Clausiliidae

Clausiliidae, also known by their common name the door snails, are a taxonomic family of small, very elongate, mostly left-handed, air-breathing land snails, sinistral terrestrial pulmonate gastropod mollusks.

Clausiliidae
Cochlodina laminata
Scientific classification
Kingdom:
Animalia
Phylum:
Mollusca
Class:
Gastropoda
(unranked):
clade Heterobranchia clade Euthyneura
clade Panpulmonata
clade Eupulmonata
clade Stylommatophora
informal group Sigmurethra
Superfamily:
Clausilioidea
Family:
Clausiliidae

Type genus
Clausilia

With about 1,300 species recent and fossil, this belong among the most diverse families of land gastropods (cf. Orthalicidae, 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
Megalophaedusa martensi is the largest species of the family Clausiliidae in the world. The individual in the photograph has a shell which is 45.5 mm long. The whole snail weighs 3.2 g live.

Almost all the species of snails in the family of door snails are left-handed, 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
Main article: clausilium
The clausilium of Clausilia dubia

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 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 (according to the values in this table).[2]

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 (according to the taxonomy of the Gastropoda by Bouchet & Rocroi, 2005).[3]

2005 taxonomy

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

subfamily Clausiliinae Gray, 1855

  • 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 (1923) - synonym: Marpessinae Wenz, 1923
  • tribe Delimini Brandt, 1956 - synonym: Papilliferini Brandt, 1961 (n.a.)
  • tribe Medorini H. Nordsieck, 1997
  • tribe Montenegrinini H. Nordsieck, 1972

subfamily Baleinae A. J. Wagner, 1913 - synonyms: Laciniariini H. Nordsieck, 1963; Tristaniinae Schileyko, 1999

subfamily † Constrictinae H. Nordsieck, 1981[4]

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[5]
  • tribe † Rillyini H. Nordsieck, 1985[6]

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 Phaedusini A. J. Wagner, 1922
  • tribe Megalophaedusini Zilch, 1954 - synonym: Zaptyxini Zilch, 1954

subfamily Serrulininae Ehrmann, 1927

Genera

Genera include:

Clausiliinae

tribe Clausiliini

  • Clausilia Draparnaud, 1805 - with two subgenera: Clausilia and Strobeliella H. Nordsieck, 1977[7]
  • Erjavecia Brusina, 1870[7]
  • Fusulus Fitzinger, 1833 - with two subgenera: Erjaveciella H. Nordsieck, 1877 and Fusulus[7]
  • Julica H. Nordsieck, 1963[7]
  • Macrogastra Hartmann, 1841 - with three subgenera: Macrogastra, Pseudovestia H. Nordsieck, 1877 and Pyrostoma Vest, 1867[7]
  • Micridyla H. Nordsieck, 1973[7]
  • Neostyriaca A. J. Wagner, 1920[7]
  • Pseudofusulus H. Nordsieck, 1977 - has the only one species: Pseudofusulus varians[7]
  • Ruthenica Lindholm, 1924[7]

tribe Gracillariini

  • Graciliaria Bielz, 1867[8]

Alopiinae

Alopiinae[9]

tribe Alopiini

  • Alopia H. & A.Adams, 1855 - with two subgenera Alopia and Kimakowiczia Szekeres, 1969
  • Herilla H. & A. Adams, 1855
  • Triloba Vest, 1867

tribe Cochlodinini

  • Cochlodina A. Férussac, 1821 - with four subgenera: Cochlodina; Cochlodinastra H. Nordsieck, 1977; Paracochlodina H. Nordsieck, 1969; Procochlodina H. Nordsieck, 1969
  • Macedonica O. Boettger, 1877

tribe Delimini

  • Barcania Brandt, 1956
  • Charpentieria Stabille, 1864 - with four subgenera: Charpentieria; Gibbularia Monterosato, 1908; Mauritanica O. Boettger, 1877; Siciliaria Vest, 1867 and Stigmatica O. Boettger, 1877
  • Delima Hartmann, 1842 - with four subgenera: Delima; Dugiana Stamol & Slapnik, 2002; Piceata O. Boettger, 1877 and Semirugata O. Boettger, 1877
  • Dilataria Vest, 1867
  • Papillifera Hartmann, 1842

