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Snail

Snail
Land snails
Although the word snail is often used for all shelled gastropods, the word "snail" can also be used in a more limited sense to mean any of several species of large, airbreathing (pulmonate) land snails. Whichever land snail species is most commonly seen or most commonly eaten in a given area will usually be referred to as "snails" by the local people. Species of land snails live in almost every kind of habitat, from deserts and mountains to marshes, woodland, and gardens. However, certain species are "anthropophilic", which means they are found most often around human habitation.

Helix pomatia The word snail is a common name for almost all members of the molluscan class Gastropoda that have coiled shells in the adult stage. When the word snail is used in a general sense, it includes sea snails, land snails and freshwater snails. Snails lacking a shell or having only a very small one are usually called slugs. Snails that have a broadly conical shell that is not coiled or appears not to be coiled are often known as limpets. The class of Gastropoda (the snails and slugs) is second only to insects in terms of total number of species. Snails are extraordinarily diverse in habitat, form, behavior, and anatomy. Therefore, what is true of one snail species may not be true of another. Snails can be found in a wide range of environments from ditches, deserts, and the abyssal depths of the sea. Although most people are familiar with terrestrial snails, land snails are in the minority. Marine snails have much greater diversity and a greater biomass. The great majority of snail species are marine. Numerous kinds can be found in fresh water and even brackish water. Many snails are herbivorous, though a few land species and many marine species are omnivores or predatory carnivores. Snails which respire using a lung belong to the group Pulmonata, while those with gills form a paraphyletic group, in other words, snails with gills are divided into a number of taxonomic groups that are not very closely related. Snails with lungs and with gills have diversified widely enough over geological time that a few species with gills can be found on land, numerous species with a lung can be found in freshwater, and a few species with a lung can be found in the sea.

Slugs
Gastropod species which lack a conspicuous shell are commonly called slugs rather than snails, although, other than having a reduced shell or no shell at all, there are really no appreciable differences between a slug and a snail except in habitat and behavior. A shell-less animal is much more maneuverable, and thus even quite large land slugs can take advantage of habitats or retreats with very little space – places that would be inaccessible to a similar-sized snail, such as under loose bark on trees or under stone slabs, logs or wooden boards lying on the ground. Taxonomic families of land slugs and sea slugs occur within numerous larger taxonomic groups of shelled species. In other words, the reduction or loss of the shell has evolved many times independently within several very different lineages of gastropods, thus the various families of slugs are very often not closely related to one another.

Biology (primarily but not exclusively of pulmonate land snails)
Physical characteristics
Most snails move by gliding along on their muscular foot, which is lubricated with mucus and covered with epithelial cilia[1]. 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

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Snail
snail shells are right-handed, 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. When the animal reaches full adult size, many species of snails build a thickened lip around the opening of the shell. At this point the animal stops growing, and begins reproducing.

Underside of a snail climbing a blade of grass, showing the muscular foot and the pneumostome on the animal’s right side aquarium. Snails move at a proverbially low speed (1 mm/s is a typical speed for adult Helix lucorum[2]). They produce mucus to aid locomotion by reducing friction, and the mucus also helps reduce the snail’s risk of mechanical injury from sharp objects. This means that they can ’walk’ over sharp objects like razors without being injured.[3] Snails 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, and 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 it by secretion. When retracted into their shells, many snails with gills (including many marine, some freshwater and some terrestrial species) are able to protect themselves with a door-like anatomical structure called an operculum. (The operculum of some sea snails has a pleasant scent when burned, so it is sometimes used as an ingredient in incense.) Snails range greatly in size. The largest land snail is the Giant African Snail or Ghana Tiger Snail (Achatina achatina; Family Achatinidae), which can measure up to 30 cm. Pomacea maculata (Family Ampullariidae), or Giant Apple Snail is the largest freshwater snail, with a diameter of up to 15 cm and a mass of over 600 g. The biggest of all snails is Syrinx aruanus, an Australian marine species which can grow up to 77.2 cm (30 inches) in length[4] and 18 kg (40lbs) in weight.[5] As the snail grows, so does its calcium carbonate shell. A snail’s shell forms a logarithmic spiral. Most

Sinistral (left-handed) species of snail from western India The shells of snails and other mollusks, and some snail egg casings, are primarily made up of calcium carbonate. Because of this, mollusks need calcium in their diet and environment to produce a strong shell. A lack of calcium, or low pH in their surroundings, can result in thin, cracked, or perforated shells. Usually a snail can repair damage to its shell over time if its living conditions improve, but severe damage can be fatal. Most 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 land snails. The eyes of most marine and freshwater snails are found at the base of the first set of tentacles.

