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Pulmonata or pulmonates is an informal group (previously an
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
, and before that, a subclass) of
snails A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
and
slug Slug, or land slug, is a common name for any apparently shell-less Terrestrial mollusc, 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 ...
s characterized by the ability to breathe air, by virtue of having a pallial lung instead of a
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
, or gills. The group includes many land and freshwater families, and several marine families. The
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
Pulmonata as traditionally defined was found to be
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
in a molecular study per Jörger ''et al.'', dating from 2010. Pulmonata are known from the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period to the present. Pulmonates have a single atrium and kidney, and a concentrated symmetrical nervous system. The mantle cavity is on the right side of the body, and lacks gills, instead being converted into a vascularised lung. Most species have a shell, but no operculum, although the group does also include several shell-less slugs. Pulmonates are hermaphroditic, and some groups possess
love dart A love dart (also known as a gypsobelum, shooting darts, or just as darts) is a sharp, calcium carbonate, calcareous or chitinous Dart (missile), dart which some hermaphroditic land snails and slugs create. Love darts are both formed and stor ...
s.


Linnean taxonomy

The taxonomy of this group according to the taxonomy of the Gastropoda (Ponder & Lindberg, 1997) was as follows: Order Pulmonata Cuvier in Blainville, 1814 - pulmonates *Suborder
Systellommatophora The Systellommatophora (synonym Gymnomorpha) is a clade of primitive, air-breathing slugs, according to the taxonomy of the Gastropoda (Bouchet & Rocroi, 2005). They are marine and terrestrial pulmonate gastropods within the Heterobranchia. T ...
Pilsbry, 1948 ***Superfamily Onchidioidea
Rafinesque Constantine Samuel Rafinesque-Schmaltz (; 22 October 178318 September 1840) was a French early 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ult ...
, 1815 ***Superfamily Otinoidea H. & A. Adams, 1855 ***Superfamily Rathouisioidea Sarasin, 1889 *Suborder Basommatophora Keferstein in Bronn, 1864 - freshwater pulmonates, pond snails ***Superfamily Acroloxoidea Thiele, 1931 ***Superfamily Amphiboloidea J.E. Gray, 1840 ***Superfamily Chilinoidea H. & A. Adams, 1855 ***Superfamily Glacidorboidea Ponder, 1986 ***Superfamily Lymnaeoidea
Rafinesque Constantine Samuel Rafinesque-Schmaltz (; 22 October 178318 September 1840) was a French early 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ult ...
, 1815 ***Superfamily Planorboidea
Rafinesque Constantine Samuel Rafinesque-Schmaltz (; 22 October 178318 September 1840) was a French early 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ult ...
, 1815 ***Superfamily Siphonarioidea J.E. Gray, 1840 *Suborder
Eupulmonata Eupulmonata is a taxonomic clade of air-breathing gastropod molluscs. The great majority of this group are land snails and slugs, but some are intertidal or inhabit coastal saltmarshes and mangroves. Possible synapomorphy of the group are gl ...
Haszprunar & Huber, 1990 **Infraorder Acteophila Dall, 1885 = formerly Archaeopulmonata ***Superfamily Melampoidea Stimpson, 1851 **Infraorder Trimusculiformes Minichev & Starobogatov, 1975 ***Superfamily Trimusculoidea Zilch, 1959 **Infraorder Stylommatophora A. Schmidt, 1856 – land snails and slugs ***Subinfraorder Orthurethra ***Superfamily Achatinelloidea Gulick, 1873 ***Superfamily Cochlicopoidea Pilsbry, 1900 ***Superfamily Partuloidea Pilsbry, 1900 ***Superfamily
Pupilloidea Pupilloidea is a superfamily of small and very small air-breathing land snails, terrestrial gastropods in the infraorder '' Pupilloidei''. They are diverse in morphology, habitat, and ecological roles. Description They are characterized by the ...
Turton, 1831 ***Subinfraorder
Sigmurethra Sigmurethra is a taxonomic category of air-breathing land snails and slugs, terrestrial pulmonate gastropod molluscs. This is an informal group which includes most land snails and slugs. The two strong synapomorphies of Sigmurethra are a long ...
***Superfamily Acavoidea Pilsbry, 1895 ***Superfamily Achatinoidea Swainson, 1840 ***Superfamily Aillyoidea Baker, 1960 ***Superfamily Arionoidea J.E. Gray in Turnton, 1840 ***Superfamily Buliminoidea Clessin, 1879 ***Superfamily Camaenoidea Pilsbry, 1895 ***Superfamily Clausilioidea Mörch, 1864 ***Superfamily Dyakioidea Gude & Woodward, 1921 ***Superfamily Gastrodontoidea Tryon, 1866 ***Superfamily
Helicoidea Helicoidea is a Taxonomy (biology), taxonomic Taxonomic rank, superfamily of air-breathing land snails, terrestrial molluscs, terrestrial pulmonate gastropod mollusks in the order (taxonomy), order Stylommatophora. Taxonomy 2017 taxonomy and l ...
Rafinesque Constantine Samuel Rafinesque-Schmaltz (; 22 October 178318 September 1840) was a French early 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ult ...
, 1815 ***Superfamily Helixarionoidea Bourguignat, 1877 ***Superfamily Limacoidea
Rafinesque Constantine Samuel Rafinesque-Schmaltz (; 22 October 178318 September 1840) was a French early 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ult ...
, 1815 ***Superfamily Oleacinoidea H. & A. Adams, 1855 ***Superfamily Orthalicoidea Albers-Martens, 1860 ***Superfamily Plectopylidoidea Moellendorf, 1900 ***Superfamily Polygyroidea Pilsbry, 1894 ***Superfamily Punctoidea Morse, 1864 ***Superfamily Rhytidoidea Pilsbry, 1893 ***Superfamily Sagdidoidera Pilsbry, 1895 ***Superfamily Staffordioidea Thiele, 1931 ***Superfamily Streptaxoidea J.E. Gray, 1806 ***Superfamily Strophocheiloidea Thiele, 1926 ***Superfamily Trigonochlamydoidea Hese, 1882 ***Superfamily Zonitoidea Mörch, 1864 ***? Superfamily Athoracophoroidea P. Fischer, 1883 = Tracheopulmonata ***? Superfamily Succineoidea Beck, 1837 = Heterurethra


