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Gastropoda

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Gastropoda
Temporal range: Cambrian–Present[1]
Gastropod collage.jpg
Various gastropods from different types: Black slug (a slug), Haliotis asinina (an abalone), Cornu aspersum (a land snail), Notarchus indicus (a seahare), Patella vulgata (a limpet), and Polycera aurantiomarginata (a nudibranch).
Scientific classification e
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Cuvier, 1795[2]
Subclasses
Diversity[3][4]
65,000 to 80,000 species
Synonyms[5]
  • Angiogastropoda - represented as Gastropoda
  • Apogastropoda - alternate representation of Gastropoda
  • Psilogastropoda - represented as Gastropoda

The gastropods (/ˈɡæstrəpɒdz/), commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (/ɡæsˈtrɒpədə/).[5]

This class comprises snails and slugs from saltwater, freshwater, and from 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.[6]

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[3][4] living snail and slug species. The anatomy, behavior, feeding, and reproductive adaptations of gastropods vary significantly from one clade or group to another, so stating many generalities for all gastropods is difficult.

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, 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 semislugs.

The marine shelled species of gastropods 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.

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Freshwater snail

Freshwater snail

Freshwater snails are gastropod mollusks that live in fresh water. There are many different families. They are found throughout the world in various habitats, ranging from ephemeral pools to the largest lakes, and from small seeps and springs to major rivers. The great majority of freshwater gastropods have a shell, with very few exceptions. Some groups of snails that live in freshwater respire using gills, whereas other groups need to reach the surface to breathe air. In addition, some are amphibious and have both gills and a lung. Most feed on algae, but many are detritivores and some are filter feeders.

Furongian

Furongian

The Furongian is the fourth and final epoch and series of the Cambrian. It lasted from 497 to 485.4 million years ago. It succeeds the Miaolingian series of the Cambrian and precedes the Lower Ordovician Tremadocian Stage. It is subdivided into three stages: the Paibian, Jiangshanian and the unnamed 10th stage of the Cambrian.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Class (biology)

Class (biology)

In biological classification, class is a taxonomic rank, as well as a taxonomic unit, a taxon, in that rank. It is a group of related taxonomic orders. Other well-known ranks in descending order of size are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order.

Anatomy

Anatomy

Anatomy is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having its beginnings in prehistoric times. Anatomy is inherently tied to developmental biology, embryology, comparative anatomy, evolutionary biology, and phylogeny, as these are the processes by which anatomy is generated, both over immediate and long-term timescales. Anatomy and physiology, which study the structure and function of organisms and their parts respectively, make a natural pair of related disciplines, and are often studied together. Human anatomy is one of the essential basic sciences that are applied in medicine.

Clade

Clade

In biological phylogenetics, a clade, also known as a monophyletic group or natural group, is a grouping of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. In the taxonomical literature, sometimes the Latin form cladus is used rather than the English form.

Abyssal zone

Abyssal zone

The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. "Abyss" derives from the Greek word ἄβυσσος, meaning bottomless. At depths of 4,000 to 6,000 metres, this zone remains in perpetual darkness. It covers 83% of the total area of the ocean and 60% of Earth's surface. The abyssal zone has temperatures around 2 to 3 °C through the large majority of its mass. Due to there being no light, there are no plants producing oxygen, which instead primarily comes from ice that had melted long ago from the polar regions. The water along the seafloor of this zone is actually devoid of oxygen, resulting in a death trap for organisms unable to quickly return to the oxygen-enriched water above. This region also contains a much higher concentration of nutrient salts, like nitrogen, phosphorus, and silica, due to the large amount of dead organic material that drifts down from the above ocean zones and decomposes. The water pressure can reach up to 76 megapascal.

Gastropod shell

Gastropod shell

The gastropod shell is part of the body of a gastropod or snail, a kind of mollusc. The shell is an exoskeleton, which protects from predators, mechanical damage, and dehydration, but also serves for muscle attachment and calcium storage. Some gastropods appear shell-less (slugs) but may have a remnant within the mantle, or in some cases the shell is reduced such that the body cannot be retracted within it (semi-slug). Some snails also possess an operculum that seals the opening of the shell, known as the aperture, which provides further protection. The study of mollusc shells is known as conchology. The biological study of gastropods, and other molluscs in general, is malacology. Shell morphology terms vary by species group.

Abalone

Abalone

Abalone is a common name for any of a group of small to very large marine gastropod molluscs in the family Haliotidae. Other common names are ear shells, sea ears, and, rarely, muttonfish or muttonshells in parts of Australia, ormer in the UK, perlemoen in South Africa, and paua in New Zealand. Abalones are marine snails. Their taxonomy puts them in the family Haliotidae, which contains only one genus, Haliotis, which once contained six subgenera. These subgenera have become alternative representations of Haliotis. The number of species recognized worldwide ranges between 30 and 130 with over 230 species-level taxa described. The most comprehensive treatment of the family considers 56 species valid, with 18 additional subspecies. The shells of abalones have a low, open spiral structure, and are characterized by several open respiratory pores in a row near the shell's outer edge. The thick inner layer of the shell is composed of nacre (mother-of-pearl), which in many species is highly iridescent, giving rise to a range of strong, changeable colors which make the shells attractive to humans as decorative objects, jewelry, and as a source of colorful mother-of-pearl.

Conch

Conch

Conch is a common name of a number of different medium-to-large-sized sea snails. Conch shells typically have a high spire and a noticeable siphonal canal.

Common periwinkle

Common periwinkle

The common periwinkle or winkle is a species of small edible whelk or sea snail, a marine gastropod mollusc that has gills and an operculum, and is classified within the family Littorinidae, the periwinkles.

Family (biology)

Family (biology)

Family is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between order and genus. A family may be divided into subfamilies, which are intermediate ranks between the ranks of family and genus. The official family names are Latin in origin; however, popular names are often used: for example, walnut trees and hickory trees belong to the family Juglandaceae, but that family is commonly referred to as the "walnut family".

Etymology

In the scientific literature, gastropods were described as "gasteropodes" by Georges Cuvier in 1795.[2] The word gastropod comes from Greek γαστήρ (gastḗr 'stomach') and πούς (poús 'foot'), a reference to the fact that the animal's "foot" is positioned below its guts.[7]

The earlier name "univalve" means one valve (or shell), in contrast to bivalves, such as clams, which have two valves or shells.

