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Temporal range: Cambrian Stage 2 – Recent
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Subkingdom: Eumetazoa
Clade: ParaHoxozoa
Clade: Bilateria
Clade: Nephrozoa
(unranked): Protostomia
(unranked): Spiralia
Superphylum: Lophotrochozoa
Phylum: Mollusca
Linnaeus, 1758

See text.

Diversity [1]
85,000 recognized living species.
Cornu aspersum (formerly Helix aspersa) - a common land snail Snail-wiki-120-Zachi-Evenor.jpg
Cornu aspersum (formerly Helix aspersa) – a common land snail

Mollusca is the second-largest phylum of invertebrate animals, after Arthropoda; members are known as molluscs or mollusks [lower-alpha 1] ( /ˈmɒləsk/ ). Around 76,000  extant species of molluscs are recognized. [3] The number of fossil species is estimated between 60,000 and 100,000 additional species. [4] The proportion of undescribed species is very high. Many taxa remain poorly studied. [5]


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

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

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

Fossilized ammonite displayed at the National Museum of the Philippines Fossilized Ammonite Mollusk displayed at Philippine National Museum.jpg
Fossilized ammonite displayed at the National Museum of the Philippines

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

A handful of mollusc species are sometimes considered hazards or pests for human activities. The bite of the blue-ringed octopus is often fatal, and that of Octopus apollyon causes inflammation that can last over a month. Stings from a few species of large tropical cone shells of the family Conidae 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.


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

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


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

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

Other characteristics that commonly appear in textbooks have significant exceptions:

 Whether characteristic is found in these classes of Molluscs
Supposed universal Molluscan characteristic [14] Aplacophora [15] :291–292 Polyplacophora [15] :292–298 Monoplacophora [15] :298–300 Gastropoda [15] :300–343 Cephalopoda [15] :343–367 Bivalvia [15] :367–403 Scaphopoda [15] :403–407
Radula, a rasping "tongue" with chitinous teethAbsent in 20% of Neomeniomorpha YesYesYesYesNoInternal, cannot extend beyond body
Broad, muscular footReduced or absentYesYesYesModified into armsYesSmall, only at "front" end
Dorsal concentration of internal organs (visceral mass)Not obviousYesYesYesYesYesYes
Large digestive ceca No ceca in some AplacophoraYesYesYesYesYesNo
Large complex metanephridia ("kidneys")NoneYesYesYesYesYesSmall, simple
One or more valves/ shellsPrimitive forms, yes; modern forms, noYesYesSnails, yes; slugs, mostly yes (internal vestigial)Octopuses, no; cuttlefish, nautilus, squid, yesYesYes
Odontophore YesYesYesYesYesNoYes


Diversity and variability of shells of molluscs on display Berlin Naturkundemuseum Muscheln.jpg
Diversity and variability of shells of molluscs on display
About 80% of all known mollusc species are gastropods (snails and slugs), including this cowry (a sea snail). Cypraea chinensis with partially extended mantle.jpg
About 80% of all known mollusc species are gastropods (snails and slugs), including this cowry (a sea snail).

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

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

Freshwater and terrestrial molluscs appear exceptionally vulnerable to extinction. Estimates of the numbers of non-marine 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 non-marine 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 non-marine species. [26]


Anatomical diagram of a hypothetical ancestral mollusc Archimollusc-en.svg
Anatomical diagram of a hypothetical ancestral mollusc

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

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

Mantle and mantle cavity

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


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

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

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

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

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

Circulatory system

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

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


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

Eating, digestion, and excretion

Snail radula at work

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= Food
= Radula

= Muscles

= Odontophore "belt" Snail radula working.png
Snail radula at work
  = Food      = Radula
  = Muscles
  = Odontophore "belt"

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

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

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

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

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

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

Nervous system

Simplified diagram of the mollusc nervous system Gastropod nervous system.gif
Simplified diagram of the mollusc nervous system

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

The acephalic molluscs (i.e., bivalves) also have this ring but it is less obvious and less important. The bivalves have only three pairs of ganglia— cerebral, pedal, and visceral— with the visceral as the largest and most important of the three functioning as the principal center of "thinking".[ citation needed ] 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.