2005 taxonomy

The taxonomy of this group according to the
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 (Gastropods are a taxonomic class of animals which consists of snails and sl ...
was as follows:


Informal Group Pulmonata

Contains the informal group Basommatophora and the clade Eupulmonata


Informal Group Basommatophora

Contains the clade Hygrophila *Superfamily Amphiboloidea **Family Amphibolidae *Superfamily Siphonarioidea **Family
Siphonariidae Siphonariidae, also known as false limpets, are a taxonomic family of small to medium-sized air-breathing sea snails, marine and brackish water pulmonate gastropod molluscs. __TOC__ Shell description Empty siphonariid limpet shells can be di ...
** † Family Acroreiidae


= Clade Hygrophila

= *Superfamily Chilinoidea **Family Chilinidae **Family Latiidae *Superfamily Acroloxoidea **Family
Acroloxidae Acroloxidae, commonly known as river limpets, are a Taxonomy (biology), taxonomic family of very small, freshwater snails, Aquatic animal, aquatic pulmonate gastropod limpet-like mollusks with a simple flattened conical gastropod shell, shell in ...
*Superfamily Lymnaeoidea **Family
Lymnaeidae Lymnaeidae, common name the pond snails, is a Taxonomy (biology), taxonomic family (biology), family of small to large air-breathing freshwater snails, Aquatic animal, aquatic pulmonate gastropod mollusks, that belong to the clade Hygrophila (cl ...
*Superfamily Planorboidea **Family
Planorbidae Planorbidae, common name the ramshorn snails or ram's horn snails, is a family (biology), family of air-breathing freshwater snails, Aquatic animal, aquatic pulmonate gastropod Mollusca, molluscs. Unlike most molluscs, the blood of ram's horn sn ...
**Family Physidae


