Heterobranchia

Cornirostra pellucida, copyright Des Beechey.

Belongs within: Gastropoda.
Contains: Cimidae, Nerinellidae, Ceritellidae, Ptygmatididae, Nerineidae, Mathildidae, Khairkhaniidae, Acteonellidae, Donaldina, Streptacididae, Valvatidae, Architectonicoidea, Prionoglossa, Ringipleura, Acteonimorpha, Euopisthobranchia, Sacoglossa, Siphonarioidea, Pylopulmonata, Acochlidimorpha, Hygrophila, Ellobioidea, Systellommatophora, Achatinina, Scolodontidae, Plectopyloidea, Punctoidea, Urocoptoidea, Rhytidoidea, Orthalicoidea, Clausilioidea, Arionoidea, Oleacinoidei, Succineoidei, Limacoidei, Helicoidei, Pupilloidea.

The Heterobranchia are a major clade of gastropods characterised by a reduction of the gill ctenidium. They also lack bolster cartilages in the radula or ciliated structures in the anterior oesophagus, and sperm cells are of the eupyrene type only (Salvini-Plawen & Steiner 1996). Many also exhibit a heterostrophic development with the whorls of the protoconch coiled in the opposite direction to those of the teleoconch. Basal members (outside the subsidiary clade Euthyneura) are mostly small, aquatic shells that were historically included in the grade 'Mesogastropoda' along with members of the Heterobranchia's sister clade, Caenogastropoda. Examples include the Valvatoidea, minute marine and freshwater hermaphroditic snails with a secondary gill, pallial tentacle, and orthostrophic (not heterostrophic) development (Salvini-Plawen & Steiner 1996). The Mathildoidea have an elongate shell with many whorls, with a teleoconch ornament of spiral and axial elements. Shells of Murchisonellidae are similarly elongate but fragile, and the animal lacks a radula. Fossil heterobranchs include the Nerineoidea of the Jurassic and Cretaceous, which had large, turriculate to oviform shells that often bore internal protrusions of the walls within the whorls.

In the Euthyneura, the ctenidium was entirely replaced by a folded gill structure called a plicatidium (further modified or replaced in more derived subgroups); it has been debated whether the plicatidium represents an evolved derivative of the ctenidium or an entirely novel structure. The euthyneurans were historically divided between the air-breathing Pulmonata and the marine Opisthobranchia but recent phylogenetic analyses have indicated that the latter grouping is paraphyletic and the former is polyphyletic. Instead, molecular studies support the exclusion of the Acteonimorpha and Ringipleura (nudibranch sea slugs and related taxa) from a clade Tectipleura uniting the remaining opisthobranchs (bubble shells, sea hares, etc.) with the 'pulmonates'. The majority of pulmonate taxa remain within a clade Eupulmonata, characterised by a contractile pneumostome and free head tentacles (Salvini-Plawen & Steiner 1996) within which the truly terrestrial lineages form the clade Geophila. The largest terrestrial lineage is the Stylommatophora, characterised by the possession of four retractable cephalic tentacles with eyes on the tips of the posterior tentacles. Molecular data supports the division of the Stylommatophora between three basal clades, treated by Bouchet et al. (2017) as the Achatinina, Scolodontidae and Helicina.

See also: I's been ejucated, now I can haz snails pleez? Kthnx.

