Hadrinae

Rhagada convicta, copyright G. & Ph. Poppe.

Belongs within: Helicoidei.

The Hadrinae are a group of land snails found in Australia and southeast Asia united by molecular analysis. Members of this clade either lack a head wart or have the head wart (positioned between and just behind the ommatophores) exposed and not eversible (Solem 1997).

<==Hadrinae [Hadridae, Sinumeloninae] BR17
    |--Micromelon Solem 1992 S97
    |--Lacustrelix Iredale 1937 S97
    |--Basedowena Iredale 1937 S97
    |--Tatemelon Solem 1993 S97
    |--Eximiorhagada Iredale 1933 S97
    |--Montanomelon Solem 1993 S97
    |--Chloritis Beck 1837 BR17, BR05 [Chloritidae]
    |    `--*C. ungulina (Linnaeus 1758) [=Helix ungulina] BR17
    |--Papuina Martens 1860 BR17, BR05 [Papuinidae, Papuininae]
    |    `--*P. lituus (Lesson 1831) [=Helix lituus] BR17
    |--Calycia Adams 1865 BR17, BR05 [Calyciidae]
    |    `--*C. crystallina (Reeve 1848) [=Bulimus crystallinus] BR17
    |--Planispira Beck 1837 BR17, BR05 [Planispiridae]
    |    `--*P. zonaria (Linnaeus 1767) [=Helix zonaria] BR17
    |--Minimelon Solem 1993 S97
    |    `--M. colmani Solem 1993 S97
    |--Falspleuroxia Solem 1997 S97
    |    `--*F. overlanderensis Solem 1997 S97
    |--Promonturconchum Solem 1997 S97
    |    `--*P. superbum Solem 1997 S97
    |--Caperantrum Solem 1997 S97
    |    `--*C. polygyrum Solem 1997 S97
    |--Cristovala Clench 1958 BR17, BR05 [Cristovalinae]
    |    `--*C. tricolor (Pfeiffer 1849) [=Helix tricolor] BR17
    |--Granulomelon Iredale 1933 S97
    |    |--G. arcigerens (Tate 1894) S97
    |    `--G. gilleni Solem 1993 S97
    |--Strepsitaurus Solem 1997 S97
    |    |--*S. rugus (Cotton 1953) [=Pleuroxia ruga] S97
    |    |--S. cardabius Solem 1997 S97
    |    |--S. milyeringus Solem 1997 S97
    |    |--S. ningaloo Solem 1997 S97
    |    `--S. williami Solem 1997 S97
    |--Plectorhagada Iredale 1933 [incl. Idamera Iredale 1939] S97
    |    |--*P. plectilis (Benson 1853) S97 (see below for synonymy)
    |    |--P. carcharias (Pfeiffer 1864) W08 (see below for synonymy)
    |    |--P. gascoynensis (Smith 1894) (see below for synonymy) S97
    |    |--P. meilgana Solem 1997 S97
    |    |--P. rovina Iredale 1939 [=P. (*Idamera) rovina] S97
    |    `--P. scolythra Solem 1997 S97
    |--Hadra Martens 1860 BR17 [incl. Micardista Iredale 1933 SS10]
    |    |--*H. bipartita (Férussac 1823) [=Helix bipartita] SS10
    |    |--H. barneyi (Cox 1873) SS10
    |    |--H. bartschi (Marshall 1927) (see below for synonymy) SS10
    |    |--H. brunodavidi Stanisic in Stanisic, Shea et al. 2010 SS10
    |    |--H. funiculata (Reeve 1854) SS10
    |    |--H. semicastanea (Pfeiffer 1849) (see below for synonymy) SS10
    |    |--H. webbi (Pilsbry 1900) [incl. H. webbi incallida Iredale 1933] SS10
    |    `--H. wilsoni Solem 1979 S97
    |--Xanthomelon Martens 1860 [Xanthomelontidae, Xanthomelontinae] BR17
    |    |--*X. pomum (Pfeiffer 1842) [=Helix pomum] SS10
    |    |--X. distractum Iredale 1938 SS10
    |    |--X. jannellei (Le Guillou 1842) (see below for synonymy) SS10
    |    |--X. magnidicum Iredale 1938 [=X. pachystylum magnidicum] SS10
    |    |--X. obliquirugosa (Smith 1894) [=Globorhagada obliquirugosa] S97
    |    |--X. pachystylum (Pfeiffer 1845) (see below for synonymy) SS10