Trochophore larva 01.png
Apical tuft (cilia)
Prototroch (cilia)
Metatroch (cilia)
/// = cilia
Trochophore larva 01.png
Trochophore larva [36]

The simplest molluscan reproductive system relies on external fertilization, but with more complex variations. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. Two gonads sit next to the coelom, a small cavity that surrounds the heart, into which they shed ova or sperm. The nephridia extract the gametes from the coelom and emit them into the mantle cavity. Molluscs that use such a system remain of one sex all their lives and rely on external fertilization. Some molluscs use internal fertilization and/or are hermaphrodites, functioning as both sexes; both of these methods require more complex reproductive systems. [16] C. obtusus is an endemic snail species of the Eastern Alps. There is strong evidence for self-fertilization in the easternmost snail populations of this species. [37]

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. [38] 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. [39]



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. [40] 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. [39]

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 . [41] Sacoglossan sea-slugs suck the sap from algae, using their one-row radula to pierce the cell walls, [42] whereas dorid nudibranchs and some Vetigastropoda feed on sponges [43] [44] and others feed on hydroids. [45] (An extensive list of molluscs with unusual feeding habits is available in the appendix of GRAHAM, A. (1955). "Molluscan diets". Journal of Molluscan Studies. 31 (3–4): 144..)


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

ClassMajor organismsDescribed living species [19] Distribution
Gastropoda [15] :300all snails and slugs including abalone, limpets, conch, nudibranchs, sea hares, sea butterflies 70,000marine, freshwater, land
Bivalvia [15] :367 clams, oysters, scallops, geoducks, mussels, rudists20,000marine, freshwater
Polyplacophora [15] :292–298 chitons 1,000rocky tidal zone and seabed
Cephalopoda [15] :343 squid, octopuses, cuttlefish, nautiluses, Spirula , belemnites†, ammonites900marine
Scaphopoda [15] :403–407tusk shells500marine 6–7,000 metres (20–22,966 ft)
Cricoconarida [ citation needed ]extinct
Aplacophora [15] :291–292worm-like molluscs320seabed 200–3,000 metres (660–9,840 ft)
Monoplacophora [15] :298–300ancient lineage of molluscs with cap-like shells31seabed 1,800–7,000 metres (5,900–23,000 ft); one species 200 metres (660 ft)
Rostroconchia [46] fossils; probable ancestors of bivalves extinct marine
Helcionelloida [47] fossils; snail-like molluscs such as Latouchella extinctmarine

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


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

Fossil record

The enigmatic Kimberella quadrata (fossil pictured) from the Ediacaran has been described as being "mollusc-like" because of its features which are shared with modern day molluscs. Kimberella quadrata 4.jpg
The enigmatic Kimberella quadrata (fossil pictured) from the Ediacaran has been described as being "mollusc-like" because of its features which are shared with modern day molluscs.

Good evidence exists for the appearance of gastropods (e.g., Aldanella ), cephalopods (e.g., Plectronoceras, Nectocaris ?) and bivalves ( Pojetaia, Fordilla ) towards the middle of the Cambrian period, c. 500  million years ago, though arguably each of these may belong only to the stem lineage of their respective classes. [49] 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. [50] Kimberella , from about 555  million years ago, has been described by some paleontologists as "mollusc-like", [51] [52] but others are unwilling to go further than "probable bilaterian", [53] [54] if that. [55]

There is an even sharper debate about whether Wiwaxia , from about 505  million years ago, was a mollusc, and much of this centers on whether its feeding apparatus was a type of radula or more similar to that of some polychaete worms. [53] [56] Nicholas Butterfield, who opposes the idea that Wiwaxia was a mollusc, has written that earlier microfossils from 515 to 510 million years ago are fragments of a genuinely mollusc-like radula. [57] This appears to contradict the concept that the ancestral molluscan radula was mineralized. [58]

The tiny Helcionellid fossil Yochelcionella is thought to be an early mollusc Yochelcionella water flow.png
The tiny Helcionellid fossil Yochelcionella is thought to be an early mollusc
Spirally coiled shells appear in many gastropods. Neptunea despecta.jpg
Spirally coiled shells appear in many gastropods.