tribe Medorini H. Nordsieck, 1997

  • Agathylla H. & A.Adams, 1855 - with two subgenera Agathylla and Agathyllina H. Nordsieck, 1969
  • Albinaria Vest, 1867 - also include former genera Carinigera Moellendorff, 1873; Cristataria Vest, 1867; Isabellaria Vest, 1867; Sericata O. Boettger, 1878.
  • Inchoatia Gittenberger & Uit de Weerd, 2006[10]
  • Lampedusa - 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

tribe Montenegrinini

  • Montenegrina O. Boettger, 1877
  • Protoherilla A. J. Wagner, 1921

Baleinae
  • 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
  • Menissa 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
  • Vestia Hesse, 1916 - with three subgenera: Brabenecia H. Nordsieck, 1974; Vestia and Vestiella H. Nordsieck, 1877

Constrictinae

Fossil subfamily Constrictinae contains genera:

  • † Constricta O. Boettger, 1877 - type genus of the subfamily[3]

Garnieriinae
  • Garnieria Bourguignat, 1877
  • Grandinenia Minato & Chen, 1984
  • Indonenia Ehrmann, 1927
  • Megalauchenia H. Nordsieck, 2007
  • Neniauchenia H. Nordsieck, 2002
  • Tropidauchenia Lindholm, 1924 - with two subgenera: Euryauchenia H. Nordsieck, 2007 and Tropidauchenia

Eualopiinae

Fossil subfamily Eualopiinae contains genera:

tribe † Eualopiini

  • † Eualopia O. Boettger, 1877 - type genus of the subfamily[3]

tribe † Rillyini

  • † Rillya Munier-Chalmas [in P. Fischer], 1883 - type genus of the tribe[3]

Laminiferinae
  • Laminifera O. Boettger, 1863 - the type genus of the family Laminiferinae.[3] Its type species is fossil.[3] (Actual status of the genus?)
  • Bofilliella Ehrmann, 1927
  • Neniatlanta Bouguignat, 1876

Mentissoideinae
  • 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
  • Bitorquata O. Boettger, 1877
  • Boettgeria O. Boettger, 1863 - with two subgenera: Boettgeriaand Loosjesiella Neubert & Groh, 1998
  • 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
  • Euxinella H. Nordsieck, 1973
  • Filosa O. Boettger, 1877
  • Galeata O. Boettger, 1877
  • Idyla H. & A. Adams, 1855 - with two subgenera: Idyla and Strigilidyla H. Nordsieck, 1994
  • Kazancia Neubert, 1992 - with two subgenera: Kazancia and Lasica Neubert, 1995
  • Macroptychia O. Boettger, 1877
  • Mentissoidea O. Boettger, 1877
  • Olympicola Hesse, 1916
  • Phrygica H. Nordsieck, 1994
  • Roseniella Thiele, 1931 - with two subgenera: Chavchetia Neubert, 1992 and Roseniella
  • Sabaeola Lindholm, 1925
  • Sprattia O. Boettger, 1883
  • Strigileuxina H. Nordsieck, 1975
  • Strumosa O. Boettger, 1877
  • Sumelia H. Nordsieck, 1994

Neniinae
  • Andinia Polinski, 1922
  • Andiniella Weyrauch, 1958
  • Audiniastra H. Nordsieck, 2005
  • Bequaertinenia Weyrauch, 1964 - with two subgenera: Bequaertinenia and Miranenia Grego & Szekeres, 2004
  • Brevinenia Neubert & H. Nordsieck, 2005
  • Columbinia Polinski, 1924 - with three subgenera: Columbinia; Paranenia Rehder, 1939 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
  • Gonionenia Pilsbry, 1926
  • Gracilinenia Polinski, 1922
  • Hemicena Pilsbry, 1949
  • Incaglaia Pilsbry, 1949 - with two subgenera: Gibbonenia Zilch, 1954 and Incaglaia
  • Incania Polinski, 1922
  • Leuconenia H. Nordsieck, 2005
  • Nenia H. & A. Adams, 1855
  • Neniaptyx H. Nordsieck, 2007
  • Neniatracta Pilsbry, 1926
  • Neniella Grego & Sekeres, 2004
  • Neniops Pilsbry, 1926
  • Nenisca Rehder, 1939
  • 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

