Tornatina planospira, copyright G. & Ph. Poppe.

Belongs within: Heterobranchia.
Contains: Thecosomata, Gymnosomata, Runcinidae, Aplysiidae, Umbraculidae, Cylichnidae, Diaphanidae, Atys, Haminoea, Retusidae, Rhizoridae, Bulla, Acteocina, Philinidae, Aglajidae, Philinoglossidae.

The Euopisthobranchia are a major clade of marine gastropods supported by molecular data, including the bubble shells, sea hares and pteropods. Subgroups include the Aplysiida, sea hares and related taxa, a group of soft-bodied marine gastropods with a reduced shell (external in Akeridae, internal in Aplysiidae) characterised by the presence of defensive opaline and purple glands in the mantle cavity, and an oesophageal gizzard containing an anterior chamber with large chitinous plates and a posterior chamber with numerous fine spines. The Pteropoda are planktonic gastropods that swim by means of broad lateral expansions of the foot (parapodia). The Cephalaspidea, bubble snails and headshield slugs, are characterised by a distinctive broadening of the head to form a shield used in digging through sand. The shield bears a central pair of light-sensitive organs, and the foot may bear lateral parapodia. The Umbraculoidea, umbrella shells, have a thin limpet-like shell and plume-like gills on the left side of the body. The Runcinoidea are minute, flattened slugs (a minute external shell may be present posteriorly) with an undivided notum covering most of the dorsum.

<==Euopisthobranchia BR17
    |--Pteropoda BR17
    |    |  i. s.: Hyalea cornea G20
    |    |--Thecosomata BR17
    |    `--Gymnosomata BR17
    |--Runcinoidea [Runcinacea, Runcinida, Runcinidea, Runciniformes] BR17
    |    |--Runcinidae BR05
    |    `--Ilbia Burn 1963 [Ilbiidae, Ilbiinae] BR05
    |         `--*I. ilbi Burn 1963 BR17
    |--Aplysiida [Anaspidea, Aplysiacea, Aplysiina, Aplysiomorpha] BR17
    |    |--Aplysiidae BR05
    |    `--Akeridae [Akerinae, Akeroidea] BR05
    |         |--Clistaxis [=Cryptaxis (preoc.)] O27
    |         |    |--*C. parvula O27
    |         |    |--C. crebripunctatus O27
    |         |    `--C. polystrigma Dall 1908 O27
    |         `--Akera Müller 1776 BR17
    |              |--*A. bullata Müller 1776 BR17
    |              |--A. carnosa G20
    |              |--A. mediojurensis Cossmann 1896 TTE93
    |              `--A. tanganyicensis Cox 1965 TTE93
    |--Umbraculoidea [Tylodinoidea, Umbraculacea, Umbraculida, Umbraculiformes, Umbraculoidei, Umbraculomorpha] BR17
    |    |--Tylodina Rafinesque 1814 [Tylodinacea, Tylodinana, Tylodinidae, Tylodininae, Tylodinoidei] BR05
    |    |    |--*T. punctulata Rafinesque 1814 BR17 [incl. T. citrina O27]
    |    |    |--T. corticalis (Tate 1889) MG-H11
    |    |    |--T. duebeni N79
    |    |    |--T. fungina Gabb 1865 O27
    |    |    |--‘Parmophorus’ patelloideus S79
    |    |    `--T. (Tylodinella) rafinesquei Philippi 1844 TTE93
    |    `--Umbraculidae TTE93
    `--Cephalaspidea [Aspidocephala, Bulliformes, Bulliformii, Bulloidei, Bullomorpha, Ectoconcha] BR17
         |  i. s.: Priscaphander JB12
         |--Newnesia Smith 1902 [Newnesiidae, Newnesioidea] BR17
         |    `--*N. antarctica Smith 1902 BR17
         |--Cylichnoidea [Diaphanacea, Diaphanida, Diaphanoidea] BR17
         |    |--Cylichnidae BR05
         |    |--Diaphanidae BR17
         |    |--Colinatys Ortea, Moro & Espinosa 2013 [Colinatydidae] BR17
         |    |    `--*C. alayoi (Espinosa & Ortea 2004) [=Atys alayoi] BR17
         |    |--Diaphanella Thiele 1912 BR17 [=Notodiaphana Thiele 1917 BR17; Notodiaphanidae BR05]
         |    |    `--*D. fragilis (Vélain 1877) [=Bulla fragilis, *Notodiaphana fragilis] BR17
         |    |--Eoscaphander Habe 1952 [Eoscaphandridae] BR17
         |    |    `--*E. fragilis Habe 1952 BR17
         |    `--Mnestia Adams & Adams 1854 [Mnestiidae] BR17
         |         `--*M. marmorata (Adams 1850) [=Bulla marmorata] BR17
         |--Haminoeidae [Atyacea, Hamineina, Haminoeoidea] BR05
         |    |  i. s.: Liloa curta (Adams 1850) MG-H11
         |    |         ‘Cylichna’ incisa Stephenson 1941 TTE93
         |    |         Aliculastrum Pilsbry 1896 TTE93
         |    |--Atys BR17
         |    |--Haminoea BR17
         |    |--Bullacta Bergh 1901 BR05 [Bullactidae, Bullactinae BR17, Bullactininae]
         |    |    |--*B. caurina (Benson 1856) [=Bullaea caurina] BR17
         |    |    `--B. exarata ZLK11
         |    `--Smaragdinellinae [Cryptophthalmidae, Cryptophthalminae, Ophthalmidae, Smaragdinellidae] BR17
         |         |--Smaragdinella Adams 1848 BR05
         |         |    `--*S. viridis (Rang in Quoy & Gaimard 1832) [=Bulla viridis] BR17
         |         `--Lathophthalmus Pruvot-Fol 1932 (see below for synonymy) BR05
         |              `--*L. olivaceus (Ehrenberg 1828) [=*Cryptophthalmus olivaceus] BR17
         |--Bulloidea BR17
         |    |--Retusidae BR17
         |    |--Rhizoridae BR17
         |    |--Bullidae [Bullacea] BR05
         |    |    |--Bulla BR05
         |    |    `--Quibulla Iredale 1929 P61, BR17
         |    |         |--*Q. botanica (Hedley 1918) BR17 [=Bulla botanica P61, Bullaria botanica F27a]
         |    |         `--Q. quoyi (Gray 1843) P61
         |    `--Tornatinidae BR17
         |         |--Acteocina BR17
         |         `--Tornatina Adams 1850 BR05
         |              |--‘Retusa’ gaimardi Finlay 1927 F27b (see below for synonymy)
         |              |--T. acrobeles (Watson 1883) [=Utriculus acrobeles] H09
         |              |--T. biplex (Adams 1850) [=Bulla biplex] H09
         |              |--T. carinata C64
         |              |--T. cerealis C64
         |              |--T. culcitella C64
         |              |--T. gracilis Adams 1850 H09
         |              |--T. leptekes (Watson 1883) [=Utriculus leptekes] H09
         |              `--T. planospira (Adams 1850) [=Bulla planospira] H09
         `--Philinoidea [Entoconcha, Philinacea] BR05
              |--Philinidae BR05
              |--Aglajidae BR05
              |--Philinoglossidae BR05
              |--Alacuppa Oskars, Bouchet & Malaquias 2015 [Alacuppidae] BR17
              |    `--*A. supracancellata (Schepman 1913) [=Atys supracancellata] BR17
              |--Colpodaspis Sars 1870 [Colpodaspididae] BR17
              |    `--*C. pusilla Sars 1870 BR17
              |--Laona Adams 1865 BR05 [Laonidae BR17, Laoninae]
              |    `--*L. zonata Adams 1865 BR17
              |--Philinorbis Habe 1950 [Philinorbidae] BR17
              |    `--*P. teramachii Habe 1950 BR17
              |--Gastropteron Kosse 1813 (see below for synonymy) BR05
              |    |--*G. meckeli Blainville 1825 BR17
              |    |--G. cinereum Dall 1925 [=G. pacificum var. cinereum] O27
              |    |--G. coccineum G20
              |    |--G. pacificum Bergh 1893 O27
              |    `--G. rubrum O27
              `--Scaphander Montfort 1810 BR05 (see below for synonymy)
                   |--*S. lignarius (Linnaeus 1758) P61, L58 [=Bulla lignaria P61; incl. S. lignarius var. curta N79]
                   |--*Tricla’ gioeni Philipsson 1788 BR17
                   |--S. librarius J64
                   |--S. multistriatus Brazier 1877 H09
                   |--S. otagoensis Dell 1956 P61
                   |--S. punctostriatus (Mighels & Adams 1841) [=Bulla punctostriatus] N79
                   `--S. willetti Dall 1919 O27

