Molluscs Of Kenya
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Mollusca is the second-largest
phylum In biology, a phylum (; plural: phyla) 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 f ...
of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
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 Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...
phylum, comprising about 23% of all the named marine organisms. Numerous molluscs also live in
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...
and terrestrial habitats. They are highly diverse, not just in size and
anatomical Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
structure, but also in behaviour and habitat. The phylum is typically divided into 7 or 8 
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
classes, of which two are entirely
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
.
Cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
molluscs, such as
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 t ...
,
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 ...
, and
octopus An octopus ( : octopuses or octopodes, see below for variants) 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, cuttle ...
es, 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
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s ( snails and
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 smal ...
s) 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 A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle may refer to: *Mantle (clothing), a cloak-like garment worn mainly by women as fashionable outerwear **Mantle (vesture), an Eastern Orthodox ve ...
with a significant cavity used for breathing and excretion, the presence of a
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
(except for
bivalves 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, bival ...
), 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 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 indep ...
-like" shell on top, which is made of proteins and
chitin Chitin ( C8 H13 O5 N)n ( ) 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 chit ...
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 The coelom (or celom) 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 r ...
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 The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
play various important roles. The generalized mollusc has two paired nerve cords, or three in
bivalve 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, bival ...
s. 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 A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
, 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,
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s, and bivalves in the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was 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 ...
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 The giant Pacific octopus (''Enteroctopus dofleini''), also known as the North Pacific giant octopus, is a large marine cephalopod belonging to the genus ''Enteroctopus''. Its spatial distribution includes the coastal North Pacific, along Mexico ...
'' 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 Schistosomiasis, also known as snail fever, bilharzia, and Katayama fever, is a disease caused by parasitic flatworms called schistosomes. The urinary tract or the intestines may be infected. Symptoms include abdominal pain, diarrhea, bloody s ...
(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 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 esti ...
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 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 ...
. The scientific study of molluscs is accordingly called
malacology Malacology is the branch of invertebrate zoology that deals with the study of the Mollusca (mollusks or molluscs), the second-largest phylum of animals in terms of described species after the arthropods. Mollusks include snails and slugs, clams, ...
. 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 A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle may refer to: *Mantle (clothing), a cloak-like garment worn mainly by women as fashionable outerwear **Mantle (vesture), an Eastern Orthodox ve ...
with a significant cavity used for
breathing Breathing (or ventilation) is the process of moving air into and from the lungs to facilitate gas exchange with the internal environment, mostly to flush out carbon dioxide and bring in oxygen. All aerobic creatures need oxygen for cellular ...
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 In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
(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 Spicules are any of various small needle-like anatomical structures occurring in organisms Spicule may also refer to: * Spicule (sponge), small skeletal elements of sea sponges * Spicule (nematode), reproductive structures found in male nematodes ...
, plates or shells. It overlaps the body with enough spare room to form a mantle cavity. * The
anus The anus (Latin, 'ring' or 'circle') 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, d ...
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 In biology, a phylum (; plural: phyla) 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 f ...
. They include snails,
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 smal ...
s and other
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s; clams and other
bivalve 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, bival ...
s;
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 t ...
s and other
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s; 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.
Cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
a such as
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 t ...
,
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 ...
, and
octopus An octopus ( : octopuses or octopodes, see below for variants) 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, cuttle ...
es 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 Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...
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
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 ...
ns. The generalized mollusc is an unsegmented, bilaterally symmetrical animal and has a single, "
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 indep ...
-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 The anus (Latin, 'ring' or 'circle') 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, d ...
, a pair of osphradia (chemical sensors) in the incoming "lane", the hindmost pair of gills and the exit openings of the nephridia ("kidneys") and
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 sper ...
s (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 Chitin ( C8 H13 O5 N)n ( ) 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 chit ...
and conchiolin (a protein hardened with calcium carbonate), except the outermost layer, which in almost all cases is all conchiolin (see
periostracum The periostracum ( ) is a thin, organic coating (or "skin") that is the outermost layer of the shell of many shelled animals, including molluscs and brachiopods. Among molluscs, it is primarily seen in snails and clams, i.e. in gastropods and ...
). 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 Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
(sometimes with traces of aragonite) to construct the eggshells. The shell consists of three layers: the outer layer (the
periostracum The periostracum ( ) is a thin, organic coating (or "skin") that is the outermost layer of the shell of many shelled animals, including molluscs and brachiopods. Among molluscs, it is primarily seen in snails and clams, i.e. in gastropods and ...
) made of organic matter, a middle layer made of columnar
calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
, and an inner layer consisting of laminated calcite, often
nacre Nacre ( , ), also known as mother of pearl, is an organicinorganic composite material produced by some molluscs as an inner shell layer; it is also the material of which pearls are composed. It is strong, resilient, and iridescent. Nacre is f ...
ous. In some forms the shell contains openings. In
abalone Abalone ( or ; via Spanish , from Rumsen ''aulón'') is a common name for any of a group of small to very large marine gastropod molluscs in the family (biology), family Haliotidae. Other common name In biology, a common name of a taxon o ...
s 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 The siphuncle is a strand of tissue passing longitudinally through the shell of a cephalopod mollusk. Only cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the living nautiluses, cuttlefish, and ...
goes through all the chambers, and the eight plates that make up the shell of