    |    |--X. prudhoensis (Smith 1894) [=Globorhagada prudhoensis] S97
    |    `--X. saginatum Iredale 1938 [=X. pachystylum saginatum] SS10
    |--Sinumelon Iredale 1930 BR05 [incl. Notobadistes Cotton & Godfrey 1932 S97]
    |    |--*S. nullaboricum (Tate 1879) BR17 (see below for synonymy)
    |    |--‘Helix’ bitaeniata Cox 1868 [=*Notobadistes bitaeniatus] S97
    |    |--S. gawleri Solem 1992 S97
    |    |--S. jimberlanensis Solem 1997 S97
    |    |--S. kalgum Iredale 1939 [incl. S. lennum Iredale 1939, S. lennum mutuum Iredale 1939] S97
    |    |--S. tarcoolanum Solem 1992 S97
    |    |--S. vagente Iredale 1939 S97
    |    `--S. wilpenensis (Tate 1894) S97
    |--Pleuroxia Ancey 1887 (see below for synonymy) S97
    |    |--*P. cyrtopleura (Pfeiffer 1862) [=Helix cyrtopleura] S97
    |    |--P. abstans Iredale 1939 [=P. (*Gantomia) abstans] S97
    |    |--P. adcockiana (Bednall 1894) S97
    |    |--P. bethana Solem 1997 S97
    |    |--P. carmeena Solem 1993 S97
    |    |--P. elfina Iredale 1939 [=P. polypleura elfina] S97
    |    |--P. everardenis (Bednall 1892) S97
    |    |--P. hinsbyi (Gude 1916) S97
    |    |--P. italowiana Solem 1992 S97
    |    |--P. oligopleura (Tate 1894) (see below for synonymy) S97
    |    |--P. phillipsiana (Angas 1873) S97
    |    |--P. polypleura (Tate 1899) [=Helix polypleura, Angasella polypleura; incl. P. commenta Iredale 1939] S97
    |    `--P. radiata (Hedley 1905) S97
    `--Rhagada Martens 1860 BR17 [incl. Bellrhagada Iredale 1938 S97, Tumegada Iredale 1939 S97; Rhagadidae, Rhagadinae]
         |  i. s.: *R. reinga (Gray 1846) BR17 [=Helix reinga BR17, Thersites (*Rhagada) reinga S97]
         |         R. angulata Solem 1997 S97
         |         R. basedowana Iredale 1939 S97
         |         R. biggeana GK12
         |         R. bulgana Solem 1997 S97
         |         R. crystalla Solem 1985 S97
         |         R. cygna Solem 1997 S97
         |         R. dampierana Solem 1997 S97
         |         R. dominica GYD14
         |         R. dringi (Pfeiffer 1846) [=Helix dringi, Bellrhagada dringi] S97
         |         R. elachystoma (von Martens 1878) [=Helix elachystoma, Thersites (Rhagada) elachystoma] S97
         |         R. felicitas GK12
         |         R. globosa Solem 1997 S97
         |         R. harti Solem 1985 S97
         |         R. herberti JHF06
         |         R. intermedia Solem 1997 S97
         |         R. kessneri GK12
         |         R. minima Solem 1997 S97
         |         R. primigena GK12
         |         R. radleyi Preston 1908 S97
         |         R. richardsonii (Smith 1874) (see below for synonymy) S97
         |         R. sheaei GK12
         |         R. supracostulata Schepman 1892 S97
         |         R. tescorum (Benson 1853) [=Helix tescorum] S97
         |         R. torulus (Férussac 1819) [=Helix torulus, Thersites (Rhagada) torulus] S97
         |--+--R. barrowensis Johnson, Stankowski et al. 2013 JS13
         |  `--+--R. convicta (Cox 1870) JS13, S97 (see below for synonymy)
         |     `--R. pilbarana Solem 1997 JS13, S97
         `--+--R. capensis Solem 1997 JS13, S97
            `--+--R. perprima Iredale 1939 JS13, S97 [=R. convicta perprima S97]
               `--R. plicata Preston 1914 JS13, S97 [=*Bellrhagada plicata S97]

