Helicoidei

Labyrinthus furcillatus, copyright Naturalis Biodiversity Center.

Belongs within: Heterobranchia.
Contains: Sagdidae, Thysanophoridae, Trissexodontidae, Sphincterochilidae, Pleurodontidae, Helicodontidae, Geomitrinae, Helicellinae, Triodopsinae, Polygyrinae, Epiphragmophora, Lysinoinae, Helminthoglyptinae, Humboldtiana, Otostomus, Helicinae, Ariantinae, Leptaxinae, Trochulinae, Amplirhagada, Austrochloritis, Sphaerospira, Figuladra, Torresitrachia, Steorra, Setobaudinia, Camaeninae, Bradybaeninae, Hadrinae.

The Helicoidei are a clade of terrestrial gastropods united by molecular phylogenetic analysis (Bouchet et al. 2017). Molecular data also support the division of the clade between two major subgroups, the less speciose Sagdoidea and the more diverse Helicoidea. The Sagdoidea are a strictly Neotropical group, with the highest diversity around the Caribbean basin, whereas Helicoidea are cosmopolitan. Members of the Helicoidea are also characterised by a thickened peristome, and the majority of families (except Sphincterochilidae and Cepolis) possess an odontognath (ribbed or striated) jaw. Many helicoids have a dart apparatus on the genitalia, but this is absent in many Camaenidae.

Bradybaenids: the little freaks
Published 19 June 2013

The wonderful assembly in the photograph above (by B. Frank) is a congregation of the Asian tramp snail Bradybaena similaris. The Bradybaenidae are a family of small snails, closely related to the garden snails of the Helicidae, that are mostly native to eastern Asia. However, a few species such as B. similaris have become widespread around the world as a result of human transportation. Not deliberate transportation of the snails themselves, of course, but transportation of plants and plant matter that have had the snails clinging to them. Also, recent phylogenetic studies have indicated that the Australasian snails hitherto included in the Camaenidae are in fact not close relatives of the North American representatives of that family, but should be placed close to or even within the Bradybaenidae (Wade et al. 2007).

Euhadra grata gratoides, from here.

You may already be familiar with the production by some species of snail of ‘love darts’, small calcareous spears that a mating snail fires into its partner. The function of the love dart is still not entirely understood, though it does seem to improve sperm uptake by the snail being darted: whether by lowering its ability to resist insemination, or because snails are mini-masochists that get off on being stabbed, I couldn’t say. Most textbooks describing the use of love darts will (at least effectively) base their description on the common garden snail Cornu aspersum (or Cantareus aspersus, or whatever the heck we’re supposed to be calling it these days), which leaves its love dart embedded in its partner’s skin. Bradybaenids whose mating behaviour has been studied, however, do things a bit differently. Instead of abandoning its dart after a single firing, bradybaenids withdraw the dart and use it to stab their partner repeatedly, making it more of a love shiv than a love dart. And when I say repeatedly, I mean repeatedly: mating pairs of Euhadra subnimbosa would, on average, stab each other with the dart over 3300 times (Koene & Chiba 2006). So vigorous is the stabbing, in fact, that the dart pierces straight through the recipient and emerges through its foot! For those with JSTOR access, a video of the process can be seen at http://www.jstor.org/stable/10.1086/508028. And trust you to go rushing to watch a film of gastropod SM.

