Clausiliidae

Clausiliidae, also known by the common name door snails, is a taxonomic family of small, very elongate, mostly left-handed, air-breathing land snails, sinistral terrestrial pulmonate gastropod mollusks.
With over 1700 recognized recent and fossil species, this is among the most diverse families of terrestrial gastropods, although the marine gastropod family Pyramidellidae is larger.
Most species of Clausiliidae have an anatomical structure known as a clausilium, which enables the snail to close off the aperture of the shell with a sliding "door".

Shell description

Almost all the species of snails in the family of door snails are sinistral, which is an uncommon feature in gastropod shells in general.
These snails have shells which are extremely high-spired, with numerous whorls.
The shells tend to be club-shaped, tapering at both ends to a rounded nub. The aperture usually has visible folds.

The clausilium

Clausiliids are also very unusual among pulmonate gastropods in that most of them have a "door" or clausilium. The clausilium is not the same thing as an operculum, which does not exist at all in most pulmonate gastropods.
The clausilium is a calcareous structure, tongue-shaped or spoon-shaped, which can close the aperture of the snail shell to protect the soft parts against predation by animals such as carnivorous beetle larvae. The narrow end of the clausilium slides in the grooves that are formed by the folds on the inside of the shell.

Anatomy

In this family, the number of haploid chromosomes lies between 21 and 30.

Taxonomy

The type genus is Clausilia Draparnaud, 1805.
The family Clausiliidae is classified within the informal group Sigmurethra, itself belonging to the clade Stylommatophora within the clade Eupulmonata.

2005 taxonomy

The taxonomy of the Gastropoda by Bouchet & Rocroi, 2005 recognizes subfamilies as follows:
subfamily Clausiliinae Gray, 1855

tribe Baleini – synonyms: Laciniariini H. Nordsieck, 1963; Tristaniinae Schileyko, 1999
tribe Clausiliini Gray, 1855 – synonym: Fusulinae Lindholm, 1924
tribe Gracillariini H. Nordsieck, 1979

subfamily Alopiinae A. J. Wagner, 1913

tribe Alopiini A. J. Wagner, 1913
tribe Cochlodinini Lindholm, 1925 – synonym: Marpessinae Wenz, 1923
tribe Delimini Brandt, 1956 – synonym: Papilliferini Brandt, 1961
tribe Medorini H. Nordsieck, 1997
tribe Montenegrinini H. Nordsieck, 1972

subfamily † Constrictinae H. Nordsieck, 1981
subfamily Garnieriinae C. Boettger, 1926

tribe Garnieriini C. Boettger, 1926
tribe Tropidaucheniini H. Nordsieck, 2002

subfamily † Eualopiinae H. Nordsieck, 1978

tribe † Eualopiini H. Nordsieck, 1978
tribe † Rillyini H. Nordsieck, 1985

subfamily Laminiferinae Wenz, 1923
subfamily Mentissoideinae Lindholm, 1924

tribe Mentissoideini Lindholm, 1924 – synonym: Euxininae I. M. Likharev, 1962
tribe Acrotomini H. Nordsieck, 1979
tribe Boettgeriini H. Nordsieck, 1979
tribe Euxinellini Neubert, 2002
tribe Filosini H. Nordsieck, 1979
tribe Olympicolini Neubert, 2002
tribe Strigileuxinini H. Nordsieck, 1994
tribe Strumosini H. Nordsieck, 1994

subfamily Neniinae Wenz, 1923 – Neniastrinae H. B. Baker, 1930
subfamily Phaedusinae A. J. Wagner, 1922

† tribe Disjunctariini H. Nordsieck, 2014
tribe Megalophaedusini Zilch, 1954 – synonym: Zaptyxini Zilch, 1954
† tribe Nordsieckiini H. Nordsieck, 2007
tribe Phaedusini A. J. Wagner, 1922
† tribe Serrulellini H. Nordsieck, 2007
tribe Serrulinini Ehrmann, 1927
tribe Synprosphymini H. Nordsieck, 2007

subfamily Serrulininae Ehrmann, 1927

Genera

Genera include:

Clausiliinae

Tribe Acrotomini H. Nordsieck, 1979Acrotoma O. Boettger, 1881 – with four subgenera: Acrotoma, Acrotomina H. Nordsieck, 1977; Bzybia H. Nordsieck, 1977 and Castelliana Suvorov, 2002Akramowskia H. Nordsieck, 1975Armenica O. Boettger, 1877 – with two subgenera: Armenica and Astrogena Szekeres, 1970Inobseratella Lindholm, 1924Phrygica H. Nordsieck, 1994Roseniella Thiele, 1931 – with two subgenera: Chavchetia Neubert, 1992 and RoseniellaScrobifera O. Boettger, 1877Sprattia O. Boettger, 1883
tribe BaleiniAlinda H. & A. Adams, 1855 – with two subgenera: Alinda and Pseudalinda O. Boettger, 1877Balea J. E. Gray, 1824Bulgarica O. Boettger, 1877 – with three subgenera Bulgarica, Pavlovicia H. Nordsieck, 1973 and Strigilecula Kennard & Woodward, 1923Lacinaria Hartmann, 1842Likharevia H. Nordsieck, 1975Mentissa H. & A.Adams, 1855Mentissella H. Nordsieck, 1973Micropontica O. Boettger, 1881 – with two subgenera: Baleopsina Lindholm, 1924 and MicroponticaMucronaria O. Boettger, 1877 – with two subgenera: Index O. Boettger, 1877 and MucronariaQuadriplicata O. Boettger, 1878Strigillaria Vest, 1867Vestia Hesse, 1916 – with three subgenera: Brabenecia H. Nordsieck, 1974; Vestia and Vestiella H. Nordsieck, 1877
Tribe Boettgeriini H. Nordsieck, 1979Boettgeria O. Boettger, 1863 – with two subgenera: Boettgeriaand Loosjesiella Neubert & Groh, 1998Macroptychia O. Boettger, 1877Sabaeola Lindholm, 1925
tribe Clausiliini

† Canalicia O. Boettger, 1863 Clausilia Draparnaud, 1805 – with two subgenera: Clausilia and Strobeliella H. Nordsieck, 1977Erjavecia Brusina, 1870Erjaveciella H. Nordsieck, 1977Fusulus Fitzinger, 1833 – with two subgenera: Erjaveciella H. Nordsieck, 1877 and Fusulus
† Hollabrunnella H. Nordsieck, 2007 Julica H. Nordsieck, 1963Macrogastra Hartmann, 1841 – with three subgenera: Macrogastra, Pseudovestia H. Nordsieck, 1877 and Pyrostoma Vest, 1867Micridyla H. Nordsieck, 1973
† Parafusulus H. Nordsieck, 2007
† Pliodiptychia H. Nordsieck, 1978 Pseudidyla O. Boettger, 1877Pseudofusulus H. Nordsieck, 1977 – has the only one species: Pseudofusulus variansRuthenica Lindholm, 1924
† Trolliella H. Nordsieck, 1981

Tribe † Emarginariini H. Nordsieck, 2007

† Emarginaria O. Boettger, 1877

Tribe Euxinellini Neubert, 2002Euxinella H. Nordsieck, 1973
Tribe Filosini H. Nordsieck, 1979Filosa O. Boettger, 1877Idyla H. & A. Adams, 1855 – with two subgenera: Idyla and Strigilidyla H. Nordsieck, 1994
tribe Gracillariini H. Nordsieck, 1979Graciliaria Bielz, 1867

† Truciella H. Nordsieck, 1978

Tribe Mentissoideini Lindholm, 1924Elia H. & A.Adams, 1855 – with four subgenera: Acroeuxina O. Boettger, 1877; Caucasica O. Boettger, 1877; Elia and Megaleuxina O. Boettger, 1877Euxina O. Boettger, 1877 – with two subgenera: Euxina and Illunellaria Lindholm, 1924Euxinastra O. Boettger, 1888 – with two subgenera: Euxinastra and Odonteuxina H. Nordsieck, 1875Galeata O. Boettger, 1877Mentissoidea O. Boettger, 1877
Tribe Olympicolini Neubert, 2002Olympicola Hesse, 1916

† Scrobiferoides Neubert, 1996

Tribe Strigileuxinini H. Nordsieck, 1994Bitorquata O. Boettger, 1877Blaeneuxina Páll-Gergely & Szekeres, 2020Strigileuxina H. Nordsieck, 1975Sumelia H. Nordsieck, 1994
Tribe Strumosini H. Nordsieck, 1994Strumosa O. Boettger, 1877

Alopiinae A. J. Wagner, 1913

Alopiinae

† Dextrospira Hrubesch, 1965

tribe AlopiiniAlopia H. & A.Adams, 1855 – with two subgenera Alopia and Kimakowiczia Szekeres, 1969Herilla H. & A. Adams, 1855Montenegrina O. Boettger, 1877Protoherilla A. J. Wagner, 1921