Internal anatomy (land snails)

The anatomy of a common snail

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Snail

Garden Snail defecating A snail breaks up its food using the radula, which is a chitinous structure containing microscopic hooks called cuticulae. With this the snail scrapes at food, which is then transferred to the digestive tract. This is why, in a quiet setting, a large land snail can be heard ’crunching’ its food: the radula is tearing away at what it is eating. The cerebral ganglia of the snail form a primitive brain divided into four sections. This structure is very much simpler than the brains of mammals, reptiles and birds, but nonetheless, snails are capable of associative learning.[6]

Two Cornu aspersa Garden snails mating snails, and marine species, such as periwinkles, have separate sexes; they are male and female. The age of sexual maturity is variable depending on species of snail, ranging from as little as 6 weeks [7] to 5 years [8]. Adverse environmental conditions may delay the onset of sexual maturity in some snails [9]. Prior to reproduction, most land snails perform a ritual courtship before mating. This may last anywhere between two and twelve hours. Prolific breeders, pulmonate land snails inseminate each other in pairs to internally fertilize their ova. Each brood may consist of up to 100 eggs. Pulmonate land snails and slugs have a reproductive opening on one side of the body, near the front, through which the outer reproductive organs are extruded so that exchange of sperm can take place. After this, fertilization occurs and the eggs develop. Garden snails bury their eggs in shallow topsoil primarily while the weather is warm and damp, usually 5 to 10 cm down, digging with their foot. Egg sizes differ between species, from a 3 mm diameter in the grove snail to a 6 mm diameter in the Giant African Land Snail. 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. The snail’s shell develops while it is still an embryo; it is, however, very weak, and needs an immediate supply of calcium. Newly hatched snails obtain this by eating the egg from which they hatched. Baby snails

Hibernation/Estivation (land snails)
Some snails hibernate during the winter (typically October through April in the Northern Hemisphere). They may also estivate 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. Some freshwater snails such as apple snails have gills and a "door" or operculum to close the shell when they withdraw. This structure functions as protection from predators as well as protecting the soft tissues from desiccation when an aquatic habitat dries out temporarily.

Reproduction (primarily land snails)

Two Helicid snails make contact prior to mating. All land snails are hermaphrodites, producing both spermatozoa and ova. Some freshwater snails, such as apple

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cannibalizing other eggs, even unhatched ones, has been recorded. Promptly after they are finished ingesting their egg casings, they crawl upwards through the small tunnel in order to digest the egg. At this stage, the young are almost completely transparent and colorless. Their shell is usually slightly smaller than the egg they hatched from, but their length when out of their shell is slightly greater than the egg diameter. After a few weeks, the snails will begin to show their first tinge of color, usually slightly blue, before they turn their adult color. Roughly three months after they have hatched, they will look like miniature versions of their mature kin. They will continue to grow, usually for two to three years, until they reach adult size, although there have been confirmed recordings of snails growing amazingly fast - becoming even bigger than their parents in little more than a month. Irrespective of their rate of growth, however, it will still take at least 1 year before they are sexually mature.[10]

Snail
Parthenogenesis has also been noted in certain species.[11][12] Freshwater pond snails do not lay their eggs in the ground, but instead they attach them to something solid, or in some genera they carry the eggs internally until they hatch, a form of vivipary.