chiton Chitons () are marine molluscs of varying size in the class Polyplacophora (), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized. They are also sometimes known as gumboots or sea cradles or coat-of-mail s ...
s 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
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 indep ...
s, 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 A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, an ...
, whose circulatory systems are
closed Closed may refer to: Mathematics * Closure (mathematics), a set, along with operations, for which applying those operations on members always results in a member of the set * Closed set, a set which contains all its limit points * Closed interval, ...
. Although molluscs are coelomates, their
coelom The coelom (or celom) 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 r ...
s are reduced to fairly small spaces enclosing the heart and gonads. The main body cavity is a hemocoel through which blood and
coelomic fluid The coelom (or celom) 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, ...
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 The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
, "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 Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
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
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
s that secrete slimy mucus, to which the food sticks. Beating
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
(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 Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
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 In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
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
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 sper ...
s sit next to the
coelom The coelom (or celom) 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 r ...
, a small cavity that surrounds the heart, into which they shed ova or
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
. 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
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
s, 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 The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical E ...
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 Xenophyophorea is a clade of foraminiferans. Members of this class are multinucleate unicellular organisms found on the ocean floor throughout the world's oceans, at depths of . They are a kind of foraminiferan that extract minerals from their s ...
'' Stannophyllum''.
Sacoglossan Sacoglossa, commonly known as the sacoglossans or the "solar-powered sea slugs", are a superorder of small sea slugs and sea snails, marine gastropod mollusks that belong to the clade Heterobranchia. Sacoglossans live by ingesting the cellular co ...
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 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. Anatomy Caudofov ...
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
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s (e.g., '' Aldanella''),
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s (e.g., '' Plectronoceras,'' '' Nectocaris'') and
bivalve 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, bival ...
s ('' Pojetaia, Fordilla'') towards the middle of the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was 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 ...
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 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 th ...
and Early
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was 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 ...
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
bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
n", 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 The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
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 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 indep ...
-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
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s. 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 ''Volborthella'' is an animal of uncertain classification, whose fossils pre-date . It has been considered for a period a cephalopod. However discoveries of more detailed fossils showed that ''Volborthella''’s small, conical shell was not secre ...
'', 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 The siphuncle is a strand of tissue passing longitudinally through the shell of a cephalopod mollusk. Only cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the living nautiluses, cuttlefish, and ...
, 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 Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish) ...
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 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 t ...
,
octopus An octopus ( : octopuses or octopodes, see below for variants) 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, cuttle ...
,
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 ...
) are abundant today. The Early Cambrian fossils '' Fordilla'' and '' Pojetaia'' are regarded as
bivalve 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, bival ...
s. "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 Hyoliths are animals with small conical shells, known as fossils from the Palaeozoic era. They are at least considered as lophotrochozoan, and possibly being lophophorates, a group which includes the brachiopods, while others consider them as be ...
are a class of extinct animals with a shell and operculum that may be molluscs. Authors who suggest they deserve their own
phylum In biology, a phylum (; plural: phyla) 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 f ...
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 "
halwaxiid Halwaxiida or halwaxiids is a proposed clade equivalent to the older orders Sachitida He 1980 and Thambetolepidea Jell 1981, loosely uniting scale-bearing Cambrian animals, which may lie in the stem group to molluscs or lophotrochozoa. Some palae ...
s" 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. The other members of the Lophotrochozoa are the
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
worms and seven marine
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phyl ...
. 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. 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 The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
-like worm. Scientists disagree about this: Giribet and colleagues concluded, in 2006, the repetition of gills and of the foot's retractor muscles were later developments, while in 2007, Sigwart concluded the ancestral mollusc was metameric, and it had a foot used for creeping and a "shell" that was mineralized. For a summary, see In one particular branch of the family tree, the shell of
conchifera Conchifera is a subphylum of the phylum Mollusca. It comprises all of the shell-bearing classes of molluscs, such as clams, tusk shells, ammonites, and monoplacophorans. The other subphylum is Aculifera. Non-monoplacophoran conchiferans emerged ...
ns is thought to have evolved from the
spicules Spicules are any of various small needle-like anatomical structures occurring in organisms Spicule may also refer to: * Spicule (sponge), small skeletal elements of sea sponges * Spicule (nematode), reproductive structures found in male nematodes ...
(small spines) of aplacophorans; but this is difficult to reconcile with the embryological origins of spicules. The molluscan shell appears to have originated from a mucus coating, which eventually stiffened into a
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
. This would have been impermeable and thus forced the development of more sophisticated respiratory apparatus in the form of gills. Eventually, the cuticle would have become mineralized, using the same genetic machinery ( engrailed) as most other bilaterian
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s. The first mollusc shell almost certainly was reinforced with the mineral aragonite. 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, although these results also lead to unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups. However, an analysis in 2009 using both morphological and molecular phylogenetics comparisons concluded the molluscs are not
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
; 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. A 2010 analysis recovered the traditional conchiferan and aculiferan groups, and showed molluscs were monophyletic, demonstrating that available data for solenogastres was contaminated. 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. Rather than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan hypothesis into question.