Systematics of Helicoidei
<==HelicoideiBR17
    |--SagdoideaBR17
    |    |--SagdidaeBP90
    |    |--Zachrysia Pilsbry 1894BP90 [ZachrysiidaeBR17]
    |    |    |--*Z. auricoma (Férussac 1821) [=Helix auricoma]BR17
    |    |    `--Z. provisoriaBP90
    |    `--SolaropsidaeBR17
    |         |--Hodopoeus crassus Pilsbry & Cockerell 1945TTE93
    |         |--Solaropsis Beck 1837 [Solaropsinae]BR05
    |         |    `--*S. pellisserpentis (Gmelin 1791) [=Helix pellisserpentis]BR17
    |         `--Caracolus Montfort 1810BR05 [CaracolinaeBR17]
    |              `--C. caracolla (Linnaeus 1758)AA88 [=Helix carocollaBR17, *C. oculatus Montfort 1810BR17]
    `--Helicoidea [Helicacea, Holopoda]BR05
         |  i. s.: Mandarina ponderosaKC11
         |         Nesiohelix samarangaeKC11
         |         SchlickumiaE99
         |           |--S. alicantensisE99
         |           |--S. aquensisE99
         |           |--S. bottiniiE99
         |           `--S. gaspardianaE99
         |         Titthodomus koeneniE99
         |         Pseudomonacha zippeiE99
         |--ThysanophoridaeBR05
         |--TrissexodontidaeBR05
         |--SphincterochilidaeBR05
         |--PleurodontidaeBR05
         |--HelicodontidaeBR05
         |--Canariella Hesse 1918BR05 [CanariellidaeBR17, Canariellinae, Canariellini]
         |    `--*C. hispidula (Lamarck 1822) [=Helix hispidula]BR17
         |--Cepolis Montfort 1810BR05
         |    |--*C. nicolsinianumBR17
         |    `--C. variansBP90
         |--GeomitridaeBR17
         |    |  i. s.: Xerotricha conspurcata (Draparnaud 1801)NB19
         |    |--GeomitrinaeBR05
         |    `--HelicellinaeBR17
         |--Polygyridae [Mesodontidae, Mesodontoidea, Polygyracea, Polygyroidea]BR17
         |    |  i. s.: Zhenjiangospira dignata Yü 1977TTE93
         |    |--TriodopsinaeBR05
         |    `--PolygyrinaeAP90
         |--TrichodiscinidaeBR17
         |    |--Trichodiscus Strebel 1880 [=Trichodiscina Martens 1892; Trichodiscininae, Trichodiscinini]BR17
         |    |    `--*T. coactiliatus (Deshayes 1838)BR17 (see below for synonymy)
         |    `--Miraverellia Baker 1922BR05 [MiraverelliinaeBR17, Miraverelliini]
         |         `--*M. sumichrasti (Crosse & Fischer 1872) [=Helix sumichrasti]BR17
         |--LabyrinthidaeBR17
         |    |--Lampadion Röding 1798BR17, BR05 [Lampadiidae]
         |    |    `--*L. labyrinthus Röding 1798BR17
         |    `--Labyrinthus Beck 1837BR17
         |         |--*L. labyrinthus (Lamarck 1792) [=Helix labyrinthus]BR17
         |         |--L. bifurcatusA66
         |         |--L. furcillatusA66
         |         `--L. leprieuriiA66
         |--ElonidaeBR05
         |    |--Klikia Pilsbry 1895 [Klikiinae, Klikiini]BR05
         |    |    `--*K. osculum (Thomä 1845) [=Helix osculum]BR17
         |    `--EloninaeBR17
         |         |--Elona Adams & Adams 1855 [Elonini]BR05
         |         |    `--*E. quimperiana (Blainville 1821) [=Helix quimperiana]BR17
         |         |--Galactochiloides Wenz 1919 [Galactochiloidini]BR17
         |         |    `--*G. nemoralites (Boubée 1831) [=Helix nemoralites]BR17
         |         `--Tropidomphalus Pilsbry 1895 [Tropidomphalini]BR17
         |              `--‘Helix’ arnoldii Thomä 1845 [=H. lepidotricha Braun 1851, *Tropidomphalus lepidotrichus]BR17
         |--Xanthonychidae [Helminthoglyptidae, Xanthonycidae, Xanthonychoidea]C06
         |    |  i. s.: Cryptostrakon Binney 1879C97
         |    |           |--*C. gabbi Binney 1879C97
         |    |           `--C. corcovadensis Cuezzo 1997C97
         |    |         Mesoglypterpes sagensis Yen 1952TTE93
         |    |--EpiphragmophoraC06
         |    |--LysinoinaeBR05
         |    |--HelmithoglyptinaeBR17
         |    |--Xanthonyx Crosse & Fischer 1867 [Xanthonychinae, Xanthonychini]BR05
         |    |    `--*X. sumichrasti (Brot 1867) [=Vitrina sumichrasti]BR17
         |    |--Echinix Thompson & Naranjo-Garcia 2012 [Echinichidae, Echinichinae]BR17
         |    |    `--*E. ochracea Thompson & Naranjo-Garcia 2012BR17
         |    |--Humboldtianinae [Humboldtianidae]BR17
         |    |    |--HumboldtianaBR05
         |    |    `--Bunnya Baker 1942BR05 [Bunnyinae, BunnyiniBR17]
         |    |         `--*B. bernardinae Baker 1942BR17
         |    `--Monadenia Pilsbry 1895BR05 [Monadeniidae, MonadeniinaeBR17]
         |         |--*M. fidelis (Gray 1843)BR17 [=Helix fidelisBR17, H. (Aglaia) fidelisC64; incl. H. nuttallianaC64]
         |         |    |--M. f. fidelisW63
         |         |    |--M. f. beryllicaW63
         |         |    `--M. f. klamathicaW63
         |         |--M. churchiW63
         |         |--M. infumataBP90
         |         |--M. setosaW63
         |         `--M. troglodytesC84
         |--Helicidae [Cochleophora, Helicinia, Scutifera]SS10
