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Mollusca

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Mollusca
Temporal range: Cambrian Stage 2 – Recent
Mollusk collage.jpg
Clockwise from top-left: Helix pomatia, a gastropod; Common octopus, a cephalopod; Tonicella lineata, a chiton; Atlantic surf clam, a bivalve.
Scientific classification e
Kingdom: Animalia
Subkingdom: Eumetazoa
Clade: ParaHoxozoa
Clade: Bilateria
Clade: Nephrozoa
(unranked): Protostomia
(unranked): Spiralia
Superphylum: Lophotrochozoa
Phylum: Mollusca
Linnaeus, 1758
Classes

See text.

Diversity[1]
85,000 recognized living species.
Cornu aspersum (formerly Helix aspersa) – a common land snail
Cornu aspersum (formerly Helix aspersa) – a common land snail

Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks[a] (/ˈmɒləsk/). Around 85,000 extant species of molluscs are recognized.[3] The number of fossil species is estimated between 60,000 and 100,000 additional species.[4] The proportion of undescribed species is very high. Many taxa remain poorly studied.[5]

Molluscs are the largest marine phylum, comprising about 23% of all the named marine organisms. Numerous molluscs also live in freshwater and terrestrial habitats. They are highly diverse, not just in size and anatomical structure, but also in behaviour and habitat. The phylum is typically divided into 7 or 8[6] taxonomic classes, of which two are entirely extinct. Cephalopod molluscs, such as squid, cuttlefish, and octopuses, are among the most neurologically advanced of all invertebrates—and either the giant squid or the colossal squid is the largest known invertebrate species. The gastropods (snails and slugs) are by far the most numerous molluscs and account for 80% of the total classified species.

The three most universal features defining modern molluscs are a mantle with a significant cavity used for breathing and excretion, the presence of a radula (except for bivalves), and the structure of the nervous system. Other than these common elements, molluscs express great morphological diversity, so many textbooks base their descriptions on a "hypothetical ancestral mollusc" (see image below). This has a single, "limpet-like" shell on top, which is made of proteins and chitin reinforced with calcium carbonate, and is secreted by a mantle covering the whole upper surface. The underside of the animal consists of a single muscular "foot". Although molluscs are coelomates, the coelom tends to be small. The main body cavity is a hemocoel through which blood circulates; as such, their circulatory systems are mainly open. The "generalized" mollusc's feeding system consists of a rasping "tongue", the radula, and a complex digestive system in which exuded mucus and microscopic, muscle-powered "hairs" called cilia play various important roles. The generalized mollusc has two paired nerve cords, or three in bivalves. The brain, in species that have one, encircles the esophagus. Most molluscs have eyes, and all have sensors to detect chemicals, vibrations, and touch. The simplest type of molluscan reproductive system relies on external fertilization, but more complex variations occur. Nearly all produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. The coelomic cavity is reduced. They have an open circulatory system and kidney-like organs for excretion.

Good evidence exists for the appearance of gastropods, cephalopods, and bivalves in the Cambrian period, 541–485.4 million years ago. However, the evolutionary history both of molluscs' emergence from the ancestral Lophotrochozoa and of their diversification into the well-known living and fossil forms are still subjects of vigorous debate among scientists.

Fossilized ammonite displayed at the National Museum of the Philippines
Fossilized ammonite displayed at the National Museum of the Philippines

Molluscs have been and still are an important food source for anatomically modern humans. Toxins that can accumulate in certain molluscs under specific conditions create a risk of food poisoning, and many jurisdictions have regulations to reduce this risk. Molluscs have, for centuries, also been the source of important luxury goods, notably pearls, mother of pearl, Tyrian purple dye, and sea silk. Their shells have also been used as money in some preindustrial societies.

A handful of mollusc species are sometimes considered hazards or pests for human activities. The bite of the blue-ringed octopus is often fatal, and that of Octopus apollyon causes inflammation that can last over a month. Stings from a few species of large tropical cone shells can also kill, but their sophisticated, though easily produced, venoms have become important tools in neurological research. Schistosomiasis (also known as bilharzia, bilharziosis, or snail fever) is transmitted to humans by water snail hosts, and affects about 200 million people. Snails and slugs can also be serious agricultural pests, and accidental or deliberate introduction of some snail species into new environments has seriously damaged some ecosystems.

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Class (biology)

Class (biology)

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

Cephalopod

Cephalopod

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

Cuttlefish

Cuttlefish

Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of buoyancy.

Colossal squid

Colossal squid

The colossal squid is part of the family Cranchiidae. It is sometimes called the Antarctic squid or giant cranch squid and is believed to be the largest squid species in terms of mass. It is the only recognized member of the genus Mesonychoteuthis and is known from only a small number of specimens. The species is confirmed to reach a mass of at least 495 kilograms (1,091 lb), though the largest specimens—known only from beaks found in sperm whale stomachs—may perhaps weigh as much as 600–700 kilograms (1,300–1,500 lb), making it the largest known invertebrate. Maximum total length has been estimated at 9–10 metres (30–33 ft). The colossal squid has the largest eyes of any known creature ever to exist, with an estimated diameter of 30–40 cm (12–16 in).

Chitin

Chitin

Chitin (C8H13O5N)n ( KY-tin) is a long-chain polymer of N-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chitin are produced each year in the biosphere. It is a primary component of cell walls in fungi (especially basidiomycetes and filamentous fungi), the exoskeletons of arthropods such as crustaceans and insects, the radulae, cephalopod beaks and gladii of molluscs and in some nematodes and diatoms. It is also synthesised by at least some fish and lissamphibians. Commercially, chitin is extracted from the shells of crabs, shrimps, shellfish and lobsters, which are major by-products of the seafood industry. The structure of chitin is comparable to cellulose, forming crystalline nanofibrils or whiskers. It is functionally comparable to the protein keratin. Chitin has proved useful for several medicinal, industrial and biotechnological purposes.

Calcium carbonate

Calcium carbonate

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

Coelom

Coelom

The coelom is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it remains undifferentiated. In the past, and for practical purposes, coelom characteristics have been used to classify bilaterian animal phyla into informal groups.

Blood

Blood

Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the circulatory system is also known as peripheral blood, and the blood cells it carries, peripheral blood cells.

Circulatory system

Circulatory system

The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, that consists of the heart and blood vessels. The circulatory system has two divisions, a systemic circulation or circuit, and a pulmonary circulation or circuit. Some sources use the terms cardiovascular system and vascular system interchangeably with the circulatory system.

Brain

Brain

A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a vertebrate's body. In a human, the cerebral cortex contains approximately 14–16 billion neurons, and the estimated number of neurons in the cerebellum is 55–70 billion. Each neuron is connected by synapses to several thousand other neurons. These neurons typically communicate with one another by means of long fibers called axons, which carry trains of signal pulses called action potentials to distant parts of the brain or body targeting specific recipient cells.

Cambrian

Cambrian

The Cambrian Period is the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago (mya) to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, and its base, are somewhat in flux. The period was established as "Cambrian series" by Adam Sedgwick, who named it after Cambria, the Latin name for 'Cymru' (Wales), where Britain's Cambrian rocks are best exposed. Sedgwick identified the layer as part of his task, along with Roderick Murchison, to subdivide the large "Transition Series", although the two geologists disagreed for a while on the appropriate categorization. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells. As a result, our understanding of the Cambrian biology surpasses that of some later periods.