Hadra bartschi (Marshall 1927) [incl. Thersites (Hadra) bartschi cepa Marshall 1927, T. (H.) dalli Marshall 1927, T. (H.) bartschi diva Marshall 1927, T. (H.) bartschi elfa Marshall 1927, T. (H.) bartschi fama Marshall 1927, T. (H.) bartschi mobiagensis Marshall 1927, T. (H.) bartschi murrayensis Marshall 1927, T. (H.) bartschi nesia Marshall 1927, T. (H.) bartschi nura Marshall 1927, T. (H.) bartschi oma Marshall 1927, T. (H.) bartschi paulensis Marshall 1927, T. (H.) bastschi quaesita Iredale 1937, T. (H.) waltoni Marshall 1927, T. (H.) bartschi yamensis Marshall 1927] SS10

Hadra semicastanea (Pfeiffer 1849) [incl. Thersites (Hadra) semicastanea alma Marshall 1927, Hadra blighi Iredale 1937, T. (H.) lizardensis Marshall 1927, T. (H.) lizardensis rada Marshall 1927, T. (H.) lizardensis suma Marshall 1927] SS10

Plectorhagada carcharias (Pfeiffer 1864) W08 [=Helix carcharias S97, H. (Dorcasia) carcharias S97, Thersites (Rhagada) carcharias S97]

Plectorhagada gascoynensis (Smith 1894) [=Helix (Trachia) gascoynensis, Planispira gascoynensis, Pleuroxia gascoynensis] S97

*Plectorhagada plectilis (Benson 1853) S97 [=Helix plectilis S97, Amplirhagada plectilis W08, H. (Dorcasia) plectilis S97, Thersites (Rhagada) plectilis S97; incl. H. paleata Reeve 1854 S97]

Pleuroxia Ancey 1887 [incl. Angasella Adams 1864 non Angasiella Crosse 1864, Angasietta Iredale 1939, Gantomia Iredale 1939] S97

Pleuroxia oligopleura (Tate 1894) [=Hadra oligopleura, Angasella oligopleura; incl. A. lemani Gude 1916, Pleuroxia lemani, P. oligopleura numba Iredale 1939] S97

Rhagada convicta (Cox 1870) JS13, S97 [=Helix convicta S97, R. (*Tumegada) convicta S97, Thersites (Rhagada) convicta S97]

Rhagada richardsonii (Smith 1874) [=Helix richardsonii, Thersites (Rhagada) richardsonii; incl. Rhagada convicta strella Iredale 1939, R. convicta tambra Iredale 1939] S97

*Sinumelon nullaboricum (Tate 1879) BR17 [=Helix nullaborica BR17, H. (Xanthomelon) nullarborica S97, Notobadistes nullarborica S97, Thersites (Badistes) nullarborica S97; incl. Sinumelon datum Iredale 1939 S97]

Xanthomelon jannellei (Le Guillou 1842) [=Helix jannellei; incl. H. pachystyla daemeli von Martens 1869, H. pachystyloides Cox 1868] SS10

Xanthomelon pachystylum (Pfeiffer 1845) [incl. X. pachystylum noscitum Iredale 1938, Thersites pachystyla subfuscozonata Cox 1894] SS10

*Type species of generic name indicated

REFERENCES

[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.

