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Molluscs


Temporal range: Cambrian–Recent
Tonicella lineata, a polyplacophoran or chiton, anterior end towards the right

Diversity
85,000[1] recognized living species


The molluscs or mollusks[note 1] /ˈmɒləsks/, compose the large phylum of invertebrate animals known as the phylum Mollusca. Around 85,000 extant species of molluscs are recognized. 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 only in size and in anatomical structure, but also in behaviour and in habitat. The phylum is typically divided into 9 or 10 taxonomic classes, of which two are entirely extinct. Cephalopod molluscs, such as squid, cuttlefish and octopus, are among the most neurologically advanced of all invertebrates—and either the giant squid or the colossal squid is the largest known invertebrate species. The gastropods (snails and slugs) are by far the most numerous molluscs in terms of classified species, and account for 80% of the total. The scientific study of molluscs is called malacology.[2]

Molluscs have such a varied range of body structures, it is difficult to find defining characteristics to apply to all modern groups. The two most universal features are a mantle with a significant cavity used for breathing and excretion, and the structure of the nervous system. As a result of this wide diversity, many textbooks base their descriptions on a hypothetical "generalized mollusc". 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 is very small, and the main body cavity is a hemocoel through which blood circulates; 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. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults.

A striking feature of molluscs is the use of the same organ for multiple functions. For example, the heart and nephridia ("kidneys") are important parts of the reproductive system, as well as the circulatory and excretory systems; in bivalves, the gills both "breathe" and produce a water current in the mantle cavity, which is important for excretion and reproduction.

Good evidence exists for the appearance of fossil forms are still subjects of vigorous debate among scientists.

Molluscs have been and still are an important food source for anatomically modern humans, but with a risk of food poisoning from toxins that accumulate in molluscs under certain conditions, and many countries 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 a money in some preindustrial societies.

Mollusc species can also represent 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 for over a month. Stings from a few species of large tropical cone shells can also kill, but their sophisticated, though easily produced, venoms have become important tools in neurological research. Schistosomiasis (also known as bilharzia, bilharziosis or snail fever) is transmitted to humans via water snail hosts, and affects about 200 million people. Snails and slugs can also be serious agricultural pests, and accidental or deliberate introduction of some snail species into new environments has seriously damaged some ecosystems.

Etymology

The words mollusc and mollusk are both derived from the French mollusque, which originated from the Latin molluscus, from mollis, soft. Molluscus was itself an adaptation of Aristotle's τὰ μαλάκια, "the soft things", which he applied to cuttlefish.[3]

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 it is now known these groups have no relation to molluscs, and very little to one another, the name Molluscoida has been abandoned.[4]

Definition

The two most universal features of the body structure of molluscs are a mantle with a significant cavity used for breathing and excretion, and the organization of the nervous system. The most abundant metallic element in molluscs is calcium.[5]

Molluscs have developed such a varied range of body structures, it is difficult to find synapomorphies (defining characteristics) to apply to all modern groups.[6] The most general characteristic of molluscs is they are unsegmented and bilaterally symmetrical.[7] The following are present in all modern molluscs:[8][9]

Other characteristics that commonly appear in textbooks have significant exceptions:

  Class
Characteristic[8] Aplacophora[10] Polyplacophora[11] Monoplacophora[12] Gastropoda[13] Cephalopoda[14] Bivalvia[15] Scaphopoda[16]
Radula, a rasping "tongue" with chitinous teeth Absent in 20% of Neomeniomorpha Yes Yes Yes Yes No Internal, cannot extend beyond body
Broad, muscular foot Reduced or absent Yes Yes Yes Modified into arms Yes Small, only at "front" end
Dorsal concentration of internal organs (visceral mass) Not obvious Yes Yes Yes Yes Yes Yes
Large digestive ceca No ceca in some Aplacophora Yes Yes Yes Yes Yes No
Large complex metanephridia ("kidneys") None Yes Yes Yes Yes Yes Small, simple

Diversity

Estimates of accepted described living species of molluscs vary from 50,000 to a maximum of 120,000 species.[1] In 1969 David Nicol estimated the probable total number of living molluscs 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 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.[21] About 200,000 living species in total are estimated,[1][22] and 70,000 fossil species,[8] 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.[23]

