Mollusca

Mollusca is the second largest phylum of invertebrate animals. The members are known as molluscs or mollusks[note 1] (/ˈmɒləsk/). Around 85,000 extant species of molluscs are recognized.[2] The number of fossil species is estimated between 60,000 and 100,000 additional species.[3]

Molluscs are the largest marine phylum, comprising about 23% of all the named marine organisms. Numerous molluscs also live in freshwater and terrestrial habitats. They are highly diverse, not just in size and in anatomical structure, but also in behaviour and in habitat. The phylum is typically divided into 8 or 9 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 and account for 80% of the total classified species.

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

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

Molluscs have been and still are an important food source for anatomically modern humans. There is a risk of food poisoning from toxins which can accumulate in certain molluscs under specific conditions, however, and because of this, 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 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.

Mollusca
Temporal range: Cambrian Stage 2–Recent
Tonicella-lineata
Tonicella lineata, a polyplacophoran or chiton, anterior end towards the right
Scientific classification
Kingdom: Animalia
Superphylum: Lophotrochozoa
Phylum: Mollusca
Linnaeus, 1758
Classes

See text.

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

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 ta malákia (lit. "the soft ones"; < μαλακός malakós "soft"), which he applied inter alia to cuttlefish.[4][5] The scientific study of molluscs is accordingly called malacology.[6]

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.[7]

Definition

The 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. Many have a calcareous shell.[8]

Molluscs have developed such a varied range of body structures, it is difficult to find synapomorphies (defining characteristics) to apply to all modern groups.[9] The most general characteristic of molluscs is they are unsegmented and bilaterally symmetrical.[10] The following are present in all modern molluscs:[11][12]

Other characteristics that commonly appear in textbooks have significant exceptions:

  Whether characteristic is found in these classes of Molluscs
Supposed universal Molluscan characteristic[11] Aplacophora[13] Polyplacophora[14] Monoplacophora[15] Gastropoda[16] Cephalopoda[17] Bivalvia[18] Scaphopoda[19]
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
One or more valves/ shells Primitive forms, yes; modern forms, no Yes Yes Snails, yes; slugs, mostly yes (internal vestigial) Octopuses, no; cuttlefish, nautilus, squid, yes Yes Yes
Odontophore Yes Yes Yes Yes Yes No Yes

Diversity

Berlin Naturkundemuseum Muscheln
Diversity and variability of shells of molluscs on display.
Cypraea chinensis with partially extended mantle
About 80% of all known mollusc species are gastropods (snails and slugs), including this cowry (a sea snail).[20]

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 mollusc species at 107,000 of which were about 12,000 fresh-water gastropods and 35,000 terrestrial. The Bivalvia would comprise about 14% of the total and the other five classes less than 2% of the living molluscs.[21] In 2009, Chapman estimated the number of described living species at 85,000.[1] Haszprunar in 2001 estimated about 93,000 named species,[22] which include 23% of all named marine organisms.[23] Molluscs are second only to arthropods in numbers of living animal species[20]—far behind the arthropods' 1,113,000 but well ahead of chordates' 52,000.[24] About 200,000 living species in total are estimated,[1][25] and 70,000 fossil species,[11] 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.[26]

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.[9] About 80% of all known mollusc species are gastropods.[20] Cephalopoda such as squid, cuttlefish, and octopuses are among the neurologically most advanced of all invertebrates.[27] The giant squid, which until recently had not been observed alive in its adult form,[28] 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.[29]

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.[30]

Hypothetical ancestral mollusc

Archimollusc-en
Anatomical diagram of a hypothetical ancestral mollusc

Because of the great range of anatomical diversity among molluscs, many textbooks start the subject of molluscan anatomy by describing what is called an archi-mollusc, hypothetical generalized mollusc, or hypothetical ancestral mollusc (HAM) to illustrate the most common features found within the phylum. The depiction is visually rather similar to modern monoplacophorans.[9][12][15][31]