Blue-ringed octopus

Blue-ringed octopus

Blue-ringed octopuses, comprising the genus Hapalochlaena, are four extremely venomous species of octopus that are found in tide pools and coral reefs in the Pacific and Indian oceans, from Japan to Australia. They can be identified by their yellowish skin and characteristic blue and black rings that change color dramatically when the animal is threatened. They eat small crustaceans, including crabs, hermit crabs, shrimp, and other small sea animals.

Etymology

The words mollusc and mollusk are both derived from the French mollusque, which originated from the Latin molluscus, from mollis, soft. Molluscus was itself an adaptation of Aristotle's τὰ μαλάκια ta malákia (the soft ones; μαλακός malakós "soft"), which he applied inter alia to cuttlefish.[7][8] The scientific study of molluscs is accordingly called malacology.[9]

The name Molluscoida was formerly used to denote a division of the animal kingdom containing the brachiopods, bryozoans, and tunicates, the members of the three groups having been supposed to somewhat resemble the molluscs. As now known, these groups have no relation to molluscs, and very little to one another, so the name Molluscoida has been abandoned.[10]

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Latin

Latin

Latin is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area around present-day Rome, but through the power of the Roman Republic it became the dominant language in the Italian region and subsequently throughout the Roman Empire. Even after the fall of Western Rome, Latin remained the common language of international communication, science, scholarship and academia in Europe until well into the 18th century, when other regional vernaculars supplanted it in common academic and political usage. For most of the time it was used, it would be considered a "dead language" in the modern linguistic definition; that is, it lacked native speakers, despite being used extensively and actively.

Aristotle

Aristotle

Aristotle was an Ancient Greek philosopher and polymath. His writings cover a broad range of subjects including physics, biology, zoology, metaphysics, logic, ethics, aesthetics, poetry, drama, music, rhetoric, psychology, linguistics, economics, politics, meteorology, geology, and government. As the founder of the Peripatetic school of philosophy in the Lyceum in Athens, he began the wider Aristotelian tradition that followed, which set the groundwork for the development of modern science.

Cuttlefish

Cuttlefish

Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of buoyancy.

Malacology

Malacology

Malacology is the branch of invertebrate zoology that deals with the study of the Mollusca, the second-largest phylum of animals in terms of described species after the arthropods. Mollusks include snails and slugs, clams, and cephalopods, along with numerous other kinds, many of which have shells. One division of malacology, conchology, is devoted to the study of mollusk shells. Malacology derives from Ancient Greek μαλακός (malakós) 'soft', and -λογία (-logía).

Brachiopod

Brachiopod

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

Bryozoa

Bryozoa

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

Tunicate

Tunicate

A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata. It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords. The subphylum was at one time called Urochordata, and the term urochordates is still sometimes used for these animals. They are the only chordates that have lost their myomeric segmentation, with the possible exception of the 'seriation of the gill slits'.

Definition

The most universal features of the body structure of molluscs are a mantle with a significant cavity used for breathing and excretion, and the organization of the nervous system. Many have a calcareous shell.[11]

Molluscs have developed such a varied range of body structures, finding synapomorphies (defining characteristics) to apply to all modern groups is difficult.[12] The most general characteristic of molluscs is they are unsegmented and bilaterally symmetrical.[13] The following are present in all modern molluscs:[14][16]

Other characteristics that commonly appear in textbooks have significant exceptions:

  Whether characteristic is found in these classes of Molluscs
Supposed universal Molluscan characteristic[14] Aplacophora[15]: 291–292  Polyplacophora[15]: 292–298  Monoplacophora[15]: 298–300  Gastropoda[15]: 300–343  Cephalopoda[15]: 343–367  Bivalvia[15]: 367–403  Scaphopoda[15]: 403–407 
Radula, a rasping "tongue" with chitinous teeth Absent in 20% of Neomeniomorpha Yes Yes Yes Yes No Internal, cannot extend beyond body
Broad, muscular foot Reduced or absent Yes Yes Yes Modified into arms Yes Small, only at "front" end
Dorsal concentration of internal organs (visceral mass) Not obvious Yes Yes Yes Yes Yes Yes
Large digestive ceca No ceca in some Aplacophora Yes Yes Yes Yes Yes No
Large complex metanephridia ("kidneys") None Yes Yes Yes Yes Yes Small, simple
One or more valves/ shells Primitive forms, yes; modern forms, no Yes Yes Snails, yes; slugs, mostly yes (internal vestigial) Octopuses, no; cuttlefish, nautilus, squid, yes Yes Yes
Odontophore Yes Yes Yes Yes Yes No Yes

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Mantle (mollusc)

Mantle (mollusc)

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

Respiration (physiology)

Respiration (physiology)

In physiology, respiration is the movement of oxygen from the outside environment to the cells within tissues, and the removal of carbon dioxide in the opposite direction that's to the environment.

Excretion

Excretion

Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after leaving the cell. Excretion is an essential process in all forms of life. For example, in mammals, urine is expelled through the urethra, which is part of the excretory system. In unicellular organisms, waste products are discharged directly through the surface of the cell.

Calcareous

Calcareous

Calcareous is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines.

Anus

Anus

The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, depending on the type of animal, includes: matter which the animal cannot digest, such as bones; food material after the nutrients have been extracted, for example cellulose or lignin; ingested matter which would be toxic if it remained in the digestive tract; and dead or excess gut bacteria and other endosymbionts.

Ventral nerve cord

Ventral nerve cord

The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, integrating sensory input and locomotor output. Because arthropods have an open circulatory system,decapitated insects can still walk, groom, and mate - illustrating that the circuitry of the ventral nerve cord is sufficient to perform complex motor programs without brain input.

Class (biology)

Class (biology)

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

Aplacophora

Aplacophora

Aplacophora is a presumably paraphyletic taxon. This is a class of small, deep-water, exclusively benthic, marine molluscs found in all oceans of the world.

Monoplacophora

Monoplacophora

Monoplacophora, meaning "bearing one plate", is a polyphyletic superclass of molluscs with a cap-like shell inhabiting deep sea environments. Extant representatives were not recognized as such until 1952; previously they were known only from the fossil record, and were thought to have become extinct 375 million years ago.

Gastropoda

Gastropoda

The gastropods, commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda.

Bivalvia

Bivalvia

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

Diversity

Diversity and variability of shells of molluscs on display
Diversity and variability of shells of molluscs on display
About 80% of all known mollusc species are gastropods (snails and slugs), including this cowry (a sea snail).[17]
About 80% of all known mollusc species are gastropods (snails and slugs), including this cowry (a sea snail).[17]

Estimates of accepted described living species of molluscs vary from 50,000 to a maximum of 120,000 species.[1] The total number of described species is difficult to estimate because of unresolved synonymy. In 1969 David Nicol estimated the probable total number of living mollusc species at 107,000 of which were about 12,000 fresh-water gastropods and 35,000 terrestrial. The Bivalvia would comprise about 14% of the total and the other five classes less than 2% of the living molluscs.[18] In 2009, Chapman estimated the number of described living mollusc species at 85,000.[1] Haszprunar in 2001 estimated about 93,000 named species,[19] which include 23% of all named marine organisms.[20] Molluscs are second only to arthropods in numbers of living animal species[17] — far behind the arthropods' 1,113,000 but well ahead of chordates' 52,000.[15]: Front endpaper  About 200,000 living species in total are estimated,[1][21] and 70,000 fossil species,[14] although the total number of mollusc species ever to have existed, whether or not preserved, must be many times greater than the number alive today.[22]

