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Helix pomatia, a species of air-breathing land snail used for escargot, is a little bit larger than the common garden snail.
Helix pomatia, a species of air-breathing land snail used for escargot, is a little bit larger than the common garden snail.
Cornu aspersum (previously Helix aspersa) – the common garden snail – in Israel
Cornu aspersum (previously Helix aspersa) – the common garden snail – in Israel
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Groups included

see #Taxonomy

Colonies of Cernuella virgata in Sicily

A land snail is any of the numerous species of snail that live on land, as opposed to the sea snails and freshwater snails. Land snail is the common name for terrestrial gastropod mollusks that have shells (those without shells are known as slugs). However, it is not always easy to say which species are terrestrial, because some are more or less amphibious between land and fresh water, and others are relatively amphibious between land and salt water.

Land snails are a polyphyletic group comprising at least nine independent evolutionary transitions to terrestrial life (the last common ancestor of all gastropods was marine).[1][2][3][4][5] The majority of land snails are pulmonates that have a lung and breathe air. Most of the non-pulmonate land snails belong to lineages in the Caenogastropoda, and tend to have a gill and an operculum. The largest clade of non-pulmonate land snails is the Cyclophoroidea, with more than 7,000 species.[6] Many of these operculate land snails live in habitats or microhabitats that are sometimes (or often) damp or wet, such as in moss.

Land snails have a strong muscular foot; they use mucus to enable them to crawl over rough surfaces and to keep their soft bodies from drying out. Like other mollusks, land snails have a mantle, and they have one or two pairs of tentacles on their head. Their internal anatomy includes a radula and a primitive brain. In terms of reproduction, many caenogastropod land snails (e.g., diplommatinids) are dioecious,[7][8] but pulmonate land snails are hermaphrodites (they have a full set of organs of both sexes) and most lay clutches of eggs in the soil. Tiny snails hatch out of the egg with a small shell in place, and the shell grows spirally as the soft parts gradually increase in size. Most land snails have shells that are right-handed in their coiling.

A wide range of different vertebrate and invertebrate animals prey on land snails. They are used as food by humans in various cultures worldwide, and are raised on farms in some areas for use as food. Land snails can also be pests because they can chew away at and destroy plants. However, their diets are just as varied as the animals themselves, and while most are primarily herbivorous or omnivorous, there are also carnivorous species. Certain land snails have been introduced to non-native areas, where they have become invasive species.

Biology

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Taxonomy

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In older taxonomy, the Gastropoda were split into three groups: Pulmonata (defined by the presence of a pallial lung, which includes the vast majority of land snails today), Opisthobranchia (a group of mainly marine gastropods with gills behind or to the right of their heart), and Prosobranchia (the rest of the gastropods; defined by having their gills in front of their heart). These groups were found to be polyphyletic, and today, the Gastropoda is split into six subclassses: Caenogastropoda, Heterobranchia, Neomphaliones, Neritimorpha, Patellogastropoda, and Vetigastropoda.[9]

Land snails evolved in three gastropod subclasses: Neritimorpha, Heterobranchia, and Caenogastropoda. Most land snails are pulmonates in the order Stylommatophora, which contains over 20,000 species of slugs and snails. However, there are also several smaller groups of land snails.[10] As the last common ancestor of all gastropods was marine, at least nine independent lineages of land snails have made the transition to land from water, most of these evolving in the Palaeozoic or the Mesozoic. This makes the land snails polyphyletic.[3]

Below are the main lineages of land snails:[3][10]

Physical characteristics and anatomy

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"The Teeth of a Snail" from Robert Hooke's Micrographia, 1665. This actually shows the jaw, against which the teeth on the radula act.
Sphincterochila boissieri in Hamakhtesh Hagadol, northern Negev. Diameter is 2.1 cm.
Amphicyclotulus amethystinus, a species of non-pulmonate land snail in the superfamily Cyclophoroidea.
The anatomy of a common snail
Underside of a snail climbing a blade of grass, showing the muscular foot and the pneumostome or respiratory pore on the animal's right side