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".[12] The visceral mass, or visceropallium, is the soft, nonmuscular metabolic region of the mollusc. It contains the body organs.[10]

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.[12] The whole soft body of bivalves lies within an enlarged mantle cavity.[10]

Shell

The mantle edge secretes a shell (secondarily absent in a number of taxonomic groups, such as the nudibranchs[10]) that consists of mainly chitin and conchiolin (a protein hardened with calcium carbonate),[12][32] except the outermost layer, which in almost all cases is all conchiolin (see periostracum).[12] Molluscs never use phosphate to construct their hard parts,[33] with the questionable exception of Cobcrephora.[34] 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.[35]

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.[10]

Foot

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

The underside consists of a muscular foot, which has adapted to different purposes in different classes.[36]: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.[12] In bivalves, the foot is adapted for burrowing into the sediment;[36]:4 in cephalopods it is used for jet propulsion,[36]:4 and the tentacles and arms are derived from the foot.[37]

Circulatory system

Most 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.[10] The blood of these molluscs contains the respiratory pigment hemocyanin as an oxygen-carrier. The heart consists of one or more pairs of atria (auricles), which receive oxygenated blood from the gills and pump it to the ventricle, which pumps it into the aorta (main artery), which is fairly short and opens into the hemocoel.[12] The atria of the heart also function as part of the excretory system by filtering waste products out of the blood and dumping it into the coelom as urine. A pair of nephridia ("little kidneys") to the rear of and connected to the coelom extracts any re-usable materials from the urine and dumps additional waste products into it, and then ejects it via tubes that discharge into the mantle cavity.[12]

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

Respiration

Most molluscs have only one pair of gills, or even only a singular gill. Generally, the gills are rather like feathers in shape, although some species have gills with filaments on only one side. They divide the mantle cavity so water enters near the bottom and exits near the top. Their filaments have three kinds of cilia, one of which drives the water current through the mantle cavity, while the other two help to keep the gills clean. If the osphradia detect noxious chemicals or possibly sediment entering the mantle cavity, the gills' cilia may stop beating until the unwelcome intrusions have ceased. Each gill has an incoming blood vessel connected to the hemocoel and an outgoing one to the heart.[12]

Eating, digestion, and excretion

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

Members of the mollusc family use intracellular digestion to function. 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.[10] 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".[12]

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.[12]

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.[12]

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.[12]

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.[10]

Nervous system

Gastropod nervous system
Simplified diagram of the mollusc nervous system

The cephalic molluscs have two pairs of main nerve cords organized around a number of paired ganglia, the visceral cords serving the internal organs and the pedal ones serving the foot. Most pairs of corresponding ganglia on both sides of the body are linked by commissures (relatively large bundles of nerves). The ganglia above the gut are the cerebral, the pleural, and the visceral, which are located above the esophagus (gullet). The pedal ganglia, which control the foot, are below the esophagus and their commissure and connectives to the cerebral and pleural ganglia surround the esophagus in a circumesophageal nerve ring or nerve collar.[12]

The acephalic molluscs (i.e., bivalves) also have this ring but it is less obvious and less important. The bivalves have only three pairs of ganglia— cerebral, pedal, and visceral— with the visceral as the largest and most important of the three functioning as the principal center of "thinking". Some such as the scallops have eyes around the edges of their shells which connect to a pair of looped nerves and which provide the ability to distinguish between light and shadow.

Reproduction

Trochophore larva[38]
Apical tuft (cilia)
Prototroch (cilia)
Stomach
Mouth
Metatroch (cilia)
Mesoderm
Anus
/// = cilia
Trochophore larva 01
Trochophore larva[38]
Trochophore larva 01

The simplest molluscan reproductive system relies on external fertilization, but with more complex variations. All produce eggs, from which may emerge trochophore larvae, more complex veliger larvae, or miniature adults. Two gonads sit next to the coelom, a small cavity that surrounds the heart, into which they shed ova or sperm. The nephridia extract the gametes from the coelom and emit them into the mantle cavity. Molluscs that use such a system remain of one sex all their lives and rely on external fertilization. Some molluscs use internal fertilization and/or are hermaphrodites, functioning as both sexes; both of these methods require more complex reproductive systems.[12]

