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
- 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 Garnieriinae C. Boettger, 1926
- tribe Garnieriini C. Boettger, 1926
- tribe Tropidaucheniini H. Nordsieck, 2002
- tribe † Eualopiini H. Nordsieck, 1978
- tribe † Rillyini H. Nordsieck, 1985
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 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
Genera
Genera include:Clausiliinae
Tribe Acrotomini H. Nordsieck, 1979- Acrotoma O. Boettger, 1881 – with four subgenera: Acrotoma, Acrotomina H. Nordsieck, 1977; Bzybia H. Nordsieck, 1977 and Castelliana Suvorov, 2002
- Akramowskia H. Nordsieck, 1975
- Armenica O. Boettger, 1877 – with two subgenera: Armenica and Astrogena Szekeres, 1970
- Inobseratella Lindholm, 1924
- Phrygica H. Nordsieck, 1994
- Roseniella Thiele, 1931 – with two subgenera: Chavchetia Neubert, 1992 and Roseniella
- Scrobifera O. Boettger, 1877
- Sprattia O. Boettger, 1883
- Alinda H. & A. Adams, 1855 – with two subgenera: Alinda and Pseudalinda O. Boettger, 1877
- Balea J. E. Gray, 1824
- Bulgarica O. Boettger, 1877 – with three subgenera Bulgarica, Pavlovicia H. Nordsieck, 1973 and Strigilecula Kennard & Woodward, 1923
- Lacinaria Hartmann, 1842
- Likharevia H. Nordsieck, 1975
- Mentissa H. & A.Adams, 1855
- Mentissella H. Nordsieck, 1973
- Micropontica O. Boettger, 1881 – with two subgenera: Baleopsina Lindholm, 1924 and Micropontica
- Mucronaria O. Boettger, 1877 – with two subgenera: Index O. Boettger, 1877 and Mucronaria
- Quadriplicata O. Boettger, 1878
- Strigillaria Vest, 1867
- Vestia Hesse, 1916 – with three subgenera: Brabenecia H. Nordsieck, 1974; Vestia and Vestiella H. Nordsieck, 1877
- Boettgeria O. Boettger, 1863 – with two subgenera: Boettgeriaand Loosjesiella Neubert & Groh, 1998
- Macroptychia O. Boettger, 1877
- Sabaeola Lindholm, 1925
- † Canalicia O. Boettger, 1863
- Clausilia Draparnaud, 1805 – with two subgenera: Clausilia and Strobeliella H. Nordsieck, 1977
- Erjavecia Brusina, 1870
- Erjaveciella H. Nordsieck, 1977
- Fusulus Fitzinger, 1833 – with two subgenera: Erjaveciella H. Nordsieck, 1877 and Fusulus
- † Hollabrunnella H. Nordsieck, 2007
- Julica H. Nordsieck, 1963
- Macrogastra Hartmann, 1841 – with three subgenera: Macrogastra, Pseudovestia H. Nordsieck, 1877 and Pyrostoma Vest, 1867
- Micridyla H. Nordsieck, 1973
- † Parafusulus H. Nordsieck, 2007
- † Pliodiptychia H. Nordsieck, 1978
- Pseudidyla O. Boettger, 1877
- Pseudofusulus H. Nordsieck, 1977 – has the only one species: Pseudofusulus varians
- Ruthenica Lindholm, 1924
- † Trolliella H. Nordsieck, 1981
- † Emarginaria O. Boettger, 1877
- Euxinella H. Nordsieck, 1973
- Filosa O. Boettger, 1877
- Idyla H. & A. Adams, 1855 – with two subgenera: Idyla and Strigilidyla H. Nordsieck, 1994