Land snails move by gliding along on their muscular foot, which is lubricated with mucus and covered with epithelial cilia.[11] This motion is powered by succeeding waves of muscular contractions that move down the ventral of the foot. This muscular action is clearly visible when a snail is crawling on the glass of a window or aquarium.[12] Snails move at a proverbially low speed (1 mm/s is a typical speed for adult Helix lucorum[13]). Snails secrete mucus externally to keep their soft bodies from drying out. They also secrete mucus from the foot to aid in locomotion by reducing friction, and to help reduce the risk of mechanical injury from sharp objects, meaning they can crawl over a sharp edge like a kitchen grater or razor without injury.[14] The mucus that land snails secrete with the foot leaves a slime trail behind them, which is often visible for some hours afterwards as a shiny "path" on the surface over which they have crawled. The mucus is produced by the suprapedal gland, located inside the foot.[15]

Snails (like all molluscs) also have a mantle, a specialized layer of tissue which covers all of the internal organs as they are grouped together in the visceral mass. The mantle also extends outward in flaps which reach to the edge of the shell and in some cases can cover the shell, and which are partially retractable. The mantle is attached to the shell, and creates the shell and makes shell growth possible by secretion.[16]

Most molluscs, including land snails, have a shell which is part of their anatomy since the larval stage. When they are active, the organs such as the lung, heart, kidney, and intestines remain inside the shell; only the head and foot emerge. The shell grows with them in size by the process of secreting calcium carbonate along the open edge and on the inner side for extra strength. Shells of land snails are made of two layers: a periostracum composed of conchiolin[17] and an inner one composed of calcium carbonate.[18] A lack of calcium, or low pH in their surroundings, can result in fragile, underdeveloped shells that are more easily damaged.[19] Usually, a snail can repair damage[20] to its shell over time if its living conditions improve, but severe damage can be fatal. When retracted into their shells, many snails with gills (including some terrestrial species) are able to protect themselves with a door-like anatomical structure called an operculum.[21] Notably, this feature is absent in the pulmonata.[22] Due to torsion, the anus and mantle cavity of gastropods are positioned above the head.[23]

Land snails range greatly in size. The largest living species is the giant African snail or Ghana Tiger Snail (Achatina achatina; Family Achatinidae), which can measure up to 30 cm.[24][25] The largest land snails of non-tropical Eurasia are endemic Caucasian snails Helix buchi and Helix goderdziana from the south-eastern Black Sea area in Georgia and Turkey; diameter of the shell of the latter may exceed 6 cm.[26] At the other end of the size spectrum is Angustopila psammion, a species with shell diameter of 0.60-0.68 mm.[27]

Most land snails bear one or two pairs of tentacles on their heads. In most land snails the eyes are carried on the first (upper) set of tentacles (called ommatophores or more informally 'eye stalks') which are usually roughly 75% of the width of the eyes. The second (lower) set of tentacles act as olfactory organs. Both sets of tentacles are retractable in stylommatophoran land snails.[28]

Digestive system

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Main article: Digestive system of gastropods
Garden snail (Cornu aspersum) defecating

The main parts of the gastropod digestive system are the buccal mass, the oesophagus, the stomach, digestive gland (or hepatopancreas), and intestine. The buccal mass consists of the mouth and pharynx. A snail breaks up its food using the radula inside its mouth. The radula is a chitinous ribbon-like structure containing rows of microscopic teeth, supported by an odontophore made of cartilage.[29][30] With this the snail scrapes at food, which is then transferred to the digestive tract. In a very quiet setting, a large land snail can be heard 'crunching' its food: the radula is tearing away at the surface of the food that the snail is eating. There is also a jaw that holds food steady while the radula tears it up.[15]