The most basic molluscan larva is a trochophore, which is planktonic and feeds on floating food particles by using the two bands of cilia around its "equator" to sweep food into the mouth, which uses more cilia to drive them into the stomach, which uses further cilia to expel undigested remains through the anus. New tissue grows in the bands of mesoderm in the interior, so the apical tuft and anus are pushed further apart as the animal grows. The trochophore stage is often succeeded by a veliger stage in which the prototroch, the "equatorial" band of cilia nearest the apical tuft, develops into the velum ("veil"), a pair of cilia-bearing lobes with which the larva swims. Eventually, the larva sinks to the seafloor and metamorphoses into the adult form. While metamorphosis is the usual state in molluscs, the cephalopods differ in exhibiting direct development: the hatchling is a 'miniaturized' form of the adult.[39]

Ecology

Feeding

Most molluscs are herbivorous, grazing on algae or filter feeders. For those grazing, two feeding strategies are predominant. Some feed on microscopic, filamentous algae, often using their radula as a 'rake' to comb up filaments from the sea floor. Others feed on macroscopic 'plants' such as kelp, rasping the plant surface with its radula. To employ this strategy, the plant has to be large enough for the mollusc to 'sit' on, so smaller macroscopic plants are not as often eaten as their larger counterparts.[40] Filter feeders are molluscs that feed by straining suspended matter and food particle from water, typically by passing the water over their gills. Most bivalves are filter feeders.

Cephalopods are primarily predatory, and the radula takes a secondary role to the jaws and tentacles in food acquisition. The monoplacophoran Neopilina uses its radula in the usual fashion, but its diet includes protists such as the xenophyophore Stannophyllum.[41] Sacoglossan sea-slugs suck the sap from algae, using their one-row radula to pierce the cell walls,[42] whereas dorid nudibranchs and some Vetigastropoda feed on sponges[43][44] and others feed on hydroids.[45] (An extensive list of molluscs with unusual feeding habits is available in the appendix of GRAHAM, A. (1955). "Molluscan diets". Journal of Molluscan Studies. 31 (3–4): 144..)

Classification

Opinions vary about the number of classes of molluscs; for example, the table below shows seven living classes,[22] 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.[13][31] Two of the commonly recognized "classes" are known only from fossils.[20]

Class Major organisms Described living species[22] Distribution
Gastropoda[46] All the snails and slugs including abalone, limpets, conch, nudibranchs, sea hares, sea butterfly 70,000 marine, freshwater, land
Bivalvia[47] clams, oysters, scallops, geoducks, mussels 20,000 marine, freshwater
Polyplacophora[14] chitons 1,000 rocky tidal zone and seabed
Cephalopoda[48] squid, octopus, cuttlefish, nautilus, spirula 900 marine
Scaphopoda[19] tusk shells 500 marine 6–7,000 metres (20–22,966 ft)
Aplacophora[13] worm-like organisms 320 seabed 200–3,000 metres (660–9,840 ft)
Monoplacophora[15] 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)
Rostroconchia[49] fossils; probable ancestors of bivalves extinct marine
Helcionelloida[50] 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.[51] Additionally, it suggests a sister taxon relationship exists between the Bivalvia and the Gastropoda. Tentaculita may also be in Mollusca (see Tentaculites).

Evolutionary history

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

Fossil record

Good evidence exists for the appearance of gastropods (e.g. Aldanella), cephalopods (e.g. Plectronoceras, ?Nectocaris) and bivalves (Pojetaia, Fordilla) towards the middle of the Cambrian period, c. 500 million years ago, though arguably each of these may belong only to the stem lineage of their respective classes.[52] However, the evolutionary history both of the emergence of molluscs from the ancestral group Lophotrochozoa, and of their diversification into the well-known living and fossil forms, is still vigorously debated.