[GK12] Gibson, L. A., & F. Köhler. 2012. Determinants of species richness and similarity of species composition of land snail communities on Kimberley islands. Records of the Western Australian Museum Supplement 81: 41–66.

[GYD14] Gibson, L. A., S. Yates & P. Doughty (eds) 2012–2014. Biodiversity Values on Selected Kimberley Islands, Australia. Records of the Western Australian Museum Supplement 81.

[JHF06] Johnson, M. S., Z. R. Hamilton & J. Fitzpatrick. 2006. Genetic diversity of Rhagada land snails on Barrow Island. Journal of the Royal Society of Western Australia 89 (2): 45–50.

[JS13] Johnson, M. S., S. Stankowski, C. S. Whisson, R. J. Teale & Z. R. Hamilton. 2013. Camaenid land snails on Barrow Island: distributions, molecular phylogenetics and taxonomic revision. Records of the Western Australian Museum Supplement 83: 159–171.

[S97] Solem, A. 1997. Camaenid land snails from Western and central Australia (Mollusca: Pulmonata: Camaenidae). VII. Taxa from Dampierland through the Nullarbor. Records of the Western Australian Museum Supplement 50: 1461–1906.

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

[W08] Wilson, B. 2008. Terrestrial gastropods of Faure Island, Shark Bay, Western Australia. Records of the Western Australian Museum Supplement 75: 21–24.

Sphaerospira

Sphaerospira mortenseni, copyright Claude & Amandine Evanno.

Belongs within: Helicoidei.

Sphaerospira is a genus of large snails found in southeastern Queensland and northeastern New South Wales characterised by a black mantle (Stanisic et al. 2010).

Characters (from Stanisic et al. 2010): Shell large to very large, globose to conically globose, with a low to elevated spire; whorls rounded, sutures impressed; protoconch with weak, crowded radial ridges, teleoconch smooth, microsculpture of very fine periostracal radial ridges; lip thickened and reflected, blue-black; umbilicus closed. Colour tawny yellow with numerous black spiral bands and a dark suffusion behind the lip, to all brown with bands visible only on early whorls; rarely all-yellow. Animal with black mantle.

<==Sphaerospira Mörch 1867 [incl. Varohadra Iredale 1933] SS10
    |--*S. fraseri (Griffith & Pidgeon 1833) (see below for synonymy) SS10
    |--S. arthuriana (Cox 1873) [incl. Varohadra halleyae Iredale 1937] SS10
    |--S. bencarlessi Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
    |--S. blomfieldi (Cox 1864) (see below for synonymy) SS10
    |--S. informis Mousson 1869 [incl. S. informis dietrichae Iredale 1937, S. informis fringilla Iredale 1937] SS10
    |--S. mortenseni (Iredale 1929) SS10
    |--S. mossmani (Brazier 1875) [=Helix (Hadra) mossmani] SS10
    |--S. oconnellensis (Cox 1871) (see below for synonymy) SS10
    |--S. rockhamptonensis (Cox 1873) (see below for synonymy) SS10
    `--S. sidneyi (Iredale 1933) [=Bentosites blomfieldi sidneyi] SS10

Sphaerospira blomfieldi (Cox 1864) [incl. Thersites concors Fulton 1904, S. blomfieldi warroensis Hedley & Musson 1892] SS10

*Sphaerospira fraseri (Griffith & Pidgeon 1833) [=Helix fraseri; incl. H. capucinus Férussac 1835, H. coarctata Férussac 1839, Sphaerospira fraseri feriarum Iredale 1937, H. fraseri flavescens Hedley 1888 non H. flavescens Pfeiffer 1848, Bentosites blomfieldi latior Iredale 1933, Sphaerospira parallela Iredale 1937, S. fraseri permuta Iredale 1937] SS10

Sphaerospira oconnellensis (Cox 1871) [incl. S. oconnellensis caroli Iredale 1933, S. oconnellensis jacksoni Iredale 1933] SS10

Sphaerospira rockhamptonensis (Cox 1873) [incl. S rockhamptonensis decreta Iredale 1937, Helix moresbyi Angas 1876, H. planibasis Brazier 1881] SS10

*Type species of generic name indicated

REFERENCES

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

Austrochloritis

Austrochloritis sp., copyright Catching the Eye.