Molluscs have more varied forms than any other animal phylum. They include snails, slugs and other gastropods; clams and other bivalves; squids and other cephalopods; and other lesser-known but similarly distinctive subgroups. The majority of species still live in the oceans, from the seashores to the abyssal zone, but some form a significant part of the freshwater fauna and the terrestrial ecosystems. Molluscs are extremely diverse in tropical and temperate regions, but can be found at all latitudes.[12] About 80% of all known mollusc species are gastropods.[17] Cephalopoda such as squid, cuttlefish, and octopuses are among the neurologically most advanced of all invertebrates.[23] The giant squid, which until recently had not been observed alive in its adult form,[24] is one of the largest invertebrates, but a recently caught specimen of the colossal squid, 10 m (33 ft) long and weighing 500 kg (1,100 lb), may have overtaken it.[25]

Freshwater and terrestrial molluscs appear exceptionally vulnerable to extinction. Estimates of the numbers of nonmarine molluscs vary widely, partly because many regions have not been thoroughly surveyed. There is also a shortage of specialists who can identify all the animals in any one area to species. However, in 2004 the IUCN Red List of Threatened Species included nearly 2,000 endangered nonmarine molluscs. For comparison, the great majority of mollusc species are marine, but only 41 of these appeared on the 2004 Red List. About 42% of recorded extinctions since the year 1500 are of molluscs, consisting almost entirely of nonmarine species.[26]

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Snail

Snail

A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name snail is also used for most of the members of the molluscan class Gastropoda that have a coiled shell that is large enough for the animal to retract completely into. When the word "snail" is used in this most general sense, it includes not just land snails but also numerous species of sea snails and freshwater snails. Gastropods that naturally lack a shell, or have only an internal shell, are mostly called slugs, and land snails that have only a very small shell are often called semi-slugs.

Slug

Slug

Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word slug is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a small internal shell, particularly sea slugs and semislugs.

Freshwater snail

Freshwater snail

Freshwater snails are gastropod mollusks that live in fresh water. There are many different families. They are found throughout the world in various habitats, ranging from ephemeral pools to the largest lakes, and from small seeps and springs to major rivers. The great majority of freshwater gastropods have a shell, with very few exceptions. Some groups of snails that live in freshwater respire using gills, whereas other groups need to reach the surface to breathe air. In addition, some are amphibious and have both gills and a lung. Most feed on algae, but many are detritivores and some are filter feeders.

Arthropod

Arthropod

Arthropods are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, often mineralised with calcium carbonate. The arthropod body plan consists of segments, each with a pair of appendages. Arthropods are bilaterally symmetrical and their body possesses an external skeleton. In order to keep growing, they must go through stages of moulting, a process by which they shed their exoskeleton to reveal a new one. Some species have wings. They are an extremely diverse group, with up to 10 million species.

Chordate

Chordate

A chordate is an animal of the phylum Chordata. All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five synapomorphies include a notochord, dorsal hollow nerve cord, endostyle or thyroid, pharyngeal slits, and a post-anal tail. The name “chordate” comes from the first of these synapomorphies, the notochord, which plays a significant role in chordate structure and movement. Chordates are also bilaterally symmetric, have a coelom, possess a circulatory system, and exhibit metameric segmentation.

Phylum

Phylum

In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla, the plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.

Clam

Clam

Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shells of equal size connected by two adductor muscles and have a powerful burrowing foot. They live in both freshwater and marine environments; in salt water they prefer to burrow down into the mud and the turbidity of the water required varies with species and location; the greatest diversity of these is in North America.

Squid

Squid

True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting these criteria. Like all other cephalopods, squid have a distinct head, bilateral symmetry, and a mantle. They are mainly soft-bodied, like octopuses, but have a small internal skeleton in the form of a rod-like gladius or pen, made of chitin.

Cephalopod

Cephalopod

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

Abyssal zone

Abyssal zone

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

Ecosystem

Ecosystem

An ecosystem consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the system through photosynthesis and is incorporated into plant tissue. By feeding on plants and on one another, animals play an important role in the movement of matter and energy through the system. They also influence the quantity of plant and microbial biomass present. By breaking down dead organic matter, decomposers release carbon back to the atmosphere and facilitate nutrient cycling by converting nutrients stored in dead biomass back to a form that can be readily used by plants and microbes.

Hypothetical ancestral mollusc

Anatomical diagram of a hypothetical ancestral mollusc
Anatomical diagram of a hypothetical ancestral mollusc

Because of the great range of anatomical diversity among molluscs, many textbooks start the subject of molluscan anatomy by describing what is called an archi-mollusc, hypothetical generalized mollusc, or hypothetical ancestral mollusc (HAM) to illustrate the most common features found within the phylum. The depiction is visually rather similar to modern monoplacophorans.[12][16][27]

The generalized mollusc is an unsegmented, bilaterally symmetrical animal and has a single, "limpet-like" shell on top. The shell is secreted by a mantle covering the upper surface. The underside consists of a single muscular "foot".[16] The visceral mass, or visceropallium, is the soft, nonmuscular metabolic region of the mollusc. It contains the body organs.[13]

Mantle and mantle cavity

The mantle cavity, a fold in the mantle, encloses a significant amount of space. It is lined with epidermis, and is exposed, according to habitat, to sea, fresh water or air. The cavity was at the rear in the earliest molluscs, but its position now varies from group to group. The anus, a pair of osphradia (chemical sensors) in the incoming "lane", the hindmost pair of gills and the exit openings of the nephridia ("kidneys") and gonads (reproductive organs) are in the mantle cavity.[16] The whole soft body of bivalves lies within an enlarged mantle cavity.[13]

Shell

The mantle edge secretes a shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs[13]) that consists of mainly chitin and conchiolin (a protein hardened with calcium carbonate),[16][28] except the outermost layer, which in almost all cases is all conchiolin (see periostracum).[16] Molluscs never use phosphate to construct their hard parts,[29] with the questionable exception of Cobcrephora.[30] While most mollusc shells are composed mainly of aragonite, those gastropods that lay eggs with a hard shell use calcite (sometimes with traces of aragonite) to construct the eggshells.[31]

The shell consists of three layers: the outer layer (the periostracum) made of organic matter, a middle layer made of columnar calcite, and an inner layer consisting of laminated calcite, often nacreous.[13]

In some forms the shell contains openings. In abalones there are holes in the shell used for respiration and the release of egg and sperm, in the nautilus a string of tissue called the siphuncle goes through all the chambers, and the eight plates that make up the shell of chitons are penetrated with living tissue with nerves and sensory structures.[32]

Foot

A 50-second video of snails (most likely Natica chemnitzi and Cerithium stercusmuscaram) feeding on the sea floor in the Gulf of California, Puerto Peñasco, Mexico

The underside consists of a muscular foot, which has adapted to different purposes (locomotion, grasping the substratum, burrowing or feeding) in different classes.[33] The foot carries a pair of statocysts, which act as balance sensors. In gastropods, it secretes mucus as a lubricant to aid movement. In forms having only a top shell, such as limpets, the foot acts as a sucker attaching the animal to a hard surface, and the vertical muscles clamp the shell down over it; in other molluscs, the vertical muscles pull the foot and other exposed soft parts into the shell.[16] In bivalves, the foot is adapted for burrowing into the sediment;[33] in cephalopods it is used for jet propulsion,[33] and the tentacles and arms are derived from the foot.[34]

Circulatory system

Most molluscs' circulatory systems are mainly open, except for cephalopods, whose circulatory systems are closed. Although molluscs are coelomates, their coeloms are reduced to fairly small spaces enclosing the heart and gonads. The main body cavity is a hemocoel through which blood and coelomic fluid circulate and which encloses most of the other internal organs. These hemocoelic spaces act as an efficient hydrostatic skeleton.[13] The blood of these molluscs contains the respiratory pigment hemocyanin as an oxygen-carrier. The heart consists of one or more pairs of atria (auricles), which receive oxygenated blood from the gills and pump it to the ventricle, which pumps it into the aorta (main artery), which is fairly short and opens into the hemocoel.[16] The atria of the heart also function as part of the excretory system by filtering waste products out of the blood and dumping it into the coelom as urine. A pair of nephridia ("little kidneys") to the rear of and connected to the coelom extracts any re-usable materials from the urine and dumps additional waste products into it, and then ejects it via tubes that discharge into the mantle cavity.[16]

Exceptions to the above are the molluscs Planorbidae or ram's horn snails, which are air-breathing snails that use iron-based hemoglobin instead of the copper-based hemocyanin to carry oxygen through their blood.