Diversity

At all taxonomic levels, gastropods are second only to the insects in terms of their diversity.[8]

Gastropods have the greatest numbers of named mollusc species. However, estimates of the total number of gastropod species vary widely, depending on cited sources. The number of gastropod species can be ascertained from estimates of the number of described species of Mollusca with accepted names: about 85,000 (minimum 50,000, maximum 120,000).[9] But an estimate of the total number of Mollusca, including undescribed species, is about 240,000 species.[10] The estimate of 85,000 molluscs includes 24,000 described species of terrestrial gastropods.[9]

Different estimates for aquatic gastropods (based on different sources) give about 30,000 species of marine gastropods, and about 5,000 species of freshwater and brackish gastropods. Many deep-sea species remain to be discovered, as only 0.0001% of the deep-sea floor has been studied biologically.[11][12] The total number of living species of freshwater snails is about 4,000.[13]

Recently extinct species of gastropods (extinct since 1500) number 444, 18 species are now extinct in the wild (but still exist in captivity), and 69 species are "possibly extinct".[14]

The number of prehistoric (fossil) species of gastropods is at least 15,000 species.[15]

In marine habitats, the continental slope and the continental rise are home to the highest diversity, while the continental shelf and abyssal depths have a low diversity of marine gastropods.[16]

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Taxonomy (biology)

Taxonomy (biology)

In biology, taxonomy is the scientific study of naming, defining (circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa and these groups are given a taxonomic rank; groups of a given rank can be aggregated to form a more inclusive group of higher rank, thus creating a taxonomic hierarchy. The principal ranks in modern use are domain, kingdom, phylum, class, order, family, genus, and species. The Swedish botanist Carl Linnaeus is regarded as the founder of the current system of taxonomy, as he developed a ranked system known as Linnaean taxonomy for categorizing organisms and binomial nomenclature for naming organisms.

Biodiversity

Biodiversity

Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic, species, and ecosystem level.

Mollusca

Mollusca

Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks. Around 85,000 extant species of molluscs are recognized. The number of fossil species is estimated between 60,000 and 100,000 additional species. The proportion of undescribed species is very high. Many taxa remain poorly studied.

Brackish water

Brackish water

Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater and fresh water together, as in estuaries, or it may occur in brackish fossil aquifers. The word comes from the Middle Dutch root brak. Certain human activities can produce brackish water, in particular civil engineering projects such as dikes and the flooding of coastal marshland to produce brackish water pools for freshwater prawn farming. Brackish water is also the primary waste product of the salinity gradient power process. Because brackish water is hostile to the growth of most terrestrial plant species, without appropriate management it is damaging to the environment.

Extinct in the wild

Extinct in the wild

A species that is extinct in the wild (EW) is one that has been categorized by the International Union for Conservation of Nature as known only by living members kept in captivity or as a naturalized population outside its historic range due to massive habitat loss.

Continental rise

Continental rise

The continental rise is a low-relief zone of accumulated sediments that lies between the continental slope and the abyssal plain. It is a major part of the continental margin, covering around 10% of the ocean floor.

Habitat

Cepaea nemoralis: a European pulmonate land snail, which has been introduced to many other countries
Cepaea nemoralis: a European pulmonate land snail, which has been introduced to many other countries

Some of the more familiar and better-known gastropods are terrestrial gastropods (the land snails and slugs). Some live in fresh water, but most named species of gastropods live in a marine environment.

Gastropods have a worldwide distribution, from the near Arctic and Antarctic zones to the tropics. They have become adapted to almost every kind of existence on earth, having colonized nearly every available medium.

In habitats where not enough calcium carbonate is available to build a really solid shell, such as on some acidic soils on land, various species of slugs occur, and also some snails with thin, translucent shells, mostly or entirely composed of the protein conchiolin.

Snails such as Sphincterochila boissieri and Xerocrassa seetzeni have adapted to desert conditions. Other snails have adapted to an existence in ditches, near deepwater hydrothermal vents, the pounding surf of rocky shores, caves, and many other diverse areas.

Gastropods can be accidentally transferred from one habitat to another by other animals, e.g. by birds.[17]

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Sea snail

Sea snail

Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the absence of a visible shell.

Sea slug

Sea slug

Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are gastropods, i.e. they are sea snails that over evolutionary time have either completely lost their shells, or have seemingly lost their shells due to having a greatly reduced or internal shell. The name "sea slug" is most often applied to nudibranchs, as well as to a paraphyletic set of other marine gastropods without obvious shells.

Land snail

Land snail

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. 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.

Semi-slug

Semi-slug

Semi-slugs, also spelled semislugs, are land gastropods whose shells are too small for them to retract into, but not quite vestigial. The shell of some semi-slugs may not be easily visible on casual inspection, because the shell may be covered over with the mantle.

Slug

Slug

Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word slug is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a small internal shell, particularly sea slugs and semislugs.

Arctic

Arctic

The Arctic is a polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada, Danish Realm (Greenland), northern Finland (Lapland), Iceland, northern Norway, Russia, northernmost Sweden and the United States (Alaska). Land within the Arctic region has seasonally varying snow and ice cover, with predominantly treeless permafrost containing tundra. Arctic seas contain seasonal sea ice in many places.

Antarctic

Antarctic

The Antarctic is a polar region around Earth's South Pole, opposite the Arctic region around the North Pole. The Antarctic comprises the continent of Antarctica, the Kerguelen Plateau and other island territories located on the Antarctic Plate or south of the Antarctic Convergence. The Antarctic region includes the ice shelves, waters, and all the island territories in the Southern Ocean situated south of the Antarctic Convergence, a zone approximately 32 to 48 km wide varying in latitude seasonally. The region covers some 20 percent of the Southern Hemisphere, of which 5.5 percent is the surface area of the Antarctica continent itself. All of the land and ice shelves south of 60°S latitude are administered under the Antarctic Treaty System. Biogeographically, the Antarctic realm is one of eight biogeographic realms of Earth's land surface.