Debate occurs about whether some Ediacaran and Early Cambrian fossils really are molluscs. Kimberella, from about 555 million years ago, has been described by some paleontologists as "mollusc-like",[53][54] but others are unwilling to go further than "probable bilaterian",[55][56] if that.[57]

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.[55][58] 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.[59] This appears to contradict the concept that the ancestral molluscan radula was mineralized.[60]

Yochelcionella water flow
The tiny Helcionellid fossil Yochelcionella is thought to be an early mollusc[50]
Neptunea despecta
Spirally coiled shells appear in many gastropods.[16]

However, the Helcionellids, which first appear over 540 million years ago in Early Cambrian rocks from Siberia and China,[61][62] are thought to be early molluscs with rather snail-like shells. Shelled molluscs therefore predate the earliest trilobites.[50] 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.[63]

Some analyses of helcionellids concluded these were the earliest gastropods.[64] 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.[16][65][66]

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

Volborthella, some fossils of which predate 530 million years ago, was long thought to be a cephalopod, but discoveries of more detailed fossils showed its shell was not secreted, but built from grains of the mineral silicon dioxide (silica), and it was not divided into a series of compartments by septa as those of fossil shelled cephalopods and the living Nautilus are. Volborthella's classification is uncertain.[67] 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 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.[68] All cephalopods with external shells except the nautiloids became extinct by the end of the Cretaceous period 65 million years ago.[69] However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.[70]

The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves.[71][72][73][74] "Modern-looking" bivalves appeared in the Ordovician period, 488 to 443 million years ago.[75] One bivalve group, the rudists, became major reef-builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event.[76] 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.[77]

Phylogeny

Lophotrochozoa

Brachiopods

Mollusca

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).[78][79] Does not include annelid worms as the analysis concentrated on fossilizable "hard" features.[78]

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,[54][55][58][59] debates arise about the relationships between the classes of living molluscs.[56] In fact, some groups traditionally classified as molluscs may have to be redefined as distinct but related.[80]

Molluscs are generally regarded members of the Lophotrochozoa,[78] 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.[81] 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.[82] 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.[83] Scientists disagree about this: Giribet and colleagues concluded, in 2006, the repetition of gills and of the foot's retractor muscles were later developments,[9] while in 2007, Sigwart concluded the ancestral mollusc was metameric, and it had a foot used for creeping and a "shell" that was mineralized.[56] 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.[82]

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.[50] Eventually, the cuticle would have become mineralized,[50] using the same genetic machinery (engrailed) as most other bilaterian skeletons.[83] The first mollusc shell almost certainly was reinforced with the mineral aragonite.[32]

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[78]

Molluscs

Solenogastres

Caudofoveata

Testaria

Polyplacophorans

Monoplacophorans

Bivalves

Scaphopods

Gastropods

Cephalopods

The "Testaria" hypothesis[78]

Morphological analyses tend to recover a conchiferan clade that receives less support from molecular analyses,[84] although these results also lead to unexpected paraphylies, for instance scattering the bivalves throughout all other mollusc groups.[85]

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.[80] A 2010 analysis recovered the traditional conchiferan and aculiferan groups, and showed molluscs were monophyletic, demonstrating that available data for solenogastres was contaminated.[86] 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.[87] Rather than eliminating unlikely relationships, the latest studies add new permutations of internal molluscan relationships, even bringing the conchiferan hypothesis into question.[88]

Human interaction

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

Uses by humans

Molluscs, especially bivalves such as clams and mussels, have been an important food source since at least the advent of anatomically modern humans, and this has often resulted in overfishing.[89] Other commonly eaten molluscs include octopuses and squids, whelks, oysters, and scallops.[90] 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.[91] Some countries regulate importation and handling of molluscs and other seafood, mainly to minimize the poison risk from toxins that can sometimes accumulate in the animals.[92]

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

Most molluscs with shells can produce pearls, but only the pearls of bivalves and some gastropods, whose shells are lined with nacre, are valuable.[16][18] The best natural pearls are produced by marine pearl oysters, Pinctada margaritifera and Pinctada mertensi, which live in the tropical and subtropical waters of the Pacific Ocean. Natural pearls form when a small foreign object gets stuck between the mantle and shell.