Belongs within: Helicoidei.

Austrochloritis is a genus of snails found in southeastern Australia with a teleoconch sculpture including periostracal setae and wavy ridgelets (Stanisic et al. 2010).

Characters (from Stanisic et al. 2010): Shell small to large, subdiscoidal to turbinate with a flat to elevated spire; whorls rounded, shouldered to angulate or keeled, sutures moderately to strongly impressed; protoconch sculpture of scattered pustules and vague rugose radial ridges, teleoconch sculpture of coarse radial ridges to smooth, with curved or straight, crowded to widely spaced periostracal setae, microsculpture of wavy periostracal ridgelets; lip reflected, often with a sulcus and swelling immediately behind; umbilicus very narrow to moderately open, V-shaped. Colour dark to light brown, lip white to brown. Animal dark grey or black to orange-brown.

<==Austrochloritis Pilsbry 1890 SS10
    |--*A. porteri (Cox 1866) [=Helix porteri] SS10
    |--A. abbotti Stanisic in Stanisic, Shea et al. 2010 SS10
    |--A. abrotonus Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. ascensa Iredale 1943 SS10
    |--A. ballengarra Stanisic in Stanisic, Shea et al. 2010 SS10
    |--A. bellengerensis (Cox 1871) [incl. Helix bellingenensis Brazier 1871] SS10
    |--A. brevipila (Pfeiffer 1849) [incl. Chloritis novocambrica Gude 1906] SS10
    |--A. congarinni Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
    |--A. cunninghamiana Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
    |--A. disjuncta (Gude 1906) SS10
    |--A. illawarra Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. iluka Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
    |--A. kanangra Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. kaputarensis Stanisic in Stanisic, Shea et al. 2010 SS10
    |--A. kiparra Stanisic in Stanisic, Shea et al. 2010 SS10
    |--A. kosciuszkoensis Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. laurieton Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. liverpoolensis Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. marksandersi Stanisic in Stanisic, Shea et al. 2010 SS10
    |--A. metuenda Iredale 1938 SS10
    |--A. monita Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. nambucca Iredale 1943 SS10
    |--A. niangala Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. nundinalis Iredale 1943 SS10
    |--A. paucisetosa Stanisic in Stanisic, Shea et al. 2010 SS10
    |--A. seaviewensis Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
    |--A. speculoris Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. stanisici Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. sydneyensis Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. timorensis Stanisic in Stanisic, Shea et al. 2010 SS10
    |--A. toonumbar Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    |--A. warrumbunglensis Shea & Griffiths in Stanisic, Shea et al. 2010 SS10
    `--A. wollemiensis Shea & Griffiths in Stanisic, Shea et al. 2010 SS10

*Type species of generic name indicated

REFERENCES

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

Steorra

Steorra bala, copyright Claude & Amandine Evanno.

Belongs within: Helicoidei.

Steorra is a genus of large snails with a banded or monochrome shell found in northeastern Queensland (Stanisic et al. 2010).

Characters (from Stanisic et al. 2010): Shell large to very large, solid to fairly thin, turbinately globose to trochoidally globose with a low to elevated conical spire; whorls rounded to subangulate, sutures impressed; protoconch with weak radial ridges, teleoconch smooth, microsculpture of crowded pustules or periostracal scales on the early whorls, and widely spaced to crowded periostracal microradial ridges on the later whorls; lip reflected, occasionally thickened; umbilicus narrowly open to closed. Colour tawny yellow with numerous dark brown spiral bands, dark brown suffusion behind lip and dark brown circum-umbilical patch, to all-over monochrome dark reddish brown; lip reflected, dark brown.

