Helicoidei

Labyrinthus furcillatus, copyright Naturalis Biodiversity Center.


Belongs within: Heterobranchia.
Contains: Sagdidae, Thysanophoridae, Trissexodontidae, Sphincterochilidae, Pleurodontidae, Geomitrinae, Helicellinae, Triodopsinae, Polygyrinae, Helicodontidae, Epiphragmophora, Lysinoinae, Helminthoglyptinae, Humboldtiana, Leptaxinae, Trochulinae, Helicinae, Ariantinae, 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. Members of the Helicoidea are also characterised by a thickened peristome. The majority of families (except Sphincterochilidae and Cepolis) also possess an odontognath (ribbed or striated) jaw. Many helicoids have a dart apparatus on the genitalia, but this is absent in many Camaenidae.

<==Helicoidei BR17
    |--Sagdoidea BR17
    |    |--Sagdidae BP90
    |    |--Zachrysia Pilsbry 1894 BP90 [Zachrysiidae BR17]
    |    |    |--*Z. auricoma (Férussac 1821) [=Helix auricoma] BR17
    |    |    `--Z. provisoria BP90
    |    `--Solaropsidae BR17
    |         |--Hodopoeus crassus Pilsbry & Cockerell 1945 TTE93
    |         |--Solaropsis Beck 1837 [Solaropsinae] BR05
    |         |    `--*S. pellisserpentis (Gmelin 1791) [=Helix pellisserpentis] BR17
    |         `--Caracolus Montfort 1810 BR05 [Caracolinae BR17]
    |              `--C. caracolla (Linnaeus 1758) AA88 [=Helix carocolla BR17, *C. oculatus Montfort 1810 BR17]
    `--Helicoidea [Helicacea, Holopoda] BR05
         |  i. s.: Mandarina ponderosa KC11
         |         Nesiohelix samarangae KC11
         |         Schlickumia E99
         |           |--S. alicantensis E99
         |           |--S. aquensis E99
         |           |--S. bottinii E99
         |           `--S. gaspardiana E99
         |         Titthodomus koeneni E99
         |         Pseudomonacha zippei E99
         |--Thysanophoridae BR05
         |--Trissexodontidae BR05
         |--Sphincterochilidae BR05
         |--Pleurodontidae BR05
         |--Canariella Hesse 1918 BR05 [Canariellidae BR17, Canariellinae, Canariellini]
         |    `--*C. hispidula (Lamarck 1822) [=Helix hispidula] BR17
         |--Cepolis Montfort 1810 BR05
         |    |--*C. nicolsinianum BR17
         |    `--C. varians BP90
         |--Geomitridae BR17
         |    |--Geomitrinae BR05
         |    `--Helicellinae BR17
         |--Polygyridae [Mesodontidae, Mesodontoidea, Polygyracea, Polygyroidea] BR17
         |    |  i. s.: Zhenjiangospira dignata Yü 1977 TTE93
         |    |--Triodopsinae BR05
         |    `--Polygyrinae BR05
         |--Trichodiscinidae BR17
         |    |--Trichodiscus Strebel 1880 [=Trichodiscina Martens 1892; Trichodiscininae, Trichodiscinini] BR17
         |    |    `--*T. coactiliatus (Deshayes 1838) [=Helix coactiliata, *Trichodiscina coactiliata] BR17
         |    `--Miraverellia Baker 1922 BR05 [Miraverelliinae BR17, Miraverelliini]
         |         `--*M. sumichrasti (Crosse & Fischer 1872) [=Helix sumichrasti] BR17
         |--Labyrinthidae BR17
         |    |--Lampadion Röding 1798 BR17, BR05 [Lampadiidae]
         |    |    `--*L. labyrinthus Röding 1798 BR17
         |    `--Labyrinthus Beck 1837 BR17
         |         |--*L. labyrinthus (Lamarck 1792) [=Helix labyrinthus] BR17
         |         |--L. bifurcatus A66
         |         |--L. furcillatus A66
         |         `--L. leprieurii A66
         |--Elonidae BR05
         |    |--Klikia Pilsbry 1895 [Klikiinae, Klikiini] BR05
         |    |    `--*K. osculum (Thomä 1845) [=Helix osculum] BR17
         |    `--Eloninae BR17
         |         |--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
         |--Helicodontidae BR05
         |--Xanthonychidae [Helminthoglyptidae, Xanthonycidae, Xanthonychoidea] C06
         |    |  i. s.: Cryptostrakon Binney 1879 C97
         |    |           |--*C. gabbi Binney 1879 C97
         |    |           `--C. corcovadensis Cuezzo 1997 C97
         |    |         Mesoglypterpes sagensis Yen 1952 TTE93
         |    |--Epiphragmophora C06
         |    |--Lysinoinae BR05
         |    |--Helmithoglyptinae BR17
         |    |--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 2012 BR17
         |    |--Humboldtianinae [Humboldtianidae] BR17
         |    |    |--Humboldtiana BR05
         |    |    `--Bunnya Baker 1942 BR05 [Bunnyinae, Bunnyini BR17]
         |    |         `--*B. bernardinae Baker 1942 BR17
         |    `--Monadenia Pilsbry 1895 BR05 [Monadeniidae, Monadeniinae BR17]
         |         |--*M. fidelis (Gray 1843) [=Helix fidelis] BR17
         |         |--M. infumata BP90
         |         `--M. troglodytes C84
         |--Hygromiidae [Helicellidae, Hygromioidea] SS10
         |    |  i. s.: Monachoides incarnatus (Müller 1774) HM07
         |    |         Xerolenta obvia (Menke 1828) HM07