         |    |  i. s.: Wladislavia Wagner 1927BR05
         |    |         Ammonoceras guayaquilenseA66
         |    |         OphiogyraA66
         |    |           |--O. stenogyraA66
         |    |           `--O. stenostreptaA66
         |    |         OtostomusA66
         |    |         Iberellus pyrenaicus (Rossmässler 1839) (see below for synonymy)A-Z04
         |    |         Dentellocaracolus Oppenheim 1890S97
         |    |           |--D. (Prothelidomus) acrochordon (Oppenheim 1895)TTE93
         |    |           `--D. damnata (Brongniart 1810)TTE93
         |    |         Eremina desertorumH84
         |    |         Loganiopharynx rarus (Boissy 1840)TTE93
         |    |         Nanhaispira eversilabia Yu & Zhang 1982TTE93
         |    |         Haplohelix anadeniaPB27
         |    |         BurungaellaPB27
         |    |           |--B. buhambaensisPB27
         |    |           |--B. imperforataPB27
         |    |           `--B. oscitansPB27
         |    |         NakuruellaPB27
         |    |           |--N. ahenaPB27
         |    |           `--N. sororPB27
         |    |         Urguessella capillataPB27
         |    |         Zingis bequaertiPB27
         |    |         Massylaea vermiculata (Müller 1774)NB19
         |    |         Assyriella thospitis Schütt & Subai 1996GN12
         |    |           |--A. t. thospitisGN12
         |    |           `--A. t. menkhorsti Gümüş & Neubert 2012GN12
         |    |--HelicinaeBR05
         |    |--AriantinaeBR05
         |    `--Murellinae [Murelleae, Murellini]BR17
         |         |--Murella Pfeiffer 1877BR05
         |         |    `--*M. muralis (Müller 1774) [=Helix muralis]BR17
         |         `--Tacheocampylaea Pfeiffer 1877 [Tacheocampylaeinae, Tacheocampylinae]BR05
         |              `--*T. raspaili (Payraudeau 1826) [=Helix raspaili]BR17
         |--Hygromiidae [Helicellidae, Hygromioidea]SS10
         |    |  i. s.: MonachoidesHM07
         |    |           |--M. incarnatus (Müller 1774)HM07
         |    |           `--M. vicinaJ63
         |    |         Xerolenta obvia (Menke 1828)HM07
         |    |         Candidula Kobelt 1871BP90
         |    |           `--C. intersecta [incl. Helicella caperata]BP90
         |    |         Xerocincta Monterosato 1892SS10
         |    |           `--*X. neglecta (Draparnaud 1805) [=Helix neglecta]SS10
         |    |         Microxeromagna Ortiz de Zarate 1950SS10
         |    |           `--M. lowei (Potiez & Michaud 1838) (see below for synonymy)SS10
         |    |         Prietocella Schileyko & Menkhorst 1997SS10
         |    |           `--*P. barbara (Linnaeus 1758)SS10 (see below for synonymy)
         |    |         ‘Helix’ australis Menke 1843S97
         |    |         XerocrassaH84
         |    |           |--X. erkeliiH84
         |    |           |--X. helleriH84
         |    |           |--H. langloisianaH84
         |    |           |--X. pilsbryiH84
         |    |           |--X. pseudojacostaH84
         |    |           |--X. seetzeniiH84 [=Trochoidea seetzeniC83]
         |    |           `--X. tuberculosaH84
         |    |         Xeropicta vestalisH84
         |    |         Archygromia durbani (Edwards 1852)TTE93
         |    |         Pseudoxerotricha subconspurcata (Sandberger 1873)TTE93
         |    |         Hemistenotrema quadrisinuosa (Boettger 1897)TTE93
         |    |--LeptaxinaeBR17
         |    |--TrochulinaeBR17
         |    `--HygromiinaeBR05
         |         |--Perforatella Schlüter 1838 [Perforatellini]BR17
         |         |    |--*P. bidentata (Gmelin 1791) [=Helix bidentata]BR17
         |         |    `--P. dibothryonJ63
         |         `--Hygromia Risso 1826 [Hygromiini]BR05
         |              |--*H. cinctella (Draparnaud 1801) [=Helix cinctella]BR17
         |              `--H. strigella Drap. 1805S01
         `--Camaenidae [Bradybaenidae, Camaenoidea, Eulotidae, Eulotinae]BR17
              |  i. s.: Multiscapta Yu 1982G88
              |           |--M. eversilabiaG88
              |           `--M. xinyuensis Gu 1988G88
              |         Exiligada Solem 1984S97
              |           `--E. negriensis Iredale 1939S88
              |         AmplirhagadaJS13
              |         Westraltrachia Iredale 1933S97
              |           |--W. commoda (Iredale 1939)S88
              |           `--W. derbyi (Cox 1892)S97
              |         Mouldingia Solem 1984S97
              |           `--M. orientalis Solem 1984S88
              |         Pleurodontites diespiter (Dall 1890)AP90
              |         Galadistes Iredale 1938SS10
              |           |--*G. liverpoolensis (Brazier 1872) [=Helix (Galaxias) liverpoolensis]SS10
              |           |--G. akubra Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--G. alleni Iredale 1943SS10