The oesophagus connects directly to the stomach, which is often relatively simple, though in pulmonates, it may be attached to an enlarged crop. The part of the stomach furthest from the oesophagus is known as a "style sac", and contains cilia that steadily draw food away from the mouth.[31] In most gastropods, the hepatopancreas combines the functions of a liver and pancreas, secreting digestive enzymes while also detoxifying food.[32] The stomach opens into a coiled intestine, which absorbs water from food and which opens directly into an anus above the head.[31]

Nervous and sensory system
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A diagram of a gastropod nervous system.
Light micrograph of a section through a snail's eye (Helix pomatia). 1 anterior chamber, 2 lens in the posterior chamber, 3 retina, 4 optic nerve
Main articles: Nervous system of gastropods and Sensory organs of gastropods

Snails have a simple nervous system, composed of up to 6 pairs of interconnected ganglia that together work to control parts of the body.[33] Three pairs of ganglia of the snail form a primitive brain. This structure is very much simpler than the brains of mammals, reptiles and birds, but nonetheless, snails are capable of associative learning.[34] Land snails also have chemoreceptors on their skin, including on their tentacles. Snails also have eyespots that can detect light, but are not advanced enough to form images.[35] In Stylommatophora, these eyespots are located on the tip of the tentacles;[28] in most others they are located at the base of the tentacles.[35] Snails also use olfaction; this they do using specialised organs on the tips of their tentacles. In Stylommatophora, the larger, upper pair of tentacles is responsible for wind perception (anemotaxis), while the lower, smaller pair of tentacles can perceive smells by contact.[36] Snails use this to track the odor of foods.[37]

Respiration and circulatory system

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Main articles: Respiratory system of gastropods and Circulatory system of gastropods

Since pulmonate snails are terrestrial rather than freshwater or marine, they have developed a simple lung for respiration, built from the mantle cavity.[38] A muscular valve, known as the pneumostome, is located on the right side of the animal and regulates the process of opening and closing the entrance of the lung.[38] When the pneumostome opens, the air can either enter or leave the lung. Non-pulmonates, such as the Cyclophoroidea, have also independently developed mantle-based lungs.[39] Like most molluscs, snails have an open circulatory system, with the blood circulating through a network of sinuses and flowing freely between organs; oxygen is carried by the pigment hemocyanin, and oxygen and carbon dioxide diffuse in and out of blood through the capillaries.[40][41] Most snails have a two-chambered heart.[42]

Shell growth

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As the snail grows, so does its calcium carbonate shell. The shell grows additively, by the addition of new calcium carbonate, which is secreted by glands located in the snail's mantle.[16] The new material is added to the edge of the shell aperture (the opening of the shell).[43] Therefore, the centre of the shell's spiral was made when the snail was younger, and the outer part when the snail was older. When the snail reaches sexual maturity, it may build a thickened lip around the shell aperture.[44]

A snail's shell forms a logarithmic spiral. Most snail shells are right-handed or dextral in coiling, meaning that if the shell is held with the apex (the tip, or the juvenile whorls) pointing towards the observer, the spiral proceeds in a clockwise direction from the apex to the opening.[45]

  • Snail with a rightwardly spiraling shell
    Snail with a rightwardly spiraling shell
  • Sinistral (left-handed) species of snail from western India
    Sinistral (left-handed) species of snail from western India

Hibernation and aestivation

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add sources
Some snails hibernate during the winter (typically October through April in the Northern Hemisphere). They may also aestivate in the summer in drought conditions. To stay moist during hibernation, a snail seals its shell opening with a dry layer of mucus called an epiphragm.