Respiration

Most molluscs have only one pair of gills, or even only a singular gill. Generally, the gills are rather like feathers in shape, although some species have gills with filaments on only one side. They divide the mantle cavity so water enters near the bottom and exits near the top. Their filaments have three kinds of cilia, one of which drives the water current through the mantle cavity, while the other two help to keep the gills clean. If the osphradia detect noxious chemicals or possibly sediment entering the mantle cavity, the gills' cilia may stop beating until the unwelcome intrusions have ceased. Each gill has an incoming blood vessel connected to the hemocoel and an outgoing one to the heart.[16]

Eating, digestion, and excretion

Snail radula at work .mw-parser-output .legend{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .legend-color{display:inline-block;min-width:1.25em;height:1.25em;line-height:1.25;margin:1px 0;text-align:center;border:1px solid black;background-color:transparent;color:black}.mw-parser-output .legend-text{}  = Food       = Radula   = Muscles   = Odontophore "belt"
Snail radula at work
  = Food       = Radula
  = Muscles
  = Odontophore "belt"

Molluscs use intracellular digestion. Most molluscs have muscular mouths with radulae, "tongues", bearing many rows of chitinous teeth, which are replaced from the rear as they wear out. The radula primarily functions to scrape bacteria and algae off rocks, and is associated with the odontophore, a cartilaginous supporting organ.[13] The radula is unique to the molluscs and has no equivalent in any other animal.

Molluscs' mouths also contain glands that secrete slimy mucus, to which the food sticks. Beating cilia (tiny "hairs") drive the mucus towards the stomach, so the mucus forms a long string called a "food string".[16]

At the tapered rear end of the stomach and projecting slightly into the hindgut is the prostyle, a backward-pointing cone of feces and mucus, which is rotated by further cilia so it acts as a bobbin, winding the mucus string onto itself. Before the mucus string reaches the prostyle, the acidity of the stomach makes the mucus less sticky and frees particles from it.[16]

The particles are sorted by yet another group of cilia, which send the smaller particles, mainly minerals, to the prostyle so eventually they are excreted, while the larger ones, mainly food, are sent to the stomach's cecum (a pouch with no other exit) to be digested. The sorting process is by no means perfect.[16]

Periodically, circular muscles at the hindgut's entrance pinch off and excrete a piece of the prostyle, preventing the prostyle from growing too large. The anus, in the part of the mantle cavity, is swept by the outgoing "lane" of the current created by the gills. Carnivorous molluscs usually have simpler digestive systems.[16]

As the head has largely disappeared in bivalves, the mouth has been equipped with labial palps (two on each side of the mouth) to collect the detritus from its mucus.[13]

Nervous system

Simplified diagram of the mollusc nervous system
Simplified diagram of the mollusc nervous system

The cephalic molluscs have two pairs of main nerve cords organized around a number of paired ganglia, the visceral cords serving the internal organs and the pedal ones serving the foot. Most pairs of corresponding ganglia on both sides of the body are linked by commissures (relatively large bundles of nerves). The ganglia above the gut are the cerebral, the pleural, and the visceral, which are located above the esophagus (gullet). The pedal ganglia, which control the foot, are below the esophagus and their commissure and connectives to the cerebral and pleural ganglia surround the esophagus in a circumesophageal nerve ring or nerve collar.[16]

The acephalic molluscs (i.e., bivalves) also have this ring but it is less obvious and less important. The bivalves have only three pairs of ganglia— cerebral, pedal, and visceral— with the visceral as the largest and most important of the three functioning as the principal center of "thinking". Some such as the scallops have eyes around the edges of their shells which connect to a pair of looped nerves and which provide the ability to distinguish between light and shadow.

Reproduction

Apical tuft (cilia)
Prototroch (cilia)
Stomach
Mouth
Metatroch (cilia)
Mesoderm
Anus
/// = cilia



 Trochophore larva[35]
Apical tuft (cilia)
Prototroch (cilia)
Stomach
Mouth
Metatroch (cilia)
Mesoderm
Anus
/// = cilia
Apical tuft (cilia)
Prototroch (cilia)
Stomach
Mouth
Metatroch (cilia)
Mesoderm
Anus
/// = cilia



 Trochophore larva[35]
Trochophore larva[35]

The simplest molluscan reproductive system relies on external fertilization, but with more complex variations. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. Two gonads sit next to the coelom, a small cavity that surrounds the heart, into which they shed ova or sperm. The nephridia extract the gametes from the coelom and emit them into the mantle cavity. Molluscs that use such a system remain of one sex all their lives and rely on external fertilization. Some molluscs use internal fertilization and/or are hermaphrodites, functioning as both sexes; both of these methods require more complex reproductive systems.[16]

The most basic molluscan larva is a trochophore, which is planktonic and feeds on floating food particles by using the two bands of cilia around its "equator" to sweep food into the mouth, which uses more cilia to drive them into the stomach, which uses further cilia to expel undigested remains through the anus. New tissue grows in the bands of mesoderm in the interior, so the apical tuft and anus are pushed further apart as the animal grows. The trochophore stage is often succeeded by a veliger stage in which the prototroch, the "equatorial" band of cilia nearest the apical tuft, develops into the velum ("veil"), a pair of cilia-bearing lobes with which the larva swims. Eventually, the larva sinks to the seafloor and metamorphoses into the adult form. While metamorphosis is the usual state in molluscs, the cephalopods differ in exhibiting direct development: the hatchling is a 'miniaturized' form of the adult.[36] The development of molluscs is of particular interest in the field of ocean acidification as environmental stress is recognized to affect the settlement, metamorphosis, and survival of larvae.[37]

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Mollusc shell

Mollusc shell

The mollusc shell is typically a calcareous exoskeleton which encloses, supports and protects the soft parts of an animal in the phylum Mollusca, which includes snails, clams, tusk shells, and several other classes. Not all shelled molluscs live in the sea; many live on the land and in freshwater.

Limpet

Limpet

Limpets are a group of aquatic snails that exhibit a conical shell shape (patelliform) and a strong, muscular foot. Limpets are members of the class Gastropoda, but are polyphyletic, meaning the various groups called "limpets" descended independently from different ancestral gastropods. This general category of conical shell is known as "patelliform" (dish-shaped). All members of the large and ancient marine clade Patellogastropoda are limpets. Within that clade, the members of the Patellidae family in particular are often referred to as "true limpets".

Gastropod shell

Gastropod shell

The gastropod shell is part of the body of a gastropod or snail, a kind of mollusc. The shell is an exoskeleton, which protects from predators, mechanical damage, and dehydration, but also serves for muscle attachment and calcium storage. Some gastropods appear shell-less (slugs) but may have a remnant within the mantle, or in some cases the shell is reduced such that the body cannot be retracted within it (semi-slug). Some snails also possess an operculum that seals the opening of the shell, known as the aperture, which provides further protection. The study of mollusc shells is known as conchology. The biological study of gastropods, and other molluscs in general, is malacology. Shell morphology terms vary by species group.

