization takes place inside the female. These species then brood the young inside their mantle cavity, eventually releasing them into the water column as veliger larvae or as crawl-away juveniles.[50]
Most of the bivalve larvae that hatch from eggs in the water column feed on
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
Some of the species in the freshwater mussel family,
Comparison with brachiopods
Brachiopods are shelled marine organisms that superficially resemble bivalves in that they are of similar size and have a hinged shell in two parts. However, brachiopods evolved from a very different ancestral line, and the resemblance to bivalves only arose because they occupy similar ecological niches. The differences between the two groups are due to their separate ancestral origins. Different initial structures have been adapted to solve the same problems, a case of convergent evolution. In modern times, brachiopods are not as common as bivalves.[53]
Both groups have a shell consisting of two valves, but the organization of the shell is quite different in the two groups. In brachiopods, the two valves are positioned on the dorsal and ventral surfaces of the body, while in bivalves, the valves are on the left and right sides of the body, and are, in most cases, mirror images of one other. Brachiopods have a lophophore, a coiled, rigid cartilaginous internal apparatus adapted for filter feeding, a feature shared with two other major groups of marine invertebrates, the bryozoans and the phoronids. Some brachiopod shells are made of calcium phosphate but most are calcium carbonate in the form of the biomineral calcite, whereas bivalve shells are always composed entirely of calcium carbonate, often in the form of the biomineral aragonite.[54]
Evolutionary history
The Cambrian explosion took place around 540 to 520 million years ago (Mya). In this geologically brief period, all the major animal phyla diverged and these included the first creatures with mineralized skeletons. Brachiopods and bivalves made their appearance at this time, and left their fossilized remains behind in the rocks.[55]
Possible early bivalves include
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
By the middle of the Paleozoic, around 400 Mya, the brachiopods were among the most abundant filter feeders in the ocean, and over 12,000 fossil species are recognized.[59] By the Permian–Triassic extinction event 250 Mya, bivalves were undergoing a huge radiation of diversity. The bivalves were hard hit by this event, but re-established themselves and thrived during the Triassic period that followed. In contrast, the brachiopods lost 95% of their species diversity.[54] The ability of some bivalves to burrow and thus avoid predators may have been a major factor in their success. Other new adaptations within various families allowed species to occupy previously unused evolutionary niches. These included increasing relative buoyancy in soft sediments by developing spines on the shell, gaining the ability to swim, and in a few cases, adopting predatory habits.[58]
For a long time, bivalves were thought to be better adapted to aquatic life than brachiopods were, outcompeting and relegating them to minor niches in later ages. These two taxa appeared in textbooks as an example of replacement by competition. Evidence given for this included the fact that bivalves needed less food to subsist because of their energetically efficient ligament-muscle system for opening and closing valves. All this has been broadly disproven, though; rather, the prominence of modern bivalves over brachiopods seems due to chance disparities in their response to extinction events.[60]
Diversity of extant bivalves
The adult maximum size of
In his 2010 treatise, Compendium of Bivalves, Markus Huber gives the total number of living bivalve species as about 9,200 combined in 106 families.[65] Huber states that the number of 20,000 living species, often encountered in literature, could not be verified and presents the following table to illustrate the known diversity:
Subclass | Superfamilies | Families | Genera | Species |
---|---|---|---|---|
Heterodonta
|
64 (incl. 1 freshwater) | 800 (16 freshwater) | 5600 (270 freshwater) | |
Arcticoidea | 2 | 6 | 13 | |
Cardioidea | 2 | 38 | 260 | |
Chamoidea
|
1 | 6 | 70 | |
Clavagelloidea | 1 | 2 | 20 | |
Crassatelloidea | 5 | 65 | 420 | |
Cuspidarioidea
|
2 | 20 | 320 | |
Cyamioidea | 3 | 22 | 140 | |
Cyrenoidea | 1 | 6 (3 freshwater) | 60 (30 freshwater) | |
Cyrenoidoidea | 1 | 1 | 6 | |
Dreissenoidea | 1 | 3 (2 freshwater) | 20 (12 freshwater) | |
Galeommatoidea | ca. 4 | about 100 | about 500 | |
Gastrochaenoidea
|
1 | 7 | 30 | |
Glossoidea | 2 | 20 | 110 | |
Hemidonacoidea
|
1 | 1 | 6 | |
Hiatelloidea
|
1 | 5 | 25 | |
Limoidea
|
1 | 8 | 250 | |
Lucinoidea
|
2 | about 85 | about 500 | |
Mactroidea | 4 | 46 | 220 | |
Myoidea
|
3 | 15 (1 freshwater) | 130 (1 freshwater) | |
Pandoroidea | 7 | 30 | 250 | |
Pholadoidea | 2 | 34 (1 freshwater) | 200 (3 freshwater) | |
Pholadomyoidea | 2 | 3 | 20 | |
Solenoidea | 2 | 17 (2 freshwater) | 130 (4 freshwater) | |
Sphaerioidea | (1 freshwater) | (5 freshwater) | (200 freshwater) | |
Tellinoidea | 5 | 110 (2 freshwater) | 900 (15 freshwater) | |
Thyasiroidea
|
1 | about 12 | about 100 | |
Ungulinoidea | 1 | 16 | 100 | |
Veneroidea | 4 | 104 | 750 | |
Verticordioidea | 2 | 16 | 160 | |
Palaeoheterodonta | 7 (incl. 6 freshwater) | 171 (170 freshwater) | 908 (900 freshwater) | |
Trigonioidea | 1 | 1 | 8 | |
Unionoidea
|
(6 freshwater) | (170 freshwater) | (900 freshwater) | |