However, the Helcionellids, which first appear over 540  million years ago in Early Cambrian rocks from Siberia and China, [59] [60] are thought to be early molluscs with rather snail-like shells. Shelled molluscs therefore predate the earliest trilobites. [47] 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. [61]

Some analyses of helcionellids concluded these were the earliest gastropods. [62] However, other scientists are not convinced these Early Cambrian fossils show clear signs of the torsion that identifies modern gastropods twists the internal organs so the anus lies above the head. [15] :300–343 [63] [64]

Nautiloid septa n siphuncle 01.png
  = Septa
  = Siphuncle
Nautiloid septa n siphuncle 01.png
Septa and siphuncle in nautiloid shell

Volborthella , some fossils of which predate 530  million years ago, was long thought to be a cephalopod, but discoveries of more detailed fossils showed its shell was not secreted, but built from grains of the mineral silicon dioxide (silica), and it was not divided into a series of compartments by septa as those of fossil shelled cephalopods and the living Nautilus are. Volborthella's classification is uncertain. [65] 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. [66] All cephalopods with external shells except the nautiloids became extinct by the end of the Cretaceous period 65  million years ago. [67] However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today. [68]

The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves. [69] [70] [71] [72] "Modern-looking" bivalves appeared in the Ordovician period, 488 to 443 million years ago. [73] One bivalve group, the rudists, became major reef-builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event. [74] 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. [75]






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

Scaphopods (tusk shells)

(snails, slugs, limpets, sea hares)

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

(spicule-covered, worm-like)

Polyplacophorans (chitons)






A possible "family tree" of molluscs (2007). [76] [77] Does not include annelid worms as the analysis concentrated on fossilizable "hard" features. [76]

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, [52] [53] [56] [57] debates arise about the relationships between the classes of living molluscs. [54] In fact, some groups traditionally classified as molluscs may have to be redefined as distinct but related. [78]

Molluscs are generally regarded members of the Lophotrochozoa, [76] a group defined by having trochophore larvae and, in the case of living Lophophorata, a feeding structure called a lophophore. The other members of the Lophotrochozoa are the annelid worms and seven marine phyla. [79] 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. [80] For example, it is uncertain whether the ancestral mollusc was metameric (composed of repeating units)—if it was, that would suggest an origin from an annelid-like worm. [81] Scientists disagree about this: Giribet and colleagues concluded, in 2006, the repetition of gills and of the foot's retractor muscles were later developments, [12] while in 2007, Sigwart concluded the ancestral mollusc was metameric, and it had a foot used for creeping and a "shell" that was mineralized. [54] In one particular branch of the family tree, the shell of conchiferans is thought to have evolved from the spicules (small spines) of aplacophorans; but this is difficult to reconcile with the embryological origins of spicules. [80]

The molluscan shell appears to have originated from a mucus coating, which eventually stiffened into a cuticle. This would have been impermeable and thus forced the development of more sophisticated respiratory apparatus in the form of gills. [47] Eventually, the cuticle would have become mineralized, [47] using the same genetic machinery (engrailed) as most other bilaterian skeletons. [81] The first mollusc shell almost certainly was reinforced with the mineral aragonite. [28]

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

Morphological analyses tend to recover a conchiferan clade that receives less support from molecular analyses, [82] although these results also lead to unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups. [83]

However, an analysis in 2009 using both morphological and molecular phylogenetics comparisons concluded the molluscs are not monophyletic; in particular, Scaphopoda and Bivalvia are both separate, monophyletic lineages unrelated to the remaining molluscan classes; the traditional phylum Mollusca is polyphyletic, and it can only be made monophyletic if scaphopods and bivalves are excluded. [78] A 2010 analysis recovered the traditional conchiferan and aculiferan groups, and showed molluscs were monophyletic, demonstrating that available data for solenogastres was contaminated. [84] 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. [85] Rather than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan hypothesis into question. [86]

Human interaction

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

Uses by humans

Molluscs, especially bivalves such as clams and mussels, have been an important food source since at least the advent of anatomically modern humans, and this has often resulted in overfishing. [87] Other commonly eaten molluscs include octopuses and squids, whelks, oysters, and scallops. [88] In 2005, China accounted for 80% of the global mollusc catch, netting almost 11,000,000 tonnes (11,000,000 long tons; 12,000,000 short tons). Within Europe, France remained the industry leader. [89] Some countries regulate importation and handling of molluscs and other seafood, mainly to minimize the poison risk from toxins that can sometimes accumulate in the animals. [90]