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
mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and Freshwater bivalve, freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other ...
s, have been an important food source since at least the advent of anatomically modern humans, and this has often resulted in overfishing. Other commonly eaten molluscs include
octopus An octopus ( : octopuses or octopodes, see below for variants) 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, cuttle ...
es and
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 t ...
s, whelks,
oysters Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not al ...
, and scallops. In 2005, China accounted for 80% of the global mollusc catch, netting almost . Within Europe, France remained the industry leader. Some countries regulate importation and handling of molluscs and other
seafood Seafood is any form of sea life regarded as food by humans, prominently including fish and shellfish. Shellfish include various species of molluscs (e.g. bivalve molluscs such as clams, oysters and mussels, and cephalopods such as octopus an ...
, mainly to minimize the poison risk from toxins that can sometimes accumulate in the animals. Most molluscs with shells can produce pearls, but only the pearls of
bivalve 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, bival ...
s and some
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s, whose shells are lined with
nacre Nacre ( , ), also known as mother of pearl, is an organicinorganic composite material produced by some molluscs as an inner shell layer; it is also the material of which pearls are composed. It is strong, resilient, and iridescent. Nacre is f ...
, are valuable. The best natural pearls are produced by marine pearl oysters, '' Pinctada margaritifera'' and ''Pinctada mertensi'', which live in the tropical and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
waters of the Pacific Ocean. Natural pearls form when a small foreign object gets stuck between the
mantle A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle may refer to: *Mantle (clothing), a cloak-like garment worn mainly by women as fashionable outerwear **Mantle (vesture), an Eastern Orthodox ve ...
and shell. The two methods of culturing
pearls A pearl is a hard, glistening object produced within the soft tissue (specifically the mantle (mollusc), mantle) of a living animal shell, shelled mollusk or another animal, such as fossil conulariids. Just like the shell of a mollusk, a pea ...
insert either "seeds" or beads into oysters. The "seed" method uses grains of ground shell from freshwater
mussel Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and Freshwater bivalve, freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other ...
s, and overharvesting for this purpose has endangered several freshwater mussel species in the southeastern United States. The pearl industry is so important in some areas, significant sums of money are spent on monitoring the health of farmed molluscs. 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. 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. 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. 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 The chlamys (Ancient Greek: χλαμύς : chlamýs, genitive: χλαμύδος : chlamydos) was a type of an ancient Greek cloak.
(or cloaks) made from ''lana pinna''. Apparently, only the ruling classes were allowed to wear these chlamys." 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 Social status is the level of social value a person is considered to possess. More specifically, it refers to the relative level of respect, honour, assumed competence, and deference accorded to people, groups, and organizations in a society. Stat ...
displays at important occasions, such as weddings. When used for commercial transactions, they functioned as
commodity In economics, a commodity is an economic good, usually a resource, that has full or substantial fungibility: that is, the market treats instances of the good as equivalent or nearly so with no regard to who produced them. The price of a comm ...
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.