Protobranchia | 10 | 49 | 700 | |
Manzanelloidea | 1 | 2 | 20 | |
Nuculanoidea | 6 | 32 | 460 | |
Nuculoidea | 1 | 8 | 170 | |
Sareptoidea | 1 | about 5 | 10 | |
Solemyoidea | 1 | 2 | 30 | |
Pteriomorphia | 25 | 240 (2 freshwater) | 2000 (11 freshwater) | |
Anomioidea | 2 | 9 | 30 | |
Arcoidea | 7 | 60 (1 freshwater) | 570 (6 freshwater) | |
Dimyoidea
|
1 | 3 | 15 | |
Limoidea
|
1 | 8 | 250 | |
Mytiloidea | 1 | 50 (1 freshwater) | 400 (5 freshwater) | |
Ostreoidea | 2 | 23 | 80 | |
Pectinoidea | 4 | 68 | 500 | |
Pinnoidea
|
1 | 3 (+) | 50 | |
Plicatuloidea
|
1 | 1 | 20 | |
Pterioidea | 5 | 9 | 80 |
Distribution
The bivalves are a highly successful class of invertebrates found in aquatic habitats throughout the world. Most are
Bivalves inhabit the tropics, as well as temperate and boreal waters. A number of species can survive and even flourish in extreme conditions. They are abundant in the Arctic, about 140 species being known from that zone.
Some freshwater bivalves have very restricted ranges. For example, the Ouachita creekshell mussel,
Behaviour
Most bivalves adopt a sedentary or even
Other bivalves, such as mussels, attach themselves to hard surfaces using tough byssus threads made of collagen and elastin proteins.[77] Some species, including the true oysters, the jewel boxes, the jingle shells, the thorny oysters and the kitten's paws, cement themselves to stones, rock or larger dead shells.[78] 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.[79]
Bivalves filter large amounts of water to feed and breathe but they are not permanently open. They regularly shut their valves to enter a resting state, even when they are permanently submerged. In oysters, for example, their behaviour follows very strict circatidal and circadian rhythms according to the relative positions of the moon and sun. During neap tides, they exhibit much longer closing periods than during spring tides.[80]
Although many non-sessile bivalves use their muscular foot to move around, or to dig, members of the freshwater family
Predators and defence
The thick shell and rounded shape of bivalves make them awkward for potential predators to tackle. Nevertheless, a number of different creatures include them in their diet. Many species of
Invertebrate predators include crustaceans, starfish and octopuses. Crustaceans crack the shells with their pincers and starfish use their water vascular system to force the valves apart and then insert part of their stomach between the valves to digest the bivalve's body. It has been found experimentally that both crabs and starfish preferred molluscs that are attached by byssus threads to ones that are cemented to the substrate. This was probably because they could manipulate the shells and open them more easily when they could tackle them from different angles.[78] 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.[88] Certain carnivorous gastropod snails such as whelks (Buccinidae) and murex snails (Muricidae) feed on bivalves by boring into their shells. A dog whelk (Nucella) drills a hole with its radula assisted by a shell-dissolving secretion. The dog whelk then inserts its extendible proboscis and sucks out the body contents of the victim, which is typically a blue mussel.[89]
Razor shells can dig themselves into the sand with great speed to escape predation. When a Pacific razor clam (Siliqua patula) is laid on the surface of the beach, it can bury itself completely in seven seconds [90] and the Atlantic jackknife clam, Ensis directus, can do the same within fifteen seconds.[91] Scallops and file clams can swim by opening and closing their valves rapidly; water is ejected on either side of the hinge area and they move with the flapping valves in front.[92] Scallops have simple eyes around the margin of the mantle and can clap their valves shut to move sharply, hinge first, to escape from danger.[92] Cockles can use their foot to move across the seabed or leap away from threats. The foot is first extended before being contracted suddenly when it acts like a spring, projecting the animal forwards.[93]
In many bivalves that have
File shells such as Limaria fragilis can produce a noxious secretion when stressed. It has numerous tentacles which fringe its mantle and protrude some distance from the shell when it is feeding. If attacked, it sheds tentacles in a process known as autotomy. The toxin released by this is distasteful and the detached tentacles continue to writhe which may also serve to distract potential predators.[97]
Mariculture
Many juveniles are further reared off the seabed in suspended rafts, on floating trays or cemented to ropes. Here they are largely free from bottom-dwelling predators such as starfish and crabs but more labour is required to tend them. They can be harvested by hand when they reach a suitable size. Other juveniles are laid directly on the seabed at the rate of 50 to 100 kilograms (110 to 220 lb) per hectare. They grow on for about two years before being harvested by dredging. Survival rates are low at about 5%.[99]
The
Similar techniques are used in different parts of the world to cultivate other species including the
Production of bivalve molluscs by mariculture in 2010 was 12,913,199 tons, up from 8,320,724 tons in 2000. Culture of clams, cockles and ark shells more than doubled over this time period from 2,354,730 to 4,885,179 tons. Culture of mussels over the same period grew from 1,307,243 to 1,812,371 tons, of oysters from 3,610,867 to 4,488,544 tons and of scallops from 1,047,884 to 1,727,105 tons.[102]