              |           |--G. intervenens Iredale 1938SS10
              |           |--G. marcescens (Cox 1868)SS10
              |           |--G. molong Shea & Griffiths in Stanisic, Shea et al. 2010SS10
              |           |--G. pilligaensis Shea & Griffiths in Stanisic, Shea et al. 2010SS10
              |           |--G. pustulosa Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--‘Helix’ stutchburyi Pfeiffer 1857SS10
              |         Mussonena Iredale 1938SS10
              |           |--*M. spinei (Cox 1868) (see below for synonymy)SS10
              |           |--M. boonah Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           |--M. carnarvon Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--M. martinlogari Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--M. maxima Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--M. nogoa Stanisic in Stanisic, Shea et al. 2010SS10
              |         Nannochloritis Iredale 1938SS10
              |           `--*N. layardi (Gude 1906) (see below for synonymy)SS10
              |         Calvigenia Iredale 1938SS10
              |           |--*C. blackmani (Cox 1868) [=Helix blackmani]SS10
              |           |--C. cognata (Gude 1907) [=Chloritis cognata, Gloreugenia cognata]SS10
              |           |--C. cootha Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           `--C. mucida (Pfeiffer 1857) [=Helix mucida, Ramogenia mucida]SS10
              |         AustrochloritisSS10
              |         Squamagenia Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           |--*S. yabba Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           `--S. separanda (Iredale 1938) [=Austrochloritis separanda]SS10
              |         Meridolum Iredale 1942SS10
              |           |--*M. jervisensis (Quoy & Gaimard 1832) [=Helix jervisensis]SS10
              |           |--M. corneovirens (Pfeiffer 1851) (see below for synonymy)SS10
              |           |--M. gilbertiBRW98
              |           |--M. grayi (Pfeiffer 1847)SS10
              |           |--M. gulosum (Gould 1846) (see below for synonymy)SS10
              |           |--M. marshalli McLauchlan 1951SS10
              |           |--M. maryae Clark 2009SS10
              |           |--M. middenense (McLauchlan 1954)SS10
              |           `--M. sheai Clark 2009SS10
              |         Temporena Iredale 1933SS10
              |           |--*T. whartoni (Cox 1871) [=Helix whartoni]SS10
              |           |--T. juliafoxae Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--T. sardalabiata (Cox 1871) (see below for synonymy)SS10
              |         Noctepuna Iredale 1933SS10
              |           |--*N. poiretiana (Reeve 1852) [=Helix poiretiana]SS10
              |           |    |--N. p. poiretianaSS10
              |           |    `--N. p. clenchi McMichael 1958SS10
              |           |--N. cerea (Hedley 1894)SS10
              |           |--N. mayana (Hedley 1899)SS10
              |           `--N. muensis (Hedley 1912) [=Papuina muensis]SS10
              |         Lamprellia Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*L. zebina (Brazier 1878) [=Helix zebina]SS10
              |           `--L. angulata Stanisic in Stanisic, Shea et al. 2010SS10
              |         SphaerospiraSS10
              |         FiguladraSS10
              |         Neveritis Iredale 1938SS10
              |           |--*N. poorei (Gude 1907) [=Chloritis poorei; incl. C. thales Gude 1907]SS10
              |           |--N. aridorum (Cox 1866)SS10
              |           `--N. misella (Gude 1907)SS10
              |         Posorites Iredale 1933SS10
              |           |--*P. fucata (Pfeiffer 1853) [=Helix fucata]SS10
              |           |--P. conscendens (Cox 1866)SS10
              |           `--P. turneri (Shirley 1921)SS10
              |         Thersites Pfeiffer 1855SS10
              |           |--*T. richmondiana (Reeve 1852) (see below for synonymy)SS10
              |           |--T. darlingtoni Clench & Archer 1938SS10
              |           |--T. mitchellae (Cox 1864) [incl. Annakelea peregrans Iredale 1937]SS10
              |           `--T. novaehollandiae (Gray 1834) [incl. Helix dupuyana Pfeiffer 1851]SS10
              |         Papuexul Iredale 1933SS10
              |           `--*P. bidwilli (Reeve 1853) [=Helix bidwilli]SS10
              |         Patrubella Iredale 1938SS10
              |           `--*P. buxtoni (Brazier 1880) [=Helix (Planispira) buxtoni]SS10
              |         Kandoschloritis Shea & Griffiths in Stanisic, Shea et al. 2010SS10
              |           `--*K. pustulosus Shea & Griffiths in Stanisic, Shea et al. 2010SS10