Calcium carbonate

Calcium carbonate

Calcium carbonate is a chemical compound with the chemical formula CaCO3. It is a common substance found in rocks as the minerals calcite and aragonite and is the main component of eggshells, gastropod shells, shellfish skeletons and pearls. Things containing much calcium carbonate or resembling it are described as calcareous. Calcium carbonate is the active ingredient in agricultural lime and is created when calcium ions in hard water react with carbonate ions to create limescale. It has medical use as a calcium supplement or as an antacid, but excessive consumption can be hazardous and cause hypercalcemia and digestive issues.

Conchiolin

Conchiolin

Conchiolins are complex proteins which are secreted by a mollusc's outer epithelium.

Rocky shore

Rocky shore

A rocky shore is an intertidal area of seacoasts where solid rock predominates. Rocky shores are biologically rich environments, and are a useful "natural laboratory" for studying intertidal ecology and other biological processes. Due to their high accessibility, they have been well studied for a long time and their species are well known.

Cave

Cave

A cave or cavern is a natural void in the ground, specifically a space large enough for a human to enter. Caves often form by the weathering of rock and often extend deep underground. The word cave can refer to smaller openings such as sea caves, rock shelters, and grottos, that extend a relatively short distance into the rock and they are called exogene caves. Caves which extend further underground than the opening is wide are called endogene caves.

Bird

Bird

Birds are a group of warm-blooded vertebrates constituting the class Aves, characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight skeleton. Birds live worldwide and range in size from the 5.5 cm (2.2 in) bee hummingbird to the 2.8 m common ostrich. There are about ten thousand living species, more than half of which are passerine, or "perching" birds. Birds have wings whose development varies according to species; the only known groups without wings are the extinct moa and elephant birds. Wings, which are modified forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in some birds, including ratites, penguins, and diverse endemic island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly seabirds and some waterbirds, have further evolved for swimming.

Anatomy

The anatomy of a common air-breathing land snail: much of this anatomy does not apply to gastropods in other clades or groups.
The anatomy of a common air-breathing land snail: much of this anatomy does not apply to gastropods in other clades or groups.

Snails are distinguished by an anatomical process known as torsion, where the visceral mass of the animal rotates 180° to one side during development, such that the anus is situated more or less above the head. This process is unrelated to the coiling of the shell, which is a separate phenomenon. Torsion is present in all gastropods, but the opisthobranch gastropods are secondarily untorted to various degrees.[18][19]

Torsion occurs in two stages. The first, mechanistic stage, is muscular, and the second is mutagenetic. The effects of torsion are primarily physiological; the organism develops an asymmetrical growth, with the majority occurring on the left side. This leads to the loss of right-paired appendages (e.g., ctenidia (comb-like respiratory apparatus), gonads, nephridia, etc.). Furthermore, the anus becomes redirected to the same space as the head. This is speculated to have some evolutionary function, as prior to torsion, when retracting into the shell, first the posterior end would get pulled in, and then the anterior. Now, the front can be retracted more easily, perhaps suggesting a defensive purpose.

However, this "rotation hypothesis" is being challenged by the "asymmetry hypothesis" in which the gastropod mantle cavity originated from one side only of a bilateral set of mantle cavities.[20]

Gastropods typically have a well-defined head with two or four sensory tentacles with eyes, and a ventral foot, which gives them their name (Greek gaster, stomach, and pous, foot). The foremost division of the foot is called the propodium. Its function is to push away sediment as the snail crawls. The larval shell of a gastropod is called a protoconch.

The principal characteristic of the Gastropoda is the asymmetry of their principal organs. The essential feature of this asymmetry is that the anus generally lies to one side of the median plane.; The ctenidium (gill-combs), the osphradium (olfactory organs), the hypobranchial gland (or pallial mucous gland), and the auricle of the heart are single or at least are more developed on one side of the body than the other ; Furthermore, there is only one genital orifice, which lies on the same side of the body as the anus.[21]

Shell

The shell of Zonitoides nitidus, a small land snail, has dextral coiling, which is typical (but not universal) of gastropod shells.
The shell of Zonitoides nitidus, a small land snail, has dextral coiling, which is typical (but not universal) of gastropod shells.

Most shelled gastropods have a one piece shell (with exceptional bivalved gastropods), typically coiled or spiraled, at least in the larval stage. This coiled shell usually opens on the right-hand side (as viewed with the shell apex pointing upward). Numerous species have an operculum, which in many species acts as a trapdoor to close the shell. This is usually made of a horn-like material, but in some molluscs it is calcareous. In the land slugs, the shell is reduced or absent, and the body is streamlined.

Some gastropods have adult shells which are bottom heavy due to the presence of a thick, often broad, convex ventral callus deposit on the inner lip and adapical to the aperture which may be important for gravitational stability.[22]

Body wall

Some sea slugs are very brightly colored. This serves either as a warning, when they are poisonous or contain stinging cells, or to camouflage them on the brightly colored hydroids, sponges and seaweeds on which many of the species are found.

Lateral outgrowths on the body of nudibranchs are called cerata. These contain an outpocketing of digestive gland called the diverticula.

Sensory organs and nervous system

The upper pair of tentacles on the head of Helix pomatia have eye spots, but the main sensory organs of the snail are sensory receptors for olfaction, situated in the epithelium of the tentacles.
The upper pair of tentacles on the head of Helix pomatia have eye spots, but the main sensory organs of the snail are sensory receptors for olfaction, situated in the epithelium of the tentacles.

The sensory organs of gastropods include olfactory organs, eyes, statocysts and mechanoreceptors.[23] Gastropods have no hearing.[23]

In terrestrial gastropods (land snails and slugs), the olfactory organs, located on the tips of the four tentacles, are the most important sensory organ.[23] The chemosensory organs of opisthobranch marine gastropods are called rhinophores.

The majority of gastropods have simple visual organs, eye spots either at the tip or base of the tentacles. However, "eyes" in gastropods range from simple ocelli that only distinguish light and dark, to more complex pit eyes, and even to lens eyes.[24] In land snails and slugs, vision is not the most important sense, because they are mainly nocturnal animals.[23]

The nervous system of gastropods includes the peripheral nervous system and the central nervous system. The central nervous system consists of ganglia connected by nerve cells. It includes paired ganglia: the cerebral ganglia, pedal ganglia, osphradial ganglia, pleural ganglia, parietal ganglia and the visceral ganglia. There are sometimes also buccal ganglia.[23]

Digestive system

The radula of a gastropod is usually adapted to the food that a species eats. The simplest gastropods are the limpets and abalones, herbivores that use their hard radula to rasp at seaweeds on rocks.