The two methods of culturing pearls insert either "seeds" or beads into oysters. The "seed" method uses grains of ground shell from freshwater mussels, and overharvesting for this purpose has endangered several freshwater mussel species in the southeastern United States.[18] The pearl industry is so important in some areas, significant sums of money are spent on monitoring the health of farmed molluscs.[93]

Meister von San Vitale in Ravenna
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.[94] 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.[95][96] 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.[97] Procopius, writing on the Persian wars circa 550 CE, "stated that the five hereditary satraps (governors) of Armenia who received their insignia from the Roman Emperor were given chlamys (or cloaks) made from lana pinna. Apparently, only the ruling classes were allowed to wear these chlamys."[98]

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.[99] 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.[100]

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

Harmful to humans

Stings and bites

Hapalochlaena lunulata
The blue-ringed octopus's rings are a warning signal; this octopus is alarmed, and its bite can kill.[102]

Some molluscs sting or bite, but deaths from mollusc venoms total less than 10% of those from jellyfish stings.[103]

All octopuses are venomous,[104] 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,[102] 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,[105] 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.[106]

Textile cone
Live cone snails can be dangerous to shell collectors, but are useful to neurology researchers.[107]

All species of cone snails are venomous and can sting painfully when handled, although many species are too small to pose much of a risk to humans, and only a few fatalities have been reliably reported. Their venom is a complex mixture of toxins, some fast-acting and others slower but deadlier.[107][103][108] 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.[107][109]

Disease vectors

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

Schistosomiasis (also known as bilharzia, bilharziosis or snail fever), a disease caused by the fluke worm Schistosoma, is "second only to malaria as the most devastating parasitic disease in tropical countries. An estimated 200 million people in 74 countries are infected with the disease – 100 million in Africa alone."[110] 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.[111]

Pests

Some species of molluscs, particularly certain snails and slugs, can be serious crop pests,[112] 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.[113]

Notes

  1. ^ The formerly dominant spelling mollusk is still used in the U.S. — see the reasons given in Gary Rosenberg's "Mollusckque - Mollusk vs Mollusc". For the spelling mollusc, see the reasons given in: Brusca & Brusca. Invertebrates (2nd ed.).

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Bibliography

  • Ruppert, E.E.; Fox, R.S.; Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks / Cole. ISBN 978-0-03-025982-1.

Further reading

  • Sturm, C.; Pearce, T.A. & Valdes, A. The Mollusks: A Guide to their Study, Collection, and Preservation. Universal Publishers. 2006. 454 pages. ISBN 1581129300
  • Trigo, J.E.; Díaz Agras, G.J.; García-Álvarez, O.L.; Guerra, A.; Moreira, J.; Pérez, J.; Rolán, E.; Troncoso, J.S. & Urgorri, V. (2018). Troncoso, J.S., Trigo, J.E. & Rolán, E., ed. Guía de los Moluscos Marinos de Galicia. Vigo: Servicio de Publicacións da Universidade de Vigo. 836 pages. ISBN 978-84-8158-787-6

External links

Arthur William Baden Powell

Arthur William Baden Powell (4 April 1901 – 1 July 1987) was a New Zealand malacologist, naturalist and palaeontologist, a major influence in the study and classification of New Zealand molluscs through much of the 20th century. He was known to his friends and family by his third name, "Baden".

The name Baden had been a given name in a Powell family since 1731, when Susannah Powell née Thistlethwayte (1696–1762) gave to her child (1731–1792) the maiden name of her mother, Susannah Baden (1663–1692). The name Baden, particularly when associated with the surname Powell, became famous in 1900–1901, the year Arthur William Baden Powell was born, because of the Siege of Mafeking, the most famous British action in the Second Boer War, which turned the British Commander of the besieged, Robert Baden-Powell, into a national hero. Throughout the British Empire, babies were named after him. No family connection has yet been established between Arthur William Baden Powell and Robert Baden-Powell.