† Protriloba H. Nordsieck, 2014 Triloba Vest, 1867

tribe Cochlodinini Lindholm, 1925 Cochlodina A. Férussac, 1821 – with four subgenera: Cochlodina; Cochlodinastra H. Nordsieck, 1977; Paracochlodina H. Nordsieck, 1969; Procochlodina H. Nordsieck, 1969Macedonica O. Boettger, 1877
tribe Delimini R. Brandt, 1956Barcania Brandt, 1956Charpentieria Stabille, 1864 – with subgenera: CharpentieriaDelima Hartmann, 1842 – with four subgenera: Delima; Dugiana Stamol & Slapnik, 2002; Piceata O. Boettger, 1877 and Semirugata O. Boettger, 1877Dilataria Vest, 1867Gibbularia Cecconi, 1908Mauritanica O. Boettger, 1879Papillifera Hartmann, 1842Sicania Tomlin, 1929Stigmatica O. Boettger, 1877
tribe Medorini H. Nordsieck, 1997Agathylla H. & A.Adams, 1855 – with two subgenera Agathylla and Agathyllina H. Nordsieck, 1969Albinaria Vest, 1867 – also includes former genusSericata O. Boettger, 1878.Carinigera Möllendorff, 1873Cristataria Vest, 1867Inchoatia Gittenberger & Uit de Weerd, 2006Isabellaria Vest, 1867Lampedusa O. Boettger, 1877 – with two subgenera Imitatrix Westerlund, 1884 and LampedusaLeucostigma A. J. Wagner, 1919Medora H. & A. Adams, 1855Muticaria Lindholm, 1925Strigilodelima A. J. Wagner, 1924Vallatia E. Gittenberger & Uit de Weerd, 2006

Constrictinae

Fossil subfamily Constrictinae contains genera:

† Constricta O. Boettger, 1877 – type genus of the subfamily
† Regiclausilia H. Nordsieck, 1981

Eualopiinae

Fossil subfamily Eualopiinae contains genera:

† Ambiguella H. Nordsieck, 2007 Formosana O. Boettger, 1877

Tribe † Eualopiini H. Nordsieck, 1978

† Eualopia O. Boettger, 1877 – type genus of the subfamily
† Monoptychia H. Nordsieck, 1972

Tribe † Rillyini † H. Nordsieck, 1985

† Neniopsilla H. Nordsieck, 2020
† Neniopsis Wenz, 1920
† Omanillya H. Nordsieck in Harzhauser et al., 2016
† Pararillya H. Nordsieck, 2002
† Proalbinaria O. Boettger in Oppenheim, 189
† Rillya Munier-Chalmas , 1883 – type genus of the tribe
† Neniopsilla H. Nordsieck, 2020
† Neniopsis Wenz, 1920
† Omanillya H. Nordsieck in Harzhauser et al., 2016
† Pararillya H. Nordsieck, 2002
† Proalbinaria O. Boettger in Oppenheim, 1895

Garnieriinae

tribe Garnieriini C. Boettger, 1926Garnieria Bourguignat, 1877
tribe Tropidaucheniini H. Nordsieck, 2002Euryauchenia H. Nordsieck, 2007Grandinenia Minato & Chen, 1984Indonenia Ehrmann, 1927Megalauchenia H. Nordsieck, 2007Tropidauchenia Lindholm, 1924 -

Laminiferinae

Bofilliella Ehrmann, 1927Neniatlanta Bouguignat, 1876Margaritiphaedusa H. Nordsieck, 2001Minatoia Hunyadi & Szekeres, 2016
Tribe Laminiferini Wenz, 1923

† Baboria Cossmann, 1898
† Laminifera O. Boettger, 1863 – the type genus of the family Laminiferinae. Its type species is fossil.
† Laminiplica H. Nordsieck, 1978
† Omanifera H. Nordsieck in Harzhauser et al., 2016

Tribe † Oospiroidesini H. Nordsieck, 2007

† Oospiroides Wenz, 1920

Tribe † Polloneriini H. Nordsieck, 2007

† Polloneria Sacco, 1886

Mentissoideinae

synonym of the tribe Mentissoideini Lindholm, 1924

Neniinae Wenz, 1923

Tribe Neniini Wenz, 1923Gonionenia Pilsbry, 1926Nenia H. & A. Adams, 1855Neniops Pilsbry, 1926Paranenia Rehder, 1939