         |    |         Candidula Kobelt 1871 BP90
         |    |           `--C. intersecta [incl. Helicella caperata] BP90
         |    |         Xerocincta Monterosato 1892 SS10
         |    |           `--*X. neglecta (Draparnaud 1805) [=Helix neglecta] SS10
         |    |         Microxeromagna Ortiz de Zarate 1950 SS10
         |    |           `--M. lowei (Potiez & Michaud 1838) (see below for synonymy) SS10
         |    |         Prietocella Schileyko & Menkhorst 1997 SS10
         |    |           `--*P. barbara (Linnaeus 1758) SS10 (see below for synonymy)
         |    |         ‘Helix’ australis Menke 1843 S97
         |    |         Xerocrassa H84
         |    |           |--X. erkelii H84
         |    |           |--X. helleri H84
         |    |           |--H. langloisiana H84
         |    |           |--X. pilsbryi H84
         |    |           |--X. pseudojacosta H84
         |    |           |--X. seetzenii H84
         |    |           `--X. tuberculosa H84
         |    |         Xeropicta vestalis H84
         |    |         Archygromia durbani (Edwards 1852) TTE93
         |    |         Pseudoxerotricha subconspurcata (Sandberger 1873) TTE93
         |    |         Hemistenotrema quadrisinuosa (Boettger 1897) TTE93
         |    |--Leptaxinae BR17
         |    |--Trochulinae BR17
         |    `--Hygromiinae BR05
         |         |--Perforatella Schlüter 1838 [Perforatellini] BR17
         |         |    `--*P. bidentata (Gmelin 1791) [=Helix bidentata] BR17
         |         `--Hygromia Risso 1826 [Hygromiini] BR05
         |              |--*H. cinctella (Draparnaud 1801) [=Helix cinctella] BR17
         |              `--H. strigella Drap. 1805 S01
         |--Helicidae [Cochleophora, Helicinia, Scutifera] SS10
         |    |  i. s.: Wladislavia Wagner 1927 BR05
         |    |         Ammonoceras guayaquilense A66
         |    |         Ophiogyra A66
         |    |           |--O. stenogyra A66
         |    |           `--O. stenostrepta A66
         |    |         Otostomus A66
         |    |           |  i. s.: O. bartletti Adams 1866 A66
         |    |           |         O. canaliculatus A66
         |    |           |         O. pulcherrimus Adams 1866 A66
         |    |           |         O. scitus Adams 1866 A66
         |    |           |--O. (Leiostracus) maculatus A66
         |    |           |--O. (Leptomerus) heloicus A66
         |    |           `--O. (Mormus) A66
         |    |                |--O. (M.) exornatus A66
         |    |                `--O. (M.) toraylii A66
         |    |         Iberellus pyrenaicus (Rossmässler 1839) (see below for synonymy) A-Z04
         |    |         Dentellocaracolus Oppenheim 1890 S97
         |    |           |--D. (Prothelidomus) acrochordon (Oppenheim 1895) TTE93
         |    |           `--D. damnata (Brongniart 1810) TTE93
         |    |         Eremina desertorum H84
         |    |         Loganiopharynx rarus (Boissy 1840) TTE93
         |    |         Nanhaispira eversilabia Yu & Zhang 1982 TTE93
         |    |         Haplohelix anadenia PB27
         |    |         Burungaella PB27
         |    |           |--B. buhambaensis PB27
         |    |           |--B. imperforata PB27
         |    |           `--B. oscitans PB27
         |    |         Nakuruella PB27
         |    |           |--N. ahena PB27
         |    |           `--N. soror PB27
         |    |         Urguessella capillata PB27
         |    |         Zingis bequaerti PB27
         |    |--Helicinae BR05
         |    |--Ariantinae BR05
         |    `--Murellinae [Murelleae, Murellini] BR17
         |         |--Murella Pfeiffer 1877 BR05
         |         |    `--*M. muralis (Müller 1774) [=Helix muralis] BR17
         |         `--Tacheocampylaea Pfeiffer 1877 [Tacheocampylaeinae, Tacheocampylinae] BR05
         |              `--*T. raspaili (Payraudeau 1826) [=Helix raspaili] BR17
         `--Camaenidae [Bradybaenidae, Camaenoidea, Eulotidae, Eulotinae] BR17
              |  i. s.: Multiscapta Yu 1982 G88
              |           |--M. eversilabia G88
              |           `--M. xinyuensis Gu 1988 G88
              |         Exiligada Solem 1984 S97
              |           `--E. negriensis Iredale 1939 S88
              |         Amplirhagada S97
              |         Westraltrachia Iredale 1933 S97
              |           |--W. commoda (Iredale 1939) S88
              |           `--W. derbyi (Cox 1892) S97
              |         Mouldingia Solem 1984 S97
              |           `--M. orientalis Solem 1984 S88
              |         Pleurodontites diespiter (Dall 1890) AP90
              |         Galadistes Iredale 1938 SS10