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.[6] About 80% of all known mollusc species are gastropods.[17] Cephalopoda such as squid, cuttlefish, and octopuses are among the neurologically most advanced of all invertebrates.[24] The giant squid, which until recently had not been observed alive in its adult form,[25] 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.[26]

Freshwater and terrestrial molluscs appear exceptionally vulnerable to extinction. Estimates of the numbers of nonmarine molluscs vary widely, partly because many regions have not been thoroughly surveyed. There is also a shortage of specialists who can identify all the animals in any one area to species. However, in 2004 the IUCN Red List of Threatened Species included nearly 2,000 endangered nonmarine molluscs. For comparison, the great majority of mollusc species are marine, but only 41 of these appeared on the 2004 Red List. About 42% of recorded extinctions since the year 1500 are of molluscs, consisting almost entirely of nonmarine species.[27]

A "generalized mollusc"

Further information: Mollusc shell

Because of the great range of anatomical diversity among molluscs, many textbooks start the subject by describing a hypothetical "generalized mollusc" to illustrate the most common features found within the phylum. The depiction is rather similar to modern monoplacophorans, and some suggest it may resemble very early molluscs.[6][9][12][28]

The generalized mollusc is bilaterally symmetrical 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".[9] The visceral mass, or visceropallium, is the soft, nonmuscular metabolic region of the mollusc. It contains the body organs.[7]

Mantle and mantle cavity

The mantle cavity, a fold in the mantle, encloses a significant amount of space. It is lined with epidermis, and is exposed, according to habitat, to sea, fresh water or air. The cavity was at the rear in the earliest molluscs, but its position now varies from group to group. The anus, a pair of osphradia (chemical sensors) in the incoming "lane", the hindmost pair of gills and the exit openings of the nephridia ("kidneys") and gonads (reproductive organs) are in the mantle cavity.[9] The whole soft body of bivalves lies within an enlarged mantle cavity.[7]

Shell

Main article: Mollusc shell

The mantle edge secretes a shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs[7]) that consists of mainly chitin and conchiolin (a protein hardened with calcium carbonate),[9][29] except the outermost layer in almost all cases is all conchiolin (see periostracum).[9] Molluscs never use phosphate to construct their hard parts,[30] with the questionable exception of Cobcrephora.[31] 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.[32]

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.[7]

Foot

File:SeaSnails.ogg The underside consists of a muscular foot, which has adapted to different purposes in different classes.[33]:4 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.[9] In bivalves, the foot is adapted for burrowing into the sediment;[33]:4 in cephalopods it is used for jet propulsion,[33]:4 and the tentacles and arms are derived from the foot.[34]

Circulation

Molluscs' circulatory systems are mainly open. 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.[7] The blood 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.[9]

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 coleom 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.[9]

Respiration

Most molluscs have only one pair of gills, or even only one 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.[9]

Eating, digestion, and excretion

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

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.[7] 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".[9]

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.[9]

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.[9]

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.[9]

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.[7]

Nervous system

Molluscs have two pairs of main nerve cords (three in bivalves) the visceral cords serving the internal organs and the pedal ones serving the foot. Both pairs run below the level of the gut, and include ganglia as local control centers in important parts of the body. Most pairs of corresponding ganglia on both sides of the body are linked by commissures (relatively large bundles of nerves). The only ganglia above the gut are the cerebral ganglia, which sit above the esophagus (gullet) and handle "messages" from and to the eyes. The pedal ganglia, which control the foot, are just below the esophagus and their commissure and connections to the cerebral ganglia encircle the esophagus in a nerve ring.[9]

The brain, in species that have one, encircles the esophagus. Most molluscs have a head with eyes, and all have a pair of sensor-containing tentacles, also on the head, that detect chemicals, vibrations and touch.[9]

Reproduction

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

The simplest molluscan reproductive system relies on external fertilization, but with more complex variations. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. Two gonads sit next to the coelom, a small cavity that surrounds the heart and sheds ova or sperm into the coloem, from which the nephridia extract them 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.[9]

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. Whilst metamorphosis is the usual state in molluscs, the cephalopods differ in exhibiting direct development: the hatchling is a 'miniaturized' form of the adult.[36]

