Mollusca

Mollusca
Mollusca
Temporal range: Cambrian–Recent
Tonicella lineata, a polyplacophoran or chiton, anterior end towards the right
Scientific classification e
Kingdom: Animalia
Superphylum: Lophotrochozoa
Phylum: Mollusca
Linnaeus, 1758
Classes
Diversity
85,000[1] recognized living species

The Mollusca (pronounced /məˈlʌskə/), common name molluscs or mollusks[note 1] (pronounced /ˈmɒləsks/), is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest marine phylum, comprising about 23% of all the named marine organisms. Numerous molluscs also live in freshwater and terrestrial habitats. Molluscs are highly diverse, not only in size and in anatomical structure, but also in behaviour and in habitat. The phylum is typically divided into nine or ten 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.

Molluscs have such a varied range of body structures that it is difficult to find defining characteristics that 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 that covers 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 – molluscs' circulatory systems are mainly open. The "generalized" mollusc's feeding system consists of a rasping "tongue" called a 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 that detect chemicals, vibrations and touch. The simplest type of molluscan reproductive system relies on external fertilization, but there are more complex variations. 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.

There is good evidence for the appearance of gastropods, cephalopods and bivalves in the Cambrian period 542 to 488.3 million years ago. However the evolutionary history both of molluscs' emergence from the ancestral Lophotrochozoa and of their diversification into the well-known living and fossil forms are still subjects of vigorous debate among scientists.

Molluscs have been and still are an important food source for anatomically modern humans. However there is a risk of food-poisoning from toxins that accumulate in molluscs under certain conditions, and many countries have regulations that aim to minimize 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 pre-industrial societies, although shell "currencies" have severe limitations compared with government-backed money.

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.

Contents

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 τᾲ μαλάκια, "the soft things", which he applied to cuttlefish.[2] The scientific study of molluscs is known as malacology.[3]

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.[4]

Molluscs have developed such a varied range of body structures that it is difficult to find synapomorphies (defining characteristics) that apply to all modern groups.[5] The most general characteristic of molluscs is that they are unsegmented and bilaterally symmetrical.[6] The following are present in all modern molluscs:[7][8]

  • The dorsal part of the body wall is a mantle (or pallium) which secretes calcareous spicules, plates or shells. It overlaps the body with enough spare room to form a mantle cavity.
  • The anus and genitals open into the mantle cavity.
  • There are two pairs of main nerve cords.[8]

Other characteristics that commonly appear in textbooks have significant exceptions:

  Class
Characteristic[7] Aplacophora[9] Polyplacophora[10] Monoplacophora[11] Gastropoda[12] Cephalopoda[13] Bivalvia[14] Scaphopoda[15]
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

About 80% of all known mollusc species are gastropods (snails and slugs), including the cowry (a sea snail) pictured here.[16]

Estimates of accepted described living species of molluscs vary from 50,000 to a maximum of 120,000 species.[1] In 2009 Chapman estimated the number of described living species at 85,000.[1] Haszprunar in 2001 estimated about 93,000 named species,[17] which include 23% of all named marine organisms.[18] Molluscs are second only to arthropods in numbers of living animal species[16]—far behind the arthropods' 1,113,000 but well ahead of chordates' 52,000.[19] It has been estimated that there are about 200,000 living species in total,[1][20] and 70,000 fossil species,[7] although the total number of mollusc species that ever existed, whether or not preserved, must be many times greater than the number alive today.[21]

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 sub-groups. 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.[5] About 80% of all known mollusc species are gastropods.[16] Cephalopoda such as squid, cuttlefish and octopus are among the neurologically most advanced of all invertebrates.[22] The giant squid, which until recently had not been observed alive in its adult form,[23] is one of the largest invertebrates. However a recently caught specimen of the colossal squid, 10 metres (33 ft) long and weighing 500 kilograms (1,100 lb), may have overtaken it.[24]

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. 42% of recorded extinctions since the year 1500 are of molluscs, almost entirely non-marine species.[25]

A "generalized mollusc"

Further information: Mollusc shell
1
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    Digestive & excretory system
    Circulatory & respiratory
    Central nervous system
    Reproductive system
 1 Radula
 2 Mouth
 3 Shell
 4 Stomach
 5 Gonad
 6 Heart
 7 Coelom
 9 Mantle
10 Mantle cavity
11 Anus
12 Gill
13 Foot
15 Pedal nerve cord
16 Gut
17 Visceral nerve cord
18 Nerve ring
A generalized mollusc[26]