              |           |--*G. liverpoolensis (Brazier 1872) [=Helix (Galaxias) liverpoolensis] SS10
              |           |--G. akubra Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--G. alleni Iredale 1943 SS10
              |           |--G. intervenens Iredale 1938 SS10
              |           |--G. marcescens (Cox 1868) SS10
              |           |--G. molong Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
              |           |--G. pilligaensis Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
              |           |--G. pustulosa Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--‘Helix’ stutchburyi Pfeiffer 1857 SS10
              |         Mussonena Iredale 1938 SS10
              |           |--*M. spinei (Cox 1868) (see below for synonymy) SS10
              |           |--M. boonah Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |           |--M. carnarvon Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--M. martinlogari Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--M. maxima Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--M. nogoa Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Nannochloritis Iredale 1938 SS10
              |           `--*N. layardi (Gude 1906) (see below for synonymy) SS10
              |         Calvigenia Iredale 1938 SS10
              |           |--*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. 2010 SS10
              |           `--C. mucida (Pfeiffer 1857) [=Helix mucida, Ramogenia mucida] SS10
              |         Austrochloritis SS10
              |         Squamagenia Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |           |--*S. yabba Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |           `--S. separanda (Iredale 1938) [=Austrochloritis separanda] SS10
              |         Meridolum Iredale 1942 SS10
              |           |--*M. jervisensis (Quoy & Gaimard 1832) [=Helix jervisensis] SS10
              |           |--M. corneovirens (Pfeiffer 1851) (see below for synonymy) SS10
              |           |--M. grayi (Pfeiffer 1847) SS10
              |           |--M. gulosum (Gould 1846) (see below for synonymy) SS10
              |           |--M. marshalli McLauchlan 1951 SS10
              |           |--M. maryae Clark 2009 SS10
              |           |--M. middenense McLauchlan 1954 SS10
              |           `--M. sheai Clark 2009 SS10
              |         Temporena Iredale 1933 SS10
              |           |--*T. whartoni (Cox 1871) [=Helix whartoni] SS10
              |           |--T. juliafoxae Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--T. sardalabiata (Cox 1871) (see below for synonymy) SS10
              |         Noctepuna Iredale 1933 SS10
              |           |--*N. poiretiana (Reeve 1852) [=Helix poiretiana] SS10
              |           |    |--N. p. poiretiana SS10
              |           |    `--N. p. clenchi McMichael 1958 SS10
              |           |--N. cerea (Hedley 1894) SS10
              |           |--N. mayana (Hedley 1899) SS10
              |           `--N. muensis (Hedley 1912) [=Papuina muensis] SS10
              |         Lamprellia Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*L. zebina (Brazier 1878) [=Helix zebina] SS10
              |           `--L. angulata Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Sphaerospira SS10
              |         Figuladra SS10
              |         Neveritis Iredale 1938 SS10
              |           |--*N. poorei (Gude 1907) [=Chloritis poorei; incl. C. thales Gude 1907] SS10
              |           |--N. aridorum (Cox 1866) SS10
              |           `--N. misella (Gude 1907) SS10
              |         Posorites Iredale 1933 SS10
              |           |--*P. fucata (Pfeiffer 1853) [=Helix fucata] SS10
              |           |--P. conscendens (Cox 1866) SS10
              |           `--P. turneri (Shirley 1921) SS10
              |         Thersites Pfeiffer 1855 SS10
              |           |--*T. richmondiana (Reeve 1852) (see below for synonymy) SS10
              |           |--T. darlingtoni Clench & Archer 1938 SS10
              |           |--T. mitchellae (Cox 1864) [incl. Annakelea peregrans Iredale 1937] SS10
              |           `--T. novaehollandiae (Gray 1834) [incl. Helix dupuyana Pfeiffer 1851] SS10
              |         Papuexul Iredale 1933 SS10
              |           `--*P. bidwilli (Reeve 1853) [=Helix bidwilli] SS10
              |         Patrubella Iredale 1938 SS10
              |           `--*P. buxtoni (Brazier 1880) [=Helix (Planispira) buxtoni] SS10
              |         Kandoschloritis Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