Ecology

Feeding

Most molluscs are herbivorous, grazing on algae. 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.[37]

Cephalopods are primarily (perhaps entirely) 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.[38] Sacoglossan sea-slugs suck the sap from algae, using their one-row radula to pierce the cell walls,[39] whereas dorid nudibranchs and some Vetigastropoda feed on sponges[40][41] and others feed on hydroids.[42] (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 eight 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.[10][28] Two of the commonly recognized "classes" are known only from fossils.[17]

Class Major organisms Described living species[19] Distribution
Caudofoveata[10] worm-like organisms 120 seabed 200–3,000 metres (660–9,840 ft)
Solenogastres[10] worm-like organisms 200 seabed 200–3,000 metres (660–9,840 ft)
Polyplacophora[11] chitons 1,000 rocky tidal zone and seabed
Monoplacophora[12] An ancient lineage of molluscs with cap-like shells 31 seabed 1,800–7,000 metres (5,900–23,000 ft); one species 200 metres (660 ft)
Gastropoda[43] All the snails and slugs including abalone, limpets, conch, nudibranchs, sea hares, sea butterfly 70,000 marine, freshwater, land
Cephalopoda[44] squid, octopus, cuttlefish, nautilus 900 marine
Bivalvia[45] clams, oysters, scallops, geoducks, mussels 20,000 marine, freshwater
Scaphopoda[16] tusk shells 500 marine 6–7,000 metres (20–22,966 ft)
Rostroconchia[46] fossils; probable ancestors of bivalves extinct marine
Helcionelloida[47] fossils; snail-like organisms such as Latouchella extinct marine

Classification into higher taxa for these groups has been and remains problematic. A phylogenetic study suggests the Polyplacophora form a clade with a monophyletic Aplacophora.[48] Additionally, it suggests a sister taxon relationship exists between the Bivalvia and the Gastropoda.

Evolution

Fossil record

Good evidence exists for the appearance of fossil forms, is still vigorously debated.

Debate occurs about whether some

However, the

Some analyses of helcionellids concluded these were the earliest gastropods.[59] However, other scientists are not convinced these Early Cambrian fossils show clear signs of the torsion that identifies modern gastropods twists the internal organs so the anus lies above the head.[13][60][61]

    = Septa
    = Siphuncle
Septa and siphuncle in nautiloid shell

The Early Cambrian fossils Even so, bivalves remain abundant and diverse.

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

Phylogeny

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

The phylogeny (evolutionary "family tree") of molluscs is a controversial subject. In addition to the debates about whether Kimberella and any of the "halwaxiids" were molluscs or closely related to molluscs,[50][51][53][54] debates arise about the relationships between the classes of living molluscs.[52] In fact, some groups traditionally classified as molluscs may have to be redefined as distinct but related.[75]

Molluscs are generally regarded members of the Lophotrochozoa,[73] a group defined by having trochophore larvae and, in the case of living Lophophorata, a feeding structure called a lophophore. The other members of the Lophotrochozoa are the annelid worms and seven marine phyla.[76] The diagram on the right summarizes a phylogeny presented in 2007.

Because the relationships between the members of the family tree are uncertain, it is difficult to identify the features inherited from the last common ancestor of all molluscs.[77] For example, it is uncertain whether the ancestral mollusc was metameric (composed of repeating units)—if it was, that would suggest an origin from an annelid-like worm.[78] Scientists disagree about this: Giribet and colleagues concluded, in 2006, the repetition of gills and of the foot's retractor muscles were later developments,[6] while in 2007, Sigwart concluded the ancestral mollusc was metameric, and it had a foot used for creeping and a "shell" that was mineralized.[52] In one particular branch of the family tree, the shell of conchiferans is thought to have evolved from the spicules (small spines) of aplacophorans; but this is difficult to reconcile with the embryological origins of spicules.[77]

The molluscan shell appears to have originated from a mucus coating, which eventually stiffened into a cuticle. This would have been impermeable and thus forced the development of more sophisticated respiratory apparatus in the form of gills.[47] Eventually, the cuticle would have become mineralized,[47] using the same genetic machinery (engrailed) as most other bilaterian skeletons.[78] The first mollusc shell almost certainly was reinforced with the mineral aragonite.[79]