<==Heterobranchia (see below for synonymy)
    |--Valvatoidea [Valvatoideae, Valvatoidei] S-PS96
    |    |  i. s.: Palaeocarboninia Bandel & Heidelberger 2002 B02
    |    |           `--P. jankei Bandel & Heidelberger 2002 FB04
    |    |         Carboninia Bandel 1996 B02
    |    |         Bandellina Schröder 1995 B02
    |    |--Valvatidae BR05
    |    |--Hyalogyrina Marshall 1988 [Hyalogryinidae, Hyalogyrinidae] BR05
    |    |    `--*H. glabra Marshall 1988 BR17
    |    |--Provalvata Bandel 1991 [Provalvatidae] BR05
    |    |    |--‘Valvata (Cincinna)’ helicelloides Huckreide 1967 (see below for synonymy) B91
    |    |    `--P. sabaudiensis (Maillard 1884) [=Valvata sabaudiensis; incl. V. (Cincinna) proavia Huckriede 1967] B91
    |    `--Cornirostra Ponder 1990 [Cornirostridae] BR05
    |         |--*C. pellucida (Laseron 1954) [=Microdiscula pellucida] BR17
    |         `--C. floridana Bieler & Mikkelsen in Bieler, Ball & Mikkelsen 1998 BC01
    `--+--+--Architectonicoidea BR05
       |  `--Prionoglossa S11
       `--Euthyneura (see below for synonymy) S11
            |  i. s.: Pneumodermopsis (Pneumodermopsis) paucidens (Boas 1866) H01
            |         Oscanius tuberculatus M62
            |         Aplysiella virescens PP64
            |         Tjaernoeia Warén & Bouchet 1988 BR05 [Tjaernoeidae, Tjaernoeiidae BR17]
            |           `--*T. monterosati (Grillo 1877) [=Fossarus monterosati] BR17
            |         Aclesia Rang 1828 G79
            |         Strategus Cooper 1862 C64
            |         Navarehus Cooper 1863 C64
            |           `--N. inermis (Cooper 1862) [=Strategus inermis] C64
            |         Doridella obscura F68
            |         Polycerella F68
            |           |--P. conyma F68
            |           |--P. davenportii F68
            |           `--P. emertoni F68
            |--Ringipleura BR17
            |--Acteonimorpha BR17
            `--Tectipleura BR17
                 |--Euopisthobranchia BR17
                 `--Panpulmonata (see below for synonymy) BR17
                      |  i. s.: Promenetus B97
                      |           |--P. exacuous (Say 1821) B97
                      |           `--P. umbilicatellus (Cockerell 1887) P88
                      |         Prophysa Bandel 1991 B91
                      |           `--*P. bristovii (de Loriol 1865) [=Physa bristovii; incl. Ph. wealdiana] B91
                      |         Proplanorbarius Bandel 1991 B91
                      |           `--*P. sanctusclaudius Bandel 1991 B91
                      |--Sacoglossa BR17
                      `--+--Siphonarioidea BR17
                         |--Pylopulmonata BR17
                         `--+--Acochlidimorpha BR17
                            `--+--Hygrophila BR17
                               `--Eupulmonata (see below for synonymy) BR17
                                    |--Ellobioidea BR17
                                    `--Geophila BR17
                                         |--Systellommatophora BR17
                                         `--Stylommatophora (see below for synonymy) BR17
                                              |  i. s.: Phasis BR05
                                              |         Scalaxis Pilsbry 1909 BR05 [Scalaxidae BR17, Scalaxinae]
                                              |           `--*S. rillyensis (de Boissy 1848) [=Achatina rillyensis] BR17
                                              |         Cherusciola Huckriede 1967 B91
                                              |           |--*C. nitida Huckriede 1967 B91
                                              |           `--C. mojonia Bandel 1991 B91
                                              |--Achatinina BR17
                                              |--Scolodontidae BR17
                                              `--Helicina [Aulacopoda, Holopoda, Subnuda] BR17
                                                   |  i. s.: Coelociontidae [Coelociidae, Coelociontoidea] BR17
                                                   |           |--Perrieria Tapparone Canefri 1878 BR17, BR05 (see below for synonymy)
                                                   |           |    `--*P. clausiliaeformis Tapparone-Canefri 1878 BR17
                                                   |           `--Coelocion Pilsbry 1904 [Coelociontinae] SS10
                                                   |                |--*C. australis (Forbes 1852) BR17 (see below for synonymy)
                                                   |                |--C. circumumbilicata Stanisic in Stanisic, Shea et al. 2010 SS10
                                                   |                `--C. craigeddiei Stanisic in Stanisic, Shea et al. 2010 SS10
                                                   |         Papilloderma Wiktor, Martin & Castillejo 1990 BR05 (see below for synonymy)
                                                   |           `--*P. altonagai Wiktor, Martin & Castillejo 1990 BR17
                                                   |         Plectopyloidea BR17
                                                   |         Punctoidea BR17
                                                   |         Testacellidae (see below for synonymy) BR17
                                                   |           |--Parmacellina vitrinaeformis Sandberger 1872 TTE93
                                                   |           `--Testacella Lamarck 1801 BR17 (see below for synonymy)
                                                   |                |--*T. haliotidea Draparnaud 1801 SS10 (see below for synonymy)
                                                   |                `--T. maugei [=Testacellus maugei] G40
                                                   |         Urocoptoidea BR17
                                                   |         Apula E99
                                                   |           |--A. amberti E99
                                                   |           |--A. escoffierae E99
                                                   |           `--A. koehnei E99
                                                   |         Mesodontopsis E99
                                                   |           |--M. chaixi E99
                                                   |           |--M. doderleini E99
                                                   |           |--M. herlacensis E99
                                                   |           |--M. ludovici E99
                                                   |           `--M. nehringi E99
                                                   |         Puisseguria E99
                                                   |           |--P. idanica E99
                                                   |           |--P. kowalczyki E99
                                                   |           `--P. zilchi E99
                                                   |         Frechenia E99
                                                   |           |--F. ducrosti E99
                                                   |           |--F. nayliesi E99
                                                   |           |--F. quadrifasciata E99
                                                   |           `--F. reichenbachi E99
                                                   |--Rhytidoidea BR17
                                                   |--Orthalicoidea BR17
                                                   |--Clausilioidea BR17
                                                   |--Arionoidea BR17
                                                   |--Oleacinoidei BR17
                                                   |--Succineoidei BR17
                                                   |--Limacoidei BR17
                                                   |--Helicoidei BR17
                                                   `--Pupilloidea BR17