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.[5][8][11][27]

The generalized mollusc has a single, "limpet-like" shell on top. The shell is secreted by a mantle that covers the upper surface. The underside consists of a single muscular "foot".[8] The visceral mass, or visceropallium, is the soft, non-muscular metabolic region of the mollusc. It contains the body organs.[6]

Mantle and mantle cavity

The mantle cavity is a fold in the mantle that encloses a significant amount of space. It is lined with epidermis. It 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.[8] The whole soft body of bivalves lies within an enlarged mantle cavity.[6]

Shell

Main article: Mollusc shell

The mantle edge secretes a shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs[6]) that consists of mainly chitin and conchiolin (a protein) hardened with calcium carbonate),[8][28] except that the outermost layer in almost all cases is all conchiolin (see periostracum).[8] 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, that is often nacreous.[6]

Foot

The underside consists of a muscular foot, which has adapted to different purposes in different classes.[32]: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 that have 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.[8] In bivalves, the foot is adapted for burrowing into the sediment;[32]:4 in cephalopods it is used for jet propulsion,[32]:4 and the tentacles and arms are derived from the foot.[33]

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.[6] 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.[8]

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.[8]

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 that 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.[8]

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. This radula is associated with the odontophore, a cartilaginous supporting organ[6]

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 that the mucus forms a long string.[8]

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 that 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.[8]

The particles are sorted by yet another group of cilia, which send the smaller particles, mainly minerals, to the prostyle so that 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.[8]

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 is in the part of the mantle cavity that is swept by the outgoing "lane" of the current created by the gills. Carnivorous molluscs usually have simpler digestive systems.[8]

As the head has largely disappeared in bivalves, their mouth has been equipped with labial palps (two on each side of the mouth) to collect the detritus from its mucus.[6]

Nervous system

Simplified diagram of the mollusc 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.[8]

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.[8]

Reproduction

Apical tuft (cilia)
Prototroch (cilia)
Stomach
Mouth
Metatroch (cilia)
Mesoderm
Anus
/// = cilia
Trochophore larva[34]

The simplest molluscan reproductive system relies on external fertilization, but there are 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 shed 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.[8]

The most basic molluscan larva is a trochophore, which is planktonic and feeds on floating food particles by using the two bands of cilia round 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 that 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.[35]

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 itself with its radula. To employ this strategy, the plant has to be large enough for the mollusc to 'sit' on; therefore smaller macroscopic plants enjoy less molluscan herbivory than their larger counterparts.[36] Naturally, there are exceptions; the 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.[37] Sacoglossan nudibranchs suck the sap from algae, using their one-row radula to pierce the cell walls,[38] whereas dorid nudibranchs and some Vetigastrpods feed on sponges[39][40] and others feed on hydroids.[41] (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. http://mollus.oxfordjournals.org/content/31/3-4/144.short. .)

Classification

Opinions vary about the number of classes of molluscs—for example the table below shows eight living classes,[17] 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.[9][27] Two of the commonly recognized "classes" are known only from fossils.[16]

Class Major organisms Described living species[17] Distribution
Caudofoveata[9] worm-like organisms 120 seabed 200–3,000 metres (660–9,800 ft)
Solenogastres[9] worm-like organisms 200 seabed 200–3,000 metres (660–9,800 ft)
Polyplacophora[10] chitons 1,000 rocky tidal zone and seabed
Monoplacophora[11] 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[42] All the snails and slugs including abalone, limpets, conch, nudibranchs, sea hares, sea butterfly 70,000 marine, freshwater, land
Cephalopoda[43] squid, octopus, cuttlefish, nautilus 900 marine
Bivalvia[44] clams, oysters, scallops, geoducks, mussels 20,000 marine, freshwater
Scaphopoda[15] tusk shells 500 marine 6–7,000 metres (20–23,000 ft)
Rostroconchia †[45] fossils; probable ancestors of bivalves extinct marine
Helcionelloida †[46] 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 that the Polyplacophora form a clade with a monophyletic Aplacophora.[47] Additionally it suggests that a sister taxon relationship exists between the Bivalvia and the Gastropoda.