Habitat

Habitat

In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical manifestation of its ecological niche. Thus "habitat" is a species-specific term, fundamentally different from concepts such as environment or vegetation assemblages, for which the term "habitat-type" is more appropriate.

Anus

Anus

The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, depending on the type of animal, includes: matter which the animal cannot digest, such as bones; food material after the nutrients have been extracted, for example cellulose or lignin; ingested matter which would be toxic if it remained in the digestive tract; and dead or excess gut bacteria and other endosymbionts.

Gill

Gill

A gill is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are kept moist. The microscopic structure of a gill presents a large surface area to the external environment. Branchia is the zoologists' name for gills.

Gonad

Gonad

A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sperm in the form of spermatozoa. The female gonad, the ovary, produces egg cells. Both of these gametes are haploid cells. Some hermaphroditic animals have a type of gonad called an ovotestis.

Chitin

Chitin

Chitin (C8H13O5N)n ( KY-tin) is a long-chain polymer of N-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chitin are produced each year in the biosphere. It is a primary component of cell walls in fungi (especially basidiomycetes and filamentous fungi), the exoskeletons of arthropods such as crustaceans and insects, the radulae, cephalopod beaks and gladii of molluscs and in some nematodes and diatoms. It is also synthesised by at least some fish and lissamphibians. Commercially, chitin is extracted from the shells of crabs, shrimps, shellfish and lobsters, which are major by-products of the seafood industry. The structure of chitin is comparable to cellulose, forming crystalline nanofibrils or whiskers. It is functionally comparable to the protein keratin. Chitin has proved useful for several medicinal, industrial and biotechnological purposes.

Conchiolin

Conchiolin

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

Calcium carbonate

Calcium carbonate

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

Cobcrephora

Cobcrephora

Cobcrephora is a genus of that resembles the Palaeoloricates, known from the Silurian of Gotland. Its interpretation as a polyplacophoran is widely challenged.

Aragonite

Aragonite

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

Ecology

Feeding

Most molluscs are herbivorous, grazing on algae or filter feeders. For those grazing, two feeding strategies are predominant. Some feed on microscopic, filamentous algae, often using their radula as a 'rake' to comb up filaments from the sea floor. Others feed on macroscopic 'plants' such as kelp, rasping the plant surface with its radula. To employ this strategy, the plant has to be large enough for the mollusc to 'sit' on, so smaller macroscopic plants are not as often eaten as their larger counterparts.[38] Filter feeders are molluscs that feed by straining suspended matter and food particles from water, typically by passing the water over their gills. Most bivalves are filter feeders, which can be measured through clearance rates. Research has demonstrated that environmental stress can affect the feeding of bivalves by altering the energy budget of organisms.[37]

Cephalopods are primarily predatory, and the radula takes a secondary role to the jaws and tentacles in food acquisition. The monoplacophoran Neopilina uses its radula in the usual fashion, but its diet includes protists such as the xenophyophore Stannophyllum.[39] Sacoglossan sea-slugs suck the sap from algae, using their one-row radula to pierce the cell walls,[40] whereas dorid nudibranchs and some Vetigastropoda feed on sponges[41][42] and others feed on hydroids.[43] (An extensive list of molluscs with unusual feeding habits is available in the appendix of GRAHAM, A. (1955). "Molluscan diets". Journal of Molluscan Studies. 31 (3–4): 144..)

Classification

Opinions vary about the number of classes of molluscs; for example, the table below shows seven living classes,[19] and two extinct ones. Although they are unlikely to form a clade, some older works combine the Caudofoveata and Solenogasters into one class, the Aplacophora.[27][15]: 291–292  Two of the commonly recognized "classes" are known only from fossils.[17]

Class Major organisms Described living species[19] Distribution
Gastropoda [15]: 300  all snails and slugs including abalone, limpets, conch, nudibranchs, sea hares, sea butterflies 70,000 marine, freshwater, land
Bivalvia [15]: 367  clams, oysters, scallops, geoducks, mussels, rudists 20,000 marine, freshwater
Polyplacophora [15]: 292–298  chitons 1,000 rocky tidal zone and seabed
Cephalopoda [15]: 343  squid, octopuses, cuttlefish, nautiluses, Spirula, belemnites†, ammonites 900 marine
Scaphopoda [15]: 403–407  tusk shells 500 marine 6–7,000 metres (20–22,966 ft)
Cricoconarida extinct
Aplacophora [15]: 291–292  worm-like molluscs 320 seabed 200–3,000 metres (660–9,840 ft)
Monoplacophora [15]: 298–300  ancient lineage of molluscs with cap-like shells 31 seabed 1,800–7,000 metres (5,900–23,000 ft); one species 200 metres (660 ft)
Rostroconchia[44] fossils; probable ancestors of bivalves extinct marine
Helcionelloida[45] fossils; snail-like molluscs such as Latouchella extinct marine

Classification into higher taxa for these groups has been and remains problematic. A phylogenetic study suggests the Polyplacophora form a clade with a monophyletic Aplacophora.[46] Additionally, it suggests a sister taxon relationship exists between the Bivalvia and the Gastropoda. Tentaculita may also be in Mollusca (see Tentaculites).

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List of mollusc orders

List of mollusc orders

List of mollusc orders illustrates the 97 orders in the phylum Mollusca, the largest marine animal phylum. 85,000 extant species are described, making up 23% of described marine organisms.

Class (biology)

Class (biology)

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

Caudofoveata

Caudofoveata

Caudofoveata is a small class within the phylum Mollusca, also known as Chaetodermomorpha. The class is often combined with Solenogastres and termed Aplacophora, but some studies have cast doubt on the monophyly of this group.

Aplacophora

Aplacophora

Aplacophora is a presumably paraphyletic taxon. This is a class of small, deep-water, exclusively benthic, marine molluscs found in all oceans of the world.

Gastropoda

Gastropoda

The gastropods, commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda.

Abalone

Abalone

Abalone is a common name for any of a group of small to very large marine gastropod molluscs in the family Haliotidae. Other common names are ear shells, sea ears, and, rarely, muttonfish or muttonshells in parts of Australia, ormer in the UK, perlemoen in South Africa, and paua in New Zealand. Abalones are marine snails. Their taxonomy puts them in the family Haliotidae, which contains only one genus, Haliotis, which once contained six subgenera. These subgenera have become alternative representations of Haliotis. The number of species recognized worldwide ranges between 30 and 130 with over 230 species-level taxa described. The most comprehensive treatment of the family considers 56 species valid, with 18 additional subspecies. The shells of abalones have a low, open spiral structure, and are characterized by several open respiratory pores in a row near the shell's outer edge. The thick inner layer of the shell is composed of nacre (mother-of-pearl), which in many species is highly iridescent, giving rise to a range of strong, changeable colors which make the shells attractive to humans as decorative objects, jewelry, and as a source of colorful mother-of-pearl.

Limpet

Limpet

Limpets are a group of aquatic snails that exhibit a conical shell shape (patelliform) and a strong, muscular foot. Limpets are members of the class Gastropoda, but are polyphyletic, meaning the various groups called "limpets" descended independently from different ancestral gastropods. This general category of conical shell is known as "patelliform" (dish-shaped). All members of the large and ancient marine clade Patellogastropoda are limpets. Within that clade, the members of the Patellidae family in particular are often referred to as "true limpets".