Clade Eupulmonata

Contains the clades Systellommatophora and Stylommatophora *Superfamily Trimusculoidea **Family Trimusculidae *Superfamily Otinoidea **Family Otinidae **Family Smeagolidae *Superfamily
Ellobioidea Ellobiidae, common name the hollow-shelled snails, is a family (biology), family of small air-breathing Land snail, land, Sea snail, marine, and brackish snails; pulmonate gastropod mollusks in the clade Eupulmonata.Bouchet, P. (2012). Ellobiida ...
**Family Ellobiidae


= Clade

Systellommatophora The Systellommatophora (synonym Gymnomorpha) is a clade of primitive, air-breathing slugs, according to the taxonomy of the Gastropoda (Bouchet & Rocroi, 2005). They are marine and terrestrial pulmonate gastropods within the Heterobranchia. T ...
( = Gymnomorpha)

= *Superfamily Onchidioidea **Family Onchidiidae *Superfamily Veronicelloidea **Family
Veronicellidae The Veronicellidae, also known by their common name the leatherleaf slugs, are a family of pulmonate terrestrial slugs. The herbivorous molluscs occur mainly in the tropical and subtropical areas of America, Asia and Africa. They act as interme ...
**Family Rathouisiidae


= Clade Stylommatophora

= Contains the subclades Elasmognatha, Orthurethra and the informal group Sigmurethra


= Subclade Elasmognatha

= *Superfamily Succineoidea **Family Succineidae *Superfamily Athoracophoroidea **Family Athoracophoridae


= Subclade Orthurethra

= *Superfamily Partuloidea **Family
Partulidae Partulidae is a family (biology), family of air-breathing land snails, Terrestrial molluscs, terrestrial gastropod mollusks in the Taxonomic rank, superfamily Pupilloidea.MolluscaBase eds. (2021). MolluscaBase. Partulidae Pilsbry, 1900. Accessed ...
**Family Draparnaudiidae *Superfamily Achatinelloidea **Family
Achatinellidae Achatinellidae is a family of tropical air-breathing land snails, terrestrial pulmonate gastropod mollusks in the superfamily Pupilloidea. Taxonomy It was previously the only family in the superfamily Achatinelloidea (according to the tax ...
*Superfamily Cochlicopoidea **Family Cochlicopidae **Family Amastridae *Superfamily
Pupilloidea Pupilloidea is a superfamily of small and very small air-breathing land snails, terrestrial gastropods in the infraorder '' Pupilloidei''. They are diverse in morphology, habitat, and ecological roles. Description They are characterized by the ...
**Family
Pupillidae Pupillidae is a family of mostly minute, air-breathing, land snails, terrestrial pulmonate gastropod mollusks or micromollusks in the superfamily Pupilloidea.MolluscaBase eds. (2021). MolluscaBase. Pupillidae W. Turton, 1831. Accessed through: ...
**Family Argnidae **Family Chondrinidae ** † Family Cylindrellinidae **Family Lauriidae **Family
Orculidae Orculidae is a family (biology), family of mostly minute, air-breathing, land snails; Terrestrial molluscs, terrestrial pulmonate gastropod mollusks in the superfamily Pupilloidea.MolluscaBase eds. (2021). MolluscaBase. Orculidae Pilsbry, 1918. A ...
**Family Pleurodiscidae **Family Pyramidulidae **Family Spelaeodiscidae **Family Strobilopsidae **Family Valloniidae **Family Vertiginidae *Superfamily Enoidea **Family
Enidae Enidae is a family of air-breathing land snails, terrestrial pulmonate gastropod mollusks.MolluscaBase eds. (2020). MolluscaBase. Enidae B. B. Woodward, 1903 (1880). Accessed through: World Register of Marine Species at: http://www.marinespecies ...
**Family Cerastidae


= Informal Group

Sigmurethra Sigmurethra is a taxonomic category of air-breathing land snails and slugs, terrestrial pulmonate gastropod molluscs. This is an informal group which includes most land snails and slugs. The two strong synapomorphies of Sigmurethra are a long ...