Reproduction

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The use of love darts by the land snail Monachoides vicinus is a form of sexual selection
Snails mating in Los Angeles
Two helicid snails make contact prior to mating.
Two Helix pomatia snails mating
See also: mating of gastropods

The great majority of land snails are hermaphrodites with a full set of reproductive organs of both sexes, able to produce both spermatozoa and ova. A few groups of land snails such as the Pomatiidae, which are distantly related to periwinkles, have separate sexes: male and female. The age of sexual maturity varies depending on species of snail, ranging from as little as 6 weeks[46] to 5 years.[47]

Most pulmonate air-breathing land snails perform courtship behaviors before mating. The courtship may last anywhere between two and twelve hours.[48] In around eight different families of pulmonate land snails and slugs, prior to mating one or more love darts are fired into the body of the partner. Love darts help to increase the chances of a snail's paternity, not only through mechanical stimulation but also by injecting mucus and hormones.[49][50]

Pulmonate land snails are prolific breeders and inseminate each other in pairs to internally fertilize their ova via a reproductive opening on one side of the body, near the front, through which the outer reproductive organs are extruded so that sperm can be exchanged. Fertilization then occurs and the eggs develop.[50] Each brood may consist of up to 400 eggs, in the case of the Giant African Land Snail.[51] Egg sizes differ between species, from a 3 mm diameter in the grove snail[52] to a 5.5 mm diameter in the giant African land snail.[53] After 2 to 4 weeks of favorable weather, these eggs hatch and the young emerge. Snails may lay eggs as often as once a month.[54]

There have been hybridizations of snail species; although these do not occur commonly in the wild, in captivity they can be coaxed into doing so. Parthenogenesis has been reported only in one species of slug,[55] but many species can self-fertilise.[56] Cylindrus obtusus is a prominent endemic snail species of the Eastern Alps. There is strong evidence for selfing (self-fertilization) in the easternmost snail populations as indicated by microsatellite data.[57] Compared to western populations, in the eastern population mucous gland structures employed in sexual reproduction are highly variable and deformed suggesting that in selfing organisms these structures have reduced function.[57]

Lifespan

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Most species of land snail are annual, others are known to live 2 or 3 years,[58][59] but some of the larger species may live over 10 years in the wild.[60] For instance, 10-year old individuals of the Roman snail Helix pomatia are probably not uncommon in natural populations.[61] Populations of some threatened species may be dependent on a pool of such long-lived adults.[62] In captivity, the lifespan of snails can be much longer than in the wild, for instance up to 25 years in H. pomatia.[63]

Diet

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add sources
In the wild, snails eat a variety of different foods. Terrestrial snails are usually herbivorous, however some species are predatory carnivores or omnivores, including the genus Powelliphanta, which includes the largest carnivorous snails in the world, native to New Zealand.[64] Prominent predatory snail families include the Spiraxidae, Haplotrematidae and Rhytididae. The diet of most land snails can include leaves, stems, soft bark, fruit, vegetables, fungi and algae. They may have a specialized crop of symbiotic bacteria that aid in digestion, especially with the breakdown of the polysaccharide cellulose into simple sugars. Some species can cause damage to agricultural crops and garden plants, and are therefore often regarded as pests.

Ecology and habitat

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Snails easily suffer moisture loss. Snails are most active at night[65] and after rainfall.[66] During unfavourable conditions, a snail remains inside its shell, usually under rocks or other hiding places, to avoid being discovered by predators. In dry climates, snails often bury themselves or hide in shaded areas, only coming out during the rainy season, e.g. Sphincterochila spp.[67]

Predators and parasites

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The larva of a glowworm (Lampyris noctiluca) attacking and eating a land snail

Many predators, both specialist and generalist, feed on snails. Some animals, such as the song thrush, break the shell of the snail by hammering it against a hard object, such as stone, to expose its edible insides.[68] Other predators, such as some species of frogs, circumvent the need to break snail shells by simply swallowing the snail whole, shell and all.[69]

In an attempt to protect themselves against predators, land snails retract their soft parts into their shell when they are resting; some bury themselves. Land snails have many natural predators, including members of all the land vertebrate groups, three examples being thrushes, hedgehogs and Pareas snakes. Invertebrate predators of land snails include ground beetles, leeches, certain land flatworms such as Platydemus manokwari,[70] carnivorous snails such as the rosy wolf snail and decollate snail, and even the predatory caterpillar Hyposmocoma molluscivora.[71]