Saltwater pearl oyster farm in Seram, Indonesia Pearl farm (Seram, Indonesia).jpg
Saltwater pearl oyster farm in Seram, Indonesia

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

The two methods of culturing pearls insert either "seeds" or beads into oysters. The "seed" method uses grains of ground shell from freshwater mussels, and overharvesting for this purpose has endangered several freshwater mussel species in the southeastern United States. [15] :367–403 The pearl industry is so important in some areas, significant sums of money are spent on monitoring the health of farmed molluscs. [91]

Byzantine Emperor Justinian I clad in Tyrian purple and wearing numerous pearls Meister von San Vitale in Ravenna 004.jpg
Byzantine Emperor Justinian I clad in Tyrian purple and wearing numerous pearls

Other luxury and high-status products were made from molluscs. Tyrian purple, made from the ink glands of murex shells, "fetched its weight in silver" in the fourth century BC, according to Theopompus. [92] 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. [93] [94] 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. [95] Procopius, writing on the Persian wars circa 550 CE, "stated that the five hereditary satraps (governors) of Armenia who received their insignia from the Roman Emperor were given chlamys (or cloaks) made from lana pinna. Apparently, only the ruling classes were allowed to wear these chlamys." [96]

Mollusc shells, including those of cowries, were used as a kind of money (shell money) in several preindustrial societies. However, these "currencies" generally differed in important ways from the standardized government-backed and -controlled money familiar to industrial societies. Some shell "currencies" were not used for commercial transactions, but mainly as social status displays at important occasions, such as weddings. [97] When used for commercial transactions, they functioned as commodity money, as a tradable commodity whose value differed from place to place, often as a result of difficulties in transport, and which was vulnerable to incurable inflation if more efficient transport or "goldrush" behavior appeared. [98]


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. [99] Potamopyrgus antipodarum is used by some water treatment plants to test for estrogen-mimicking pollutants from industrial agriculture. Several species of mollusca have been used as bioindicators of environmental stresses that can cause DNA damage. These species include the American oyster Crassostrea virginica, [100] zebra mussels (Dreissena polymorpha) [101] [102] and the blue mussel Mytilus edulis. [103]

Harm to humans

Stings and bites

The blue-ringed octopus's rings are a warning signal; this octopus is alarmed, and its bite can kill. Hapalochlaena lunulata.JPG
The blue-ringed octopus's rings are a warning signal; this octopus is alarmed, and its bite can kill.

Some molluscs sting or bite, but deaths from mollusc venoms total less than 10% of those from jellyfish stings. [105]

All octopuses are venomous, [106] 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, [104] but their venom kills 25% of human victims. Another tropical species, Octopus apollyon , causes severe inflammation that can last for over a month even if treated correctly, [107] and the bite of Octopus rubescens can cause necrosis that lasts longer than one month if untreated, and headaches and weakness persisting for up to a week even if treated. [108]

Live cone snails can be dangerous to shell collectors, but are useful to neurology researchers. Textile cone.JPG
Live cone snails can be dangerous to shell collectors, but are useful to neurology researchers.

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. [109] [105] [110] 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. [109] [111]

Disease vectors

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

Schistosomiasis (also known as bilharzia, bilharziosis or snail fever), a disease caused by the fluke worm Schistosoma , is "second only to malaria as the most devastating parasitic disease in tropical countries. An estimated 200 million people in 74 countries are infected with the disease – 100 million in Africa alone." [112] 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. [113]


Some species of molluscs, particularly certain snails and slugs, can be serious crop pests, [114] and when introduced into new environments, can unbalance local ecosystems. One such pest, the giant African snail Achatina fulica , has been introduced to many parts of Asia, as well as to many islands in the Indian Ocean and Pacific Ocean. In the 1990s, this species reached the West Indies. Attempts to control it by introducing the predatory snail Euglandina rosea proved disastrous, as the predator ignored Achatina fulica and went on to extirpate several native snail species instead. [115]

See also

Explanatory notes

  1. The formerly dominant U.K. spelling mollusk is still used in the U.S.—see the reasons given by Gary Rosenberg (1996). [2] For the spelling mollusc, see the reasons given in: Brusca & Brusca. Invertebrates (2nd ed.)..