Many marine gastropods are burrowers, and have a siphon that extends out from the mantle edge. Sometimes the shell has a siphonal canal to accommodate this structure. A siphon enables the animal to draw water into their mantle cavity and over the gill. They use the siphon primarily to "taste" the water to detect prey from a distance. Gastropods with siphons tend to be either predators or scavengers.

Respiratory system

Almost all marine gastropods breathe with a gill, but many freshwater species, and the majority of terrestrial species, have a pallial lung. The respiratory protein in almost all gastropods is hemocyanin, but one freshwater pulmonate family, the Planorbidae, have hemoglobin as the respiratory protein.

In one large group of sea slugs, the gills are arranged as a rosette of feathery plumes on their backs, which gives rise to their other name, nudibranchs. Some nudibranchs have smooth or warty backs with no visible gill mechanism, such that respiration may likely take place directly through the skin.

Circulatory system

Gastropods have open circulatory system and the transport fluid is hemolymph. Hemocyanin is present in the hemolymph as the respiratory pigment.

Excretory system

The primary organs of excretion in gastropods are nephridia, which produce either ammonia or uric acid as a waste product. The nephridium also plays an important role in maintaining water balance in freshwater and terrestrial species. Additional organs of excretion, at least in some species, include pericardial glands in the body cavity, and digestive glands opening into the stomach.

Reproductive system

Mating behaviour of Elysia timida
Mating behaviour of Elysia timida

Courtship is a part of mating behavior in some gastropods, including some of the Helicidae. Again, in some land snails, an unusual feature of the reproductive system of gastropods is the presence and utilization of love darts.

In many marine gastropods other than the opisthobranchs, there are separate sexes (dioecious/gonochoric); most land gastropods, however, are hermaphrodites.

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Clade

Clade

In biological phylogenetics, a clade, also known as a monophyletic group or natural group, is a grouping of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. In the taxonomical literature, sometimes the Latin form cladus is used rather than the English form.

Anus

Anus

The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, depending on the type of animal, includes: matter which the animal cannot digest, such as bones; food material after the nutrients have been extracted, for example cellulose or lignin; ingested matter which would be toxic if it remained in the digestive tract; and dead or excess gut bacteria and other endosymbionts.

Opisthobranchia

Opisthobranchia

Opisthobranchs is now an informal name for a large and diverse group of specialized complex gastropods which used to be united in the subclass Opisthobranchia. That taxon is no longer considered to represent a monophyletic grouping.

Ctenidium (mollusc)

Ctenidium (mollusc)

A ctenidium is a respiratory organ or gill which is found in many molluscs. This structure exists in bivalves, cephalopods, Polyplacophorans (chitons), and in aquatic gastropods such as freshwater snails and marine snails. Some aquatic gastropods possess one ctenidium known as monopectinate and others have a pair of ctenidia known as bipectinate.

Gonad

Gonad

A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sperm in the form of spermatozoa. The female gonad, the ovary, produces egg cells. Both of these gametes are haploid cells. Some hermaphroditic animals have a type of gonad called an ovotestis.

Nephridium

Nephridium

The nephridium is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys. Nephridia remove metabolic wastes from an animal's body. Nephridia come in two basic categories: metanephridia and protonephridia. All nephridia- and kidney- having animals belong to the clade Nephrozoa.

Greek language

Greek language

Greek is an independent branch of the Indo-European family of languages, native to Greece, Cyprus, southern Italy, southern Albania, and other regions of the Balkans, the Black Sea coast, Asia Minor, and the Eastern Mediterranean. It has the longest documented history of any Indo-European language, spanning at least 3,400 years of written records. Its writing system is the Greek alphabet, which has been used for approximately 2,800 years; previously, Greek was recorded in writing systems such as Linear B and the Cypriot syllabary. The alphabet arose from the Phoenician script and was in turn the basis of the Latin, Cyrillic, Armenian, Coptic, Gothic, and many other writing systems.

Foot

Foot

The foot is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is a separate organ at the terminal part of the leg made up of one or more segments or bones, generally including claws and or nails.

Protoconch

Protoconch

A protoconch is an embryonic or larval shell which occurs in some classes of molluscs, e.g., the initial chamber of an ammonite or the larval shell of a gastropod. In older texts it is also called "nucleus". The protoconch may sometimes consist of several whorls, but when this is the case, the whorls show no growth lines.

Osphradium

Osphradium

The osphradium is a pigmented chemosensory epithelium patch in the mantle cavity present in six of the eight extant classes of molluscs, on or adjacent to the ctenidia (gills). The main function of this organ is disputed but it is believed to be used to test incoming water for silt and possible food particles or, in some species, for sensing the presence of light.

Hypobranchial gland

Hypobranchial gland

The hypobranchial gland is a glandular structure which is part of the anatomy of many mollusks, including several different families of gastropods, and also many protobranch bivalves. This gland produces mucus as well as biologically active compounds. The cephalopod ink sac is a modified hypobranchial gland.

Auricle (anatomy)

Auricle (anatomy)

The auricle or auricula is the visible part of the ear that is outside the head. It is also called the pinna, a term that is used more in zoology.

Life cycle

Egg strings of an Aplysia species.
Egg strings of an Aplysia species.

Courtship is a part of the behavior of mating gastropods with some pulmonate families of land snails creating and utilizing love darts, the throwing of which have been identified as a form of sexual selection.[25]

The main aspects of the life cycle of gastropods include:

  • Egg laying and the eggs of gastropods
  • The embryonic development of gastropods
  • The larvae or larval stadium: some gastropods may be trochophore and/or veliger
  • Estivation and hibernation (each of these are present in some gastropods only)
  • The growth of gastropods
  • Courtship and mating in gastropods: fertilization is internal or external according to the species. External fertilization is common in marine gastropods.

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Reproductive system of gastropods

Reproductive system of gastropods

The reproductive system of gastropods varies greatly from one group to another within this very large and diverse taxonomic class of animals. Their reproductive strategies also vary greatly, see Mating of gastropods.