Use as food
Bivalves have been an important source of food for humans at least since Roman times[103] and empty shells found in middens at archaeological sites are evidence of earlier consumption.[87] Oysters, scallops, clams, ark clams, mussels and cockles are the most commonly consumed kinds of bivalve, and are eaten cooked or raw. In 1950, the year in which the Food and Agriculture Organization (FAO) started making such information available, world trade in bivalve molluscs was 1,007,419 tons.[104] By 2010, world trade in bivalves had risen to 14,616,172 tons, up from 10,293,607 tons a decade earlier. The figures included 5,554,348 (3,152,826) tons of clams, cockles and ark shells, 1,901,314 (1,568,417) tons of mussels, 4,592,529 (3,858,911) tons of oysters and 2,567,981 (1,713,453) tons of scallops.[104] China increased its consumption 400-fold during the period 1970 to 1997.[105]
It has been known for more than a century that consumption of raw or insufficiently cooked shellfish can be associated with infectious diseases. These are caused either by bacteria naturally present in the sea such as
Viral and bacterial infections
In 1816 in France, a physician, J. P. A. Pasquier, described an outbreak of
Since the 1970s, outbreaks of oyster-
Paralytic shellfish poisoning
Paralytic shellfish poisoning (PSP) is primarily caused by the consumption of bivalves that have accumulated toxins by feeding on toxic dinoflagellates, single-celled protists found naturally in the sea and inland waters. Saxitoxin is the most virulent of these. In mild cases, PSP causes tingling, numbness, sickness and diarrhoea. In more severe cases, the muscles of the chest wall may be affected leading to paralysis and even death. In 1937, researchers in California established the connection between blooms of these phytoplankton and PSP.[109] The biotoxin remains potent even when the shellfish are well-cooked.[109] In the United States, there is a regulatory limit of 80 µg/g of saxitoxin equivalent in shellfish meat.[109]
Amnesic shellfish poisoning
Amnesic shellfish poisoning (ASP) was first reported in eastern Canada in 1987. It is caused by the substance domoic acid found in certain diatoms of the genus Pseudo-nitzschia. Bivalves can become toxic when they filter these microalgae out of the water. Domoic acid is a low-molecular weight amino acid that is able to destroy brain cells causing memory loss, gastroenteritis, long-term neurological problems or death. In an outbreak in the western United States in 1993, finfish were also implicated as vectors, and seabirds and mammals suffered neurological symptoms.[109] In the United States and Canada, a regulatory limit of 20 µg/g of domoic acid in shellfish meat is set.[110]
Ecosystem services
When they live in polluted waters, bivalve molluscs have a tendency to accumulate substances such as
There are limitations to the use of bivalves as bioindicators. The level of pollutants found in the tissues varies with species, age, size, time of year and other factors. The quantities of pollutants in the water may vary and the molluscs may reflect past rather than present values. In a study near Vladivostok it was found that the level of pollutants in the bivalve tissues did not always reflect the high levels in the surrounding sediment in such places as harbours. The reason for this was thought to be that the bivalves in these locations did not need to filter so much water as elsewhere because of the water's high nutritional content.[113]
A study of nine different bivalves with widespread distributions in tropical marine waters concluded that the mussel,
Crushed shells, available as a by-product of the seafood canning industry, can be used to remove pollutants from water. It has been found that, as long as the water is maintained at an alkaline pH, crushed shells will remove cadmium, lead and other heavy metals from contaminated waters by swapping the calcium in their constituent aragonite for the heavy metal, and retaining these pollutants in a solid form.[116] The rock oyster (Saccostrea cucullata) has been shown to reduce the levels of copper and cadmium in contaminated waters in the Persian Gulf. The live animals acted as biofilters, selectively removing these metals, and the dead shells also had the ability to reduce their concentration.[117]
Other uses
Conchology is the scientific study of mollusc shells, but the term conchologist is also sometimes used to describe a collector of shells. Many people pick up shells on the beach or purchase them and display them in their homes. There are many private and public collections of mollusc shells, but the largest one in the world is at the Smithsonian Institution, which houses in excess of 20 million specimens.[118]
Shells are used decoratively in many ways. They can be pressed into concrete or plaster to make decorative paths, steps or walls and can be used to embellish picture frames, mirrors or other craft items. They can be stacked up and glued together to make ornaments. They can be pierced and threaded onto necklaces or made into other forms of jewellery. Shells have had various uses in the past as body decorations, utensils, scrapers and cutting implements. Carefully cut and shaped shell tools dating back 32,000 years have been found in a cave in Indonesia. In this region, shell technology may have been developed in preference to the use of stone or bone implements, perhaps because of the scarcity of suitable rock materials.[119]
The