tribe Phaedusini

  • Acanthophaedusa H. Nordsieck, 2007
  • 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
  • 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 five subgenera: Dentiphaedusa H. Nordsieck, 2003; Euphaedusa; Papilliphaedusa H. Nordsieck, 2003; Tauphaedusa H. Nordsieck, 2003 and Telophaedusa H. Nordsieck, 2003
  • Fuchsiana Gredler, 1887
  • Hemiphaedusa O. Boettger, 1877 - with 10 subgenera: Dendrophaedusa H. Nordsieck, 2002; Hemiphaedusa; Hemiphaedusoides H. Nordsieck, 2001; Hemizaptyx Pilsbry, 1905; Labyrinthiphaedusa H. Nordsieck, 2001; Margaritiphaedusa H. Nordsieck, 2001; Notoptychia Ehrmann, 1927; Pinguiphaedusa Azuma, 1982; Placeophaedusa Minato, 1994 and Selenophaedusa Lindholm, 1924
  • 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. (Neophaedusa is a synonym of Megalophaedusa).[11]
  • Miraphaedusa H. Nordsieck, 2005
  • Oospira Blanford, 1872[12] - with five subgenera: Atractophaedusa Ehrmann, 1927; Formosana O. Boettger, 1877; Formosanella H. Nordsieck, 2003; Oospira; Paraformosella H. Nordsieck, 2003 and Siphonophaedusa Lindholm, 1924
  • Paraphaedusa O. Boettger, 1877
  • Phaedusa H. & A.Adams, 1855 - with four subgenera: Metaphaedusa H. Nordsieck, 2001; Phaedusa; Pseudophaedusa Tomiyama, 1984 and Stereophaedusa O. Boettger, 1877
  • Reinia Kobelt, 1876 - with three subgenera: Parareinia H. Nordsieck, 1998; Pictophaedusa Azuma, 1982 and Reinia
  • Sinigena Lindholm, 1925
  • Stereozaptyx Pilsbry, 1905
  • Streptodera Lindholm, 1925
  • Zaptyx Pilsbry, 1901 - with two subgenera: Prozaptyx Loosjes, 1950 and Zaptyx

tribe Megalophaedusini

  • Leptacme Ehrmann, 1927[12]

tribe ?

  • Diceratoptyx Pilsbry, 1905
  • Heterozaptyx Pilsbry, 1906
  • Lindholmiella Ehrmann, 1927
  • Metazaptyx Pilsbry, 1905
  • Oligozaptx Pilsbry, 1905
  • Parazaptyx Pilsbry, 1905
  • Pliciphaedusa H. Nordsieck, 1998
  • Pulchraptyx Minato, 1981
  • Renschiphaedusa Loosjes & Loosjes-van Bemmel, 1973
  • Selenoptyx Pilsbry, 1908
  • Serriphaedusa H. Nordsieck, 2001
  • Synprosphyma Germain, 1919 - with two subgenera; Excussispira Lindholm, 1925 and Synprosphyma
  • Thaumatoptyx Pilsbry, 1908
  • Tyrannophaedusa Pilsbry, 1901 - with two subgenera: Aulacophaedusa Azuma, 1982 and Tyrannophaedusa
  • Tyrannozaptyx Käufel, 1930
  • Zaptychopsis Ehrmann, 1927

Serrulininae
  • Caspiophaedusa Lindholm, 1924
  • Dobatia H. Nordsieck, 1973
  • Graecophaedusa Rähle, 1982
  • Laeviphaedusa Likharev & Stelkov, 1965
  • Microphaedusa H. Nordsieck, 1978
  • Nothoserrulina Németh & Szekeres, 1995
  • Pamphylica Németh & Szekeres, 1995
  • Pravispira Lindholm, 1924
  • Pontophaedusa Lindholm, 1924
  • Pontophaedusella H. Nordsieck, 1994
  • Sciocochlea C. Boettger, 1935
  • Serrulina Mousson, 1873
  • Serrulinella H. Nordsieck, 1984
  • Truncatophaedusa Majoros, Németh & Szili-Kovács 1994
  • Tsoukatosia Gittenberger, 2000 - with two subgenera: Agiosspeleikos A. & P. Reischütz, 2003 and Tsoukatosia

Conservation

Although non-marine molluscs appear to be exceptionally vulnerable to extinction,[13] the IUCN Red list listed only 9 species[14] from this family.

gollark: I doubt it matters. The real issue is that I have absolutely no idea how it's meant to work.
gollark: Yep.
gollark: Unfortunately, environments are hard.
gollark: I copy some keys out of the global environment, then apply the FS changes, then `setfenv` everything to the new env.
gollark: What I want to do is `load` a new environment and stuff, and then have functions depending on global stuff (like `loadfile` using `fs.open` or whatever) use the functions in that environment.