Mating of gastropods

Mating of gastropods

The mating of gastropods is a vast and varied topic, because the taxonomic class Gastropoda is very large and diverse, a group comprising sea snails and sea slugs, freshwater snails and land snails and slugs. Gastropods are second only to the class Insecta in terms of total number of species. Some gastropods have separate sexes, others are hermaphroditic. Some hermaphroditic groups have simultaneous hermaphroditism, whereas some sequential hermaphroditism. In addition, numerous very different mating strategies are used within different taxa.

Mating

Mating

In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. Fertilization is the fusion of two gametes. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization. Mating may also lead to external fertilization, as seen in amphibians, fishes and plants. For most species, mating is between two individuals of opposite sexes. However, for some hermaphroditic species, copulation is not required because the parent organism is capable of self-fertilization (autogamy); for example, banana slugs.

Love dart

Love dart

A love dart is a sharp, calcareous or chitinous dart which some hermaphroditic land snails and slugs create. Love darts are both formed and stored internally in a dart sac. These darts are made in sexually mature animals only, and are used as part of the sequence of events during courtship, before actual mating takes place. Darts are quite large compared to the size of the animal: in the case of the semi-slug genus Parmarion, the length of a dart can be up to one fifth that of the semi-slug's foot.

Sexual selection

Sexual selection

Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with, and compete with members of the same sex for access to members of the opposite sex. These two forms of selection mean that some individuals have greater reproductive success than others within a population, for example because they are more attractive or prefer more attractive partners to produce offspring. Successful males benefit from frequent mating and monopolizing access to one or more fertile females. Females can maximise the return on the energy they invest in reproduction by selecting and mating with the best males.

Trochophore

Trochophore

A trochophore is a type of free-swimming planktonic marine larva with several bands of cilia.

Veliger

Veliger

A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells.

Hibernation

Hibernation

Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most commonly occurs during winter months.

Feeding behavior

A Pomacea maculata floating and eating a carrot
A Pomacea maculata floating and eating a carrot

The diet of gastropods differs according to the group considered. Marine gastropods include some that are herbivores, detritus feeders, predatory carnivores, scavengers, parasites, and also a few ciliary feeders, in which the radula is reduced or absent. Land-dwelling species can chew up leaves, bark, fruit and decomposing animals while marine species can scrape algae off the rocks on the seafloor. Certain species such as the Archaeogastropda maintain horizontal rows of slender marginal teeth. In some species that have evolved into endoparasites, such as the eulimid Thyonicola doglieli, many of the standard gastropod features are strongly reduced or absent.

A few sea slugs are herbivores and some are carnivores. The carnivorous habit is due to specialisation. Many gastropods have distinct dietary preferences and regularly occur in close association with their food species.

Some predatory carnivorous gastropods include, for example: Cone shells, Testacella, Daudebardia, Turrids, Ghost slug and others.

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Pomacea maculata

Pomacea maculata

Pomacea maculata is a species of large freshwater snail with an operculum, an aquatic gastropod mollusk in the family Ampullariidae, the apple snails.

Herbivore

Herbivore

A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthparts adapted to rasping or grinding. Horses and other herbivores have wide flat teeth that are adapted to grinding grass, tree bark, and other tough plant material.

Detritus

Detritus

In biology, detritus is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts communities of microorganisms that colonize and decompose it. In terrestrial ecosystems it is present as leaf litter and other organic matter that is intermixed with soil, which is denominated "soil organic matter". The detritus of aquatic ecosystems is organic material that is suspended in the water and accumulates in depositions on the floor of the body of water; when this floor is a seabed, such a deposition is denominated "marine snow".

Carnivore

Carnivore

A carnivore, or meat-eater, is an animal or plant whose food and energy requirements derive from animal tissues whether through hunting or scavenging.

Scavenger

Scavenger

Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding behavior. Scavengers play an important role in the ecosystem by consuming dead animal and plant material. Decomposers and detritivores complete this process, by consuming the remains left by scavengers.

Radula

Radula

The radula is an anatomical structure used by mollusks for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food enters the esophagus. The radula is unique to the mollusks, and is found in every class of mollusk except the bivalves, which instead use cilia, waving filaments that bring minute organisms to the mouth.

Eulimidae

Eulimidae

Eulimidae is a family of very small parasitic sea snails, marine gastropod mollusks in the superfamily Vanikoroidea.

Thyonicola dogieli

Thyonicola dogieli

Thyonicola dogieli is a parasitic species of gastropod mollusc in the family Eulimidae. It parasitises sea cucumbers in the northeastern Pacific Ocean.

Sea slug

Sea slug

Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are gastropods, i.e. they are sea snails that over evolutionary time have either completely lost their shells, or have seemingly lost their shells due to having a greatly reduced or internal shell. The name "sea slug" is most often applied to nudibranchs, as well as to a paraphyletic set of other marine gastropods without obvious shells.

Testacella

Testacella

Testacella is genus of small to medium-large, predatory, air-breathing, land slugs.

Daudebardia

Daudebardia

Daudebardia are small air-breathing land snails or semi-slugs, terrestrial pulmonate gastropods in the family Oxychilidae, the glass snails.

Turrid

Turrid

Turrid, plural turrids, is a common name for a very large group of predatory sea snails, marine gastropod mollusks which until recently were all classified in the family Turridae. However, recently the family was discovered to be polyphyletic and therefore was split into a number of families.

Genetics

Gastropods exhibit an important degree of variation in mitochondrial gene organization when compared to other animals.[26] Main events of gene rearrangement occurred at the origin of Patellogastropoda and Heterobranchia, whereas fewer changes occurred between the ancestors of Vetigastropoda (only tRNAs D, C and N) and Caenogastropoda (a large single inversion, and translocations of the tRNAs D and N).[26] Within Heterobranchia, gene order seems relatively conserved, and gene rearrangements are mostly related with transposition of tRNA genes.[26]

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Mutation

Mutation

In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mitosis, or meiosis or other types of damage to DNA, which then may undergo error-prone repair, cause an error during other forms of repair, or cause an error during replication. Mutations may also result from insertion or deletion of segments of DNA due to mobile genetic elements.

Patellogastropoda

Patellogastropoda

The Patellogastropoda, common name true limpets and historically called the Docoglossa, are members of a major phylogenetic group of marine gastropods, treated by experts either as a clade or as a taxonomic order.