Evolution

Fossil record

There is good evidence for the appearance of gastropods, cephalopods and bivalves in the Cambrian period 542 to 488.3 million years ago. 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.

There is debate about whether some Ediacaran and Early Cambrian fossils really are molluscs. Kimberella, from about 555 million years ago, has been described as "mollusc-like",[48][49] but others are unwilling to go further than "probable bilaterian".[50][51] 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.[50][52] 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.[53] This appears to contradict the concept that the ancestral molluscan radula was mineralized.[54]

The tiny Helcionellid fossil Yochelcionella is thought to be an early mollusc[46]
Spirally coiled shells appear in many gastropods[12]

However, the Helcionellids, which first appear over 540 million years ago in Early Cambrian rocks from Siberia and China,[55][56] are thought to be early molluscs with rather snail-like shells. Shelled molluscs therefore predate the earliest trilobites.[46] 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. There have been suggestions that the tiny specimens were juveniles and the larger ones adults.[57]

Some analyses of helcionellids concluded that these were the earliest gastropods.[58] However other scientists are not convinced that Early Cambrian fossils show clear signs of the torsion that identifies modern gastropods twists the internal organs so that the anus lies above the head.[12][59][60]

    = Septa
    = Siphuncle
Septa and siphuncle in nautiloid shell

For a long time it was thought that Volborthella, some fossils of which pre-date 530 million years ago, was a cephalopod. However discoveries of more detailed fossils showed that Volborthella’s shell was not secreted but built from grains of the mineral silicon dioxide (silica), and that 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.[61] The Late Cambrian fossil Plectronoceras is now thought to be the earliest clearly cephalopod fossil, as its shell had septa and a siphuncle, a strand of tissue that Nautilus uses to remove water from compartments that 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.[62] All cephalopods with external shells except the nautiloids became extinct by the end of the Cretaceous period 65 million years ago.[63] However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.[64]

The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves.[65][66][67][68] "Modern-looking" bivalves appeared in the Ordovician period, 488 to 443 million years ago.[69] One bivalve group, the rudists, became major reef-builders in the Cretaceous, but became extinct in the Cretaceous-Tertiary extinction.[70] Even so, bivalves remain abundant and diverse.

The Hyolitha is a class of extinct animals with a shell and operculum that may be molluscs. Authors who suggest that they deserve their own phylum do not comment on the position of this phylum in the tree of life[71]

Phylogeny

Lophotrochozoa

Brachiopods






Bivalves


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



Gastropods
(snails, slugs, limpets, sea hares)



Cephalopods
(nautiloids, ammonites, squid, etc.)


Scaphopods (tusk shells)







Aplacophorans
(spicule-covered, worm-like)


Polyplacophorans (chitons)




Halwaxiids

Wiwaxia


Halkieria



Orthrozanclus



Odontogriphus



A possible "family tree" of molluscs (2007).[72][73] Does not include annelid worms as the analysis concentrated on fossilizable "hard" features.[72]

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,[49][50][52][53] there are debates about the relationships between the classes of living molluscs.[51] In fact some groups traditionally classifed as molluscs may have to be redefined as distinct but related.[74]

Molluscs are generally regarded members of the Lophotrochozoa,[72] 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.[75] 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.[76] 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.[77] Scientists disagree about this: Giribet and colleagues concluded in 2006 that the repetition of gills and of the foot's retractor muscles were later developments, [5] while in 2007 Sigwart concluded that the ancestral mollusc was metameric, and that it had a foot used for creeping and a "shell" that was mineralized.[51] In one particular one branch of the family tree, the shell of conchiferans is thought to have evolved from the spicules (small spines) of aplacophorans; however this is difficult to reconcile with the embryological origins of spicules.[76]

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.[46] Eventually, the cuticle would have become mineralized,[46] using the same genetic machinery (engrailed) as most other bilaterian skeletons.[77] The first mollusc shell almost certainly was reinforced with the mineral aragonite.[78]

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

Molluscs
Aculifera


Solenogastres


Caudofoveata



Polyplacophorans


Conchifera

Monoplacophorans



Bivalves


Scaphopods


Gastropods


Cephalopods




The "Aculifera" hypothesis[72]
Molluscs


Solenogastres


Caudofoveata

Testaria

Polyplacophorans



Monoplacophorans



Bivalves


Scaphopods


Gastropods


Cephalopods






The "Testaria" hypothesis[72]