Sea silk is a fine fabric woven from the byssus threads of bivalves, particularly the pen shell (Pinna nobilis). It used to be produced in the Mediterranean region where these shells are endemic. It was an expensive fabric and overfishing has much reduced populations of the pen shell.[124]
Crushed shells are added as a calcareous supplement to the diet of laying poultry. Oyster shell and cockle shell are often used for this purpose and are obtained as a by-product from other industries.[125]
Pearls and mother-of-pearl
Mother-of-pearl or nacre is the naturally occurring lustrous layer that lines some mollusc shells. It is used to make pearl buttons and in artisan craftwork to make organic jewellery. It has traditionally been inlaid into furniture and boxes, particularly in China. It has been used to decorate musical instruments, watches, pistols, fans and other products. The import and export of goods made with nacre are controlled in many countries under the International Convention of Trade in Endangered Species of Wild Fauna and Flora.[126]
A
Symbolism
The scallop is the symbol of
Bivalvian taxonomies
For the past two centuries no consensus has existed on bivalve
Since the year 2000, taxonomic studies using
Practical taxonomy of R.C. Moore
R.C. Moore, in Moore, Lalicker, and Fischer, 1952, Invertebrate Fossils, gives a practical and useful classification of pelecypods (Bivalvia) even if somewhat antiquated, based on shell structure, gill type, and hinge teeth configuration. Subclasses and orders given are:
- Subclass:Prionodesmacea
- Order
- Paleoconcha
- Taxodonta: Many teeth (e.g. order Nuculida)
- Schizodonta: Big bifurcating teeth (e.g. Trigonia spp.)
- Isodonta: Equal teeth (e.g. Spondylus spp.)
- Dysodonta: Absent teeth and ligaments joins the valves.
- Subclass:Teleodesmacea
- Order
- Lower Ordovician – Recent]
- Upper Cretaceous]
- Desmodonta: Hinge-teeth absent or irregular with ligaments (e.g. family Anatinidae).
Prionodesmacea have a prismatic and nacreous shell structure, separated mantle lobes, poorly developed siphons, and hinge teeth that are lacking or unspecialized. Gills range from protobranch to eulamellibranch. Teleodesmacea on the other hand have a porcelanous and partly nacreous shell structure; Mantle lobes that are generally connected, well developed siphons, and specialized hinge teeth. In most, gills are eulamellibranch.
1935 taxonomy
In his 1935 work Handbuch der systematischen Weichtierkunde (Handbook of Systematic Malacology), Johannes Thiele introduced a mollusc taxonomy based upon the 1909 work by Cossmann and Peyrot. Thiele's system divided the bivalves into three orders. Taxodonta consisted of forms that had taxodont dentition, with a series of small parallel teeth perpendicular to the hinge line. Anisomyaria consisted of forms that had either a single adductor muscle or one adductor muscle much smaller than the other. Eulamellibranchiata consisted of forms with ctenidial gills. The Eulamellibranchiata was further divided into four suborders: Schizodonta, Heterodonta, Adapedonta and Anomalodesmata.[142][143]
Taxonomy based upon hinge tooth morphology
The systematic layout presented here follows Newell's 1965 classification based on hinge tooth morphology (all taxa marked † are extinct) :[133]
Subclass | Order |
---|---|
Palaeotaxodonta
|
Nuculoida (nut shells)
|
Cryptodonta | † Praecardioida
Solemyoida
|
Pteriomorphia | ark shells )
Limoida (file shells)
mussels )
oysters , formerly included in Pterioida)
† Praecardioida
pen shells )
|
Palaeoheterodonta | Trigonioida (Neotrigonia is the only extant genus)
freshwater mussels )
|
Heterodonta
|
† Cycloconchidae
† Hippuritoida
)
Veneroida (hard-shell clams, cockles, razor shells )
|
Anomalodesmata | Pholadomyoida
|
The monophyly of the subclass Anomalodesmata is disputed. The standard view now is that it resides within the subclass Heterodonta.[134][137][144]
Taxonomy based upon gill morphology
An alternative systematic scheme exists using gill morphology.[145] This distinguishes between Protobranchia, Filibranchia and Eulamellibranchia. The first corresponds to Newell's Palaeotaxodonta and Cryptodonta, the second to his Pteriomorphia, with the last corresponding to all other groups. In addition, Franc separated the Septibranchia from his eulamellibranchs because of the morphological differences between them. The septibranchs belong to the superfamily Poromyoidea and are carnivorous, having a muscular septum instead of filamentous gills.[146]
2010 taxonomy
In May 2010, a new taxonomy of the Bivalvia was published in the journal Malacologia. In compiling this, the authors used a variety of phylogenetic information including molecular analysis, anatomical analysis, shell morphology and shell microstructure as well as bio-geographic, paleobiogeographic and stratigraphic information. In this classification 324 families are recognized as valid, 214 of which are known exclusively from fossils and 110 of which occur in the recent past, with or without a fossil record.[139] This classification has since been adopted by WoRMS.[147]
Proposed classification of Class Bivalvia (under the redaction of Rüdiger Bieler, Joseph G. Carter and Eugene V. Coan) (all taxa marked † are extinct) :[148]
Grade Euprotobranchia
Subclass Infraclass
Infraclass Euheterodonta
|
Subclass Palaeoheterodonta
Subclass Protobranchia
|
Subclass Pteriomorphia
Infraclass Eupteriomorphia
|
References
- ^ Jell, Peter A. (1980). "Earliest known pelecypod on Earth — a new Early Cambrian genus from South Australia". Alcheringa: An Australasian Journal of Palaeontology. 4 (3): 233–239. .