              |           `--*K. pustulosus Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
              |         Ramogenia Iredale 1938 SS10
              |           `--R. challengeri (Gude 1906) [incl. Chloritis obnubila Gude 1907, *R. obnubila] SS10
              |         Trachygenia Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*T. lanuginosa (Gude 1907) [=Chloritis lanuginosa, Ramogenia lanuginosa] SS10
              |           |--T. capiosetosa Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--T. microlepida Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--T. praecursoris (Hedley 1912) (see below for synonymy) SS10
              |         Eurytrachia Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*E. mucosa (Cox 1868) (see below for synonymy) SS10
              |           `--E. costulata Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |         Tolgachloritis Iredale 1933 SS10
              |           |--*T. jacksoni (Hedley 1912) [=Chloritis jacksoni] SS10
              |           |--T. campbelli (Iredale 1938) [=Mussonena campbelli] SS10
              |           |--T. costata Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--T. kondaparinga Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Gloreugenia Iredale 1933 SS10
              |           |--*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. 2010 SS10
              |           |--*B. hedleyi (Fulton 1907) [=Chloritis (Austrochloritis) hedleyi, Gloreugenia hedleyi] SS10
              |           `--B. gerrardstirlingi Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Crookshanksia Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--*C. frankstanisici Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Trozena Iredale 1938 SS10
              |           `--*T. morata Iredale 1938 SS10
              |         Protolinitis Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*P. terriirwinae Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--P. pusilla (Hedley 1912) SS10
              |         Aetholitis Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*A. agamemnon (Gude 1906) [=Chloritis agamemnon] SS10
              |           |--A. astaeus (Gude 1906) SS10
              |           |--A. francisae Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--A. gerdesae Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--A. janetzkiae Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Obsteugenia Iredale 1933 SS10
              |           `--*O. inflecta (Hedley 1912) [=Chloritis inflecta; incl. C. bilabiata Odhner 1917] SS10
              |         Trachiopsis Pilsbry 1893 SS10
              |           |--T. strangulata (Hombron & Jaquinot 1841) (see below for synonymy) SS10
              |           |--T. musgravensis Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |           |--T. railtoni Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--T. setosa Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Chloritisanax Iredale 1933 SS10
              |           `--*C. banneri (Pfeiffer 1863) [=Helix banneri] SS10
              |         Pommerhelix Clark 2009 SS10
              |           |--*P. carmelae Clark 2009 SS10
              |           |--P. depressa (Hedley 1901) SS10
              |           |--P. duralensis (Cox 1868) SS10
              |           |--P. exocarpi (Cox 1868) SS10
              |           |--P. insularis Clark 2009 SS10
              |           |--P. mastersi (Cox 1864) SS10
              |           |--P. monacha (Pfeiffer 1859) [incl. Meridolum bowdenae McLauchlan 1954] SS10
              |           `--P. stanisici Clark 2009 SS10
              |         Sauroconcha Zhang & Shea 2008 SS10
              |           `--*S. caperteeana Zhang & Shea 2008 SS10
              |         Petraconcha Clark 2009 SS10
              |           `--*P. griffithsi Clark 2009 SS10
              |         Ponderconcha Clark 2009 SS10
              |           |--*P. murphyi Clark 2009 SS10
              |           |--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. 2010 SS10
              |         Meridistes Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--*M. woolnoughi Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Stanisicia Clark 2009 SS10
              |           `--*S. macintyrensis Clark 2009 SS10
              |         Pallidelix Iredale 1933 SS10
              |           |--*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. 2010 SS10
              |         Ventopelita Iredale 1943 SS10
              |           `--*V. leucocheilus (Cox 1868) (see below for synonymy) SS10
              |         Moretonistes Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--*M. mansueta (Reeve 1854) (see bleow for synonymy) SS10
