Mollusc

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

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  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.

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.

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 ''ta malákia'' (the soft ones; < ''malakós'' "soft"), which he applied '' inter alia ''to cuttlefish. The scientific study of molluscs is accordingly called malacology.

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.

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.

Molluscs have developed such a varied range of body structures, finding synapomorphies (defining characteristics) to apply to all modern groups is difficult. The most general characteristic of molluscs is they are unsegmented and bilaterally symmetrical. The following are present in all modern molluscs:
* The dorsal part of the body wall is a mantle (or pallium) which secretes calcareous spicules, plates or shells. It overlaps the body with enough spare room to form a mantle cavity.
* The anus and genitals open into the mantle cavity.
* There are two pairs of main nerve cords.

Other characteristics that commonly appear in textbooks have significant exceptions:

Diversity

Estimates of accepted described living species of molluscs vary from 50,000 to a maximum of 120,000 species. 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. In 2009, Chapman estimated the number of described living mollusc species at 85,000.; (online edition). Haszprunar in 2001 estimated about 93,000 named species, which include 23% of all named marine organisms. Molluscs are second only to arthropods in numbers of living animal species — far behind the arthropods' 1,113,000 but well ahead of chordates' 52,000. About 200,000 living species in total are estimated, and 70,000 fossil species, 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.

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. About 80% of all known mollusc species are gastropods. Cephalopoda such as squid, cuttlefish, and octopuses are among the neurologically most advanced of all invertebrates. The giant squid, which until recently had not been observed alive in its adult form, is one of the largest invertebrates, but a recently caught specimen of the colossal squid, long and weighing , may have overtaken it.

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.

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.

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". The visceral mass, or visceropallium, is the soft, nonmuscular metabolic region of the mollusc. It contains the body organs.

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. The whole soft body of bivalves lies within an enlarged mantle cavity.

Shell

The mantle edge secretes a shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs) that consists of mainly chitin and conchiolin (a protein hardened with calcium carbonate), except the outermost layer, which in almost all cases is all conchiolin (see periostracum). Molluscs never use phosphate to construct their hard parts, with the questionable exception of '' Cobcrephora''.
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.

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.

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.

Foot

The underside consists of a muscular foot, which has adapted to different purposes (locomotion, grasping the substratum, burrowing or feeding) in different classes. 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. In bivalves, the foot is adapted for burrowing into the sediment; page 4 in cephalopods it is used for jet propulsion, and the tentacles and arms are derived from the foot.

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. 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. 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.

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.

Eating, digestion, and excretion

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. 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".

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.

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.

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.

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.

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''.

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

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.

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. 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.

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.
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.

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''. Sacoglossan sea-slugs suck the sap from algae, using their one-row radula to pierce the cell walls, whereas dorid nudibranchs and some Vetigastropoda feed on sponges and others feed on hydroids. (An extensive list of molluscs with unusual feeding habits is available in the appendix of .)

Classification

Opinions vary about the number of classes of molluscs; for example, the table below shows seven living classes, 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. Two of the commonly recognized "classes" are known only from fossils.

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. Additionally, it suggests a sister taxon relationship exists between the Bivalvia and the Gastropoda. Tentaculita may also be in Mollusca (see ''Tentaculites'').

Evolution

Fossil record

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. , though arguably each of these may belong only to the stem lineage of their respective classes. 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 , has been described by some paleontologists as "mollusc-like", but others are unwilling to go further than "probable bilaterian", if that.

There is an even sharper debate about whether '' Wiwaxia'', from about , 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. Nicholas Butterfield, who opposes the idea that ''Wiwaxia'' was a mollusc, has written that earlier microfossils from are fragments of a genuinely mollusc-like radula. This appears to contradict the concept that the ancestral molluscan radula was mineralized.

However, the Helcionellids, which first appear over in Early Cambrian rocks from Siberia and China, are thought to be early molluscs with rather snail-like shells. Shelled molluscs therefore predate the earliest trilobites. 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.

Some analyses of helcionellids concluded these were the earliest gastropods. 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.

''Volborthella'', some fossils of which predate , 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. 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. All cephalopods with external shells except the nautiloids became extinct by the end of the Cretaceous period . However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.

The Early Cambrian fossils '' Fordilla'' and '' Pojetaia'' are regarded as bivalves. "Modern-looking" bivalves appeared in the Ordovician period, . One bivalve group, the rudists, became major reef-builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event. 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.

Phylogeny

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, debates arise about the relationships between the classes of living molluscs. In fact, some groups traditionally classified as molluscs may have to be redefined as distinct but related.

Molluscs are generally regarded members of the Lophotrochozoa, a group defined by having trochophore larvae and, in the case of living Lophophorata, a feeding structure called a lophophore

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