- JSTOR 1304610. Archived from the originalon 10 November 2016. Retrieved 1 December 2015.
- ^ Linnaeus, Carolus (1758). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata (in Latin). Vol. 1. Laurentii Salvii. p. 645.
- ^ Parker, Sybil (1984). McGraw-Hill Dictionary of Scientific and Technical Terms. McGraw-Hill Education.
- S2CID 132821312.
- ^ "The Phylum Brachiopoda". Earthlife Web. Retrieved 5 May 2012.
- ^ "Ostracoda". Oxford Dictionaries. Oxford University Press. Archived from the original on 8 September 2011. Retrieved 1 July 2012.
- ^ Webb, J. (1979). "A reappraisal of the palaeoecology of conchostracans (Crustacea: Branchiopoda)". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 158 (2): 259–275.
- ISBN 978-81-315-0104-7.)
{{cite book}}
: CS1 maint: multiple names: authors list (link - ISBN 978-0-03-030504-7.
- ^ a b Wells, Roger M. (1998). "Class Bivalvia". Invertebrate Paleontology Tutorial. State University of New York College at Cortland. Archived from the original on 28 February 2010. Retrieved 11 April 2012.
- ISBN 978-81-315-0104-7.)
{{cite book}}
: CS1 maint: multiple names: authors list (link - ^ a b
Kennedy, W. J.; Taylor, J. D.; Hall, A. (1969). "Environmental and biological controls on bivalve shell mineralogy". Biological Reviews. 44 (4): 499–530. S2CID 29279688.
- ISBN 0-85238-234-0.
- ISBN 978-0-7603-2593-3.
- ^ a b c Morton, Brian. "Bivalve: The mantle and musculature". Encyclopædia Britannica. Retrieved 5 May 2012.
- ISBN 0-85238-234-0.
- ISBN 0-85238-234-0.
- ISBN 0-85238-234-0.
- ^ Edmondson, C. H. (1962). "Teredinidae, ocean travelers" (PDF). Occasional Papers of Bernice P. Bishop Museum. 23 (3): 45–59.
- ISBN 978-81-315-0104-7.)
{{cite book}}
: CS1 maint: multiple names: authors list (link - ISBN 0-85238-234-0.
- ISBN 0-85238-234-0.
- ISBN 0-85238-234-0.
- ISBN 0-85238-234-0.
- ISBN 978-81-315-0104-7.)
{{cite book}}
: CS1 maint: multiple names: authors list (link - ^ a b Cofrancesco, Alfred F. (2002). "Nervous System and Sense Organs in Bivalves". Zebra Mussel Research Program. Archived from the original on 15 April 2012. Retrieved 5 May 2012.
- ^ a b Morton, Brian. "Mollusk: The nervous system and organs of sensation". Encyclopædia Britannica. Retrieved 8 July 2012.
- .
- ^
Morton, B. (2008). "The evolution of eyes in the Bivalvia: new insights". American Malacological Bulletin. 26 (1–2): 35–45. S2CID 85866382.
- S2CID 122110847.
- ISBN 978-0-03-030504-7.
- ISBN 0-85238-234-0.
- ^ a b Vaughan, Burton (2008). "The Bivalve, Poromya granulata". Archerd Shell Collection. Archived from the original on 7 April 2018. Retrieved 3 April 2012.
- ^ Morton, Brian. "Bivalve: The respiratory system". Encyclopædia Britannica. Retrieved 8 July 2012.
- ISBN 0-85238-234-0.
- ^ ISSN 2296-7745.
- ^ ISBN 978-0-03-030504-7.
- ISBN 0-85238-234-0.
- ^ Lützen, J.; Berland, B.; Bristow, G. A. (2011). "Morphology of an endosymbiotic bivalve, Entovalva nhatrangensis (Bristow, Berland, Schander & Vo, 2010) (Galeommatoidea)" (PDF). Molluscan Research. 31 (2): 114–124.
- ISBN 978-0-03-030504-7.
- ^ Morton, Brian. "Bivalve: The digestive system and nutrition". Encyclopædia Britannica. Retrieved 7 May 2012.
- .
- ^ Morton, Brian. "Bivalve: The excretory system". Encyclopædia Britannica. Retrieved 7 May 2012.
- ISBN 0-85238-234-0.
- ^ a b c
Dorit, Robert L.; Walker, Warren F. Jr.; Barnes, Robert D. (1991). Zoology. Saunders College Publishing. p. 682. ISBN 978-0-03-030504-7.