Lifespan
The lifespan of snails varies from species to species. In the wild, Achatinidae snails live around 5 to 7 years and Helix snails live about 2 to 3 years. Aquatic Apple Snails live only a year or so. Most deaths are due to predators or parasites. In captivity, their lifespan is much longer, ranging from 10 to 15 years for most species. On occasions, snails have lived beyond this lifespan, up to 30 years.[13]

Diet
In the wild, snails eat a variety of different foods. Terrestrial snails are herbivorous. Their diet can include leaves, stems, soft bark, fruit, vegetables and algae. Some species can cause damage to agricultural crops and garden plants, and are therefore often regarded as pests. Aquatic snails eat other varieties of food such as plankton, algae, plants, and other microscopic organisms that live underwater.

Predators
Land snails have many natural predators, including members of all major vertebrate groups, decollate snails, ground beetles, leeches, and even the predatory caterpillar Hyposmocoma molluscivora. The Botia family of freshwater fish also feed on freshwater snails by sucking them out of their shells. In the pulmonate marsh snail, Succinea putris, there is a parasitic flatworm, Leucochloridium paradoxum, which prevents the snail from retracting its enlarged and parasitized eye stalk, which thus makes the snail much more likely to be eaten by a bird, the final host of the worm. Humans also pose great dangers to snails in the wild. Pollution and habitat destruction have caused the extinction of a number of snail species in recent years.[14][15]

The egg mass of a freshwater pulmonate snail, attached to a solid surface.

Snails in cuisine
Snails have been eaten for thousands of years, beginning in 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.[16][17] It should be noted that wild-caught land snails that are undercooked can harbor a parasite (Angiostrongylus cantonensis) that may cause a rare kind of meningitis.[18]

Freshwater species mating There have been hybridizations of snails; although these do not occur commonly in the wild, in captivity they can be coaxed into doing so.

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Snail
flour and offered water for at least a week. This process is thought to cleanse the snails. Snails are also popular in Portuguese cuisine where they are called in Portuguese caracóis, and served in cheap snack houses and taverns, usually stewed (with different mixtures of white wine, garlic, piri piri, oregano, coriander or parsley, and sometimes chouriço). Bigger varieties, called caracoletas, are generally grilled and served with a butter sauce, but other dishes also exist such as feijoada de caracóis. Overall, Portugal consumes about 4,000 tonnes of snails each year.[20]

A snail farm in Provence Specialized snail caviar is growing in popularity in European cuisine.[19]

Cooked French escargots Traditional Spanish cuisine also uses snails ("caracoles"), consuming several species such as Helix aspersa, Helix punctata, Helix pisana or Helix alonensis among others. Small to medium-size varieties are usually cooked in several spicy sauces or even in soups, while the bigger ones may be reserved for other dishes such as the "arroz con conejo y caracoles" (a paella-style rice with snails and rabbit meat, very popular in the inner regions of south-eastern Spain). Snails are very popular in Catalonia, where they are called "caragols" or "cargols." In fact, a snail celebration, the "Aplec del cargol," takes place in Lleida each May, drawing more than 200,000 visitors from abroad. Popular Catalonian recipes for snails are a la llauna, grilled inside their own shells and then eaten after dipping them in garlic mayonnaise, or a la gormanda, boiled in tomato and onion sauce. In Greece, snails are especially popular in the island of Crete, but are also eaten in many parts of the country and can even be found in supermarkets, sometimes placed alive near partly refrigerated vegetables. In this regard, snails are one of the few live organisms sold at supermarkets as food. They are eaten either boiled with vinegar added, or sometimes cooked alive in a casserole with tomato, potatoes and squashes. Limpets and sea snails also find their way to the Greek table around the country. Another snail cooking method is the Kohli Bourbouristi (χοχλιοί μπου(ρ)μπουριστοί)[21] a traditional Cretan dish, which consists of fried snails in olive oil with salt, vinegar and rosemary.