Peruiniinae H. Nordsieck, 2005

Andinia Polinski, 1922Audiniastra H. Nordsieck, 2005Andiniella Weyrauch, 1958Bequaertinenia Weyrauch, 1964 – with two subgenera: Bequaertinenia and Miranenia Grego & Szekeres, 2004Brevinenia Neubert & H. Nordsieck, 2005Columbinia Polinski, 1924 – with subgenera: Columbinia and Steatonenia Pilsbry, 1926Cyclonenia H. Nordsieck, 1999Cylindronenia Ehrmann in Pilsbry, 1949 – with two subgenera: Cylindronenia and Cylindroneniella H. Nordsieck, 2007Ehrmanniella Zilch, 1949Gracilinenia Polinski, 1922Hemicena Pilsbry, 1949Incaglaia Pilsbry, 1949 – with two subgenera: Gibbonenia Zilch, 1954 and IncaglaiaIncania Polinski, 1922Leuconenia H. Nordsieck, 2005Neniaptyx H. Nordsieck, 2007Neniatracta Pilsbry, 1926Neniella Grego & Sekeres, 2004Parabalea Ancey, 1882Peruinia Polinski, 1922Pfeifferiella Weyrauch, 1957Pseudogracilinenia Loosjes & Loosjes-van Bemmel, 1984Steeriana Jousseaume, 1900Symptychiella H. Nordsieck, 1999 – with two subgenera: Divnenia H. Nordsieck, 2005 and SymptychiellaTemesa H. & A. Adams, 1855Zilchiella Weyrauch, 1957

Phaedusinae A. J. Wagner, 1922

Selenophaedusa Lindholm, 1924Siciliaria Vest, 1867
Tribe † Disjunctariini H. Nordsieck, 2014

† Disjunctaria O. Boettger, 1877

Tribe † Nordsieckiini H. Nordsieck, 2007

† Nordsieckia Truc, 1972
† Serruluna H. Nordsieck, 1981

tribe Phaedusini A. J. Wagner, 1922Acanthophaedusa H. Nordsieck, 2007Atractophaedusa Ehrmann, 1927Bacillophaedusa Grego & Szekeres, 2011Bathyptychia Lindholm, 1925 – with three subgenera: Bathyptychia; Brachyptychia H. Nordsieck, 2001 and Strictiphaedusa H. Nordsieck, 2001Castanophaedusa Páll-Gergely & Szekeres, 2017Celsiphaedusa H. Nordsieck, 2001Changphaedusa Motochin & Ueshima, 2017Cylindrophaedusa O. Boettger, 1877 – with three subgenera: Cylindrophaedusa and Montiphaedusa H. Nordsieck, 2002Dautzenbergiella Lindholm, 1924 – with two subgenera: Dautzenbergiella and Mansuyiella H. Nordsieck, 2003Euphaedusa O. Boettger, 1877 – with subgenera: Dentiphaedusa H. Nordsieck, 2003; Euphaedusa; and Telophaedusa H. Nordsieck, 2003Fuchsiana Gredler, 1887Hemiphaedusa O. Boettger, 1877 – with subgenera: Dendrophaedusa H. Nordsieck, 2002; Hemiphaedusa; Hemiphaedusoides H. Nordsieck, 2001; Hemizaptyx Pilsbry, 1905; Labyrinthiphaedusa H. Nordsieck, 2001; Pinguiphaedusa Azuma, 1982; Placeophaedusa Minato, 1994)Juttingia Loosjes, 1965 – with two subgenera: Juttingia and Pseudohemiphaedusa H. Nordsieck, 2002Liparophaedusa Lindholm, 1924Loosjesia H. Nordsieck, 2002Luchuphaedusa Pilsbry, 1901 – with two subgenera: Luchuphaedusa and Nesiophaedusa Pilsbry, 1905Macrophaedusa Moellendorff, 1883Macrophaedusella H. Nordsieck, 2001Megalophaedusa O. Boettger, 1877 – with four subgenera: Megalophaedusa; Mesophaedusa Ehrmann, 1929; Mesozaptyx Kuroda, 1963 and Mundiphaedusa Minato, 1979..Messageriella Páll-Gergely & Szekeres, 2017Miraphaedusa H. Nordsieck, 2005Musaphaedusa H. Nordsieck, 2018Nannophaedusa H. Nordsieck, 2012Notoptychia Ehrmann, 1927Oospira Blanford, 1872 – with subgenera Formosanella H. Nordsieck, 2003; Oospira; Paraformosella H. Nordsieck, 2003 and Siphonophaedusa Lindholm, 1924Papilliphaedusa H. Nordsieck, 2003Paraphaedusa O. Boettger, 1877Phaedusa H. & A.Adams, 1855 – with subgenera: Metaphaedusa H. Nordsieck, 2001; Phaedusa; Pseudophaedusa Tomiyama, 1984 Probosciphaedusa Z.-Y. Chen, 2021Reinia Kobelt, 1876 – with three subgenera: Parareinia H. Nordsieck, 1998; Pictophaedusa Azuma, 1982 and ReiniaSerriphaedusa H. Nordsieck, 2001Sinigena Lindholm, 1925Solitariphaedusa Motochin & Ueshima, 2017Stereophaedusa O. Boettger, 1877Streptodera Lindholm, 1925Tauphaedusa H. Nordsieck, 2003Tosaphaedusa Ehrmann, 1929Zaptyx Pilsbry, 1901 – with two subgenera: Prozaptyx Loosjes, 1950 and Zaptyx
Tribe † Serrulellini H. Nordsieck, 2007