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,[80] although these results also lead to unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups.[81]

However, an analysis in 2009 using both morphological and molecular phylogenetics comparisons concluded the molluscs are not monophyletic; in particular, Scaphopoda and Bivalvia are both separate, monophyletic lineages unrelated to the remaining molluscan classes; the traditional phylum Mollusca is polyphyletic, and it can only be made monophyletic if scaphopods and bivalves are excluded.[75] A 2010 analysis managed to recover the traditional conchiferan and aculiferan groups, but similarly concluded the molluscs are not monophyletic, this time suggesting the solenogastres are more closely related to the nonmolluscan taxa used as an outgroup than to other molluscs.[82] 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.[83] Rather than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan hypothesis into question.[84]

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. Their outlandish forms have helped conjure up tales of mythological sea monsters, such as the kraken. A number of species of molluscs can bite or sting humans, and some have become agricultural pests.

Uses by humans

Further information: Seashell

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.[85] Other commonly eaten molluscs include octopuses and squids, whelks, oysters, and scallops.[86] 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.[87] Some countries regulate importation and handling of molluscs and other seafood, mainly to minimize the poison risk from toxins that accumulate in the animals.[88]


Most molluscs with shells can produce pearls, but only the pearls of bivalves and some gastropods, whose shells are lined with nacre, are valuable.[13][15] 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 USA.[15] The pearl industry is so important in some areas, significant sums of money are spent on monitoring the health of farmed molluscs.[89]


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.[90] 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.[91][92] 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.[93] 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 (Pinna "wool," or byssus). Apparently, only the ruling classes were allowed to wear these chlamys."[94]

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.[95] 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.[96]

Stings and bites


A risk of food poisoning from toxins that accumulate in molluscs occurs under certain conditions, and many countries have regulations that aim to minimize this risk. Blue-ringed octopus bites are often fatal, and the bite of other octopuses can cause unpleasant symptoms. Stings from a few species of large tropical cone shells can also kill. However, the sophisticated venoms of these cone snails have become important tools in neurological research and show promise as sources of new medications.

When handled alive, a few species of molluscs can sting or bite and, with some species, this can present a serious risk to the human handling the animal. To put this into perspective, though, deaths from mollusc venoms are less than 10% of the number of deaths from jellyfish stings.[98]

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


All species of cone snails are venomous and can sting when handled, although many species are too small to pose much of a risk to humans. These carnivorous gastropods feed on marine invertebrates (and in the case of larger species, on fish). Their venom is based on a huge array of toxins, some fast-acting and others slower but deadlier; they can afford to do this because their toxins require less time and energy to be produced compared with those of snakes or spiders.[102] Many painful stings have been reported, and a few fatalities, although some of the reported fatalities may be exaggerations.[98] Only a few larger species of cone snails which can capture and kill fish are likely to be seriously dangerous to humans.[103] The effects of individual cone-shell toxins on victims' nervous systems are so precise as to be useful tools for research in neurology, and the small size of their molecules makes it easy to synthesize them.[102][104]

The traditional belief that a giant clam can trap the leg of a person between its valves, thus causing drowning, is a myth.[105]

Pests


Schistosomiasis (also known as bilharzia, bilharziosis or snail fever) is transmitted to humans via water snail hosts, and affects about 200 million people. A few species of snails and slugs are serious agricultural pests; in addition, accidental or deliberate introduction of various snail species into new territory has resulted in serious damage to some natural ecosystems.

Schistosomiasis 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."[106] The parasite has 13 known species, of which two infect humans. The parasite itself is not a mollusc, but all the species have freshwater snails as intermediate hosts.[107]

Some species of molluscs, particularly certain snails and slugs, can be serious crop pests,[108] 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.[109]

Despite its name, Molluscum contagiosum is a viral disease, and is unrelated to molluscs.[110]

Footnotes

Citations

Further reading

  • ; http://www.springerlink.com/content/l665628020163255/;

External links

  • Researchers complete mollusk evolutionary tree; 26 October 2011
  • Hardy's Internet Guide to Marine Gastropods
  • Rotterdam Natural History Museum Shell Image Gallery

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