- Graciliaria Bielz, 1867
- † Truciella H. Nordsieck, 1978
- Elia H. & A.Adams, 1855 – with four subgenera: Acroeuxina O. Boettger, 1877; Caucasica O. Boettger, 1877; Elia and Megaleuxina O. Boettger, 1877
- Euxina O. Boettger, 1877 – with two subgenera: Euxina and Illunellaria Lindholm, 1924
- Euxinastra O. Boettger, 1888 – with two subgenera: Euxinastra and Odonteuxina H. Nordsieck, 1875
- Galeata O. Boettger, 1877
- Mentissoidea O. Boettger, 1877
- Olympicola Hesse, 1916
- † Scrobiferoides Neubert, 1996
- Bitorquata O. Boettger, 1877
- Blaeneuxina Páll-Gergely & Szekeres, 2020
- Strigileuxina H. Nordsieck, 1975
- Sumelia H. Nordsieck, 1994
- Strumosa O. Boettger, 1877
Alopiinae A. J. Wagner, 1913
Alopiinae- † Dextrospira Hrubesch, 1965
- Alopia H. & A.Adams, 1855 – with two subgenera Alopia and Kimakowiczia Szekeres, 1969
- Herilla H. & A. Adams, 1855
- Montenegrina O. Boettger, 1877
- Protoherilla A. J. Wagner, 1921
- † Protriloba H. Nordsieck, 2014
- Triloba Vest, 1867
- Cochlodina A. Férussac, 1821 – with four subgenera: Cochlodina; Cochlodinastra H. Nordsieck, 1977; Paracochlodina H. Nordsieck, 1969; Procochlodina H. Nordsieck, 1969
- Macedonica O. Boettger, 1877
- Barcania Brandt, 1956
- Charpentieria Stabille, 1864 – with subgenera: Charpentieria
- Delima Hartmann, 1842 – with four subgenera: Delima; Dugiana Stamol & Slapnik, 2002; Piceata O. Boettger, 1877 and Semirugata O. Boettger, 1877
- Dilataria Vest, 1867
- Gibbularia Cecconi, 1908
- Mauritanica O. Boettger, 1879
- Papillifera Hartmann, 1842
- Sicania Tomlin, 1929
- Stigmatica O. Boettger, 1877
- Agathylla H. & A.Adams, 1855 – with two subgenera Agathylla and Agathyllina H. Nordsieck, 1969
- Albinaria Vest, 1867 – also includes former genusSericata O. Boettger, 1878.
- Carinigera Möllendorff, 1873
- Cristataria Vest, 1867
- Inchoatia Gittenberger & Uit de Weerd, 2006
- Isabellaria Vest, 1867
- Lampedusa O. Boettger, 1877 – with two subgenera Imitatrix Westerlund, 1884 and Lampedusa
- Leucostigma A. J. Wagner, 1919
- Medora H. & A. Adams, 1855
- Muticaria Lindholm, 1925
- Strigilodelima A. J. Wagner, 1924
- Vallatia 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
- † Eualopia O. Boettger, 1877 – type genus of the subfamily
- † Monoptychia H. Nordsieck, 1972
- † 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, 1926
- Garnieria Bourguignat, 1877
- tribe Tropidaucheniini H. Nordsieck, 2002
- Euryauchenia H. Nordsieck, 2007
- Grandinenia Minato & Chen, 1984
- Indonenia Ehrmann, 1927
- Megalauchenia H. Nordsieck, 2007
- Tropidauchenia Lindholm, 1924 -
Laminiferinae
- Bofilliella Ehrmann, 1927
- Neniatlanta Bouguignat, 1876
- Margaritiphaedusa H. Nordsieck, 2001
- Minatoia Hunyadi & Szekeres, 2016
- † 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
- † Oospiroides Wenz, 1920
