Pulmonata

Pulmonata; Temporal range: Carboniferous–recent PreꞒ Ꞓ O S D C P T J K Pg N
	Various examples of Pulmonata
Scientific classification
Kingdom:	Animalia
Phylum:	Mollusca
Class:	Gastropoda
Subclass:	Heterobranchia
Informal group:	Pulmonata; Cuvier, 1814
Taxonomic subdivisions
	informal group Basommatophora; clade Eupulmonata;

Pulmonata, or "pulmonates", is an informal group (previously an order, and before that a subclass) 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.

The taxon Pulmonata as traditionally defined was found to be polyphyletic in a molecular study per Jörger et al., dating from 2010.^[1]

Pulmonata are known from the Carboniferous Period to the present.^[2]

Pulmonates have a single atrium and kidney, and a concentrated, symmetrical, nervous system. The mantle cavity is located 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 darts.^[3]

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 Pilsbry, 1948
- - Superfamily Onchidioidea Rafinesque, 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, 1815
  - Superfamily Planorboidea Rafinesque, 1815
  - Superfamily Siphonarioidea J.E. Gray, 1840
Suborder Eupulmonata Haszprunar & Huber, 1990
- Infraorder Acteophila Dall, 1885 = formerly Archaeopulmonata
  - Superfamily Melampoidea Stimpson, 1851
- Infraorder Trimusculiformes Minichev & Starobogatov, 1975
  - Superfamily Trimusculoidea Zilch, 1959

Shells of pulmonate stylommatophoran snails in a museum collection

An artistic but scientifically incorrect version of various European land snails and slugs (one species here is not a pulmonate), their food plants and fungi, and a beetle that eats mollusks, bottom right.

- Infraorder Stylommatophora A. Schmidt, 1856 - land snails
  - Subinfraorder Orthurethra
  - Superfamily Achatinelloidea Gulick, 1873
  - Superfamily Cochlicopoidea Pilsbry, 1900
  - Superfamily Partuloidea Pilsbry, 1900
  - Superfamily Pupilloidea Turton, 1831
  - Subinfraorder Sigmurethra
  - 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 Rafinesque, 1815
  - Superfamily Helixarionoidea Bourguignat, 1877
  - Superfamily Limacoidea Rafinesque, 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

Examples of Pulmonata: Achatina fulica top right, Bielzia coerulans top left, Praticolella berlandieriana center right, Megalobulimus oblongus in the center, Euglandina rosea center left, Helix pomatia bottom right & Ashmunella levettei bottom left

The taxonomy of this group according to the taxonomy of the Gastropoda (Bouchet & Rocroi, 2005)^[4] 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
- † Family Acroreiidae

Clade Hygrophila

Superfamily Chilinoidea
- Family Chilinidae
- Family Latiidae
Superfamily Acroloxoidea
- Family Acroloxidae
Superfamily Lymnaeoidea
- Family Lymnaeidae
Superfamily Planorboidea
- Family Planorbidae
- Family Physidae

Clade Eupulmonata

Contains the clades Systellommatophora and Stylommatophora

Superfamily Trimusculoidea
- Family Trimusculidae
Superfamily Otinoidea
- Family Otinidae
- Family Smeagolidae
Superfamily Ellobioidea
- Family Ellobiidae

Clade Systellommatophora ( = Gymnomorpha)

Superfamily Onchidioidea
- Family Onchidiidae
Superfamily Veronicelloidea
- Family Veronicellidae
- 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
- Family Draparnaudiidae
Superfamily Achatinelloidea
- Family Achatinellidae
Superfamily Cochlicopoidea
- Family Cochlicopidae
- Family Amastridae
Superfamily Pupilloidea
- Family Pupillidae
- Family Argnidae
- Family Chondrinidae
- † Family Cylindrellinidae
- Family Lauriidae
- Family Orculidae
- Family Pleurodiscidae
- Family Pyramidulidae
- Family Spelaeodiscidae
- Family Strobilopsidae
- Family Valloniidae
- Family Vertiginidae
Superfamily Enoidea
- Family Enidae
- Family Cerastidae