Powell was born at Wellington, New Zealand, on 4 April 1901. His schooling was in Auckland, and he trained in printing at the Elam School of Fine Arts. This training, and his interest in conchology, set him on his life's work. He started writing scientific papers on mollusca in 1921, and became one of the few experts in New Zealand shellfish.

Powell married Isabel Essie Gittos on 19 December 1928, at Devonport in Auckland. They had a son.

He was appointed to the Auckland War Memorial Museum as palaeontologist and conchologist in 1929, working on some lesser-known mollusc families. He also studied New Zealand’s big land snails, the Paryphanta, and the Placostylus flax snails. From 1932 Powell participated in dredging expeditions on the British research ship Discovery II exploring coastal Northland and discovering large numbers of new species. Other field trips from the 1930s to 1960 took him to Stewart Island, the Chatham Islands, the Kermadec Islands and the Antarctica and Subantarctic region, resulting in many important papers.

Powell was a fellow of the Royal Society of New Zealand from 1940 and was the recipient of the Hector Memorial Medal and Prize in 1947. He also received an honorary DSc in 1956 from the University of New Zealand and was appointed a Commander of the Order of the British Empire for services to marine science in the 1981 New Year Honours.His wife died in 1976. Two years later, he married Ida Madoline Worthy (née Hayes) at Whangarei. Powell died on 1 July 1987 in Auckland.

Taxa named after him include:

Powelliphanta O'Connor, 1945

Antimargarita powelli Aldea, Zelaya & Troncoso, 2009

Falsilunatia powelli Dell, 1956

Philine powelli Rudman, 1970

Zeacolpus pagoda powelli Marwick, 1957The World Register of Marine Species mentions 837 marine taxa, named by Powell. Many have become synonyms.

Caenogastropoda

Caenogastropoda (from Ancient Greek caeno- meaning "recent") is a taxonomic clade, a large diverse group which are mostly sea snails and other marine gastropod mollusks, but also includes some freshwater snails and some land snails.

Caenogastropoda contains many families of shelled marine molluscs – including the periwinkles, cowries, wentletraps, moon snails, murexes, cone snails and turrids – and constitutes about 60% of all living gastropods.

Calliostomatidae

Calliostomatidae is a family of sea snails within the superfamily Trochoidea and the clade Vetigastropoda.

Cerithiopsidae

Cerithiopsidae are a family of very small and minute sea snails, marine gastropod molluscs or micromollusks in the informal group Ptenoglossa. Gastropods in this family are known as cerithiopsids.

These tiny snails have shells that are very high-spired and consist of multiple whorls.

Euthyneura

Euthyneura is a taxonomic clade of snails and slugs, which includes species from freshwater, marine, aquatic and terrestrial gastropod mollusks in the clade Heterobranchia.

Euthyneura are considered the crown group of Gastropoda, and are characterised by several autapomorphies, but are named for euthyneury. They are considered to be the most successful and diverse group of Gastropoda. Within this taxon, the Gastropoda have reached their peak in species richness and ecological diversity. This obvious evolutionary success can probably be attributed to several factors. Marine Opisthobranchia, e.g., have evolved several clades specialised on less used food resources such as sponges or cnidarians. A key innovation in the evolution of Pulmonata was the colonization of freshwater and terrestrial habitats.Various phylogenetic studies focused on Euthyneura: Dayrat et al. (2001), Dayrat & Tillier (2002) and Grande et al. (2004). Morphological analyses by Dayrat and Tillier (2002) demonstrated the need to explore new datasets in order to critically analyse the phylogeny of this controversial group of gastropods. Klussmann-Kolb et al. (2008) traced an evolutionary scenario regarding colonisation of different habitats based on phylogenetic hypothesis and they showed that traditional classification of Euthyneura needs to be reconsidered.