<==Steorra Stanisic in Stanisic, Shea et al. 2010 SS10
    |--*S. mourilyani (Brazier 1875) (see below for synonymy) SS10
    |--S. bala (Brazier 1878) [=Helix bala] SS10
    |--S. bellaria (Iredlae 1937) [=Gnarosophia bellaria] SS10
    |--S. cookensis (Brazier 1875) (see below for synonymy) SS10
    |--S. crookshanksi Stanisic in Stanisic, Shea et al. 2010 SS10
    |--S. estherlilleyae Stanisic in Stanisic, Shea et al. 2010 SS10
    |--S. jimfergusoni Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
    |--S. mazee (Brazier 1878) [=Helix mazee] SS10
    |--S. mulgravensis (Brazier 1872) (see below for synonymy) SS10
    |--S. rawnesleyi (Brazier 1873) [=Helix (Camaena) rawnesleyi] SS10
    |--S. tomsoni (Brazier 1876) [=Helix (Hadra) tomsoni] SS10
    `--S. worsfoldi Stanisic in Stanisic, Shea et al. 2010 SS10

Steorra cookensis (Brazier 1875) [=Helix (Hadra) cookensis; incl. H. bebias Brazier 1878, Pallidelix spurlingi Iredale 1937] SS10

*Steorra mourilyani (Brazier 1875) [=Helix (Hadra) mourilyani; incl. Varohadra banfieldi Iredale 1937, H. (H.) hanni Brazier 1876, H. (H.) hilli Brazier 1875, Varohadra hubbardi Iredale 1937, H. (H.) johnstonei Brazier 1875, V. russelli Iredale 1937] SS10

Steorra mulgravensis (Brazier 1872) [=Helix mulgravensis; incl. Gnarosophia palmensis austrina Iredale 1933, H. palmensis meridionalis Brazier 1881, G. mitifica Iredale 1933, H. incei multifasciata Cox 1864 non H. multifasciata Weinland & Anton 1857, H. (Hadra) palmensis Brazier 1876] SS10

*Type species of generic name indicated

REFERENCES

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

Figuladra

Pale banded snail Figuladra mattea, from the Queensland Museum.

Belongs within: Helicoidei.

Figuladra is a genus of snails found in southeastern Queensland possessing a globose, almost smooth shell and red mantle (Stanisic et al. 2010).

Characters (from Stanisic et al. 2010): Shell large to very large, loosely to tightly coiled, depressedly globose to trochoidally globose with a dome-shaped spire; whorls rounded and often laterally compressed, sutures impressed; protoconch with weak, crowded radial ridges, teleoconch smooth, microsculpture of extremely fine, periostracal microradial threads; lip reflected; umbilicus narrowly open to almost closed. Colour yellowish with few to numerous brown spiral bands to monochrome yellow; lip white, often with dark brown suffusion behind it and extending to umbilical area. Animal pinkish orange with red mantle.

<==Figuladra Iredale 1933 SS10
    |--F. incei (Philippi 1846) SS10
    |    |--F. i. incei (see below for synonymy) SS10
    |    |--*F. i. curtisiana (Pfeiffer 1864) (see below for synonymy) SS10
    |    `--F. i. lessoni (Pfeiffer 1846) (see below for synonymy) SS10
    |--F. appendiculata (Reeve 1854) [incl. Helix thatcheri Cox 1870] SS10
    |--F. barneyae Stanisic in Stanisic, Shea et al. 2010 SS10
    |--F. bayensis (Brazier 1875) [=Helix (Hadra) bayensis] SS10
    |--F. challisi (Cox 1873) SS10
    |--F. mattea (Iredale 1933) SS10
    |--F. muirorum Stanisic in Stanisic, Shea et al. 2010 SS10
    |--F. narelleae Stanisic & Potter in Stanisic, Shea et al. 2010 SS10
    |--F. pallida (Hedley & Musson 1892) [=Helix rockhamptonensis pallida, Hadra rockhamptonensis pallida] SS10
    |--F. reducta (Iredale 1937) [=Varohadra bayensis reducta] SS10
    `--F. volgiola (Iredale 1933) [incl. Helix andersoni Cox 1872 non Blanford 1869] SS10