Heterobranchia incertae sedis:
  Micromelo undatus GO06
  Tricarilda striatissima Gründel 1999 GK02
  Pseudoceratodes BR05
  Cimidae BR17
  Dolomitella Bandel 1994 [Dolomitellidae] BR05
    `--*D. semiornata (Kittl 1894) [=Hypsipleura semiornata] BR17
  Kuskokwimia Frýda & Blodgett 2001 [Kuskokwimiidae] FB01
    `--*K. moorei Frýda & Blodgett 2001 FB01
  Misurinella Bandel 1994 [Misurinellidae] BR05
    `--*M. sinistrorsa (Kittl 1894) [=Euchrysalis sinistrorsa] BR17
  Nerineoidea [Nerineacea, Nerineida, Nerineina] FB01
    |  i. s.: Brouzetia Y56
    |--Nerinellidae BR05
    |--Ceritellidae BR05
    |--Ptygmatididae BR17
    |--Nerineidae BR05
    |--Eunerinea Cox 1947 [Eunerineidae, Eunerineidae] BR17
    |    `--*E. castor (d’Orbigny 1850) [=Nerinea castor] BR05
    |--Pseudonerinea de Loriol 1890 BR05 [Pseudonerineidae BR17]
    |    |--*P. blauenensis de Loriol 1890 BR17
    |    |--P. stantoni Allison 1955 TTE93
    |    `--P. sturtoni Allison 1955 TTE93
    `--Itieriidae [Itieriacea, Itieriinae] BR17
         |--Itieria Matheron 1842 BR05
         |    `--*I. cabaneti Matheron 1842 BR17
         `--Itruvia Stoliczka 1867 TTE93, BR05 [Itruviidae]
              |--*I. canaliculata (d’Orbigny 1842) [=Pyramidella canaliculata] BR17
              `--I. carinata TTE93
  Mathildoidea [Mathildoidei] FB01
    |  i. s.: Turristylus Blaschke 1905 TE01
    |           `--*T. triadicus [=Eustylus triadicus] TE01
    |--Mathildidae BR05
    |--Gordenella Gründel 1990 [Gordenellidae] BR05
    |    `--*G. pommerana (Schmidt 1905) [=Cerithium pommeranum] BR17
    |--Trachoecus Kittl 1894 [Trachoecidae] BR05
    |    `--*T. gemmellaroi Kittl 1894 BR17
    `--Schartia Nützel & Kaim 2014 [Schartiidae, Schartiinae] BR17
         `--*S. carinata Nützel & Kaim 2014 BR17
  Damesia Holzapfel 1888 BK03
    `--*D. cretacea (Müller 1851) [=Crepidula cretacea] KC60
  Khairkhaniidae P08
  Loxebala Bandel 1996 B02
    `--*L. vallandroensis Bandel 1996 B02
  Haszprunariella Kiel & Bandel 2001 B02
  Acteonellidae BR17
  Murchisonelloidea BR17
    |--Donaldina BR17
    `--Murchisonellidae [Ebalidae] BR05
         |--Falsoebala Gründel 1998 B02
         |--Murchisonella Mörch 1875 [Murchisonellinae, Murchisonellini] BR05
         |    `--*M. spectrum (Mörch 1875) [=Murchisonia (*Murchisonella) spectrum] B02
         `--Ebala Gray 1847 [Ebalinae] BR05
              |--*E. nitidissima (Montagu 1803) [=Turbo nitidissimus] BR17
              |--E. cassiana Bandel 1996 B02
              |--E. elegantissima [=Turbo elegantissimus] BR17
              `--E. liassica Schröder 1995 B02
  Rhodope Koelliker 1847 BR05 (see below for synonymy)
    `--*R. veranii Koelliker 1847 BR17
  Streptacidoidea FB01
    |  i. s.: Heteroloxonema Frýda 2000 FB04
    |           `--*H. moniliformis (Goldfuss 1844) FB04 (see below for synonymy)
    |--Streptacididae B02
    |--Cassianebala Bandel 1996 [Cassianebalidae] BR05
    |    `--*C. speciensis Bandel 1996 B02
    `--Neodonaldina Bandel 1996 TE01 [Neodonaldinidae B02]
         `--*N. elongata (Zardini 1978) [=Spirocyclina elongata] TE01
  Neamphitomaria Bandel 1988 TTE93