Heterobranchia

Heterobranchia

Heterobranchia, the heterobranchs, is a taxonomic clade of snails and slugs, which includes marine, aquatic and terrestrial gastropod mollusks.

Vetigastropoda

Vetigastropoda

Vetigastropoda is a major taxonomic group of sea snails, marine gastropod mollusks that form a very ancient lineage. Taxonomically the Vetigastropoda are sometimes treated as an order, although they are treated as an unranked clade in Bouchet and Rocroi, 2005.

Caenogastropoda

Caenogastropoda

Caenogastropoda is a taxonomic subclass of molluscs in the class Gastropoda. It is a large diverse group which are mostly sea snails and other marine gastropod mollusks, but also includes some freshwater snails and some land snails. The subclass is the most diverse and ecologically successful of the gastropods.

Chromosomal inversion

Chromosomal inversion

An inversion is a chromosome rearrangement in which a segment of a chromosome becomes inverted within its original position. An inversion occurs when a chromosome undergoes a two breaks within the chromosomal arm, and the segment between the two breaks inserts itself in the opposite direction in the same chromosome arm. The breakpoints of inversions often happen in regions of repetitive nucleotides, and the regions may be reused in other inversions. Chromosomal segments in inversions can be as small as 100 kilobases or as large as 100 megabases. The number of genes captured by an inversion can range from a handful of genes to hundreds of genes. Inversions can happen either through ectopic recombination, chromosomal breakage and repair, or non-homologous end joining.

Chromosomal translocation

Chromosomal translocation

In genetics, chromosome translocation is a phenomenon that results in unusual rearrangement of chromosomes. This includes balanced and unbalanced translocation, with two main types: reciprocal-, and Robertsonian translocation. Reciprocal translocation is a chromosome abnormality caused by exchange of parts between non-homologous chromosomes. Two detached fragments of two different chromosomes are switched. Robertsonian translocation occurs when two non-homologous chromosomes get attached, meaning that given two healthy pairs of chromosomes, one of each pair "sticks" and blends together homogeneously.

Geological history and evolution

Trochonema sp., an early gastropod from the Middle Ordovician of the Galena Group of Minnesota.
Trochonema sp., an early gastropod from the Middle Ordovician of the Galena Group of Minnesota.
Fossil gastropod and attached mytilid bivalves on a Jurassic limestone bedding plane of the Matmor Formation in southern Israel.
Fossil gastropod and attached mytilid bivalves on a Jurassic limestone bedding plane of the Matmor Formation in southern Israel.

The first gastropods were exclusively marine, with the earliest representatives of the group appearing in the Late Cambrian (Chippewaella, Strepsodiscus),[27] though their only gastropod character is a coiled shell, so they could lie in the stem lineage, if they are gastropods at all.[28] Earliest Cambrian organisms like Helcionella, Barskovia and Scenella are no longer considered gastropods, and the tiny coiled Aldanella of earliest Cambrian time is probably not even a mollusk.

As such, it's not until the Ordovician that the first crown-group members arise.[29] By the Ordovician period the gastropods were a varied group present in a range of aquatic habitats. Commonly, fossil gastropods from the rocks of the early Palaeozoic era are too poorly preserved for accurate identification. Still, the Silurian genus Poleumita contains fifteen identified species. Fossil gastropods were less common during the Palaeozoic era than bivalves.[29]

Most of the gastropods of the Palaeozoic era belong to primitive groups, a few of which still survive. By the Carboniferous period many of the shapes seen in living gastropods can be matched in the fossil record, but despite these similarities in appearance the majority of these older forms are not directly related to living forms. It was during the Mesozoic era that the ancestors of many of the living gastropods evolved.[29]

One of the earliest known terrestrial (land-dwelling) gastropods is Anthracopupa (=Maturipupa),[30] which is found in the Coal Measures of the Carboniferous period in Europe, but relatives of the modern land snails are rare before the Cretaceous period, when the familiar Helix first appeared.[29]

In rocks of the Mesozoic era, gastropods are slightly more common as fossils; their shells are often well preserved. Their fossils occur in ancient beds deposited in both freshwater and marine environments. The "Purbeck Marble" of the Jurassic period and the "Sussex Marble" of the early Cretaceous period, which both occur in southern England, are limestones containing the tightly packed remains of the pond snail Viviparus.[29]

Rocks of the Cenozoic era yield very large numbers of gastropod fossils, many of these fossils being closely related to modern living forms. The diversity of the gastropods increased markedly at the beginning of this era, along with that of the bivalves.[29]

Certain trail-like markings preserved in ancient sedimentary rocks are thought to have been made by gastropods crawling over the soft mud and sand. Although these trace fossils are of debatable origin, some of them do resemble the trails made by living gastropods today.[29]

Gastropod fossils may sometimes be confused with ammonites or other shelled cephalopods. An example of this is Bellerophon from the limestones of the Carboniferous period in Europe, the shell of which is planispirally coiled and can be mistaken for the shell of a cephalopod.

Gastropods are one of the groups that record the changes in fauna caused by the advance and retreat of the Ice Sheets during the Pleistocene epoch.

Cladogram

A cladogram showing the phylogenic relationships of Gastropoda with example species:[31]

Gastropoda

Panpulmonata False Limpet Shell Macro Digon3.jpgAncylus fluviatilis O.F.Müller, 1774 (4206383684).jpgGrapevinesnail 01a.jpgBiomphalaria glabrata.jpg

Euopisthobranchia Aplysia californica NHGRI-79108.jpg

Nudipleura Glaucus atlanticus 1 cropped.jpg

CaenogastropodaAmpullaria ampullacea (Linné) (2999358117).jpgAcicula lineata (Draparnaud, 1801) (4309322782).jpgEpitonium-scalare--Trp-6.pngCharonia tritonis.pngAcanthina brevidentata (Wood, 1828) (3063441884).jpgNeverita didyma.jpgOncomelania hupensis.jpg

NeritimorphaTheodoxus anatolicus (Récluz, 1841) (3000195752).jpg

VetigastropodaAngaria Delphinus Shell Macro.JPGOxystele variegata - ZooKeys-365-337-g004A.jpgLucapina suffusa (Reeve, 1850) (4947706383).jpgFissurella nimbosa Linné, 1758 (4426866577).jpg

Patellogastropoda Patelloida pustulata (Helbling, 1779) (4631972937).jpg

Cocculiniformia, Neomphalina and Lower Heterobranchia are not included in the above cladogram.