*Figuladra incei curtisiana (Pfeiffer 1864) [=Helix curtisiana; incl. Varohadra curtisiana exedra Iredale 1933, H. (Hadra) parsoni Cox 1872, H. seminigra Morelet 1864] SS10

Figuladra incei incei (Philippi 1846) [incl. Helix (Camaena) praetermissi; incl. Varohadra keppelensis degener Iredale 1937, V. keppelensis Iredale 1937] SS10

Figuladra incei lessoni (Pfeiffer 1846) [incl. Helix (Camaena) aureedensis Brazier 1872, Varohadra bernhardi Iredale 1933, H. (Hadra) yeppoonensis Beddome 1897] SS10

*Type species of generic name indicated

REFERENCES

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

Bradybaeninae

Euhadra quaesita, from Davison et al. (2005).

Belongs within: Helicoidei.

The Bradybaeninae are a group of snails found in Eurasia and Africa, characterised by a dart sac bearing two divided glands with one to two accessory sacs.

<==Bradybaeninae BR05
    |--Aegista Albers 1850 BR17 [Aegistinae, Aegistini BR05]
    |    `--*A. chinensis (Philippi 1845) [=Helix chinensis] BR17
    |--Bradybaenini BR05
    |    |--Buliminopsis Heude 1890 [Buliminopsinae] BR05
    |    |    `--*B. buliminus (Heude 1882) [=Helix buliminus] BR17
    |    |--Bradybaena Beck 1837 SS10
    |    |    `--*B. similaris (Rang 1831) BR17 [=Helix similaris SS10, H. (Dorcasia) similaris TC89]
    |    `--Fruticicola Held 1837 [=Eulota Hartmann 1840; Fruticicoleae, Fruticicolidae, Fruticicolinae] BR17
    |         |--*F. fruticum (Müller 1774) [=Helix fruticum, *Eulota fruticum] BR17
    |         `--‘Eulota (Acusta)’ ravida S01
    `--Euhadra Pilsbry 1890 [Euhadrinae, Euhadrini] BR05
         |--*E. peliomphala (Pfeiffer 1850) [=Helix peliomphala] BR17
         |--E. calizona T07
         |    |--E. c. calizona T07
         |    `--E. c. amaliae T07
         |--E. congenita UA03
         |--E. dixoni T07
         |--E. herklotsi HF01
         |--E. senckenbergiana UA03
         `--+--E. aomoriensis UA03
            `--E. quaesita UA03

*Type species of generic name indicated

REFERENCES

[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.

[HF01] Hwang, U. W., M. Friedrich, D. Tautz, C. J. Park & W. Kim. 2001. Mitochondrial protein phylogeny joins myriapods with chelicerates. Nature 413: 154–157.

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

[S01] Sturany, R. 1901. Csigák és kagylók [Mollusken]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 69–74. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[TC89] Tapparone Canefri, T. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XVIII.—Molluschi terrestri e d’acqua dolce. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 295–359.

[T07] Tsurusaki, N. 2007. The Chugoku Mountains—a hotspot of geographical differentiation of species. Taxa 22: 3–14.

[UA03] Ueshima, R., & T. Asam. 2003. Single-gene speciation by left-right reversal. Nature 425: 679.

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, 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. 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.

<==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
         |--Helicodontidae 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 AP90
         |--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
         |--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 below 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. 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 barbara SS10, Cochlicella barbara BP90; incl. H. ventricosa Draparnaud 1801 SS10, Bulimus ventricosus G40, Cochlicella 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. 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

[A66] Adams, H. 1866. List of land and freshwater shells collected by Mr. E. Bartlett on the Upper Amazons, and on the River Ucayali, eastern Peru, with descriptions of new species. Proceedings of the Zoological Society of London 1866: 440–445.