- ^ Morton, Brian. "Bivalve: The reproductive system". Encyclopædia Britannica. Retrieved 7 May 2012.
- ^ Helm, M. M.; Bourne, N.; Lovatelli, A. (2004). "Gonadal development and spawning". Hatchery culture of bivalves: a practical manual. FAO. Retrieved 8 May 2012.
- ^ Morton, Brian. "Bivalve: Reproduction and life cycles". Encyclopædia Britannica. Retrieved 7 May 2012.
- S2CID 189852952.
- .
- ISBN 978-0-313-33922-6.
- ISBN 978-0-03-030504-7.
- ^ a b
Barnes, R. S. K.; Callow, P.; Olive, P. J. W. (1988). The Invertebrates: A New Synthesis. Blackwell Scientific Publications. p. 140. ISBN 978-0-632-03125-2.
- ISBN 978-0-201-75054-6.
- .
- S2CID 49380913.
- ^ a b "Fossil Record". University of Bristol. Archived from the original on 12 July 2011. Retrieved 11 May 2012.
- ^ Brosius, L. (2008). "Fossil Brachiopods". GeoKansas. Kansas Geological Survey. Archived from the original on 5 July 2008. Retrieved 2 July 2012.
- JSTOR 2400538.
- ^ "Condylonucula maya". Extreme bivalves. Archived from the original on 15 October 2013. Retrieved 19 April 2012.
- ^ "Book review: Conchologists of America". Archived from the original on 7 August 2012. Retrieved 19 April 2012.
- ^ Grall, George. "Giant Clam: Tridacna gigas". National Geographic Society. Archived from the original on 21 June 2007. Retrieved 24 June 2012.
- S2CID 130048023.
- ISBN 978-3-939767-28-2.
- ^ Yonge, C. M. (1949). The Sea Shore. Collins. p. 228.
- ^ Bivalves Arctic Ocean Diversity. Retrieved 2012-04-21.
- ^ "Adamussium colbecki (Smith, 1902)". Antarctic Field Guide. Archived from the original on 14 October 2013. Retrieved 21 April 2012.
- ^ Rice, Tony. "Hydrothermal vents". Deep Ocean. Fathom. Archived from the original on 10 December 2008. Retrieved 21 April 2012.
- ^ Vesicomyinae (Bivalvia: Vesicomyidae) of the Kuril–Kamchatka Trench and adjacent abyssal regions
- ^ "Extreme bivalves". Museum of the Earth. Archived from the original on 15 October 2013. Retrieved 21 April 2012.
- ^ Christian, A. D. (2007). "Life History and Population Biology of the State Special Concern Ouachita Creekshell, Villosa arkansasensis (I. Lea 1862)" (PDF). Arkansas State University. Retrieved 21 April 2012.
- S2CID 20387549.
- UC Riverside. Archived from the originalon 23 June 2010. Retrieved 21 April 2012.
- ^ a b
Barnes, R. S. K.; Callow, P.; Olive, P. J. W. (1988). The Invertebrates: A New Synthesis. Blackwell Scientific Publications. pp. 132–134. ISBN 978-0-632-03125-2.
- ISBN 978-0-632-03125-2.
- ISBN 0-85238-234-0.
- ^ a b Harper, Elizabeth M. (1990). "The role of predation in the evolution of cementation in bivalves" (PDF). Palaeontology. 34 (2): 455–460. Archived from the original (PDF) on 19 October 2015. Retrieved 30 July 2017.
- ^ Barrett, John; Yonge, C. M. (1958). Collins Pocket Guide to the Sea Shore. London: William Collins Sons and Co. Ltd. p. 148.
- S2CID 25356955.
- S2CID 13546885.
- doi:10.1046/j.1365-2427.1998.00313.x.)
{{cite journal}}
: CS1 maint: numeric names: authors list (link - S2CID 87037074.
- .
- JSTOR 1376994.
- .
- ^ a b Rollins, H. B.; Sandweiss, D. H.; Brand, U.; Rollins, J. C. (1987). "Growth increment and stable isotope analysis of marine bivalves: implications for the geoarchaeological record of El Niño". Geoarchaeology. 2 (3): 181–197. .
- .
- ^ Carefoot, Tom (2010). "Learn about whelks and relatives: foods, feeding and growth". A snail's odyssey. Archived from the original on 5 July 2012. Retrieved 19 April 2012.
- ^ "Pacific razor clam". California Department of Fish and Game. 2001. Retrieved 9 May 2012.
- ^ Naylor, Martin (2005). "American jack knife clam, (Ensis directus)" (PDF). Alien species in Swedish seas and coastal areas. Archived from the original (PDF) on 4 September 2012. Retrieved 18 April 2012.
- ^ a b Carefoot, Tom (2010). "Learn about scallops: Predators and defenses". A snail's odyssey. Archived from the original on 20 January 2012. Retrieved 18 April 2012.
- ^ Bourquin, Avril (2000). "Bivalvia: The foot and locomotion". The Phylum Mollusca. Archived from the original on 24 April 2001. Retrieved 19 April 2012.
- .