Morphological analyses tend to recover a conchiferan clade that receives less support from molecular analyses,[79] although these results also lead to unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups.[80]

However, an analysis in 2009 that used both morphological and molecular phylogenetics comparisons concluded that the molluscs are not monophyletic; in particular, that Scaphopoda and Bivalvia are both separate, monophyletic lineages unrelated to the remaining molluscan classes—in other words that the traditional phylum Mollusca is polyphyletic, and that it can only be made monophyletic if scaphopods and bivalves are excluded.[74] A 2010 analysis managed to recover the traditional conchiferan and aculiferan groups, but similarly concluded that the molluscs are not monophyletic, this time suggesting that solenogastres are more closely related to the non-molluscan taxa used as an outgroup than to other molluscs.[81] Current molecular data is insufficient to constrain the molluscan phylogeny, and since the methods used to determine the confidence in clades are prone to over-estimation, it is risky to place too much emphasis even on the areas that different studies agree.[82] Rather than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan hypothesis into question.[83][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 pre-industrial 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 over-fishing.[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]

Photo of three circular metal cages in shallows, with docks, boathouses and palm trees in background
Saltwater pearl oyster farm in Seram, Indonesia

Most molluscs that have shells can produce pearls, but only the pearls of bivalves and some gastropods whose shells are lined with nacre are valuable.[12][14] The best natural pearls are produced by marine pearl oysters. Pinctada margaritifera and Pinctada mertensi, which live in the tropical and sub-tropical waters of the Pacific Ocean. Natural pearls form when a small foreign object gets stuck between the mantle and shell.

There are two methods of culturing pearls, by inserting either "seeds" or beads into oysters. The "seed" method uses grains of ground shell from freshwater mussels, and over-harvesting for this purpose has endangered several freshwater mussel species in the southeastern USA.[14] The pearl industry is so important in some areas that significant sums of money are spent on monitoring the health of farmed molluscs.[89]

Mosaic of mustachioed, curly-haired man wearing crown and surrounded by halo
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.[90] The discovery of large numbers of Murex shells on Crete suggests that the Minoans may have pioneered the extraction of "Imperial purple" during the Middle Minoan period in the 20th–18th century 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 pre-industrial 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, in other words 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

There is a risk of food poisoning from toxins that accumulate in molluscs 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.

The blue-ringed octopus's rings are a warning signal—this octopus is alarmed, and its bite can kill.[97]

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 however, 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]

Photo of cone on ocean bottom
Live cone snails can be dangerous to shell-collectors but are useful to neurology researchers[102]

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 are carnivorous gastropods that 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 the few larger species of cone snail that 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 that they are 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 drowning them, is a myth.[105]

Pests

Skin vesicles created by the penetration of Schistosoma. Source: Centers for Disease Control and Prevention

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, and 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]

Notes

Footnotes

  1. ^ Spelled mollusks in the USA, see reasons given in Rosenberg's [1]; for the spelling mollusc see the reasons given by Brusca & Brusca. Invertebrates (2nd ed.). 

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Further reading

  • Starr & Taggart (2002). Biology: The Unity and Diversity of Life. Pacific Grove, California: Thomson Learning. ISBN 0534027423. 
  • Nunn, J.D., Smith, S.M., Picton, B.E. and McGrath, D. (2002). "Checklist, atlas of distribution and bibliography for the marine mollusca of Ireland". Marine Biodiversity in Ireland and Adjacent Waters. 8. Ulster Museum. 
  • Ostroumov SA (2001). "An amphiphilic substance inhibits the mollusk capacity to filter phytoplankton cells from water.". Izvestiia Akademii nauk. Seriia biologicheskaia / Rossiiskaia akademiia nauk (1): 108–16. PMID 11236572. ; http://www.springerlink.com/content/l665628020163255/;
  • Dame, R.; Olenin, S., eds (2005). The comparative roles of suspension-feeders in ecosystems. Dordrecht: Springer. 

External links

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Mesozoa
Eumetazoa
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  • Mollusca — Mol*lus ca, prop. n. pl. [NL. See {Mollusk}.] (Zo[ o]l.) One of the grand divisions of the animal kingdom, a phylum including the classes Cephalopoda, Gastropoda, Pteropoda, Scaphopoda, and Pelecyopoda (syn. Bivalvia, formerly called… …   The Collaborative International Dictionary of English

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