In the case of the marsh snail Succinea putris, the snails can be parasitized by a microscopic flatworm of the species Leucochloridium paradoxum, which then reproduces within the snail's body. The flatworms invade the snail's eye stalks, causing them to become enlarged. Birds are attracted to and consume these eye stalks, consuming the flatworms in the process and becoming the final hosts of the flatworm.[72]

Human activity poses great dangers to land snails in the wild. Pollution and habitat destruction have caused the extinction of a considerable number of snail species in recent years.[73][74] Spread of invasive species of certain carnivorous snails has resulted in the extinction and near-extinction of other snails; see below.

As invasive species

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Several snail species are invasive for various reasons. The Giant African Land Snail (Lissachatina fulica) is a major pest in several countries (including but not limited to: Malaysia,[75] China,[76] Taiwan,[77] Trinidad and Tobago,[78] Cuba,[79] Brazil,[80] and the U.S. states of Florida[81] and Hawaii[82]) due to the pet trade. It poses a threat to its environment due to its ability to consume over 500 different species of plants, lay up to 400 eggs in one batch, and the fact that it carries rat lungworm.[81][51] The predatory Euglandina rosea was introduced to the Hawaiian islands to combat Lissachatina fulica, but ended up harming native Achatinellidae species instead, driving many of them to extinction[83] (e.g. Achatinella apexfulva).

Human relevance

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As food

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Further information: Snails as food
A snail farm in Provence

Land snails have been eaten for thousands of years, going back at least as far as the Pleistocene. Archaeological evidence of snail consumption is especially abundant in Capsian sites in North Africa, but is also found throughout the Mediterranean region in archaeological sites dating between 12,000 and 6,000 years ago.[84][85] Snail eggs, sold as snail caviar, are a specialty food that is growing in popularity in European cuisine.[86] Snails contain many nutrients. They are rich in calcium and also contain vitamin B1 and E. They contain various essential amino acids, and are low in calories and fat.[87] However, wild-caught land snails which are prepared for the table but are not thoroughly cooked, can harbor a parasite (Angiostrongylus cantonensis) that can cause a rare kind of meningitis.[88] The process of snail farming is called heliciculture. The establishment of snail farms outside of Europe has introduced several species to North America, South America, and Africa, where some escapees have established themselves as invasive species.[89][90][91][92]

Africa

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In parts of West Africa, specifically Ghana, Achatina achatina, Ghana tiger snails, are served as a delicacy.[93] Snail, called "igbin" in Yoruba language is a delicacy, widely eaten in Nigeria, especially among the Yorubas and Igbos.[94] In Igbo language, snails are called "Ejuna" or "Eju". In Cameroon, snails, usually called 'nyamangoro' and 'slow boys', are a delicacy especially to natives of the South West region of Cameroon.[95] The snails are either eaten cooked and spiced or with a favourite dish called 'eru'. In North Morocco, small snails are eaten as snacks in spicy soup (babbouche).[96] The recipe is similar to "caracoles a la andaluza", prepared in Andalusia (South Spain),[97] showing the close cultural relationship between both kinds of cuisine.

Europe

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Add citations

Oceania

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Bulime cooked in garlic butter in Ile des Pins, New Caledonia, species Placostylus fibratus

In New Caledonia, Placostylus fibratus (French: bulime) is considered a highly prized delicacy and is locally farmed to ensure supplies.[98] It is often served by restaurants prepared in the French style with garlic butter.

As pests

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Add to section lissachatina fulica (stucco/parasites), theba pisana, cantareus apertus, cornu aspersum, etc.

Prevention and control

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Main article: Molluscicide

Metaldehyde and iron phosphate can be used to exterminate snails.[99] Since copper generates electric shocks that make it difficult for snails to move, it makes a great barrier material for them.[100] note to self: expand!!!

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Add to section

See also

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References

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[edit]

Category:Mollusc common names Category:Gastropods

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