- S2CID 91784258.
- S2CID 131914929.
- S2CID 20401676.
- ^ a b FAO State of Fisheries and Aquaculture 2012
- ^ a b c "Cultured Aquatic Species Information Programme: Ostrea edulis (Linnaeus, 1758)". FAO Fisheries and Aquaculture Department. Retrieved 19 May 2012.
- ^ a b "Cultured Aquatic Species Information Programme: Crassostrea gigas (Thunberg, 1793)". FAO Fisheries and Aquaculture Department. Retrieved 19 May 2012.
- ^ "Cultured Aquatic Species". FAO Fisheries and Aquaculture Department. Retrieved 19 May 2012.
- ^ "Fishery Statistical Collections: Global Aquaculture Production". FAO Fisheries and Aquaculture Department. Retrieved 23 May 2012.
- ^ "Daily life: Roman cuisine". Oracle ThinkQuest Education Foundation. Archived from the original on 8 May 2012. Retrieved 12 May 2012.
- ^ a b "Fishery Statistical Collections: Global Production". FAO Fisheries and Aquaculture Department. Retrieved 23 May 2012.
- ^ a b c d e f
Potasman, I.; Paz, A.; Odeh, M. (2002). "Infectious outbreaks associated with bivalve shellfish consumption: a worldwide perspective". Clinical Infectious Diseases. 35 (8): 921–928. PMID 12355378.
- ^
Rippey, S. R. (1994). "Infectious diseases associated with molluscan shellfish consumption". Clinical Microbiology. 7 (4): 419–425. PMID 7834599.
- S2CID 36548443.
- PMID 1661240.
- ^ a b c d Wekell, John C.; Hurst, John; Lefebvre, Kathi A. (2004). "The origin of the regulatory limits for PSP and ASP toxins in shellfish". Journal of Shellfish Research. 23 (3): 927–930.
- ^ "Amnesic Shellfish Poisoning". Harmful Algae. Woods Hole Oceanographic Institution. 2007. Retrieved 14 May 2012.
- ^ ISBN 9783319967769 Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
- .
- ^ Shulkin, V. M.; Kavun, V. I. A. (1995). "The use of marine bivalves in heavy metal monitoring near Vladivostok, Russia". Marine Pollution Bulletin. 31 (4–12): 330–333. .
- .
- S2CID 35586676. Archived from the original(PDF) on 16 July 2011.
- ^ Reilly, Michael (27 April 2009). "Sea Shells Used to Clean Up Heavy Metals". Discovery News. Archived from the original on 31 March 2012. Retrieved 18 May 2012.
- ISSN 2008-4935. Archived from the original(PDF) on 17 October 2013.
- ^ "The magnificent shells of the Smithsonian". Smithsonian.com. Retrieved 5 May 2012.
- ^ Catling, Chris (6 July 2009). "Shell Tools Rewrite Australasian Prehistory". World Archaeology. Archived from the original on 17 March 2013. Retrieved 18 May 2012.
- ISBN 978-0-8109-3689-8.
- ^ Kuhm, H. W. (2007). "Aboriginal uses of shell". The Wisconsin Archeologist. 17 (1): 1–8.
- ^
Hesse, Rayner W.; Hesse Jr., Rayner W. (2007). Jewelrymaking Through History: An Encyclopedia. Greenwood Publishing Group. p. 35. ISBN 978-0-313-33507-5.
- ISBN 978-0-415-01306-2.
- .
- ^ "Poultry Grit, Oystershell and Wood Shavings". Ascott Smallholding Supplies Ltd. Archived from the original on 7 August 2013. Retrieved 18 May 2012.
- ^ Hodin, Jessica (20 October 2010). "Contraband Chic: Mother-of-Pearl Items Sell With Export Restrictions". New York Observer. Retrieved 18 May 2012.
- ^ "Pearl Oyster Farming and Pearl Culture". Training manual produced by the Central Marine Fisheries Research Institute at Tuticorin, India. FAO Fisheries and Aquaculture Department. 1991. Retrieved 18 May 2012.
- ^ "Fulcanelli et la façade du palais Jacques Coeur" (in French). Fulcanelli, La rue de l'alchimie à travers l'architecture, les livres et les alchimistes. Retrieved 14 June 2012.
- ^ Gilmer, Maureen (19 March 2004). "Venus honored in Roman garden shrines". Chicago Sun-Times via HighBeam Research. Archived from the original on 11 May 2013. Retrieved 21 May 2012.
- ISBN 978-0-8118-0462-2.
- ^ "The Shell global homepage". Retrieved 21 May 2012.
- ISBN 978-0-8137-3026-4.
- ^ a b
Newell, Norman D. (1969). "Bivalvia Systematics". In Moore, R.C (ed.). Treatise on Invertebrate Paleontology Part N. The Paleontological Institute. ISBN 978-0-8137-3014-1.
- ^ a b Giribet, G.; Wheeler, W. (2002). "On bivalve phylogeny: a high-level analysis of the Bivalvia (Mollusca) based on combined morphology and DNA sequence data". Invertebrate Biology. 121 (4): 271–324. .