† Serrulastra H. Nordsieck, 1981
† Serrulella H. Nordsieck, 1978

Tribe Serrulinini Ehrmann, 1927Caspiophaedusa Lindholm, 1924Cotyorica Grego & Szekeres, 2017Dobatia H. Nordsieck, 1973

† Euxinophaedusa Likharev, 1962 Graecophaedusa Rähle, 1982Laeviphaedusa Likharev & Stelkov, 1965Microphaedusa H. Nordsieck, 1978Nothoserrulina Németh & Szekeres, 1995Pamphylica Németh & Szekeres, 1995Pontophaedusa Lindholm, 1924Pontophaedusella H. Nordsieck, 1994Pravispira Lindholm, 1924Sciocochlea C. Boettger, 1935Serrulina Mousson, 1873Truncatophaedusa Majoros, Németh & Szili-Kovács 1994Tsoukatosia Gittenberger, 2000 – with two subgenera: Agiosspeleikos A. & P. Reischütz, 2003 and Tsoukatosia

Tribe Synprosphymini H. Nordsieck, 2007Excussispira Lindholm, 1925Synprosphyma A. J. Wagner, 1920

Synonyms

Diceratoptyx Pilsbry, 1905: synonym of Zaptyx Pilsbry, 1900 Heterozaptyx Pilsbry, 1906: synonym of Zaptyx Pilsbry, 1900 Kazancia Neubert, 1992: synonym of Lindholm, 1924 Leptacme Ehrmann, 1927: synonym of Oospira Grego & Szekeres, 2011 represented as Oospira W. T. Blanford, 1872Lindholmiella Ehrmann, 1927: synonym of Oospira Ehrmann, 1927 represented as Oospira W. T. Blanford, 1872Metazaptyx Pilsbry, 1905synonym of Zaptyx Pilsbry, 1900Neniauchenia H. Nordsieck, 2002: synonym of Grandinenia Minato & D.-N. Chen, 1984Neostyriaca A. J. Wagner, 1920: synonym of Clausilia A. J. Wagner, 1920 represented as Clausilia Draparnaud, 1805Oligozaptx Pilsbry, 1905: synonym of Zaptyx Pilsbry, 1905 represented as Zaptyx Pilsbry, 1900Parazaptyx Pilsbry, 1905: synonym of Zaptyx Pilsbry, 1905 represented as Zaptyx Pilsbry, 1900Pliciphaedusa H. Nordsieck, 1998: synonym of Stereophaedusa H. Nordsieck, 1998 represented as Stereophaedusa O. Boettger, 1877Pulchraptyx Minato, 1981: synonym of Zaptyx Minato, 1981 represented as Zaptyx Pilsbry, 1900Renschiphaedusa Loosjes & Loosjes-van Bemmel, 1973: synonym of Phaedusa H. Adams & A. Adams, 1855 Selenoptyx Pilsbry, 1908: synonym of Zaptyx Pilsbry, 1908 represented as Zaptyx Pilsbry, 1900Serrulinella H. Nordsieck, 1984: synonym of Serrulina H. Nordsieck, 1984 represented as Serrulina Mousson, 1873 Stereozaptyx Pilsbry, 1905: synonym of Zaptyx Pilsbry, 1905 represented as Zaptyx Pilsbry, 1900Thaumatoptyx Pilsbry, 1908: synonym of Zaptyx Pilsbry, 1908 represented as Zaptyx Pilsbry, 1900Tyrannophaedusa Pilsbry, 1901: synonym of Megalophaedusa Pilsbry, 1900 represented as Megalophaedusa O. Boettger, 1877Tyrannozaptyx Käufel, 1930: synonym of Zaptyx Käufel, 1930 represented as Zaptyx Pilsbry, 1900Zaptychopsis Ehrmann, 1927: synonym of Zaptyx Ehrmann, 1927 represented as Zaptyx Pilsbry, 1900

Conservation

Although non-marine molluscs appear to be exceptionally vulnerable to extinction, the IUCN Red list listed only 9 species from this family.