*Coelocion australis (Forbes 1852) BR17 [=Balea australis SS10; incl. C. exigua Adams & Angas 1868 SS10]

Eupulmonata [Acteophila, Actophila, Akteophila, Amphipulmonata, Ellobiiformes, Phyllovora] BR17

Euthyneura [Acera, Aceridae, Actaeonacea, Acteobranchia, Acteoniformes, Actopleura, Aiolobranchiata, Anangia, Androgyna, Angiophora, Anthobranchia, Apneumonophora, Aporobranchiata, Aptera, Aulacognatha, Aulocognatha, Bullariacea, Bulliones, Eleutherobranchia, Eleutherobranchiatae, Enterobranchiata, Haplomorpha, Hemiphyllidinae, Monostichoglossata, Notaspidea, Notobranchia, Opisthobranchia, Opisthobranchiata, Opisthopneumona, Pentaganglionata, Phlebenterata, Placoesophaga, Pleuroanthobranchia, Pomatobranchiata, Procephala, Scaphandracea, Scaphandroidei, Siphoglossa, Steganobranchia, Stegobranchia, Tectibranchia, Tetracerata, Tetradontogastra] S11

Heterobranchia [Allogastropoda, Allomorpha, Architectibranchia, Dextrobranchia, Peracliones, Pulmobranchiata, Rhinophoralia]

*Heteroloxonema moniliformis (Goldfuss 1844) FB04 [=Turritella moniliformis FB04, Loxonema moniliforme FB01]

Panpulmonata [Adelopneumona, Aeropneusta, Basommatophora, Branchiopneusta, Coelopneumonata, Coelopnoa, Cilopnoa, Eupneumona, Heliciones, Inoperculata, Limaciones, Limnophila, Limnophilidae, Petrophila, Pulmonata, Pulmonifera, Saccobranchia, Siphonariiformes, Siphonariiformii, Siphonariiones, Siphonarioidei, Thalassophila] BR17

Papilloderma Wiktor, Martin & Castillejo 1990 BR05 [Papillodermatidae, Papillodermatoidea BR17, Papillodermidae]

Perrieria Tapparone Canefri 1878 BR17, BR05 [Perrieriinae]

Rhodope Koelliker 1847 BR05 [Apneusta, Rhodopacea, Rhodopadae, Rhodopemorpha, Rhodopidae, Rhodopida, Rhodopiformes, Rhodopoidea BR17]

Stylommatophora [Achatinoinei, Agnathomorpha, Anurethra, Arioniformes, Ataeniae, Brachynephra, Dolichonephria, Geocochlides, Haplogona, Helixina, Helixinia, Holognatha, Holopodopes, Limaciformes, Limaciformii, Lissognatha, Mesurethra, Monogonophora, Nephropneusta, Oleacinina, Oxygnatha, Pupillina, Pupilloinei, Pycnognatha, Pycnognathae, Sigmurethra, Soleiferae, Stylommatophorida, Vasopulmonata] BR17

Testacella Lamarck 1801 BR17 [incl. Testacelloides Wagner 1914 SS10, Testacellus Férussac 1819 SS10]

*Testacella haliotidea Draparnaud 1801 SS10 [=Limax (Testacella) haliotoidea G20, Testacellus haliotideus G40; incl. Testacella europaea G40, Testacella galliae G40, Testacella scutulum G40]

Testacellidae [Testacellacidae, Testacellina, Testacellinae, Testacelloidea, Testacelloinei] BR17

‘Valvata (Cincinna)’ helicelloides Huckreide 1967 [=V. helicoides de Loriol 1865 non Stoliczka 1862, *Provalvata helicoides] B91

*Type species of generic name indicated

REFERENCES

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Last updated: 11 September 2021.