- ^ Bieler, R.; Mikkelsen, P. M. (2006). "Bivalvia – a look at the branches". Zoological Journal of the Linnean Society. 148 (3): 223–235. .
- ^ Mikkelsen, P. M.; Bieler, R.; Kappner, I.; Rawlings, T. A. (2006). "Phylogeny of Veneroidea (Mollusca: Bivalvia) based on morphology and molecules". Zoological Journal of the Linnean Society. 148 (3): 439–521. .
- ^ a b
Taylor, J. D.; Williams, S. T.; Glover, E. A.; Dyal, P. (November 2007). "A molecular phylogeny of heterodont bivalves (Mollusca: Bivalvia: Heterodonta): New analyses of 18S and 28S rRNA genes". Zoologica Scripta. 36 (6): 587–606. S2CID 84933022.
- ^ Taylor, John D.; Glover, Emily A.; Williams, Suzanne T. (2009). "Phylogenetic position of the bivalve family Cyrenoididae – removal from (and further dismantling of) the superfamily Lucinoidea". Nautilus. 123 (1): 9–13.
- ^ a b
Bouchet, Philippe; Rocroi, Jean-Pierre; Bieler, Rüdiger; Carter, Joseph G.; Coan, Eugene V. (2010). "Nomenclator of Bivalve Families with a Classification of Bivalve Families". S2CID 86546840.
- ^
Tëmkin, I. (2010). "Molecular phylogeny of pearl oysters and their relatives (Mollusca, Bivalvia, Pterioidea)". BMC Evolutionary Biology. 10: 342. PMID 21059254.
- ^ Taylor, John D.; Glover, Emily A.; Smith, Lisa; Dyal, Patricia; Williams, Suzanne T. (September 2011). "Molecular phylogeny and classification of the chemosymbiotic bivalve family Lucinidae (Mollusca: Bivalvia)". Zoological Journal of the Linnean Society. 163 (1): 15–49. . (subscription required)
- ^
Schneider, Jay A. (2001). "Bivalve systematics during the 20th century". Journal of Paleontology. 75 (6): 1119–1127. ISSN 0022-3360.
- ^
Ponder, W. F.; Lindberg, David R. (2008). Phylogeny and Evolution of the Mollusca. University of California Press. p. 117. ISBN 978-0-520-25092-5.
- ^ Harper, E. M.; Dreyer, H.; Steiner, G. (2006). "Reconstructing the Anomalodesmata (Mollusca: Bivalvia): morphology and molecules". Zoological Journal of the Linnean Society. 148 (3): 395–420. .
- ^ Franc, A. (1960). "Classe de Bivalves". In Grassé, P.-P. (ed.). Traité de Zoologie: Anatomie, Systématique, Biologie (in French). Vol. 5. Masson et Cie. pp. 1845–2164.
- ^ "Septibranchia". McGraw-Hill Dictionary of Scientific and Technical Terms. McGraw-Hill Companies. Retrieved 7 May 2012.
- ^ Gofas, Serge (2012). "Bivalvia". WoRMS. World Register of Marine Species. Retrieved 30 June 2012.
- ^ Carter, J. G.; Altaba, C. R.; Anderson, L. C.; Araujo, R.; Biakov, A. S.; Bogan, A. E.; Campbell, D. C.; Campbell, M.; Chen, J.; Cope, J. C. W.; Delvene. G.; Dijkstra, H. H.; Fang, Z.; Gardner, R. N.; Gavrilova, V. A.; Goncharova, I. A.; Harries, P. J.; Hartman, J. H.; Hautmann, M.; Hoeh, W. R.; Hylleberg, J.; Jiang, B.; Johnston, P.; Kirkendale, L.; Kleemann, K.; Koppka, J.; Kříž, J.; Machado, D.; Malchus, N.; Márquez-Aliaga, A.; Masse, J-P.; McRoberts, C. A.; Middelfart, P. U.; Mitchell, S.; Nevesskaja, L. A.; Özer, S.; Pojeta, J. Jr.; Polubotko, I. V.; Pons, J. M.; Popov, S.; Sánchez, T.; Sartori, A. F.; Scott, R. W.; Sey, I. I.; Signorelli, J. H.; Silantiev, V. V.; Skelton, P. W.; Steuber, T.; Waterhouse, J. B.; Wingard, G. L.; Yancey, T. (2011). "A synoptical classification of the Bivalvia (Mollusca)" (PDF). Paleontological Contributions. 4: 1–47.
Further reading
- Schneider, Jay A. (2001). "Bivalve systematics during the 20th century". Journal of Paleontology. 75 (6): 1119–1127. ISSN 0022-3360.
- Poutiers, J.-M.; Bernard, F. R. (1995). "Carnivorous bivalve molluscs (Anomalodesmata) from the tropical western Pacific Ocean, with a proposed classification and a catalogue of recent species". In Bouchet, P. (ed.). Résultats des Campagnes Musorstom. Mémoires Muséum National d'Histoire Naturelle. Vol. 167. pp. 107–188.
- Vaught, K. C. (1989). A Classification of the Living Mollusca. American Malacologists. ISBN 978-0-915826-22-3.