              |         Ramogenia Iredale 1938SS10
              |           `--R. challengeri (Gude 1906) [incl. Chloritis obnubila Gude 1907, *R. obnubila]SS10
              |         Trachygenia Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*T. lanuginosa (Gude 1907) [=Chloritis lanuginosa, Ramogenia lanuginosa]SS10
              |           |--T. capiosetosa Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--T. microlepida Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--T. praecursoris (Hedley 1912) (see below for synonymy)SS10
              |         Eurytrachia Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*E. mucosa (Cox 1868) (see below for synonymy)SS10
              |           `--E. costulata Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |         Tolgachloritis Iredale 1933SS10
              |           |--*T. jacksoni (Hedley 1912) [=Chloritis jacksoni]SS10
              |           |--T. campbelli (Iredale 1938) [=Mussonena campbelli]SS10
              |           |--T. costata Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--T. kondaparinga Stanisic in Stanisic, Shea et al. 2010SS10
              |         Gloreugenia Iredale 1933SS10
              |           |--*G. coxeni (Cox 1871) [=Helix coxeni; incl. G. carduelis Iredale 1938]SS10
              |           `--G. blackalli (Brazier 1875) [incl. Chloritis exilis Gude 1907]SS10
              |         Boriogenia Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*B. hedleyi (Fulton 1907) [=Chloritis (Austrochloritis) hedleyi, Gloreugenia hedleyi]SS10
              |           `--B. gerrardstirlingi Stanisic in Stanisic, Shea et al. 2010SS10
              |         Crookshanksia Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--*C. frankstanisici Stanisic in Stanisic, Shea et al. 2010SS10
              |         Trozena Iredale 1938SS10
              |           `--*T. morata Iredale 1938SS10
              |         Protolinitis Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*P. terriirwinae Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--P. pusilla (Hedley 1912)SS10
              |         Aetholitis Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*A. agamemnon (Gude 1906) [=Chloritis agamemnon]SS10
              |           |--A. astaeus (Gude 1906)SS10
              |           |--A. francisae Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--A. gerdesae Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--A. janetzkiae Stanisic in Stanisic, Shea et al. 2010SS10
              |         Obsteugenia Iredale 1933SS10
              |           `--*O. inflecta (Hedley 1912) [=Chloritis inflecta; incl. C. bilabiata Odhner 1917]SS10
              |         Trachiopsis Pilsbry 1893SS10
              |           |--T. strangulata (Hombron & Jaquinot 1841) (see below for synonymy)SS10
              |           |--T. musgravensis Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           |--T. railtoni Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--T. setosa Stanisic in Stanisic, Shea et al. 2010SS10
              |         Chloritisanax Iredale 1933SS10
              |           `--*C. banneri (Pfeiffer 1863) [=Helix banneri]SS10
              |         Pommerhelix Clark 2009SS10
              |           |--*P. carmelae Clark 2009SS10
              |           |--P. depressa (Hedley 1901)SS10
              |           |--P. duralensis (Cox 1868)SS10
              |           |--P. exocarpi (Cox 1868)SS10
              |           |--P. insularis Clark 2009SS10
              |           |--P. mastersi (Cox 1864)SS10
              |           |--P. monacha (Pfeiffer 1859) [incl. Meridolum bowdenae McLauchlan 1954]SS10
              |           `--P. stanisici Clark 2009SS10
              |         Sauroconcha Zhang & Shea 2008SS10
              |           `--*S. caperteeana Zhang & Shea 2008SS10
              |         Petraconcha Clark 2009SS10
              |           `--*P. griffithsi Clark 2009SS10
              |         Ponderconcha Clark 2009SS10
              |           |--*P. murphyi Clark 2009SS10
              |           |--P. gilberti (Pfeiffer 1846)SS10
              |           |--P. ianthostoma (Cockerell 1929)SS10
              |           |--P. morosa (Morelet 1853) [incl. Meridolum ascensum Iredale 1938]SS10
              |           `--P. warrumbungliana Stanisic in Stanisic, Shea et al. 2010SS10
              |         Meridistes Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--*M. woolnoughi Stanisic in Stanisic, Shea et al. 2010SS10
              |         Stanisicia Clark 2009SS10
              |           `--*S. macintyrensis Clark 2009SS10
              |         Pallidelix Iredale 1933SS10
              |           |--*P. greenhilli (Cox 1866) (see below for synonymy)SS10