Europe
Snails are eaten in several European countries, as they were in the past in the Roman Empire. Mainly three species, all from the genus Helix, are ordinarily eaten: • Helix pomatia, or edible snail, generally prepared in its shell, with parsley butter (size: 40 to 55 mm for an adult weight of 25 to 45 g.; typically found in Burgundy, France). • Helix aspersa: • Helix aspersa aspersa also known as the European brown snail, is cooked in many different ways, according to different local traditions (size: 28 to 35 mm for an adult weight of 7 to 15 g.; typically found in the Mediterranean countries of Europe and North Africa and the French Atlantic coast). • Helix aspersa maxima (size 40 to 45 mm for an average weight of 20 to 30 g.; typically found in North Africa). Snails are a delicacy in French cuisine, where they are called escargot. In an English-language menu, escargot is generally reserved for snails prepared with traditional French recipes (served in the shell with a garlic and parsley butter). Before preparing snails to eat, the snails should be fasting for three days with only water available. After three days of fasting, the snails should be fed

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In Sicily, snails (or babbaluci as they are commonly called in Sicilian) are a very popular dish as well. They are usually boiled with salt first, then served with tomato sauce or bare with oil, garlic and parsley. Snails are similarly appreciated in other Italian regions, such as Sardinia. Snails (or bebbux as they are called in Maltese) are a dish on the Mediterranean island of Malta, generally prepared and served in the Sicilian manner. In southwestern Germany there is a regional specialty of soup with snails and herbs, called "Black Forest Snail Chowder" (Badener Schneckensuepple). Heliciculture is the farming of snails. "They are protected in the wild almost everywhere (at least, the Roman Snail must not be collected any more), but the Roman Snail and the Garden Snail (Cornu aspersum) especially are cultivated on snail farms."[13] Although there is not usually considered to be a tradition of snail eating in Britain, common garden snails Helix aspersa were eaten in the Southwick area of Sunderland in the North East of England. They were collected from quarries and along the stone walls of railway embankments during the winter when the snails were hibernating and had voided the contents of their guts. It is thought that this tradition was introduced in the 19th Century by French immigrant glass workers - see Peter Gibson’s 1999 book - The People’s History: Southwick. "Snail suppers" were a feature of local pubs and Southwick working men were collecting and eating snails as late as the 1970s, though the tradition may now have died out.

Snail

A dish of cooked freshwater snails, ampullariids and viviparids from Poipet, Cambodia In parts of West Africa, specifically Ghana, snails are served as a delicacy. [1] Achatina achatina, Ghana tiger snails, are also known as some of the largest snails in the world.

Central America
Archaeological investigations in Guatemala have revealed that the diet of the Maya of the Classic Period (AD 250-900) included freshwater snails.[22]

Cultural depictions

Asia
Several different snail species are eaten in Asian cuisine.

Moche land snails (Scutalus sp.), 200 AD. Larco Museum Collection, Lima, Peru. A dish of cooked freshwater nerites from the Rajang River, Sarawak, Malaysia Due to its slowness, the snail has traditionally been seen as a symbol of laziness. In Judeo-Christian culture, it has often been viewed as a manifestation of the deadly sin of sloth.[23] Psalms 58:8 uses snail slime as a metaphorical punishment. Snails were widely noted and used in divination.[23] The Greek poet Hesiod wrote that snails signified the

Africa
Achatina fulica, the Giant East African Snail, is sliced and canned and sold to consumers as escargot.

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time to harvest by climbing the stalks, while the Aztec moon god Tecciztecatl bore a snail shell on his back. This symbolised rebirth; the snail’s penchant appearing and disappearing was analogised with the moon.[24] More recently, Carl Jung noted that the snail was representative of the self in dreams. In psychology, the soft insides are analogous to the subconscious, as the shell is the conscious.[23] In contemporary speech, the expression "a snail’s pace" is often used to describe a slow, inefficient process. The phrase "snail mail" is used to mean regular postal service delivery of paper messages as opposed to the delivery of E-mail or electronic mail, which is virtually instantaneous.

Snail

References
[1] [2]

[3]

Gallery
C. aspersa, a brown Garden snail from Europe

[4] [5] [6]

Cornu aspersa, the Garden snail, in the USA

Cornu aspersa, the Garden snail, Hampshire, UK

C. aspersa, Garden snail from England

[7] [8]

[9] [10] [11] [12] [13] [14] [15] [16] The introduced snail Theba pisana in Kadina, South Australia[25]

C. aspersa in a pool of water

Two grove White-lipped snails, CeTwo grove snail (Cepaea paea nemoralis snails, C. nemorhortensis) alis, mating