              |         Pseudodistes Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |           |--*P. biggenden Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |           `--P. reevesi Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Jimbouria Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*J. rodhobsoni Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--J. chrisalleni Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
              |         Adclarkia Stanisic 1996 SS10
              |           |--*A. dawsonensis Stanisic 1996 SS10
              |           |--A. cameroni Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--A. dulacca Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Lynfergusonia Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*L. mundubbera Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--L. taroomiana Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |         Perioinsolita Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |           `--*P. pokryszkoae Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
              |         Toombatrachia Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--*T. wheelerae Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Melostrachia Solem 1979 SS10
              |           |--*M. glomerans (Iredale 1938) [=Torresitrachia glomerans] SS10
              |           `--M. acuticostata (Fulton 1907) SS10
              |         Spurlingia Iredale 1933 SS10
              |           |--S. dunkiensis (Forbes 1851) (see below for synonymy) SS10
              |           |--S. excellens Iredale 1937 SS10
              |           |--S. gemma Iredale 1937 SS10
              |           |--S. monticola (Clench & Archer 1938) SS10
              |           |--S. portus Iredale 1937 SS10
              |           |--S. praehadra (Odhner 1917) SS10
              |           |--S. tinarooensis Iredale 1937 SS10
              |           `--S. undara Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Jacksonena Iredale 1937 SS10
              |           `--*J. rudis (Hedley 1912) [=Planispira rudis] SS10
              |         Forrestena Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--*F. delicata (Hedley 1912) [=Planispira delicata, Jacksonena delicata] SS10
              |         Zyghelix Iredale 1937 SS10
              |           `--*Z. forsteriana (Reeve 1852) (see below for synonymy) SS10
              |         Torresitrachia SS10
              |         Austrocamaena Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*A. ricnattrassi Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--A. thorntonensis Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Rhynchotrochus Möllendorff 1895 SS10
              |           |--*R. taylorianus (Adams & Reeve 1850) [=Helix tayloriana] SS10
              |           `--R. macgillivrayi (Forbes 1851) [incl. R. macgillivrayi extensior Iredale 1938] SS10
              |         Crikey Stanisic 2009 SS10
              |           `--*C. steveirwini Stanisic 2009 SS10
              |         Meliobba Iredale 1930 SS10
              |           `--*M. shafferyi Iredale 1940 SS10
              |         Gnarosophia Iredale 1933 SS10
              |           `--*G. bellendenkerensis (Brazier 1875) (see below for synonymy) SS10
              |         Bentosites Iredale 1933 SS10
              |           |--*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) (see below for synonymy) SS10
              |           |--B. gavisa Iredale 1933 (see below for synonymy) SS10
              |           `--B. hefferani Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Billordia Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*B. bridgettae Stanisic in Stanisic, Shea et al. 2010 SS10
              |           `--B. nicoletteae Stanisic in Stanisic, Shea et al. 2010 SS10
              |         Marilynessa Stanisic in Stanisic, Shea et al. 2010 SS10
              |           |--*M. yulei (Forbes 1851) (see below for synonymy) 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
              |         Steorra SS10
              |         Monteithosites Stanisic 1996 SS10
              |           `--*M. helicostracum Stanisic 1996 SS10
              |         Cristilabrum Solem 1981 S97
              |           `--C. kessneri Solem 1989 S97
              |         Quistrachia Iredale 1939 S97
              |           |  i. s.: *Q. monogramma (Ancey 1898) [=Trachia monogramma, Planispira monogramma] S97
              |           |         Q. herberti Solem 1997 S97
              |           |         Q. leptogramma (Pfeiffer 1846) (see below for synonymy) S97
              |           |--+--Q. legendrei Solem 1997 JS13, S97