Evolution of molluscs

The evolution of the molluscs is the way in which the Mollusca, one of the largest groups of invertebrate animals, evolved. This phylum includes gastropods, bivalves, scaphopods, cephalopods, and several other groups.

The fossil record of mollusks is relatively complete, and they are well represented in most fossil-bearing marine strata. Very early organisms which have dubiously been compared to molluscs include Kimberella and Odontogriphus.

Gastropoda

The gastropods (), more commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca, called Gastropoda. This class comprises snails and slugs from saltwater, from freshwater, and from the land. There are many thousands of species of sea snails and slugs, as well as freshwater snails, freshwater limpets, and land snails and slugs.

The class Gastropoda contains a vast total of named species, second only to the insects in overall number. The fossil history of this class goes back to the Late Cambrian. As of 2017, 721 families of gastropods are known, of which 245 are extinct and appear only in the fossil record, while 476 are currently extant with or without a fossil record.Gastropoda (previously known as univalves and sometimes spelled "Gasteropoda") are a major part of the phylum Mollusca, and are the most highly diversified class in the phylum, with 65,000 to 80,000 living snail and slug species. The anatomy, behavior, feeding, and reproductive adaptations of gastropods vary significantly from one clade or group to another. Therefore, it is difficult to state many generalities for all gastropods.

The class Gastropoda has an extraordinary diversification of habitats. Representatives live in gardens, woodland, deserts, and on mountains; in small ditches, great rivers and lakes; in estuaries, mudflats, the rocky intertidal, the sandy subtidal, in the abyssal depths of the oceans including the hydrothermal vents, and numerous other ecological niches, including parasitic ones.

Although the name "snail" can be, and often is, applied to all the members of this class, commonly this word means only those species with an external shell big enough that the soft parts can withdraw completely into it. Those gastropods without a shell, and those with only a very reduced or internal shell, are usually known as slugs; those with a shell into which they can partly but not completely withdraw are termed semi-slugs.

The marine shelled species of gastropod include species such as abalone, conches, periwinkles, whelks, and numerous other sea snails that produce seashells that are coiled in the adult stage—though in some, the coiling may not be very visible, for example in cowries. In a number of families of species, such as all the various limpets, the shell is coiled only in the larval stage, and is a simple conical structure after that.

Henry Augustus Pilsbry

Henry Augustus Pilsbry (7 December 1862 – 26 October 1957) was an American biologist, malacologist and carcinologist, among other areas of study. He was a dominant presence in many fields of invertebrate taxonomy for the better part of a century. For much of his career, his authority with respect to the classification of certain substantial groups of organisms was unchallenged: barnacles, chitons, North American terrestrial mollusks, and others.

Henry Suter

Henry Suter (born Hans Heinrich Suter, 9 March 1841 – 31 July 1918) was a Swiss-born New Zealand zoologist, naturalist, palaeontologist, and malacologist.

Lophotrochozoa

Lophotrochozoa (, "crest/wheel animals") is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence.

Naticidae

Naticidae, common name moon snails or necklace shells, is a family of minute to large-sized predatory sea snails, marine gastropod molluscs in the clade Littorinimorpha. The shells of the species in this family are mostly globular in shape.

Naticidae is the only family in the superfamily Naticoidea.

It has been estimated that worldwide there are about 260–270 recent species of naticid snails. This group is assumed to have originated in the late Triassic or in the early Jurassic. Members of this family can be recognized by the shape of their shells, distinct appearance or by their predatory behavior.

Neogastropoda

Neogastropoda is a clade of sea snails, both freshwater and marine gastropod mollusks.

Nudibranch

Nudibranchs () are a group of soft-bodied, marine gastropod molluscs which shed their shells after their larval stage. They are noted for their often extraordinary colours and striking forms, and they have been given colourful nicknames to match, such as "clown", "marigold", "splendid", "dancer", and "dragon". Currently, about 3,000 valid species of nudibranchs are known.The word "nudibranch" comes from the Latin nudus "naked" and the Ancient Greek βράγχια (bránkhia) "gills".