References
  1. J. E. Gray. April 14, 1855. Catalogue of Pulmonata or air-breathing Mollusca in the collection of the British Museum, Part I: 156.
  2. Barker G. M.: Gastropods on Land: Phylogeny, Diversity and Adaptive Morphology. in Barker G. M. (ed.): The biology of terrestrial molluscs. CABI Publishing, Oxon, UK, 2001, ISBN 0-85199-318-4. 1-146, cited pages: 139 and 142.
  3. Bouchet, Philippe; Rocroi, Jean-Pierre; Frýda, Jiri; Hausdorf, Bernard; Ponder, Winston; Valdés, Ángel & Warén, Anders (2005). "Classification and nomenclator of gastropod families". Malacologia. Hackenheim, Germany: ConchBooks. 47 (1–2): 1–397. ISBN 3-925919-72-4. ISSN 0076-2997.
  4. Nordsieck H. (1981). Archiv für Molluskenkunde 111(1-3): 101.
  5. Nordsieck H. (1978). Archiv für Molluskenkunde 109(1-3): 104.
  6. Nordsieck H. (1985). Heldia 1(3): 83.
  7. "Clausiliini". Fauna Europaea, last update 27 January 2011, accessed 27 April 2011.
  8. "Graciliaria". Fauna Europaea, last update 27 January 2011, accessed 27 April 2011.
  9. Uit de Weerd D. R. (2004). "Molecular phylogenetic history of eastern Mediterranean Alopiinae, a group of morphologically indeterminate land snails". Doctoral thesis, Leiden University, ISBN 90-6464-874-3.
  10. Gittenberger E. & Uit de Weerd D. R. (2009). "Summarizing data on the Inchoatia taxa, including Inchoatia megdova bruggeni subspec. nov. (Gastropoda, Pulmonata, Clausiliidae)" Zoologische Mededelingen 83 http://www.zoologischemededelingen.nl/83/nr03/a08 Archived 16 January 2014 at the Wayback Machine
  11. cf. Nordsieck H. (2006). "Species list of recent Clausiliidae".
  12. Maassen W. J. M. & Gittenberger E. (2007). "Three new clausiliid land snails from Tonkin, northern Vietnam (Gastropoda: Pulmonata: Clausiliidae)". Zoologische Mededelingen 81(1): http://www.zoologischemededelingen.nl/81/nr01/a10 Archived 7 October 2011 at the Wayback Machine
  13. Lydeard, C.; Cowie R.; Ponder, W.F.; et al. (April 2004). "The global decline of nonmarine mollusks" (PDF). BioScience. 54: 321–330. doi:10.1641/0006-3568(2004)054[0321:TGDONM]2.0.CO;2. Archived from the original (PDF) on 31 March 2007. Retrieved 20 Oct 2009.
  14. IUCN 2009. IUCN Red List of Threatened Species. Version 2009.2. <www.iucnredlist.org>. Downloaded on 5 March 2010.

Further reading
  • Maltz T. K. & Sulikowska-Drozd A. (2008) "Life Cycles of Clausiliids of Poland — Knowns and Unknowns". Annales Zoologici 58(4): 857-880. doi:10.3161/000345408X396783.
  • Nordsieck H. (2007). Worldwide Door Snails. ConchBooks, 213 pp. ISBN 978-3-939767-07-7.
  • Uit de Weerd D. R. (2004). "Molecular phylogenetic history of eastern Mediterranean Alopiinae, a group of morphologically indeterminate land snails". Doctoral thesis, Department of Biology, Faculty of Science, Leiden University. HTM, PDF.
  • Uit de Weerd D. R., Piel W. H. & Gittenberger E. (2004). "Widespread polyphyly among Alopiinae snail genera: when phylogeny mirrors biogeography more closely than morphology". Molecular Phylogenetics and Evolution 33(3): 533-548. doi:10.1016/j.ympev.2004.07.010
  • Páll-Gergely B. & Szekeres M. , 2017. New and little-known Clausiliidae (Gastropoda: Pulmonata) from Laos and southern Vietnam. Journal of Conchology 42(6): 507-519
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