[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.

[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.

[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.

[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.

[C84] Cain, A. J. 1984. Heterosematism in snails. Malacologia 25 (1): 161–172.

[C97] Cuezzo, M. G. 1997. Cryptostrakon corcovadensis, a new species of semislug from Costa Rica (Helicoidea: Xanthonychidae) with comments on the systematic position of the genus. American Malacological Bulletin 14 (1): 1–8.

[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.

[E99] Esu, D. 1999. Contribution to the knowledge of Neogene climate changes in western and central Europe by means of non-marine molluscs. In: J. Agustí, L. Rook & P. Andrews (eds) The Evolution of Neogene Terrestrial Ecosystems in Europe pp. 328–354. Cambridge University Press.

[GK12] Gibson, L. A., & F. Köhler. 2012. Determinants of species richness and similarity of species composition of land snail communities on Kimberley islands. Records of the Western Australian Museum Supplement 81: 41–66.

[GYD14] Gibson, L. A., S. Yates & P. Doughty (eds) 2012–2014. Biodiversity Values on Selected Kimberley Islands, Australia. Records of the Western Australian Museum Supplement 81.

[G40] Gray, J. E. 1840. A Manual of the Land and Fresh-water Shells of the British Islands, with figures of each of the kinds. By William Turton, M.D. A new edition, thoroughly revised and much enlarged. Longman, Orme, Brown, Green, and Longmans: London.

[G88] Gu H.-L. 1988. Fossils of land snails from Xinyu Formation (early Tertiary) of Qingjiang Basin, Jiangxi Province. Acta Palaeontologica Sinica 27 (1): 111–124.

[H84] Heller, J. 1984. Shell colours of desert landsnails. Malacologia 25 (2): 355–359.

[HM07] Honek, A., & Z. Martinkova. 2007. A field method for quantifying the grazing activity of slugs, with particular reference to Arion lusitanicus (Mollusca). Malacologia 49 (2): 273–281.

[JS13] Johnson, M. S., S. Stankowski, C. S. Whisson, R. J. Teale & Z. R. Hamilton. 2013. Camaenid land snails on Barrow Island: distributions, molecular phylogenetics and taxonomic revision. Records of the Western Australian Museum Supplement 83: 159–171.

[KC11] Kocot, K. M., J. T. Cannon, C. Todt, M. R. Citarella, A. B. Kohn, A. Meyer, S. R. Santos, C. Schander, L. L. Moroz, B. Lieb & K. M. Halanych. 2011. Phylogenomics reveals deep molluscan relationships. Nature 477: 452–456.

[PB27] Pilsbry, H. A., & J. Bequaert. 1927. The aquatic mollusks of the Belgian Congo, with a geographical and ecological account of Congo malacology. Bulletin of the American Museum of Natural History 53 (2): 69–602, pls 10–77.

[S11] Simone, L. R. L. 2011. Phylogeny of the Caenogastropoda (Mollusca), based on comparative morphology. Arquivos de Zoologia 42 (4): 161–323.

[S88] Solem, A. 1988. Non-camaenid land snails of the Kimberley and Northern Territory, Australia. I. Systematics, affinities and ranges. Invertebrate Taxonomy 4: 455–604.

[S97] Solem, A. 1997. Camaenid land snails from Western and central Australia (Mollusca: Pulmonata: Camaenidae). VII. Taxa from Dampierland through the Nullarbor. Records of the Western Australian Museum Supplement 50: 1461–1906.

[SS10] Stanisic, J., M. Shea, D. Potter & O. Griffiths. 2010. Australian Land Snails vol. 1. A. field guide to eastern Australian species. Bioculture Press: Mauritius.

[S01] Sturany, R. 1901. Csigák és kagylók [Mollusken]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 69–74. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[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.

Last updated: 26 June 2020.