- † Polloneria Sacco, 1886
Mentissoideinae
synonym of the tribe Mentissoideini Lindholm, 1924Neniinae Wenz, 1923
Tribe Neniini Wenz, 1923- Gonionenia Pilsbry, 1926
- Nenia H. & A. Adams, 1855
- Neniops Pilsbry, 1926
- Paranenia Rehder, 1939
Peruiniinae H. Nordsieck, 2005
- Andinia Polinski, 1922
- Audiniastra H. Nordsieck, 2005
- Andiniella Weyrauch, 1958
- Bequaertinenia Weyrauch, 1964 – with two subgenera: Bequaertinenia and Miranenia Grego & Szekeres, 2004
- Brevinenia Neubert & H. Nordsieck, 2005
- Columbinia Polinski, 1924 – with subgenera: Columbinia and Steatonenia Pilsbry, 1926
- Cyclonenia H. Nordsieck, 1999
- Cylindronenia Ehrmann in Pilsbry, 1949 – with two subgenera: Cylindronenia and Cylindroneniella H. Nordsieck, 2007
- Ehrmanniella Zilch, 1949
- Gracilinenia Polinski, 1922
- Hemicena Pilsbry, 1949
- Incaglaia Pilsbry, 1949 – with two subgenera: Gibbonenia Zilch, 1954 and Incaglaia
- Incania Polinski, 1922
- Leuconenia H. Nordsieck, 2005
- Neniaptyx H. Nordsieck, 2007
- Neniatracta Pilsbry, 1926
- Neniella Grego & Sekeres, 2004
- Parabalea Ancey, 1882
- Peruinia Polinski, 1922
- Pfeifferiella Weyrauch, 1957
- Pseudogracilinenia Loosjes & Loosjes-van Bemmel, 1984
- Steeriana Jousseaume, 1900
- Symptychiella H. Nordsieck, 1999 – with two subgenera: Divnenia H. Nordsieck, 2005 and Symptychiella
- Temesa H. & A. Adams, 1855
- Zilchiella Weyrauch, 1957
Phaedusinae A. J. Wagner, 1922
- Selenophaedusa Lindholm, 1924
- Siciliaria Vest, 1867
- † Disjunctaria O. Boettger, 1877
- † Nordsieckia Truc, 1972
- † Serruluna H. Nordsieck, 1981
- Acanthophaedusa H. Nordsieck, 2007
- Atractophaedusa Ehrmann, 1927
- Bacillophaedusa Grego & Szekeres, 2011
- Bathyptychia Lindholm, 1925 – with three subgenera: Bathyptychia; Brachyptychia H. Nordsieck, 2001 and Strictiphaedusa H. Nordsieck, 2001
- Castanophaedusa Páll-Gergely & Szekeres, 2017
- Celsiphaedusa H. Nordsieck, 2001
- Changphaedusa Motochin & Ueshima, 2017
- Cylindrophaedusa O. Boettger, 1877 – with three subgenera: Cylindrophaedusa and Montiphaedusa H. Nordsieck, 2002
- Dautzenbergiella Lindholm, 1924 – with two subgenera: Dautzenbergiella and Mansuyiella H. Nordsieck, 2003
- Euphaedusa O. Boettger, 1877 – with subgenera: Dentiphaedusa H. Nordsieck, 2003; Euphaedusa; and Telophaedusa H. Nordsieck, 2003
- Fuchsiana Gredler, 1887
- Hemiphaedusa 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, 2002
- Liparophaedusa Lindholm, 1924
- Loosjesia H. Nordsieck, 2002
- Luchuphaedusa Pilsbry, 1901 – with two subgenera: Luchuphaedusa and Nesiophaedusa Pilsbry, 1905
- Macrophaedusa Moellendorff, 1883
- Macrophaedusella H. Nordsieck, 2001
- Megalophaedusa O. Boettger, 1877 – with four subgenera: Megalophaedusa; Mesophaedusa Ehrmann, 1929; Mesozaptyx Kuroda, 1963 and Mundiphaedusa Minato, 1979..