Conch

Conch

Conch is a common name of a number of different medium-to-large-sized sea snails. Conch shells typically have a high spire and a noticeable siphonal canal.

Bivalvia

Bivalvia

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

Clam

Clam

Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shells of equal size connected by two adductor muscles and have a powerful burrowing foot. They live in both freshwater and marine environments; in salt water they prefer to burrow down into the mud and the turbidity of the water required varies with species and location; the greatest diversity of these is in North America.

Geoduck

Geoduck

The Pacific geoduck is a species of very large saltwater clam in the family Hiatellidae. The common name is derived from the Lushootseed (Nisqually) word gʷídəq.

Mussel

Mussel

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

Evolution

The use of love darts by the land snail Monachoides vicinus is a form of sexual selection
The use of love darts by the land snail Monachoides vicinus is a form of sexual selection

Fossil record

The enigmatic Kimberella quadrata (fossil pictured) from the Ediacaran has been described as being "mollusc-like" because of its features which are shared with modern day molluscs.
The enigmatic Kimberella quadrata (fossil pictured) from the Ediacaran has been described as being "mollusc-like" because of its features which are shared with modern day molluscs.

Good evidence exists for the appearance of gastropods (e.g., Aldanella), cephalopods (e.g., Plectronoceras, Nectocaris) and bivalves (Pojetaia, Fordilla) towards the middle of the Cambrian period, c. 500 million years ago, though arguably each of these may belong only to the stem lineage of their respective classes.[47] However, the evolutionary history both of the emergence of molluscs from the ancestral group Lophotrochozoa, and of their diversification into the well-known living and fossil forms, is still vigorously debated.

Debate occurs about whether some Ediacaran and Early Cambrian fossils really are molluscs. Kimberella, from about 555 million years ago, has been described by some paleontologists as "mollusc-like",[48][49] but others are unwilling to go further than "probable bilaterian",[50][51] if that.[52]

There is an even sharper debate about whether Wiwaxia, from about 505 million years ago, was a mollusc, and much of this centers on whether its feeding apparatus was a type of radula or more similar to that of some polychaete worms.[50][53] Nicholas Butterfield, who opposes the idea that Wiwaxia was a mollusc, has written that earlier microfossils from 515 to 510 million years ago are fragments of a genuinely mollusc-like radula.[54] This appears to contradict the concept that the ancestral molluscan radula was mineralized.[55]

The tiny Helcionellid fossil Yochelcionella is thought to be an early mollusc[45]
The tiny Helcionellid fossil Yochelcionella is thought to be an early mollusc[45]
Spirally coiled shells appear in many gastropods.[15]: 300–343 
Spirally coiled shells appear in many gastropods.[15]: 300–343 

However, the Helcionellids, which first appear over 540 million years ago in Early Cambrian rocks from Siberia and China,[56][57] are thought to be early molluscs with rather snail-like shells. Shelled molluscs therefore predate the earliest trilobites.[45] Although most helcionellid fossils are only a few millimeters long, specimens a few centimeters long have also been found, most with more limpet-like shapes. The tiny specimens have been suggested to be juveniles and the larger ones adults.[58]

Some analyses of helcionellids concluded these were the earliest gastropods.[59] However, other scientists are not convinced these Early Cambrian fossils show clear signs of the torsion that identifies modern gastropods twists the internal organs so the anus lies above the head.[15]: 300–343 [60][61]

  = Septa
  = Siphuncle



 Septa and siphuncle in nautiloid shell
  = Septa
  = Siphuncle
  = Septa
  = Siphuncle



 Septa and siphuncle in nautiloid shell
Septa and siphuncle in nautiloid shell

Volborthella, some fossils of which predate 530 million years ago, was long thought to be a cephalopod, but discoveries of more detailed fossils showed its shell was not secreted, but built from grains of the mineral silicon dioxide (silica), and it was not divided into a series of compartments by septa as those of fossil shelled cephalopods and the living Nautilus are. Volborthella's classification is uncertain.[62] The Middle Cambrian fossil Nectocaris is often interpreted as a cephalopod with 2 arms and no shell, but the Late Cambrian fossil Plectronoceras is now thought to be the earliest undisputed cephalopod fossil, as its shell had septa and a siphuncle, a strand of tissue that Nautilus uses to remove water from compartments it has vacated as it grows, and which is also visible in fossil ammonite shells. However, Plectronoceras and other early cephalopods crept along the seafloor instead of swimming, as their shells contained a "ballast" of stony deposits on what is thought to be the underside, and had stripes and blotches on what is thought to be the upper surface.[63] All cephalopods with external shells except the nautiloids became extinct by the end of the Cretaceous period 65 million years ago.[64] However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.[65]

The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves.[66][67][68][69] "Modern-looking" bivalves appeared in the Ordovician period, 488 to 443 million years ago.[70] One bivalve group, the rudists, became major reef-builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event.[71] Even so, bivalves remain abundant and diverse.

The Hyolitha are a class of extinct animals with a shell and operculum that may be molluscs. Authors who suggest they deserve their own phylum do not comment on the position of this phylum in the tree of life.[72]

Phylogeny

Lophotrochozoa

Brachiopods

Mollusca

Bivalves

Monoplacophorans
("limpet-like", "living fossils")

Scaphopods (tusk shells)

Gastropods
(snails, slugs, limpets, sea hares)

Cephalopods
(nautiloids, ammonites, octopus, squid, etc.)

Aplacophorans
(spicule-covered, worm-like)

Polyplacophorans (chitons)

† Halwaxiids

Wiwaxia

Halkieria

Orthrozanclus

Odontogriphus

A possible "family tree" of molluscs (2007).[73][74] Does not include annelid worms as the analysis concentrated on fossilizable "hard" features.[73]

The phylogeny (evolutionary "family tree") of molluscs is a controversial subject. In addition to the debates about whether Kimberella and any of the "halwaxiids" were molluscs or closely related to molluscs,[49][50][53][54] debates arise about the relationships between the classes of living molluscs.[51] In fact, some groups traditionally classified as molluscs may have to be redefined as distinct but related.[75]

Molluscs are generally regarded members of the Lophotrochozoa,[73] a group defined by having trochophore larvae and, in the case of living Lophophorata, a feeding structure called a lophophore. The other members of the Lophotrochozoa are the annelid worms and seven marine phyla.[76] The diagram on the right summarizes a phylogeny presented in 2007 without the annelid worms.