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List of marine gastropod genera in the fossil record

List of marine gastropod genera in the fossil record

This list of marine gastropod genera in the fossil record is an attempt to list all the genera of sea snails or marine gastropod mollusks which have been found in the fossil record. Nearly all of these are genera of shelled forms, since it is relatively rare for gastropods without a shell to leave any recognizable traces. It is also worth pointing out that this list of genera represents only a very tiny fraction of the number of genera that must actually have existed over the evolutionary time span: the fossil record is an extremely patchy and exceedingly incomplete pteryrecord of life on earth in earlier geological eras.

Galena Group

Galena Group

The Galena Group or Galena Limestone refers to a sedimentary sequence of Ordovician limestone that was deposited atop the Decorah Shale. It is part of the Ordovician stratigraphy of the Upper Midwestern United States. It was deposited in a calm marine environment, and is fossiliferous.

Minnesota

Minnesota

Minnesota is a state in the Upper Midwestern region of the United States. It is the 12th largest U.S. state in area and the 22nd most populous, with over 5.75 million residents. Minnesota is home to western prairies, now given over to intensive agriculture; deciduous forests in the southeast, now partially cleared, farmed, and settled; and the less populated North Woods, used for mining, forestry, and recreation. Roughly a third of the state is covered in forests, and it is known as the "Land of 10,000 Lakes" for having over 14,000 bodies of fresh water covering at least ten acres. More than 60% of Minnesotans live in the Minneapolis–Saint Paul metropolitan area, known as the "Twin Cities", the state's main political, economic, and cultural hub. With a population of about 3.7 million, the Twin Cities is the 16th largest metropolitan area in the U.S. Other minor metropolitan and micropolitan statistical areas in the state include Duluth, Mankato, Moorhead, Rochester, and St. Cloud.

Matmor Formation

Matmor Formation

The Matmor Formation is a geologic formation of up to 100 metres (330 ft) thick, that is exposed in Hamakhtesh Hagadol in southern Israel. The Matmor Formation contains fossils from a Jurassic equatorial shallow marine environment. Bivalves, gastropods, sponges, corals, echinoderms, and sclerobionts are present in the Matmor Formation to various degrees. The stratigraphy of the Matmor Formation consists of alternating layers of limestone and marl.

Israel

Israel

Israel, officially the State of Israel, is a country in Western Asia. Situated in the Southern Levant, it is bordered by Lebanon to the north, by Syria to the northeast, by Jordan to the east, by the Red Sea to the south, by Egypt to the southwest, by the Mediterranean Sea to the west, and by the Palestinian territories — the West Bank along the east and the Gaza Strip along the southwest. Tel Aviv is the economic and technological center of the country, while its seat of government is in its proclaimed capital of Jerusalem, although Israeli sovereignty over East Jerusalem is unrecognized internationally.

Furongian

Furongian

The Furongian is the fourth and final epoch and series of the Cambrian. It lasted from 497 to 485.4 million years ago. It succeeds the Miaolingian series of the Cambrian and precedes the Lower Ordovician Tremadocian Stage. It is subdivided into three stages: the Paibian, Jiangshanian and the unnamed 10th stage of the Cambrian.

Chippewaella

Chippewaella

Chippewaella patellitheca is a stem-gastropod mollusc from Furongian-aged strata of Late Cambrian Wisconsin. According to Peter J. Wagner, it is the most basal gastropod.

Cambrian

Cambrian

The Cambrian Period is the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago (mya) to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, and its base, are somewhat in flux. The period was established as "Cambrian series" by Adam Sedgwick, who named it after Cambria, the Latin name for 'Cymru' (Wales), where Britain's Cambrian rocks are best exposed. Sedgwick identified the layer as part of his task, along with Roderick Murchison, to subdivide the large "Transition Series", although the two geologists disagreed for a while on the appropriate categorization. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells. As a result, our understanding of the Cambrian biology surpasses that of some later periods.

Helcionella

Helcionella

Helcionella is a genus of helcionellid, a fossil marine invertebrate animal that is considered to be a mollusk and may possibly be a gastropod. The shells of these animals are about a centimetre in size. The tip of the shell extends beyond the rear extremity of the shell's aperture, and the shell is "endogastric" in shape.

Barskovia

Barskovia

Barskovia is a torted conical shell known from earliest Cambrian small skeletal fossils, interpreted as a helcionelloid.

Aldanella

Aldanella

Aldanella is an extinct paleozoic mollusc that was assigned to the Gastropod stem group but may also belong to a paraphyletic "Monoplacophora".

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Taxonomy

A group of fossil shells of Turritella cingulifera from the Pliocene of Cyprus
A group of fossil shells of Turritella cingulifera from the Pliocene of Cyprus
Five views of a shell of a Fulguropsis species
Five views of a shell of a Fulguropsis species
Microphoto (35x) of Gastropoda sp. from Holocene sediments of Amuq Plain SSE Turkey
Microphoto (35x) of Gastropoda sp. from Holocene sediments of Amuq Plain SSE Turkey

Since Darwin, biological taxonomy has attempted to reflect the phylogeny of organisms, i.e., the tree of life. The classifications used in taxonomy attempt to represent the precise interrelatedness of the various taxa. However, the taxonomy of the Gastropoda is constantly being revised and so the versions shown in various texts can differ in major ways.

In the older classification of the gastropods, there were four subclasses:[32]

The taxonomy of the Gastropoda is still under revision, and more and more of the old taxonomy is being abandoned, as the results of DNA studies slowly become clearer. Nevertheless, a few of the older terms such as "opisthobranch" and "prosobranch" are still sometimes used in a descriptive way.

New insights based on DNA sequencing of gastropods have produced some revolutionary new taxonomic insights. In the case of the Gastropoda, the taxonomy is now gradually being rewritten to embody strictly monophyletic groups (only one lineage of gastropods in each group). Integrating new findings into a working taxonomy remain challenging. Consistent ranks within the taxonomy at the level of subclass, superorder, order, and suborder have already been abandoned as unworkable. Ongoing revisions of the higher taxonomic levels are expected in the near future.

Convergent evolution, which appears to exist at especially high frequency in gastropods, may account for the observed differences between the older phylogenies, which were based on morphological data, and more recent gene-sequencing studies.