- Messageriella Páll-Gergely & Szekeres, 2017
- Miraphaedusa H. Nordsieck, 2005
- Musaphaedusa H. Nordsieck, 2018
- Nannophaedusa H. Nordsieck, 2012
- Notoptychia Ehrmann, 1927
- Oospira Blanford, 1872 – with subgenera Formosanella H. Nordsieck, 2003; Oospira; Paraformosella H. Nordsieck, 2003 and Siphonophaedusa Lindholm, 1924
- Papilliphaedusa H. Nordsieck, 2003
- Paraphaedusa O. Boettger, 1877
- Phaedusa H. & A.Adams, 1855 – with subgenera: Metaphaedusa H. Nordsieck, 2001; Phaedusa; Pseudophaedusa Tomiyama, 1984
- Probosciphaedusa Z.-Y. Chen, 2021
- Reinia Kobelt, 1876 – with three subgenera: Parareinia H. Nordsieck, 1998; Pictophaedusa Azuma, 1982 and Reinia
- Serriphaedusa H. Nordsieck, 2001
- Sinigena Lindholm, 1925
- Solitariphaedusa Motochin & Ueshima, 2017
- Stereophaedusa O. Boettger, 1877
- Streptodera Lindholm, 1925
- Tauphaedusa H. Nordsieck, 2003
- Tosaphaedusa Ehrmann, 1929
- Zaptyx Pilsbry, 1901 – with two subgenera: Prozaptyx Loosjes, 1950 and Zaptyx
- † Serrulastra H. Nordsieck, 1981
- † Serrulella H. Nordsieck, 1978
- Caspiophaedusa Lindholm, 1924
- Cotyorica Grego & Szekeres, 2017
- Dobatia H. Nordsieck, 1973
- † Euxinophaedusa Likharev, 1962
- Graecophaedusa Rähle, 1982
- Laeviphaedusa Likharev & Stelkov, 1965
- Microphaedusa H. Nordsieck, 1978
- Nothoserrulina Németh & Szekeres, 1995
- Pamphylica Németh & Szekeres, 1995
- Pontophaedusa Lindholm, 1924
- Pontophaedusella H. Nordsieck, 1994
- Pravispira Lindholm, 1924
- Sciocochlea C. Boettger, 1935
- Serrulina Mousson, 1873
- Truncatophaedusa Majoros, Németh & Szili-Kovács 1994
- Tsoukatosia Gittenberger, 2000 – with two subgenera: Agiosspeleikos A. & P. Reischütz, 2003 and Tsoukatosia
- Excussispira Lindholm, 1925
- Synprosphyma 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, 1872
- Lindholmiella Ehrmann, 1927: synonym of Oospira Ehrmann, 1927 represented as Oospira W. T. Blanford, 1872
- Metazaptyx Pilsbry, 1905synonym of Zaptyx Pilsbry, 1900
- Neniauchenia H. Nordsieck, 2002: synonym of Grandinenia Minato & D.-N. Chen, 1984
- Neostyriaca A. J. Wagner, 1920: synonym of Clausilia A. J. Wagner, 1920 represented as Clausilia Draparnaud, 1805
- Oligozaptx Pilsbry, 1905: synonym of Zaptyx Pilsbry, 1905 represented as Zaptyx Pilsbry, 1900
- Parazaptyx Pilsbry, 1905: synonym of Zaptyx Pilsbry, 1905 represented as Zaptyx Pilsbry, 1900
- Pliciphaedusa H. Nordsieck, 1998: synonym of Stereophaedusa H. Nordsieck, 1998 represented as Stereophaedusa O. Boettger, 1877
- Pulchraptyx Minato, 1981: synonym of Zaptyx Minato, 1981 represented as Zaptyx Pilsbry, 1900
- Renschiphaedusa 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, 1900
- Serrulinella 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, 1900
- Thaumatoptyx Pilsbry, 1908: synonym of Zaptyx Pilsbry, 1908 represented as Zaptyx Pilsbry, 1900
- Tyrannophaedusa Pilsbry, 1901: synonym of Megalophaedusa Pilsbry, 1900 represented as Megalophaedusa O. Boettger, 1877
- Tyrannozaptyx Käufel, 1930: synonym of Zaptyx Käufel, 1930 represented as Zaptyx Pilsbry, 1900
- Zaptychopsis Ehrmann, 1927: synonym of Zaptyx Ehrmann, 1927 represented as Zaptyx Pilsbry, 1900