Nudibranchs are often casually called sea slugs, as they are a family of opistobranchs (sea slugs), with the phylum Mollusca (molluscs), but many sea slugs belong to several taxonomic groups which are not closely related to nudibranchs. A number of these other sea slugs, such as the photosynthetic Sacoglossa and the colourful Aglajidae, are often confused with nudibranchs.

Seguenzioidea

Seguenzioidea is a superfamily of minute to medium-sized sea snails, marine gastropod mollusks in the clade Vetigastropoda.

Solariellidae

Solariellidae is a family of small sea snails, marine gastropod mollusks in the superfamily Trochoidea (according to the taxonomy of the Gastropoda by Bouchet & Rocroi, 2005).This family has no subfamilies.

This family consists of extremely diverse species and needs taxonomic revision. The relationship between genera are often uncertain. Many species remain undescribed or have been recently described.

Treatise on Invertebrate Paleontology

The Treatise on Invertebrate Paleontology (or TIP) published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and covering every phylum, class, order, family, and genus of fossil and extant (still living) invertebrate animals. The prehistoric invertebrates are described as to their taxonomy, morphology, paleoecology, stratigraphic and paleogeographic range. However, genera with no fossil record whatsoever have just a very brief listing.

Publication of the decades-long Treatise on Invertebrate Paleontology is a work-in-progress; and therefore it is not yet complete: For example, there is no volume yet published regarding the post-Paleozoic era caenogastropods (a molluscan group including the whelk and periwinkle). Furthermore, every so often, previously published volumes of the Treatise are revised.

Trochidae

The Trochidae, common name top-snails or top-shells, are a taxonomic family of very small to large sea snails, marine gastropod molluscs in the clade Vetigastropoda (according to the taxonomy of the Gastropoda by Bouchet & Rocroi from 2005).

This family is commonly known as the "top-snails" because in many species the shell resembles a toy spinning top.

Trochoidea (superfamily)

Trochoidea is a superfamily of small to very large vetigastropod sea snails with gills and an operculum. Species within this superfamily have nacre as the inner shell layer. The families within this superfamily include the Trochidae, the top snails. This superfamily is the largest vetigastropodan superfamily, containing more than 2,000 species.This taxon is not the same as a pulmonate land snail genus which is spelled the same way: Trochoidea (genus).

Winston Ponder

Winston F. Ponder (born 1941) is a noted malacologist from New Zealand who has named and described many marine and freshwater animals, especially micromolluscs. He graduated with an MSc, PhD and DSc from the University of Auckland, New Zealand.

Ponder was the principal research scientist in the malacology section of the Australian Museum, Sydney, Australia and helped to build up the museum's mollusc collection so that it became one of the most extensive of its kind in the world. Ponder retired from this post after a long career of more than forty years of research on molluscs, and is now an Honorary Fellow.

He has been the president of the Society of Australian Systematic Biologists, and was the managing editor of the journal Molluscan Research of the Malacological Society of Australasia. for 8 years.

Early in his career, in 1964, he worked on Antarctic collections together with Richard Dell and Alan Beu, resulting in a major monograph on the Antarctic bivalves, chitons and scaphopods.

Ponder is the author of more than 300 research publications. Many of these are on the subjects of the freshwater molluscs of Australia, and on invertebrate conservation. One major contribution was a taxonomy of the Gastropoda, which he published together with David R. Lindberg in 1997. This was the last major publication on the taxonomy of the Gastropoda that was based on the morphology of snails and slugs (their internal and external shapes and forms), and did not take into account any analysis of their DNA or RNA.

In 2008, again with David Lindberg, he edited the book "Phylogeny and Evolution of the Mollusca" in which 36 experts provided an up-to-date review on the evolutionary history of the Mollusca, based on reinvestigation of morphological characters, molecular data and the fossil record.

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