Bouchet & Rocroi (2005)[3][33] made sweeping changes in the systematics, resulting in a taxonomy that is a step closer to the evolutionary history of the phylum. The Bouchet & Rocroi classification system is based partly on the older systems of classification, and partly on new cladistic research.

In the past, the taxonomy of gastropods was largely based on phenetic morphological characters of the taxa. The recent advances are more based on molecular characters from DNA[34] and RNA research. This has made the taxonomical ranks and their hierarchy controversial. The debate about these issues is not likely to end soon.

In the Bouchet, Rocroi et al. taxonomy, the authors have used unranked clades for taxa above the rank of superfamily (replacing the ranks suborder, order, superorder and subclass), while using the traditional Linnaean approach for all taxa below the rank of superfamily. Whenever monophyly has not been tested, or is known to be paraphyletic or polyphyletic, the term "group" or "informal group" has been used. The classification of families into subfamilies is often not well resolved, and should be regarded as the best possible hypothesis.

In 2004, Brian Simison and David R. Lindberg showed possible diphyletic origins of the Gastropoda based on mitochondrial gene order and amino acid sequence analyses of complete genes.[35]

In the 2017 issue of the Malacologia journal (available online from 4 January 2018), a significantly updated version of the 2005 "Bouchet & Rocroi" taxonomy was published in the paper "Revised Classification, Nomenclator and Typification of Gastropod and Monoplacophoran Families".[36]

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Taxonomy of the Gastropoda (Bouchet & Rocroi, 2005)

Taxonomy of the Gastropoda (Bouchet & Rocroi, 2005)

The taxonomy of the Gastropoda as it was revised in 2005 by Philippe Bouchet and Jean-Pierre Rocroi is a system for the scientific classification of gastropod mollusks. The paper setting out this taxonomy was published in the journal Malacologia. The system encompasses both living and extinct groups, as well as some fossils whose classification as gastropods is uncertain.

Changes in the taxonomy of gastropods since 2005

Changes in the taxonomy of gastropods since 2005

This overview lists proposed changes in the taxonomy of gastropods at the family level and above since 2005, when the taxonomy of the Gastropoda by Bouchet & Rocroi (2005) was published. In other words, these are recent updates in the way various groups of snails and slugs are classified.

Pliocene

Pliocene

The Pliocene is the epoch in the geologic time scale that extends from 5.333 million to 2.58 million years ago. It is the second and most recent epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch. Prior to the 2009 revision of the geologic time scale, which placed the four most recent major glaciations entirely within the Pleistocene, the Pliocene also included the Gelasian Stage, which lasted from 2.588 to 1.806 million years ago, and is now included in the Pleistocene.

Cyprus

Cyprus

Cyprus, officially the Republic of Cyprus, is an island country located south of the Anatolian Peninsula in the eastern Mediterranean Sea. It is geographically in Western Asia, but its cultural ties and geopolitics are overwhelmingly Southeastern European. Cyprus is the third-largest and third-most populous island in the Mediterranean. It is located north of Egypt, east of Greece, south of Turkey, and west of Lebanon and Syria. Its capital and largest city is Nicosia. The northeast portion of the island is de facto governed by the self-declared Turkish Republic of Northern Cyprus.

Fulguropsis

Fulguropsis

Fulguropsis is a genus of sea snails, marine gastropod molluscs in the family Busyconidae, the crown conches and their allies.

Charles Darwin

Charles Darwin

Charles Robert Darwin was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended from a common ancestor is now generally accepted and considered a fundamental concept in science. In a joint publication with Alfred Russel Wallace, he introduced his scientific theory that this branching pattern of evolution resulted from a process he called natural selection, in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding. Darwin has been described as one of the most influential figures in human history and was honoured by burial in Westminster Abbey.

Opisthobranchia

Opisthobranchia

Opisthobranchs is now an informal name for a large and diverse group of specialized complex gastropods which used to be united in the subclass Opisthobranchia. That taxon is no longer considered to represent a monophyletic grouping.

Prosobranchia

Prosobranchia

Prosobranchia was a large taxonomic subclass of sea snails, land snails and freshwater snails. This taxon of gastropods dates back to the 1920s. It has however been proven to be polyphyletic. Generally speaking in biology taxonomy is required to reflect phylogeny, in other words the classification of a group must reflect its evolutionary descent, as far as that is known, so the taxon Prosobranchia is no longer considered suitable to be used.

Pulmonata

Pulmonata

Pulmonata or pulmonates, is an informal group of snails and slugs characterized by the ability to breathe air, by virtue of having a pallial lung instead of a gill, or gills. The group includes many land and freshwater families, and several marine families.

Taxonomy (biology)

Taxonomy (biology)

In biology, taxonomy is the scientific study of naming, defining (circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa and these groups are given a taxonomic rank; groups of a given rank can be aggregated to form a more inclusive group of higher rank, thus creating a taxonomic hierarchy. The principal ranks in modern use are domain, kingdom, phylum, class, order, family, genus, and species. The Swedish botanist Carl Linnaeus is regarded as the founder of the current system of taxonomy, as he developed a ranked system known as Linnaean taxonomy for categorizing organisms and binomial nomenclature for naming organisms.

DNA

DNA

Deoxyribonucleic acid is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of all known organisms and many viruses. DNA and ribonucleic acid (RNA) are nucleic acids. Alongside proteins, lipids and complex carbohydrates (polysaccharides), nucleic acids are one of the four major types of macromolecules that are essential for all known forms of life.

Convergent evolution

Convergent evolution

Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last common ancestor of those groups. The cladistic term for the same phenomenon is homoplasy. The recurrent evolution of flight is a classic example, as flying insects, birds, pterosaurs, and bats have independently evolved the useful capacity of flight. Functionally similar features that have arisen through convergent evolution are analogous, whereas homologous structures or traits have a common origin but can have dissimilar functions. Bird, bat, and pterosaur wings are analogous structures, but their forelimbs are homologous, sharing an ancestral state despite serving different functions.

Source: "Gastropoda", Wikipedia, Wikimedia Foundation, (2023, March 20th), https://en.wikipedia.org/wiki/Gastropoda.

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References

This article incorporates CC-BY-2.0 text from the reference.[26]

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