Product Code Database
Mollusca is a phylum of protostome invertebrate , whose members are known as molluscs or mollusks (). Around 76,000 extant taxon species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The number of additional fossil species is estimated between 60,000 and 100,000, (2025). 9780674019720, Harvard University Press. ISBN 9780674019720 and the proportion of undescribed species is very high. Many taxa remain poorly studied.
Molluscs are the largest marine biology phylum, comprising about 23% of all the named . They are highly diverse, not just in size and anatomical structure, but also in behaviour and habitat, as numerous groups are freshwater and even terrestrial species. The phylum is typically divided into 7 or 8 taxonomic classes, of which two are entirely extinct. Cephalopod molluscs, such as squid, cuttlefish, and , are among the most neurobiology of all —and either the giant squid or the colossal squid is the largest known extant invertebrate species. The (, and abalone) are by far the most diverse class and account for 80% of the total classified molluscan species.
The four most universal features defining modern molluscs are a soft body composed almost entirely of 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" gastropod shell on top, which is made of 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 , 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 . The brain, in species that have one, encircles the esophagus. Most molluscs have , 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 , 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, , and bivalves in the Cambrian period, 541–485.4 million years ago. However, the evolutionary history both of molluscs' emergence from the ancestral Lophotrochozoa and of their diversification into the well-known living and fossil forms are still subjects of vigorous debate among scientists. Molluscs have been and still are an important food source for 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 , mother of pearl, Tyrian purple dye, and sea silk. Their shells have also been Shell money 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 Enteroctopus dofleini causes inflammation that can last over a month. Stings from a few species of large tropical 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 .
The name Molluscoida was formerly used to denote a division of the animal kingdom containing the , , and , the members of the three groups having been supposed to somewhat resemble the molluscs. As now known, Brachipoda, Bryozoa and Mollusca are all part of the Lophotrochozoa but have very little relation to the Tunicata, so the name Molluscoida has been abandoned.
Molluscs have developed such a varied range of body structures, finding synapomorphy (defining characteristics) to apply to all modern groups is difficult. The most general characteristic of molluscs is they are unsegmented and bilaterally symmetrical. (2025). 9780199549443, Oxford University Press. ISBN 9780199549443 The following are present in all modern molluscs: (2025). 9780878930975, Sinauer Associates. ISBN 9780878930975
| Whether characteristic is found in these classes of Molluscs | ||||||
Molluscs have more varied forms than any other animal phylum. They include , and other ; and other ; and other ; 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 . Molluscs are extremely diverse in Tropics and temperate regions, but can be found at all . About 80% of all known mollusc species are gastropods. such as squid, cuttlefish, and are among the most neurologically advanced of all invertebrates. The giant squid, which until recently had not been observed alive in its adult form, is one of the largest , surpassed in weight but not in length by the colossal squid.
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.
The generalized mollusc is an unsegmented, bilaterally symmetrical animal and has a single, "limpet-like" gastropod 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.
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 .
In some forms the shell contains openings. In abalone 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 are penetrated with living tissue with nerves and sensory structures. (2006). 9781139458474, Cambridge University Press. ISBN 9781139458474 The earliest-derived living mollusca, the Polyplacophora (chitons) and shell-less vermiform Aplacophora, remains contentious despite many developmental and molecular studies of these organisms. Both studies investigated molluscan phylogeny through synthesis of paleontological and neontological data, and the other being large and small-subunit nuclear rRna sequences. Another article describing the chitons, with their shells, this time, is one by D K Jacobs and others, where shell evolution of molluscs and chitons are examined to visualize the evolution of the chiton, from the Polyplacophora, to the Aplacophora, and others.
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.
Molluscs' mouths also contain that secrete slimy mucus, to which the food sticks. Beating cilia (tiny "hairs") drive the mucus towards the stomach, so the mucus forms a long string called a "food string".
At the tapered rear end of the stomach and projecting slightly into the hindgut is the prostyle, a backward-pointing cone of feces and mucus, which is rotated by further cilia so it acts as a bobbin, winding the mucus string onto itself. Before the mucus string reaches the prostyle, the acidity of the stomach makes the mucus less sticky and frees particles from it.
The particles are sorted by yet another group of cilia, which send the smaller particles, mainly minerals, to the prostyle so eventually they are excreted, while the larger ones, mainly food, are sent to the stomach's cecum (a pouch with no other exit) to be digested. The sorting process is by no means perfect.
Periodically, circular muscles at the hindgut's entrance pinch off and excrete a piece of the prostyle, preventing the prostyle from growing too large. The anus, in the part of the mantle cavity, is swept by the outgoing "lane" of the current created by the gills. Carnivorous molluscs usually have simpler digestive systems.
As the head has largely disappeared in bivalves, the mouth has been equipped with labial palps (two on each side of the mouth) to collect the detritus from its mucus.
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.
The most basic molluscan larva is a trochophore, which is 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 metamorphosis 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.
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 such as the xenophyophore Stannophyllum. 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 and others feed on hydroids. (An extensive list of molluscs with unusual feeding habits is available in the appendix of )
| marine, freshwater, land |
| marine, freshwater |
| rocky tidal zone and seabed |
| marine |
| marine |
| seabed |
| seabed ; one species |
| marine |
| marine |
| extinct |
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 worms and seven marine phylum. 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-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 is thought to have evolved from the (small spines) of ; but this is difficult to reconcile with the embryology origins of spicules.
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. Eventually, the cuticle would have become mineralized,using the same genetic machinery (engrailed) as most other bilaterian . The first mollusc shell almost certainly was reinforced with the mineral aragonite.