= *Superfamily Clausilioidea **Family
Clausiliidae Clausiliidae, also known by the common name door snails, is a Taxonomy (biology), taxonomic family (biology), family of small, very elongate, mostly Gastropod shell#Chirality in gastropods, left-handed, air-breathing land snails, sinistral terre ...
** † Family Anadromidae ** † Family Filholiidae ** † Family Palaeostoidae *Superfamily Orthalicoidea **Family Orthalicidae **Family Cerionidae **Family Coelociontidae ** † Family Grangerellidae **Family Megaspiridae **Family Placostylidae **Family Urocoptidae *Superfamily Achatinoidea **Family
Achatinidae Achatinidae (New Latin, from Greek "''agate''") is a family of medium to large sized tropical land snails, terrestrial pulmonate gastropod mollusks from Africa. Well known species include '' Achatina achatina'' the Giant African Snail, and '' L ...
**Family
Ferussaciidae Ferussaciidae is a family of air-breathing land snails, terrestrial molluscs, terrestrial pulmonate gastropod mollusks in the Taxonomic rank, superfamily Achatinoidea.MolluscaBase eds. (2020). MolluscaBase. Ferussaciidae Bourguignat, 1883. Access ...
**Family Micractaeonidae **Family Subulinidae *Superfamily Aillyoidea **Family Aillyidae *Superfamily Testacelloidea **Family Testacellidae **Family Oleacinidae **Family Spiraxidae *Superfamily Papillodermatoidea **Family Papillodermatidae *Superfamily Streptaxoidea **Family
Streptaxidae Streptaxidae is a family of carnivorous air-breathing land snails, terrestrial pulmonate gastropod mollusks in the clade Stylommatophora. Streptaxidae are monophyletic while all subfamilies and several genera are polyphyletic. Six Streptaxi ...
*Superfamily Rhytidoidea **Family Rhytididae **Family
Chlamydephoridae ''Chlamydephorus'' is a genus of air-breathing land slugs, terrestrial pulmonate gastropod mollusks in the family Chlamydephoridae. It is the only genus within the family Chlamydephoridae.Herbert D. G. (1997). "The terrestrial slugs of KwaZul ...
**Family Haplotrematidae **Family Scolodontidae *Superfamily Acavoidea **Family Acavidae **Family Caryodidae **Family Dorcasiidae **Family Macrocyclidae **Family Megomphicidae **Family
Strophocheilidae Strophocheilidae is a Taxonomy (biology), taxonomic family (biology), family of air-breathing land snails, terrestrial molluscs, terrestrial pulmonate gastropod mollusks in the superfamily Acavoidea (according to the Taxonomy of the Gastropoda (B ...
*Superfamily Punctoidea **Family Punctidae ** † Family Anostomopsidae **Family
Charopidae Charopidae is a Taxonomy (biology), taxonomic family (biology), family of small air-breathing land snails (and semi-slugs such as ''Otoconcha dimidiata''), terrestrial molluscs, terrestrial pulmonate gastropod mollusks in the superfamily Punctoi ...
**Family Cystopeltidae **Family Discidae **Family Endodontidae **Family Helicodiscidae **Family Oreohelicidae **Family Thyrophorellidae *Superfamily Sagdoidea **Family Sagdidae


= limacoid clade

= *Superfamily Staffordioidea **Family Staffordiidae *Superfamily Dyakioidea **Family Dyakiidae *Superfamily Gastrodontoidea **Family Gastrodontidae **Family Chronidae **Family Euconulidae **Family
Oxychilidae Oxychilidae is a taxonomic family of air-breathing land snails, terrestrial pulmonate gastropod mollusks in the superfamily Gastrodontoidea. Distribution The distribution of Oxychilidae includes the Nearctic, western-Palearctic, eastern-Pale ...
**Family Pristilomatidae **Family Trochomorphidae **Fossil taxa probably belonging to the Gastrodontoidea ***Subfamily † Archaeozonitinae ***Subfamily † Grandipatulinae ***Subfamily † Palaeoxestininae *Superfamily Parmacelloidea **Family Parmacellidae **Family
Milacidae Milacidae is a family (biology), family of air-breathing, keel (slug), keeled, land slugs. These are shell-less terrestrial molluscs, terrestrial gastropod mollusks in the superfamily Parmacelloidea. This family has no subfamilies (according to ...
**Family Trigonochlamydidae *Superfamily Zonitoidea **Family Zonitidae *Superfamily Helicarionoidea **Family Helicarionidae **Family Ariophantidae **Family Urocyclidae *Superfamily Limacoidea **Family Limacidae **Family
Agriolimacidae ''Agriolimacidae'' is a family of small and medium-sized land slugs, or shell-less snails, terrestrial pulmonate gastropod mollusks. Distribution Distribution of Limacidae is Holarctic, this include: Nearctic, western Palearctic and eastern P ...
**Family Boettgerillidae **Family Vitrinidae


