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