Related Research Articles

<span class="mw-page-title-main">Octopus</span> Soft-bodied eight-limbed order of molluscs

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

<span class="mw-page-title-main">Squid</span> Superorder of cephalopod molluscs

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

<span class="mw-page-title-main">Cephalopod</span> Class of mollusks

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

<span class="mw-page-title-main">Gastropoda</span> Class of molluscs

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

<span class="mw-page-title-main">Bivalvia</span> Class of molluscs

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

The radula is an anatomical structure used by mollusks 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 enters the esophagus. The radula is unique to the mollusks, and is found in every class of mollusk except the bivalves, which instead use cilia, waving filaments that bring minute organisms to the mouth.

<span class="mw-page-title-main">Chiton</span> Class (Polyplacophora) of marine molluscs

Chitons are marine molluscs of varying size in the class Polyplacophora, formerly known as Amphineura. About 940 extant and 430 fossil species are recognized.

<span class="mw-page-title-main">Monoplacophora</span> Superclass of molluscs

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

<span class="mw-page-title-main">Coleoidea</span> Subclass of cephalopods

Coleoidea or Dibranchiata, is one of the two subclasses of cephalopods, containing all the various taxa popularly thought of as "soft-bodied" or "shell-less". Unlike its extant sister group Nautiloidea, whose members have a rigid outer shell for protection, the coleoids have at most an internal shell called cuttlebone or gladius that is used for buoyancy or as muscle anchorage. Some species, notably incirrate octopuses, have lost their cuttlebone altogether, while in some it has been replaced by a chitinous support structure. A unique trait of the group is the ability to edit their own RNA.

<span class="mw-page-title-main">Siphon (mollusc)</span> Anatomical structure which is part of the body of some aquatic molluscs

A siphon is an anatomical structure which is part of the body of aquatic molluscs in three classes: Gastropoda, Bivalvia and Cephalopoda.

<span class="mw-page-title-main">Tusk shell</span> Class of elephant tusk shell molluscs

The tusk shells or tooth shells, technically the Scaphopoda, are members of a class of shelled marine mollusc with worldwide distribution, and are the only class of exclusively infaunal marine molluscs. Shells of species within this class range in length 0.5–18 cm (0.20–7.09 in). Members of the order Dentaliida tend to be larger than those of the order Gadilida.

<span class="mw-page-title-main">Aplacophora</span> Class of molluscs

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

<span class="mw-page-title-main">Evolution of molluscs</span> The origin and diversification of molluscs through geologic time

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

<i>Odontogriphus</i> Genus of soft-bodied animals from middle Cambrian

Odontogriphus is a genus of soft-bodied animals known from middle Cambrian Lagerstätte. Reaching as much as 12.5 centimetres (4.9 in) in length, Odontogriphus is a flat, oval bilaterian which apparently had a single muscular foot and a "shell" on its back that was moderately rigid but of a material unsuited to fossilization.

<span class="mw-page-title-main">Marine invertebrates</span> Marine animals without a vertebrate column

Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.

<span class="mw-page-title-main">Helcionellid</span> Extinct order of molluscs

Helcionellid or Helcionelliformes is an order of small fossil shells that are universally interpreted as molluscs, though no sources spell out why this taxonomic interpretation is preferred. These animals are first found about 540 to 530 million years ago in the late Nemakit-Daldynian age, which is the earliest part of the Cambrian period. A single species persisted to the Early Ordovician. These fossils are component of the small shelly fossils (SSF) assemblages.

<span class="mw-page-title-main">Mollusc shell</span> Exoskeleton of an animal in the phylum Mollusca

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

<span class="mw-page-title-main">Brachiopod</span> Phylum of marine animals also known as lamp shells

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

<span class="mw-page-title-main">Digestive system of gastropods</span>

The digestive system of gastropods has evolved to suit almost every kind of diet and feeding behavior. Gastropods as the largest taxonomic class of the mollusca are very diverse: the group includes carnivores, herbivores, scavengers, filter feeders, and even parasites.

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


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