Classification into higher taxa for molluscan classes has been and remains problematic. Numerous different clades have been proposed but few have received strong support. Traditionally, Mollusca is split into two subphyla, Conchifera and Aculifera, based on the presence of a shell. The "Testaria" hypothesis is similar, but includes chitons with the rest of the conchiferans. Some studies completely reject the proposal, instead favoring a "Serialia" hypothesis which classifies chitons and monoplacophorans as closely related.
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 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.
A 2025 analysis, using newly sequenced genomes, supports the Aculifera-Conchifera hypothesis. Scaphopoda was recovered sister to Bivalvia, and Cephalopoda as sister to the Bivalvia-Scaphopoda-Gastropoda clade.
Debate occurs about whether some Ediacaran and Early Cambrian fossils really are molluscs. Kimberella, from about , has been described by some paleontologists as "mollusc-like", but others are unwilling to go further than "probable ", if that.
There is an even sharper debate about whether Wiwaxia, from about , was a mollusc, and much of this centers on whether its feeding apparatus was a type of radula or more similar to that of some polychaete worms. Nicholas Butterfield, who opposes the idea that Wiwaxia was a mollusc, has written that earlier from are fragments of a genuinely mollusc-like radula. This appears to contradict the concept that the ancestral molluscan radula was mineralized.
Some analyses of helcionellids concluded these were the earliest . However, other scientists are not convinced these Early Cambrian fossils show clear signs of the torsion that identifies modern gastropods twists the internal organs so the anus lies above the head.
Volborthella, some fossils of which predate , was long thought to be a cephalopod, but discoveries of more detailed fossils showed its shell was not secreted, but built from grains of the mineral silicon dioxide (silica), and it was not divided into a series of compartments by septa as those of fossil shelled cephalopods and the living Nautilus are. Volborthellas classification is uncertain. The Middle Cambrian fossil Nectocaris is often interpreted as a cephalopod with 2 arms and no shell, but the Late Cambrian fossil Plectronoceras is now thought to be the earliest undisputed cephalopod fossil, as its shell had septa and a siphuncle, a strand of tissue that Nautilus uses to remove water from compartments it has vacated as it grows, and which is also visible in fossil ammonite shells. However, Plectronoceras and other early cephalopods crept along the seafloor instead of swimming, as their shells contained a "ballast" of stony deposits on what is thought to be the underside, and had stripes and blotches on what is thought to be the upper surface. All cephalopods with external shells except the became extinct by the end of the Cretaceous period . However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.
The Early Cambrian fossils Fordilla and Pojetaia are regarded as . "Modern-looking" bivalves appeared in the Ordovician period, . One bivalve group, the , became major reef-builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event. Even so, bivalves remain abundant and diverse.
The Hyolitha are a class of extinct animals with a shell and operculum that may be molluscs. Authors who suggest they deserve their own phylum do not comment on the position of this phylum in the tree of life.
Most molluscs with shells can produce pearls, but only the pearls of and some , whose shells are lined with nacre, are valuable. The best natural pearls are produced by marine , 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 , 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-Social 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 Before Christ, according to Theopompus.The fourth-century Before Christ historian Theopompus, cited by Athenaeus (12:526) around 200 BC; according to 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 textile produced from the long silky threads (byssus) secreted by several bivalve molluscs, particularly Pinna nobilis, to attach themselves to the sea bed. Webster's Third New International Dictionary (Unabridged) 1976. G. & C. Merriam Co., p. 307. Procopius, writing on the Persian wars circa 550 Common Era, "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."
Mollusc shells, including those of , 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. When used for commercial transactions, they functioned as a means of exchange similar to money in ordinary trade, a tradable good 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.
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, causes severe inflammation that can last for over a month even if treated correctly, 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.
All species of 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 , 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 makes it easy to synthesize them.
Evolutionary history
Good evidence exists for the appearance of (e.g., Aldanella), (e.g., Plectronoceras, Nectocaris?) and ( Pojetaia, Fordilla) towards the middle of the Cambrian period, c. , though arguably each of these may belong only to the stem lineage of their respective classes. However, the evolutionary history both of the emergence of molluscs from the ancestral group Lophotrochozoa, and of their diversification into the well-known living and fossil forms, is still vigorously debated.
However, the , 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 . 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.
fossil Yochelcionella is thought to be an early mollusc.]] .]]
Human interaction
For millennia, molluscs have been a source of food for humans, as well as important luxury goods, notably , 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. Some species of molluscs can bite or sting humans, and some have become agricultural pests.
Uses by humans
Molluscs, especially bivalves such as and , 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 and , , oysters, and scallops. (2025). 9780443056277, Elsevier Health Sciences. ISBN 9780443056277 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, mainly to minimize the poison risk from that can sometimes accumulate in the animals.
(2025). 9780521541107, Cambridge University Press. ISBN 9780521541107 Particularly chapters "Boom and slump for the cowrie trade" (pages 64–79) and "The cowrie as money: transport costs, values and inflation" (pages 125–147)
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.
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, zebra mussels ( Dreissena polymorpha) and the blue mussel Mytilus edulis.
Harm to humans
Stings and bites
Some molluscs sting or bite, but deaths from mollusc venoms total less than 10% of those from jellyfish stings.
Disease vectors
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." 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.
Pests
Some species of molluscs, particularly certain snails and slugs, can be serious crop pests, and when introduced into new environments, can unbalance local . 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.
See also
Explanatory notes
Further reading
(2025). 9781581129304, Universal. ISBN 9781581129304
(2025). 9788481587876, Servicio de Publicacións da Universidade de Vigo. ISBN 9788481587876
External links
|
|