              |           |--P. bennetti (Brazier 1872) [incl. Helix (Xanthomelon) daintreei Brazier 1875]SS10
              |           `--P. chinchilla Stanisic in Stanisic, Shea et al. 2010SS10
              |         Ventopelita Iredale 1943SS10
              |           `--*V. leucocheilus (Cox 1868) (see below for synonymy)SS10
              |         Moretonistes Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--*M. mansueta (Reeve 1854) (see below for synonymy)SS10
              |         Pseudodistes Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           |--*P. biggenden Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           `--P. reevesi Stanisic in Stanisic, Shea et al. 2010SS10
              |         Jimbouria Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*J. rodhobsoni Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--J. chrisalleni Shea & Griffiths in Stanisic, Shea et al. 2010SS10
              |         Adclarkia Stanisic 1996SS10
              |           |--*A. dawsonensis Stanisic 1996SS10
              |           |--A. cameroni Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--A. dulacca Stanisic in Stanisic, Shea et al. 2010SS10
              |         Lynfergusonia Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*L. mundubbera Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--L. taroomiana Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |         Perioinsolita Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |           `--*P. pokryszkoae Stanisic & Potter in Stanisic, Shea et al. 2010SS10
              |         Toombatrachia Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--*T. wheelerae Stanisic in Stanisic, Shea et al. 2010SS10
              |         Melostrachia Solem 1979SS10
              |           |--*M. glomerans (Iredale 1938) [=Torresitrachia glomerans]SS10
              |           `--M. acuticostata (Fulton 1907)SS10
              |         Spurlingia Iredale 1933SS10
              |           |--S. dunkiensis (Forbes 1851) (see below for synonymy)SS10
              |           |--S. excellens Iredale 1937SS10
              |           |--S. gemma Iredale 1937SS10
              |           |--S. monticola (Clench & Archer 1938)SS10
              |           |--S. portus Iredale 1937SS10
              |           |--S. praehadra (Odhner 1917)SS10
              |           |--S. tinarooensis Iredale 1937SS10
              |           `--S. undara Stanisic in Stanisic, Shea et al. 2010SS10
              |         Jacksonena Iredale 1937SS10
              |           `--*J. rudis (Hedley 1912) [=Planispira rudis]SS10
              |         Forrestena Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--*F. delicata (Hedley 1912) [=Planispira delicata, Jacksonena delicata]SS10
              |         Zyghelix Iredale 1937SS10
              |           `--*Z. forsteriana (Reeve 1852) (see below for synonymy)SS10
              |         TorresitrachiaS97
              |         Austrocamaena Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*A. ricnattrassi Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--A. thorntonensis Stanisic in Stanisic, Shea et al. 2010SS10
              |         Rhynchotrochus Möllendorff 1895SS10
              |           |--*R. taylorianus (Adams & Reeve 1850) [=Helix tayloriana]SS10
              |           `--R. macgillivrayi (Forbes 1851) [incl. R. macgillivrayi extensior Iredale 1938]SS10
              |         Crikey Stanisic 2009SS10
              |           `--*C. steveirwini Stanisic 2009SS10
              |         Meliobba Iredale 1930SS10
              |           `--*M. shafferyi Iredale 1940SS10
              |         Gnarosophia Iredale 1933SS10
              |           `--*G. bellendenkerensis (Brazier 1875) (see below for synonymy)SS10
              |         Bentosites Iredale 1933SS10
              |           |--*B. macleayi (Cox 1865) [=Helix macleayi; incl. B. macleayi wardiana Iredale 1933]SS10
              |           |--B. coxi (Crosse 1866) (see below for synonymy)SS10
              |           |--B. etheridgei (Brazier 1877)SS10
              |           |--B. fortasse (Iredale 1933) [=Varohadra volgiola fortasse; incl. V. probleema Iredale 1933]SS10
              |           |--B. gavisa Iredale 1933 (see below for synonymy)SS10
              |           `--B. hefferani Stanisic in Stanisic, Shea et al. 2010SS10
              |         Billordia Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*B. bridgettae Stanisic in Stanisic, Shea et al. 2010SS10
              |           `--B. nicoletteae Stanisic in Stanisic, Shea et al. 2010SS10
              |         Marilynessa Stanisic in Stanisic, Shea et al. 2010SS10
              |           |--*M. yulei (Forbes 1851) [=Helix yulei; incl. H. basalis Pilsbry 1890, H. rainbirdi Cox 1870]SS10