Gastropteron Kosse 1813 [=Gasteropteron (l. c.), Gastropterum Agassiz 1847; Gasteropteridae, Gastropteridae, Gastropterinae, Gastropteroidae] BR05

Lathophthalmus Pruvot-Fol 1932 [=Cryptophthalmus Ehrenberg 1828 non Rafinesque 1814; Lathophthalminae] BR05

‘Retusa’ gaimardi Finlay 1927 F27b [=Bulla voluta Quoy & Gaimard 1833 non Gmelin 1791 F27b, Acteocina voluta O27, *Tornatina voluta BR17]

Scaphander Montfort 1810 BR05 [incl. Gioenia D00, Tricla Philipsson 1788 (nom. rej.) D00, BR05; Scaphandridae BR17, Scaphandroidea, Triclidae]

*Type species of generic name indicated


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

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[D00] Draparnaud, C. 1800. Observations sur la Gioenia. Bulletin des Sciences, par la Societé Philomathique de Paris 2 (39): 113–114.

[F27a] Finlay, H. J. 1927a. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[F27b] Finlay, H. J. 1927b. New specific names for austral Mollusca. Transactions and Proceedings of the New Zealand Institute 57: 488–533.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

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

[J64] Jeffreys, J. G. 1864. Report of the committee appointed for exploring the coasts of Shetland by means of the dredge. Report of the British Association for the Advancement of Science 33: 70–81.

[JB12] Johnson, M. E., & B. G. Baarli. 2012. Development of intertidal biotas through Phanerozoic time. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 63–128. Springer.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[N79] Norman, A. M. 1879. The Mollusca of the fiords near Bergen, Norway. Journal of Conchology 2: 8–77.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 1. Stanford University Press: Stanford University (California).

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.

[S79] Smith, E. A. 1879. Notes on the species of the genus Scutus. Journal of Conchology 2: 252–264.

[TTE93] Tracey, S., J. A. Todd & D. H. Erwin. 1993. Mollusca: Gastropoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 131–167. Chapman & Hall: London.

[ZLK11] Zou, S., Q. Li & L. Kong. 2011. Additional gene data and increased sampling give new insights into the phylogenetic relationships of Neogastropoda, within the caenogastropod phylogenetic framework. Molecular Phylogenetics and Evolution 61: 425–435.

Last updated: 5 September 2021.

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