= other Sigmurethra

= Two superfamilies belongs to clade Sigmurethra, but they are not in the limacoid clade. *Superfamily Arionoidea **Family Arionidae **Family Anadenidae **Family Ariolimacidae **Family Binneyidae **Family Oopeltidae **Family Philomycidae *Superfamily
Helicoidea Helicoidea is a Taxonomy (biology), taxonomic Taxonomic rank, superfamily of air-breathing land snails, terrestrial molluscs, terrestrial pulmonate gastropod mollusks in the order (taxonomy), order Stylommatophora. Taxonomy 2017 taxonomy and l ...
**Family
Helicidae Helicidae is a large, diverse family of western Palaearctic, medium to large-sized, air-breathing land snails, sometimes called the "typical snails." It includes some of the largest European land snails, several species are common in anthropoge ...
**Family Bradybaenidae **Family
Camaenidae Camaenidae is a Family (biology), family of air-breathing land snails, terrestrial molluscs, terrestrial pulmonate gastropod mollusks in the superfamily Helicoidea, the typical snails and their allies. This is one of the most diverse families i ...
**Family Cepolidae **Family Cochlicellidae **Family Elonidae **Family Epiphragmophoridae **Family Halolimnohelicidae **Family Helicodontidae **Family Helminthoglyptidae **Family Humboldtianidae **Family
Hygromiidae Hygromiidae is a taxonomic family of small to medium-sized air-breathing land snails, terrestrial pulmonate gastropod mollusks in the superfamily Helicoidea.Helicellidae Ihering, 1909 Anatomy Some snails in genera within this family create and ...
**Family Monadeniidae **Family Pleurodontidae **Family Polygyridae **Family Sphincterochilidae **Family Thysanophoridae **Family Trissexodontidae **Family Xanthonychidae


2010 taxonomy

Jörger ''et al.'' (2010) analyzed major groups within the
Heterobranchia Heterobranchia, the heterobranchs (meaning "different gill"), is a taxonomic clade of snails and slugs, which includes marine, aquatic, and terrestrial gastropod molluscs. Heterobranchia is one of the main clades of gastropods. Currently Het ...
using genetic data and found that Pulmonata as traditionally defined was
polyphyletic A polyphyletic group is an assemblage that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
, for instance some pulmonates were more closely related to
Sacoglossa Sacoglossa are a superorder of small sea slugs and sea snails, marine gastropod mollusks that belong to the clade Heterobranchia known as sacoglossans. There are 284 valid species recognized within this superorder. Sacoglossans live by ingestin ...
and Acochlidia. They proposed the more inclusive taxon
Panpulmonata Panpulmonata is a taxonomic clade of snails and slugs in the clade Heterobranchia within the clade Euthyneura. Panpulmonata was established as a new taxon by Jörger et al. in October 2010. The older name "Pulmonata" referred to a group of euth ...
to unite the clades Siphonarioidea,
Sacoglossa Sacoglossa are a superorder of small sea slugs and sea snails, marine gastropod mollusks that belong to the clade Heterobranchia known as sacoglossans. There are 284 valid species recognized within this superorder. Sacoglossans live by ingestin ...
, Glacidorboidea, Pyramidelloidea, Amphiboloidea, Hygrophila, Acochlidia and
Eupulmonata Eupulmonata is a taxonomic clade of air-breathing gastropod molluscs. The great majority of this group are land snails and slugs, but some are intertidal or inhabit coastal saltmarshes and mangroves. Possible synapomorphy of the group are gl ...
.


References


Further reading

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External links


Pulmonate snails & slugs in captivity
{{Authority control Obsolete gastropod taxa Carboniferous first appearances Taxa named by Georges Cuvier