Because the relationships between the members of the family tree are uncertain, it is difficult to identify the features inherited from the last common ancestor of all molluscs.[77] For example, it is uncertain whether the ancestral mollusc was metameric (composed of repeating units)—if it was, that would suggest an origin from an annelid-like worm.[78] Scientists disagree about this: Giribet and colleagues concluded, in 2006, the repetition of gills and of the foot's retractor muscles were later developments,[12] while in 2007, Sigwart concluded the ancestral mollusc was metameric, and it had a foot used for creeping and a "shell" that was mineralized.[51] In one particular branch of the family tree, the shell of conchiferans is thought to have evolved from the spicules (small spines) of aplacophorans; but this is difficult to reconcile with the embryological origins of spicules.[77]

The molluscan shell appears to have originated from a mucus coating, which eventually stiffened into a cuticle. This would have been impermeable and thus forced the development of more sophisticated respiratory apparatus in the form of gills.[45] Eventually, the cuticle would have become mineralized,[45] using the same genetic machinery (engrailed) as most other bilaterian skeletons.[78] The first mollusc shell almost certainly was reinforced with the mineral aragonite.[28]

The evolutionary relationships within the molluscs are also debated, and the diagrams below show two widely supported reconstructions:

Morphological analyses tend to recover a conchiferan clade that receives less support from molecular analyses,[79] although these results also lead to unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups.[80]

However, an analysis in 2009 using both morphological and molecular phylogenetics comparisons concluded the molluscs are not monophyletic; in particular, Scaphopoda and Bivalvia are both separate, monophyletic lineages unrelated to the remaining molluscan classes; the traditional phylum Mollusca is polyphyletic, and it can only be made monophyletic if scaphopods and bivalves are excluded.[75] A 2010 analysis recovered the traditional conchiferan and aculiferan groups, and showed molluscs were monophyletic, demonstrating that available data for solenogastres was contaminated.[81] Current molecular data are insufficient to constrain the molluscan phylogeny, and since the methods used to determine the confidence in clades are prone to overestimation, it is risky to place too much emphasis even on the areas of which different studies agree.[82] Rather than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan hypothesis into question.[83]

Discover more about Evolution related topics

Evolution of molluscs

Evolution of molluscs

The evolution of the molluscs is the way in which the Mollusca, one of the largest groups of invertebrate animals, evolved. This phylum includes gastropods, bivalves, scaphopods, cephalopods, and several other groups. The fossil record of mollusks is relatively complete, and they are well represented in most fossil-bearing marine strata. Very early organisms which have dubiously been compared to molluscs include Kimberella and Odontogriphus.

Evolution of cephalopods

Evolution of cephalopods

The cephalopods have a long geological history, with the first nautiloids found in late Cambrian strata, and purported stem-group representatives present in the earliest Cambrian lagerstätten.

Love dart

Love dart

A love dart is a sharp, calcareous or chitinous dart which some hermaphroditic land snails and slugs create. Love darts are both formed and stored internally in a dart sac. These darts are made in sexually mature animals only, and are used as part of the sequence of events during courtship, before actual mating takes place. Darts are quite large compared to the size of the animal: in the case of the semi-slug genus Parmarion, the length of a dart can be up to one fifth that of the semi-slug's foot.

Monachoides vicinus

Monachoides vicinus

Monachoides vicinus is a species of air-breathing land snail, a terrestrial pulmonate gastropod mollusk in the family Hygromiidae, the hairy snails and their allies.

Kimberella

Kimberella

Kimberella is an extinct genus of bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity with this group is contentious.

Ediacaran

Ediacaran

The Ediacaran Period is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and the beginning of the Phanerozoic Eon. It is named after the Ediacara Hills of South Australia.

Aldanella

Aldanella

Aldanella is an extinct paleozoic mollusc that was assigned to the Gastropod stem group but may also belong to a paraphyletic "Monoplacophora".

Cephalopod

Cephalopod

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

Fordilla

Fordilla

Fordilla is an extinct genus of early bivalves, one of two genera in the extinct family Fordillidae. The genus is known solely from Early Cambrian fossils found in North America, Greenland, Europe, the Middle East, and Asia. The genus currently contains three described species, Fordilla germanica, Fordilla sibirica, and the type species Fordilla troyensis.

Cambrian

Cambrian

The Cambrian Period is the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago (mya) to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, and its base, are somewhat in flux. The period was established as "Cambrian series" by Adam Sedgwick, who named it after Cambria, the Latin name for 'Cymru' (Wales), where Britain's Cambrian rocks are best exposed. Sedgwick identified the layer as part of his task, along with Roderick Murchison, to subdivide the large "Transition Series", although the two geologists disagreed for a while on the appropriate categorization. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells. As a result, our understanding of the Cambrian biology surpasses that of some later periods.

Lophotrochozoa

Lophotrochozoa

Lophotrochozoa is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, brachiopods, and platyhelminthes.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Human interaction

For millennia, molluscs have been a source of food for humans, as well as important luxury goods, notably pearls, mother of pearl, Tyrian purple dye, sea silk, and chemical compounds. Their shells have also been used as a form of currency in some preindustrial societies. A number of species of molluscs can bite or sting humans, and some have become agricultural pests.

Uses by humans

Molluscs, especially bivalves such as clams and mussels, have been an important food source since at least the advent of anatomically modern humans, and this has often resulted in overfishing.[84] Other commonly eaten molluscs include octopuses and squids, whelks, oysters, and scallops.[85] In 2005, China accounted for 80% of the global mollusc catch, netting almost 11,000,000 tonnes (11,000,000 long tons; 12,000,000 short tons). Within Europe, France remained the industry leader.[86] Some countries regulate importation and handling of molluscs and other seafood, mainly to minimize the poison risk from toxins that can sometimes accumulate in the animals.[87]

Saltwater pearl oyster farm in Seram, Indonesia
Saltwater pearl oyster farm in Seram, Indonesia

Most molluscs with shells can produce pearls, but only the pearls of bivalves and some gastropods, whose shells are lined with nacre, are valuable.[15]: 300–343, 367–403  The best natural pearls are produced by marine pearl oysters, Pinctada margaritifera and Pinctada mertensi, which live in the tropical and subtropical waters of the Pacific Ocean. Natural pearls form when a small foreign object gets stuck between the mantle and shell.

The two methods of culturing pearls insert either "seeds" or beads into oysters. The "seed" method uses grains of ground shell from freshwater mussels, and overharvesting for this purpose has endangered several freshwater mussel species in the southeastern United States.[15]: 367–403  The pearl industry is so important in some areas, significant sums of money are spent on monitoring the health of farmed molluscs.[88]

Byzantine Emperor Justinian I clad in Tyrian purple and wearing numerous pearls
Byzantine Emperor Justinian I clad in Tyrian purple and wearing numerous pearls

Other luxury and high-status products were made from molluscs. Tyrian purple, made from the ink glands of murex shells, "fetched its weight in silver" in the fourth century BC, according to Theopompus.[89] The discovery of large numbers of Murex shells on Crete suggests the Minoans may have pioneered the extraction of "imperial purple" during the Middle Minoan period in the 20th–18th centuries BC, centuries before the Tyrians.[90][91] Sea silk is a fine, rare, and valuable fabric produced from the long silky threads (byssus) secreted by several bivalve molluscs, particularly Pinna nobilis, to attach themselves to the sea bed.[92] Procopius, writing on the Persian wars circa 550 CE, "stated that the five hereditary satraps (governors) of Armenia who received their insignia from the Roman Emperor were given chlamys (or cloaks) made from lana pinna. Apparently, only the ruling classes were allowed to wear these chlamys."[93]

Mollusc shells, including those of cowries, were used as a kind of money (shell money) in several preindustrial societies. However, these "currencies" generally differed in important ways from the standardized government-backed and -controlled money familiar to industrial societies. Some shell "currencies" were not used for commercial transactions, but mainly as social status displays at important occasions, such as weddings.[94] When used for commercial transactions, they functioned as commodity money, as a tradable commodity whose value differed from place to place, often as a result of difficulties in transport, and which was vulnerable to incurable inflation if more efficient transport or "goldrush" behavior appeared.[95]

Bioindicators

Bivalve molluscs are used as bioindicators to monitor the health of aquatic environments in both fresh water and the marine environments. Their population status or structure, physiology, behaviour or the level of contamination with elements or compounds can indicate the state of contamination status of the ecosystem. They are particularly useful since they are sessile so that they are representative of the environment where they are sampled or placed.[96] Potamopyrgus antipodarum is used by some water treatment plants to test for estrogen-mimicking pollutants from industrial agriculture.