              |           |--M. findera (Iredale 1937) [=Varohadra findera]SS10
              |           |--M. macneilli (Iredale 1937) [=Varohadra macneilli]SS10
              |           |--M. starena (Iredale 1937) [=Varohadra starena]SS10
              |           `--M. thorogoodi (Iredale 1937) [=Varohadra thorogoodi]SS10
              |         SteorraSS10
              |         Monteithosites Stanisic 1996SS10
              |           `--*M. helicostracum Stanisic 1996SS10
              |         Cristilabrum Solem 1981S97
              |           |--C. bubulumBRW98
              |           |--C. buryillumBRW98
              |           |--C. grossumBRW98
              |           |--C. kessneri Solem 1989S97
              |           |--C. monodonBRW98
              |           |--C. primumBRW98
              |           `--C. simplexBRW98
              |         Quistrachia Iredale 1939JS13
              |           |  i. s.: *Q. monogramma (Ancey 1898)JS13 [=Trachia monogrammaJS13, Planispira monogrammaS97]
              |           |         Q. herberti Solem 1997S97
              |           |         Q. leptogramma (Pfeiffer 1846) (see below for synonymy)S97
              |           |--+--Q. legendrei Solem 1997JS13, S97
              |           |  `--Q. turneri Solem 1997JS13, S97
              |           `--+--Q. montebelloensis (Preston 1914) (see below for synonymy)JS13
              |              `--+--Q. lefroyi Solem 1997JS13, S97
              |                 `--Q. warroorana Solem 1997JS13, S97
              |         SetobaudiniaS97
              |         Kimboraga Iredale 1933S97
              |           |--K. cascadensisGK12
              |           |--K. exanimaGK12
              |           |--K. glabraGK12
              |           `--K. wulalamGYD14
              |         Globorhagada Iredale 1933S97
              |           |--G. confusaGK12
              |           |--G. uwinsensisGK12
              |           |--G. wunandarraGK12
              |           |--G. wurroolguGK12
              |           `--G. yoowadanGK12
              |         Mesodontrachia Solem 1985S97
              |         Parglogenia Iredale 1938S97
              |         Ningbingia Solem 1981S97
              |         Turgenitubulus Solem 1981S97
              |           `--T. pagodula Solem 1985S97
              |         Prymnbriareus Solem 1981S97
              |         Ordtrachia Solem 1984S97
              |         Prototrachia Solem 1984S97
              |         Baudinella Thiele 1931S97
              |           |--B. boongareensisGK12
              |           |--B. occidentalisGK12
              |           |--B. setobaudinioidesGK12
              |           |--B. thieleiGK12
              |           `--B. tuberculataGK12
              |         Retroterra Solem 1985S97
              |           |--R. aequabilisGK12
              |           `--R. discoideaGK12
              |         Kendrickia Solem 1985S97
              |         Carinotrachia Solem 1985S97
              |           |--C. admiraleGK12
              |           |--C. carsoniana Solem 1985S97
              |           `--C. elevataGK12
              |         Australocosmica augustaeGK12
              |         Kimberleydiscus fasciatusGK12
              |         Kimberleymelon tealeiGK12
              |         KimberleytrachiaGK12
              |           |--K. achernariaGK12
              |           |--K. aequumGK12
              |           |--K. alphacentauriGK12
              |           |--K. amplirhagadoidesGK12
              |           |--K. canopiGK12
              |           |--K. chartaceaGK12
              |           |--K. crawfordiGK12
              |           |--K. crucisGK12
              |           `--K. somniatorGK12
              |         Molema stankowskiGK12
              |         Olympus nimbus Simone 2010S11
              |         Cathaica fasciola (Draparnaud 1805) [=Helix fasciola; incl. H. pyrrhozona]S01
              |         Kanabohelix kanabensis (White 1876)TTE93
              |         Coneulota diarti (Cossmann 1907)TTE93
              |--CamaeninaeS97
              |--BradybaeninaeBR17
              |--HadrinaeBR17
              `--HelicostylinaeBR05
                   |  i. s.: ‘Helix’ pyrostomaC84
                   |--Helicostyla Férussac 1821BR05
                   |    `--*H. mirabilis (Férussac 1821)BR17 [=Helix mirabilisBR17, Helix galactites Lamarck 1822S41]
                   |--Pfeifferia Gray 1853 [Pfeifferiana, Pfeifferiinae]BR05
                   |    `--*P. micans (Pfeiffer 1845) [=Helix micans]BR17
                   `--Cochlostyla Férussac 1821 [Cochlostylidae, Cochlostylinae]BR05
                        `--*C. ovoidea (Bruguière 1789) [=Bulimus ovoideus]BR17