Informal Group Sigmurethra

Superfamily Clausilioidea
- Family Clausiliidae
- † 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
- Family Ferussaciidae
- Family Micractaeonidae
- Family Subulinidae
Superfamily Aillyoidea
- Family Aillyidae
Superfamily Testacelloidea
- Family Testacellidae
- Family Oleacinidae
- Family Spiraxidae
Superfamily Papillodermatoidea
- Family Papillodermatidae
Superfamily Streptaxoidea
- Family Streptaxidae
Superfamily Rhytidoidea
- Family Rhytididae
- Family Chlamydephoridae
- Family Haplotrematidae
- Family Scolodontidae
Superfamily Acavoidea
- Family Acavidae
- Family Caryodidae
- Family Dorcasiidae
- Family Macrocyclidae
- Family Megomphicidae
- Family Strophocheilidae
Superfamily Punctoidea
- Family Punctidae
- † Family Anastomopsidae
- Family Charopidae
- 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
- Family Pristilomatidae
- Family Trochomorphidae
- Fossil taxa probably belonging to the Gastrodontoidea
  - Subfamily † Archaeozonitinae
  - Subfamily † Grandipatulinae
  - Subfamily † Palaeoxestininae
Superfamily Parmacelloidea
- Family Parmacellidae
- Family Milacidae
- Family Trigonochlamydidae
Superfamily Zonitoidea
- Family Zonitidae
Superfamily Helicarionoidea
- Family Helicarionidae
- Family Ariophantidae
- Family Urocyclidae
Superfamily Limacoidea
- Family Limacidae
- Family Agriolimacidae
- 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
- Family Helicidae
- Family Bradybaenidae
- Family Camaenidae
- Family Cepolidae
- Family Cochlicellidae
- Family Elonidae
- Family Epiphragmophoridae
- Family Halolimnohelicidae
- Family Helicodontidae
- Family Helminthoglyptidae
- Family Humboldtianidae
- Family Hygromiidae
- Family Monadeniidae
- Family Pleurodontidae
- Family Polygyridae
- Family Sphincterochilidae
- Family Thysanophoridae
- Family Trissexodontidae
- Family Xanthonychidae

2010 taxonomy

Jörger et al. (2010)^[1] analyzed major groups within the Heterobranchia using genetic data and found that Pulmonata as traditionally defined was polyphyletic, for instance some pulmonates were more closely related to Sacoglossa and Acochlidia. They proposed the more inclusive taxon Panpulmonata to unite the clades Siphonarioidea, Sacoglossa, Glacidorboidea, Pyramidelloidea, Amphiboloidea, Hygrophila, Acochlidia and Eupulmonata.^[1]

References

^ ^a ^b ^c Jörger, Katharina M; Stöger, Isabella; Kano, Yasunori; Fukuda, Hiroshi; Knebelsberger, Thomas; Schrödl, Michael (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". BMC Evolutionary Biology. 10 (1): 323. doi:10.1186/1471-2148-10-323. PMC 3087543. PMID 20973994.
^ (in Czech) Pek I., Vašíček Z., Roček Z., Hajn. V. & Mikuláš R. 1996. Základy zoopaleontologie. Olomouc, 264 pp., ISBN 80-7067-599-3.
^ Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. p. 377. ISBN 0-03-056747-5.
^ Bouchet, Philippe; Rocroi, Jean-Pierre; Frýda, Jiri; Hausdorf, Bernard; Ponder, Winston; Valdés, Ángel & Warén, Anders (2005). "Classification and nomenclator of gastropod families". Malacologia. Hackenheim, Germany: ConchBooks. 47 (1–2): 1–397. ISBN 3-925919-72-4. ISSN 0076-2997.

External links

Pulmonate snails & slugs in captivity

[Jörger_2010-1] Jörger, Katharina M; Stöger, Isabella; Kano, Yasunori; Fukuda, Hiroshi; Knebelsberger, Thomas; Schrödl, Michael (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". BMC Evolutionary Biology. 10 (1): 323. doi:10.1186/1471-2148-10-323. PMC 3087543. PMID 20973994.

[2] (in Czech) Pek I., Vašíček Z., Roček Z., Hajn. V. & Mikuláš R. 1996. Základy zoopaleontologie. Olomouc, 264 pp., ISBN 80-7067-599-3.

[IZ-3] Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. p. 377. ISBN 0-03-056747-5.

[4] Bouchet, Philippe; Rocroi, Jean-Pierre; Frýda, Jiri; Hausdorf, Bernard; Ponder, Winston; Valdés, Ángel & Warén, Anders (2005). "Classification and nomenclator of gastropod families". Malacologia. Hackenheim, Germany: ConchBooks. 47 (1–2): 1–397. ISBN 3-925919-72-4. ISSN 0076-2997.

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