Harmful to humans

Stings and bites

The blue-ringed octopus's rings are a warning signal; this octopus is alarmed, and its bite can kill.[97]
The blue-ringed octopus's rings are a warning signal; this octopus is alarmed, and its bite can kill.[97]

Some molluscs sting or bite, but deaths from mollusc venoms total less than 10% of those from jellyfish stings.[98]

All octopuses are venomous,[99] but only a few species pose a significant threat to humans. Blue-ringed octopuses in the genus Hapalochlaena, which live around Australia and New Guinea, bite humans only if severely provoked,[97] but their venom kills 25% of human victims. Another tropical species, Octopus apollyon, causes severe inflammation that can last for over a month even if treated correctly,[100] and the bite of Octopus rubescens can cause necrosis that lasts longer than one month if untreated, and headaches and weakness persisting for up to a week even if treated.[101]

Live cone snails can be dangerous to shell collectors, but are useful to neurology researchers.[102]
Live cone snails can be dangerous to shell collectors, but are useful to neurology researchers.[102]

All species of cone snails are venomous and can sting painfully when handled, although many species are too small to pose much of a risk to humans, and only a few fatalities have been reliably reported. Their venom is a complex mixture of toxins, some fast-acting and others slower but deadlier.[102][98][103] The effects of individual cone-shell toxins on victims' nervous systems are so precise as to be useful tools for research in neurology, and the small size of their molecules makes it easy to synthesize them.[102][104]

Disease vectors

Skin vesicles created by the penetration of Schistosoma. (Source: CDC)
Skin vesicles created by the penetration of Schistosoma. (Source: CDC)

Schistosomiasis (also known as bilharzia, bilharziosis or snail fever), a disease caused by the fluke worm Schistosoma, is "second only to malaria as the most devastating parasitic disease in tropical countries. An estimated 200 million people in 74 countries are infected with the disease – 100 million in Africa alone."[105] The parasite has 13 known species, two of which infect humans. The parasite itself is not a mollusc, but all the species have freshwater snails as intermediate hosts.[106]

Pests

Some species of molluscs, particularly certain snails and slugs, can be serious crop pests,[107] and when introduced into new environments, can unbalance local ecosystems. One such pest, the giant African snail Achatina fulica, has been introduced to many parts of Asia, as well as to many islands in the Indian Ocean and Pacific Ocean. In the 1990s, this species reached the West Indies. Attempts to control it by introducing the predatory snail Euglandina rosea proved disastrous, as the predator ignored Achatina fulica and went on to extirpate several native snail species instead.[108]

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Molluscs in culture

Molluscs in culture

Culture consists of the social behaviour and norms in human societies transmitted through social learning. Molluscs play a variety of roles in culture, including but not limited to art and literature, with both practical interactions—whether useful or harmful—and symbolic uses.

Pearl

Pearl

A pearl is a hard, glistening object produced within the soft tissue of a living shelled mollusk or another animal, such as fossil conulariids. Just like the shell of a mollusk, a pearl is composed of calcium carbonate in minute crystalline form, which has deposited in concentric layers. The ideal pearl is perfectly round and smooth, but many other shapes, known as baroque pearls, can occur. The finest quality of natural pearls have been highly valued as gemstones and objects of beauty for many centuries. Because of this, pearl has become a metaphor for something rare, fine, admirable and valuable.

Sea silk

Sea silk

Sea silk is an extremely fine, rare, and valuable fabric that is made from the long silky filaments or byssus secreted by a gland in the foot of pen shells. The byssus is used by the clam to attach itself to the sea bed.

Currency

Currency

A currency is a standardization of money in any form, in use or circulation as a medium of exchange, for example banknotes and coins. A more general definition is that a currency is a system of money in common use within a specific environment over time, especially for people in a nation state. Under this definition, the British Pound Sterling (£), euros (€), Japanese yen (¥), and U.S. dollars (US$) are examples of (government-issued) fiat currencies. Currencies may act as stores of value and be traded between nations in foreign exchange markets, which determine the relative values of the different currencies. Currencies in this sense are either chosen by users or decreed by governments, and each type has limited boundaries of acceptance; i.e., legal tender laws may require a particular unit of account for payments to government agencies.

Seashell

Seashell

A seashell or sea shell, also known simply as a shell, is a hard, protective outer layer usually created by an animal or organism that lives in the sea. The shell is part of the body of the animal. Empty seashells are often found washed up on beaches by beachcombers. The shells are empty because the animal has died and the soft parts have decomposed or been eaten by another animal.

List of edible molluscs

List of edible molluscs

This is a partial list of edible molluscs. Molluscs are a large phylum of invertebrate animals, many of which have shells. Edible molluscs are harvested from saltwater, freshwater, and the land, and include numerous members of the classes Gastropoda (snails), Bivalvia, Cephalopoda, and Polyplacophora (chitons).

Clam

Clam

Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shells of equal size connected by two adductor muscles and have a powerful burrowing foot. They live in both freshwater and marine environments; in salt water they prefer to burrow down into the mud and the turbidity of the water required varies with species and location; the greatest diversity of these is in North America.

Mussel

Mussel

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

Octopus

Octopus

An octopus is a soft-bodied, eight-limbed mollusc of the order Octopoda. The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, an octopus is bilaterally symmetric with two eyes and a beaked mouth at the center point of the eight limbs. The soft body can radically alter its shape, enabling octopuses to squeeze through small gaps. They trail their eight appendages behind them as they swim. The siphon is used both for respiration and for locomotion, by expelling a jet of water. Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse of all invertebrates.

Squid

Squid

True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting these criteria. Like all other cephalopods, squid have a distinct head, bilateral symmetry, and a mantle. They are mainly soft-bodied, like octopuses, but have a small internal skeleton in the form of a rod-like gladius or pen, made of chitin.

Seafood

Seafood

Seafood is any form of sea life regarded as food by humans, prominently including fish and shellfish. Shellfish include various species of molluscs, crustaceans, and echinoderms. Historically, marine mammals such as cetaceans as well as seals have been eaten as food, though that happens to a lesser extent in modern times. Edible sea plants such as some seaweeds and microalgae are widely eaten as sea vegetables around the world, especially in Asia.

Indonesia

Indonesia

Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guinea. Indonesia is the world's largest archipelagic state and the 14th-largest country by area, at 1,904,569 square kilometres. With over 275 million people, Indonesia is the world's fourth-most populous country and the most populous Muslim-majority country. Java, the world's most populous island, is home to more than half of the country's population.

Source: "Mollusca", Wikipedia, Wikimedia Foundation, (2023, February 14th), https://en.wikipedia.org/wiki/Mollusca.

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Explanatory notes
  1. ^ The formerly dominant U.K. spelling mollusk is still used in the U.S.—see the reasons given by Gary Rosenberg (1996).[2] For the spelling mollusc, see the reasons given in: Brusca & Brusca. Invertebrates (2nd ed.)..
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Further reading
  • Sturm, C.; Pearce, T.A. & Valdes, A. The Mollusks: A Guide to Their Study, Collection, and Preservation. Universal Publishers. 2006. 454 pages. ISBN 1581129300
  • Trigo, J.E.; Díaz Agras, G.J.; García-Álvarez, O.L.; Guerra, A.; Moreira, J.; Pérez, J.; Rolán, E.; Troncoso, J.S. & Urgorri, V. (2018). Troncoso, J.S., Trigo, J.E. & Rolán, E., ed. Guía de los Moluscos Marinos de Galicia. Vigo: Servicio de Publicacións da Universidade de Vigo. 836 pages. ISBN 978-84-8158-787-6
External links

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