Bentosites coxi (Crosse 1866) [incl. Helix cerata Cox 1868, H. cerea Cox 1864, H. (Helicostyla) croftoni Cox 1872, H. forbesii Cox 1864]SS10

Bentosites gavisa Iredale 1933 [incl. B. birchi Iredale 1933, Helix gratiosa Cox 1871 non Studer 1820]SS10

*Eurytrachia mucosa (Cox 1868) [=Helix mucosa, Trachiopsis mucosa; incl. Planispira truculenta Hedley 1912, P. tuckeri Pilsbry 1894]SS10

*Gnarosophia bellendenkerensis (Brazier 1875) [=Helix bellendenkerensis; incl. H. beddomei Brazier 1878 non Brazier 1876, Thersites castanea Odhner 1917, Gnarosophia humoricola Iredale 1937, T. pterinus Clench & Archer 1938]SS10

Iberellus pyrenaicus (Rossmässler 1839) [=Helix hispanica var. pyrenaica; incl. H. companyonii Aleron in Companyo 1837 (n. n.), Iberellus companyonii, H. minoricensis Mittre 1842]A-Z04

Meridolum corneovirens (Pfeiffer 1851) [incl. Helix mabillei Crosse 1864, H. mabillei minor Crosse 1864, H. mulgoae Cox 1868]SS10

Meridolum gulosum (Gould 1846) [incl. Helix cailleti Crosse 1864, H. coriaria Pfeiffer 1847, H. cailleti pallidior Crosse 1864, H. scotti Cox 1864]SS10

Microxeromagna lowei (Potiez & Michaud 1838) [=Helix lowei, Helix (Xerophila) armillata Lowe 1852; incl. Helicella (Candidula) mayeri Gude 1914, Helix subsecta Tate 1879, Helix vestita Rambur 1868, Cernuella vestita, *Microxeromagna vestita]SS10

*Moretonistes mansueta (Reeve 1854) [=Helix mansueta, Ventopelita mansueta; incl. H. yatalaensis Cox 1873]SS10

*Mussonena spinei (Cox 1868) [=Helix spinei; incl. H. hystrix Cox 1868 non Mighels in Pfeiffer 1846, Chloritis munda Gude 1907]SS10

*Nannochloritis layardi (Gude 1906) [=Chloritis layardi; incl. Austrochloritis blighiana Iredale 1938, A. senticula Iredale 1938]SS10

*Pallidelix greenhilli (Cox 1866) [=Helix greenhilli; incl. H. expeditionis Cox 1868, Galadistes expeditionis]SS10

*Prietocella barbara (Linnaeus 1758)SS10 [=Helix barbaraSS10, Cochlicella barbaraBP90; incl. H. ventricosa Draparnaud 1801SS10, Bulimus ventricosusG40, Cochlicella ventricosaBP90, H. (Cochlicella) ventrosa Férussac 1821SS10]

Quistrachia leptogramma (Pfeiffer 1846) [=Helix leptogramma, Globorhagada leptogramma, Thersites (Rhagada) leptogramma]S97

Quistrachia montebelloensis (Preston 1914) [=Rhagada montebelloensis, Globorhagada montebelloensis; incl. Quistrachia barrowensis Solem 1997]JS13

Spurlingia dunkiensis (Forbes 1851) [incl. S. cadmus Iredale 1937, Helix (Hadra) coxenae Brazier 1875, S. helga Iredale 1937, H. nicomede Brazier 1878, *S. nicomede, Thersites trachydermon Clench & Archer 1938]SS10

Temporena sardalabiata (Cox 1871) [=Helix sardalabiata; incl. Varohadra saxicola Iredale 1937, H. (Hadra) stephensoniana Brazier 1872]SS10

*Thersites richmondiana (Reeve 1852) [=Helix richmondiana; incl. H. richmondiana decolorata Pilsbry 1890, Annakelea tympanum Iredale 1937]SS10

Trachiopsis strangulata (Hombron & Jaquinot 1841) [incl. Helix cyclostoma Le Guillou 1842, H. (Trachia) dentoni Ford 1889, H. tuckeri Pfeiffer 1846, *Trachiopsis tuckeri]SS10

Trachygenia praecursoris (Hedley 1912) [=Chloritis cognata praecursoris, Gloreugenia praecursoris; incl. Austrochloritis fringilla Iredale 1938]SS10

*Trichodiscus coactiliatus (Deshayes 1838)BR17 [=Helix coactiliataBR17, Averellia (*Trichodiscina) coactiliataJ68, BR17]

*Ventopelita leucocheilus (Cox 1868) [=Helix leucocheilus; incl. H. leucocheilus lismorensis Pilsbry 1890, H. mariae Cox 1864]SS10

*Zyghelix forsteriana (Reeve 1852) [=Helix forsteriana; incl. Thersites (Hadra) forsteriana ada Marshall 1927, H. (Hadra) darwini Brazier 1872, H. hetaera Pfeiffer 1860, H. forsteriana major Pfeiffer 1859]SS10

*Type species of generic name indicated

References

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[A-Z04] Alonso-Zarazaga, M. A. 2004. Precisiones sobre un artículo nomenclatural acerca de la validez “taxonómica” de Iberellus companyonii (Aleron, 1837) recientemente publicado (Mollusca, Pulmonata, Helicidae). Graellsia 60 (1): 101–105.

[AA88] Auffenberg, K., & T. Auffenberg. 1988. Density, spatial distribution, activity patterns, and biomass of the land snail, Geophorus bothropoma Moellendorff (Prosobranchia: Helicinidae). Nautilus 102 (1): 40–45.

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