Judas' ear shell Ellobium aurisjudae, copyright Ron Yeo.

Belongs within: Heterobranchia.
Contains: Siphonarioidea, Hygrophila, Systellommatophora, Stylommatophora, Melampinae, Pythiinae.

The Pulmonata are a group of gastropods characterised by modification of the mantle cavity into a vascularised 'pallial lung' allowing direct gas exchange with the air, and the concommitant loss of the gills. As such, the group includes the majority of terrestrial gastropods, forming the clade Eupulmonata. It also includes a number of primarily aquatic lineages commonly grouped as the Basommatophora, though this latter group is of uncertain monophyly.

Pulmonata [Adelopneumona, Aeropneusta, Archaeopulmonata, Limnophila, Limnophilidae, Petrophila, Pulmonifera]
    |  i. s.: Promenetus B97
    |           |--P. exacuous (Say 1821) P88
    |           `--P. umbilicatellus (Cockerell 1887) P88
    |--Basommatophora BR05
    |    |--Siphonarioidea BR05
    |    |--Hygrophila BR05
    |    `--Amphibolidae [Amphibolacea, Amphibolida, Amphiboliformes, Amphiboliformii, Amphiboloidea] BR05
    |         |--Ampullacera Quoy & Gaimard 1832 [Ampullacerae, Ampullaceridae] BR05
    |         |--Amphibola Schumacher 1817 BR05
    |         |    `--A. crenata O81
    |         `--Salinator Hedley 1900 [Salinatoridae] BR05
    |              |--S. fragilis (Lamarck 1822) [=Ampullaria fragilis] H09
    |              `--S. solida GO06
    `--Eupulmonata [Acteophila, Actophila, Akteophila] BR05
         |--Systellommatophora BR05
         |--Stylommatophora BR05
         |--Otinoidea BR05
         |    |--Otina Gray 1847 [Otinidae, Otininae] BR05
         |    `--Smeagol Climo 1980 [Smeagolidae, Smeagolina] BR05
         |         `--S. manneringi Climo 1980 APS-P86
         |--Trimusculidae [Trimusculoidea] BR05
         |    |--Trimusculus Schmidt 1818 [Trimusculinae] BR05
         |    `--Gadinia Gray 1824 BR05 (see below for synonymy)
         |         |--*G. peruviana O27
         |         |--G. nivea Hutton 1878 [incl. Hipponyx hexagonus] F27
         |         `--G. reticulata (Sowerby 1835) [=Mouretia reticulata] O27
         `--Ellobiidae [Auriculoidea, Ellobiacea, Ellobioidea] BR05
              |  i. s.: Marinula M54
              |           |--M. filholi HS01
              |           |--M. patula (Lowe 1835) [=Melampus patulus] H09
              |           |--M. striata Odhner 1924 [incl. Cremnobates parva Swainson 1855] F27
              |           `--M. xanthostoma M54
              |         Phytia Gray 1821 O27
              |           |--*P. denticulata [=Alexia denticulata] O27
              |           `--P. setifer Cooper 1872 O27
              |         Allochroa layardi (Adams & Adams 1855) T13
              |--Melampinae BR05
              |--Pythiinae W97
              |--Zaptychius Walcott 1883 [Zaptychiinae] BR05
              |    `--Z. quanshengxigouensis ZZ88
              |--Pedipedinae W97
              |    |--Microtralia alba (Gassies 1865) [=Ellobium (Auriculodes) album] W97
              |    `--Pedipes Férussac 1821 BR05
              |         |--*P. afer O27
              |         |--P. liratus Binney 1860 O27
              |         `--P. unisulcatus Cooper 1866 O27
              |--Carychiinae [Carychiadae, Carychiidae] BR05
              |    |--Zospeum Bourguignat 1856 [Zospeidae] BR05
              |    |    `--Z. exiguum (Kuščer 1932) BV86
              |    `--Carychium Müller 1773 BR05
              |         |--C. exile BP90
              |         |    |--C. e. exile BP90
              |         |    `--C. e. canadense BP90
              |         |--C. floridanum BP90
              |         |--C. nannodes BP90
              |         `--C. stygium BP90
              `--Ellobiinae W97
                   |--Auricula Lamarck 1799 [Amphibiae, Auriculacea, Auriculea, Auriculidae, Auriculinae] BR05
                   |--Leucophytia Winckworth 1949 [Leucophytiidae] BR05
                   `--Ellobium Röding 1798 W97
                        |--*E. aurisjudae (Linnaeus 1758) [=Bulla aurisjudae] W97
                        |--E. aurismidae Linnaeus 1767 [=Bulla aurismidae] W97
                        |--E. chinense (Pfeiffer 1855) W97
                        |--E. helvaceum (Pfeiffer 1876) [=Auriculus helvaceus] H09
                        |--E. polita (Metcalfe 1851) W97
                        `--E. semisculptum Adams & Adams 1854 [=E. (Auriculodes) semisculptum] W97

Gadinia Gray 1824 BR05 [incl. Mouretia Sowerby 1834 O27, Rowellia Cooper 1865 O27; Gadiniadae, Gadiniidae, Gadiniinae, Gadinioidea]

*Type species of generic name indicated


[APS-P86] Arnaud, P. M., Cl. Poizat & L. v. Salvini-Plawen. 1986. Marine-interstitial Gastropoda (including one freshwater interstitial species).  In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 153–176. E. J. Brill/Dr. W. Backhuys: Leiden.

[BV86] Bole, J., & F. Velkovrh. 1986. Mollusca from continental subterranean aquatic habitats. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 177–208. E. J. Brill/Dr. W. Backhuys: Leiden.

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[B97] Brown, K. M. 1997. Temporal and spatial patterns of abundance in the gastropod assemblage of a macrophyte bed. American Malacological Bulletin 14 (1): 27–33.

[BP90] Burch, J. B., & T. A. Pearce. 1990. Terrestrial Gastropoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 201–309. John Wiley & Sones: New York.

[F27] Finlay, H. J. 1927. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[GO06] Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrödl & M. K. Nishiguchi. 2006. Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons. Proceedings of the National Academy of Sciences of the USA 103 (20): 7723–7728.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[M54] Macpherson, J. H. 1954. Molluscs (sea shells and snails). In: Willis, J. M. (ed.) The Archipelago of the Recherche pp. 55–63. Australian Geographical Society: Melbourne.

[O81] O’Brien, C. O. 1981. A. A. Book of New Zealand Wildlife: A guide to the native and introduced animals of New Zealand. Lansdowne Press: Auckland.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. II pt I. Stanford University Press: Stanford University (California).

[P88] Pip, E. 1988. Niche congruency of freshwater gastropods in central North America with respect to six water chemistry parameters. Nautilus 102 (2): 65–72.

[T13] Taylor, C. K. 2013. Annotated bibliography for Barrow Island terrestrial invertebrates. Records of the Western Australian Museum Supplement 83: 135–144.

[W97] Wells, F. E. 1997. A review of the northern Australian species of the genera Cassidula and Ellobium (Gastropoda: Ellobiidae). In: Hanley, J. R., G. Caswell et al. (eds) Proceedings of the Sixth International Marine Biological Workshop. The marine flora and fauna of Darwin Harbour, Northern Territory, Australia pp. 213–229. Museums and Art Galleries of the Northern Territory and the Australian Marine Sciences Association: Darwin.

[ZZ88] Zhu G.-X. & Zhou Y.-Z. 1988. Fresh-water molluscan fauna across Jurassic-Cretaceous boundary in southeast Jilin. Acta Palaeontologica Sinica 27 (6): 713–721.

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