              |           |  `--Q. turneri Solem 1997 JS13, S97
              |           `--+--Q. montebelloensis (Preston 1914) (see below for synonymy) JS13
              |              `--+--Q. lefroyi Solem 1997 JS13, S97
              |                 `--Q. warroorana Solem 1997 JS13, S97
              |         Setobaudinia S97
              |         Kimboraga Iredale 1933 S97
              |           |--K. cascadensis GK12
              |           |--K. exanima GK12
              |           |--K. glabra GK12
              |           `--K. wulalam GYD14
              |         Globorhagada Iredale 1933 S97
              |           |--G. confusa GK12
              |           |--G. uwinsensis GK12
              |           |--G. wunandarra GK12
              |           |--G. wurroolgu GK12
              |           `--G. yoowadan GK12
              |         Mesodontrachia Solem 1985 S97
              |         Parglogenia Iredale 1938 S97
              |         Ningbingia Solem 1981 S97
              |         Turgenitubulus Solem 1981 S97
              |           `--T. pagodula Solem 1985 S97
              |         Prymnbriareus Solem 1981 S97
              |         Ordtrachia Solem 1984 S97
              |         Prototrachia Solem 1984 S97
              |         Baudiniella Thiele 1931 S97
              |         Retroterra Solem 1985 S97
              |           |--R. aequabilis GK12
              |           `--R. discoidea GK12
              |         Kendrickia Solem 1985 S97
              |         Carinotrachia Solem 1985 S97
              |           |--C. admirale GK12
              |           |--C. carsoniana Solem 1985 S97
              |           `--C. elevata GK12
              |         Australocosmica augustae GK12
              |         Baudinella GK12
              |           |--B. boongareensis GK12
              |           |--B. occidentalis GK12
              |           |--B. setobaudinioides GK12
              |           |--B. thielei GK12
              |           `--B. tuberculata GK12
              |         Kimberleydiscus fasciatus GK12
              |         Kimberleymelon tealei GK12
              |         Kimberleytrachia GK12
              |           |--K. achernaria GK12
              |           |--K. aequum GYD14
              |           |--K. alphacentauri GYD14
              |           |--K. amplirhagadoides GK12
              |           |--K. canopi GK12
              |           |--K. chartacea GK12
              |           |--K. crawfordi GK12
              |           |--K. crucis GK12
              |           `--K. somniator GK12
              |         Molema stankowski GK12
              |         Olympus nimbus Simone 2010 S11
              |         Cathaica fasciola (Draparnaud 1805) [=Helix fasciola; incl. H. pyrrhozona] S01
              |         Kanabohelix kanabensis (White 1876) TTE93
              |         Coneulota diarti (Cossmann 1907) TTE93
              |--Camaeninae BR05
              |--Bradybaeninae BR17
              |--Hadrinae BR17
              `--Helicostylinae BR05
                   |  i. s.: ‘Helix’ pyrostoma C84
                   |--Helicostyla Férussac 1821 BR05
                   |    `--*H. mirabilis (Férussac 1821) [=Helix mirabilis] BR17
                   |--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 fortasse (Iredale 1933) [=Varohadra volgiola fortasse; incl. V. probleema Iredale 1933] 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

*Marilynessa yulei (Forbes 1851) [=Helix yulei; incl. H. basalis Pilsbry 1890, H. rainbirdi Cox 1870] SS10

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. yatalensis 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 barbara SS10, Cochlicella barbara BP90; incl. H. ventricosa Draparnaud 1801 SS10, C. ventricosa BP90, H. (Cochlicella) ventrosa Férussac 1821 SS10]

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

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

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

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[AP90] Auffenberg, K., & R. W. Portell. 1990. A new fossil land snail (Gastropoda: Pulmonata: Polygyridae) from the Middle Miocene of northern Florida. Nautilus 103 (4): 143–148.

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

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[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.

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[C06] Cuezzo, M. G. 2006. Systematic revision and cladistic analysis of Epiphragmophora Doering from Argentina and southern Bolivia (Gastropoda: Stylommatophora: Xanthonychidae). Malacologia 49 (1): 121–188.

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