Giant East Brazilian African Freshwater Snail, Snail snail in the Brazil (Achatina genus Physa, fulica) the pouch snail

[17]

See also
• • • • Snail racing Pasilalinic-sympathetic compass Love dart Gastropod shell [18] [19] [20]

"integument (mollusks)."Encyclopædia Britannica. 2009. Encyclopædia Britannica 2006 Ultimate Reference Suite DVD G.A. Pavlova (01 May 2001). "Effects of serotonin, dopamine and ergometrine on locomotion in the pulmonate mollusc Helix lucorum" (pdf). Journal of Experimental Biology 204 (9): 1625–1633. PMID 11398751. http://jeb.biologists.org/cgi/reprint/204/9/1625. Retrieved on 2006-05-24. "Snails for kids and teachers : All about snails". Kiddyhouse.com : The resource center for kids and teachers. http://www.kiddyhouse.com/Snails/. Retrieved on 2008-08-08. Conchologists of America FAQ no. 6 Fun Animal Facts Christie Sahley, Alan Gelperin, and Jerry W. Rudy (January 1, 1982). "One-Trial Associative Learning Modifies Food Odor Preferences of a Terrestrial Mollusc". Proceedings of the National Academy of Sciences (National Academy of Sciences) 78 (1): 640–642. doi:10.1073/pnas.78.1.640. PMID 16592960. http://digital.library.adelaide.edu.au/dspace/ handle/2440/37983 http://www.columbia.edu/itc/cerc/danoff-burg/ invasion_bio/inv_spp_summ/ Euglandina_rosea.html http://www.jstage.jst.go.jp/article/aez/37/1/ 37_199/_article/-char/en Snails www.blackwell-synergy.com/doi/pdf/10.1111/ j.1420-9101.2004.00791.x http://links.jstor.org/ sici?sici=0014-3820(199208)46%3A4%3C907%3APIAFSR%3E2.0.CO% ^ The Roman snail or escargot (Helix pomatia) Centre for Biological Diversity: snail extinction update Hawaii’s Extinct Species – Snails Prehistoric edible land snails in the circumMediterranean: the archaeological evidence., D. Lubell. In J-J. Brugal & J. Desse (eds.), Petits Animaux et Sociétés Humaines. Du Complément Alimentaire Aux Ressources Utilitaires. XXIVe rencontres internationales d’archéologie et d’histoire d’Antibes, pp. 77-98. Antibes: Éditions APDCA.] Are land snails a signature for the MesolithicNeolithic transition? In, M. Budja (ed.), Neolithic Studies 11. Documenta Praehistorica XXXI: 1-24. D. Lubell. Division of Parasitic Diseases - Angiostrongylus Infection Fact Sheet Snail caviar! The new gourmet frontier According to the article Comemos três a quatro mil toneladas de caracóis por ano, by Joana Ferreira da

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Costa, in the 26.08.2007 edition (n.6358) of the daily newspaper Público. [21] Popping Fried Snails - Kohli Bourbouristi Recipe [22] (Spanish) Foias, Antonia E. (2000). "Entre la política y economía: Resultados preliminares de las primeras temporadas del Proyecto Arqueológico Motul de San José." (PDF online publication). XIII Simposio de Investigaciones Arqueológicas en Guatemala, 1999 (edited by J.P. Laporte, H. Escobedo, B. Arroyo and A.C. de Suasnávar) (Museo Nacional de Arqueología y Etnología, Guatemala): 771–799. http://www.asociaciontikal.com/ pdf/60.99_-_Foias.pdf. Retrieved on 2009-03-01. , page 777.

Snail
[23] ^ de Vries, Ad (1976). Dictionary of Symbols and Imagery. Amsterdam: North-Holland Publishing Company. pp. 430. ISBN 0-7204-8021-3. [24] Cooper, JC (1992). Symbolic and Mythological Animals. London: Aquarian Press. pp. 213. ISBN 1-85538-118-4. [25] [http://www.dpiw.tas.gov.au/inter.nsf/ Attachments/LBUN-74L3NM

External links
• predatory snails on the UF / IFAS Featured Creatures Web site. • Land Snail Ecology • Snailworlds-Timelaps

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