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. Potamopyrgus antipodarum is used by some water treatment plants to test for estrogen-mimicking pollutants from industrial agriculture.


Harmful to humans


Stings and bites

Some molluscs sting or bite, but deaths from mollusc venoms total less than 10% of those from jellyfish stings. All octopuses are venomous, 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, but their venom kills 25% of human victims. Another tropical species, ''
Octopus apollyon The giant Pacific octopus (''Enteroctopus dofleini''), also known as the North Pacific giant octopus, is a large marine cephalopod belonging to the genus ''Enteroctopus''. Its spatial distribution includes the coastal North Pacific, along Mexico ...
'', causes severe inflammation that can last for over a month even if treated correctly, and the bite of '' Octopus rubescens'' can cause
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated dige ...
that lasts longer than one month if untreated, and headaches and weakness persisting for up to a week even if treated. 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. 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.


Disease vectors

Schistosomiasis Schistosomiasis, also known as snail fever, bilharzia, and Katayama fever, is a disease caused by parasitic flatworms called schistosomes. The urinary tract or the intestines may be infected. Symptoms include abdominal pain, diarrhea, bloody s ...
(also known as bilharzia, bilharziosis or snail fever), a disease caused by the fluke worm ''
Schistosoma ''Schistosoma'' is a genus of trematodes, commonly known as blood flukes. They are parasitic flatworms responsible for a highly significant group of infections in humans termed '' schistosomiasis'', which is considered by the World Health Organ ...
'', 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." 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 In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...
.


Pests

Some species of molluscs, particularly certain snails and slugs, can be serious crop pests, and when introduced into new environments, can unbalance local ecosystems. One such pest, the giant African snail ''
Achatina fulica ''Lissachatina fulica'' is a species of large land snail that belongs in the subfamily Achatininae of the family Achatinidae. It is also known as the Giant African land snail.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 ''Lissachatina fulica'' is a species of large land snail that belongs in the subfamily Achatininae of the family Achatinidae. It is also known as the Giant African land snail.Land snail A land snail is any of the numerous species of snail that live on land, as opposed to the sea snails and freshwater snails. ''Land snail'' is the common name for terrestrial gastropod mollusks that have shells (those without shells are known as ...
* Sea slug *
Sea snail Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the ...


Explanatory notes


References


Further reading

* Sturm, C.; Pearce, T.A. & Valdes, A. ''The Mollusks: A Guide to Their Study, Collection, and Preservation''. Universal Publishers. 2006. 454 pages. * 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.


External links

*
Researchers complete mollusk evolutionary tree; 26 October 2011

Hardy's Internet Guide to Marine Gastropods

Rotterdam Natural History Museum
Shell Image Gallery
Mussel Watch Programme
* Online biomonitoring of bivalve activity, 24/7
MolluSCAN ''eye''
{{Authority control Articles containing video clips Animal phyla Controversial taxa Extant Cambrian first appearances