Theodoxus

''Theodoxus'' is a genus of nerites, small water snails with an operculum, some of which live in freshwater, and some in both freshwater and brackish water, aquatic gastropod mollusks in the family Neritidae, the nerites.Bouchet, P. (2014). Theodoxus Montfort, 1810. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=153531 on 2014-11-17.

Distribution

The distribution of the genus ''Theodoxus'' includes Europe and northern Africa and also extends east to southern Iran. No other species within Neritidae have sympatrical distribution with ''Theodoxus''. The distribution of the genus ''Theodoxus'' is an exception within Neritidae, because Neritidae live primarily in the southern hemisphere. Species within ''Theodoxus'' are the only Neritidae snails, that live in temperate climate.

Bunje & Lindberg (2007) presented the first phylogenetic hypothesis of the clade ''Theodoxus''.Bunje P. M. & Lindberg D. R. (2007). "Lineage divergence of a freshwater snail clade associated with post-Tethys marine basin development". ''Molecular Phylogenetics and Evolution'' 42(2): 373–387. . The evolution of the genus ''Theodoxus'' was affected by separating of Mediterranean Sea and Paratethys sea in Miocene.

Description

The shell in this genus is semiovular with a flat apertural plain. There is no umbilicus. The columella and inner whorls are dissolved.

Species in the genus ''Theodoxus '' are highly variable in size, in color pattern of the periostracum, in details of the operculum and in the radula, and all these factors can make identification to species level very challenging.Zettler M. L., Frankowski J., Bochert R. & Röhner M. (2004). "Morphological and ecological features of ''Theodoxus fluviatilis'' (Linnaeus, 1758) from Baltic brackish water and German freshwater populations". ''Journal of Conchology'' 38(3): 305-316
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Ecology

These animals live on stones, and often also under stones, in up to 5–6 m depth or deeper, feeding on algal covers. ''Theodoxus '' needs rough surfaces in order to be able to digest its food, so a stony substrate is necessary. Green algae are not consumed; ''Theodoxus '' has no cellulases. These snails lay egg capsules containing 30-70 eggs each, usually on the shells of other ''Theodoxus '' animals; only one juvenile grows, the other eggs serve as food.

Species

There were no systematic review of the genus ''Theodoxus'' as of 2007. Bunje (2004) noted at least 34 extant taxa at species level in the genus ''Theodoxus''. IUCN Red List (2015) provided conservation status for 23 species of ''Theodoxus''.

Species within the genus ''Theodoxus'' include:
; Subgenus ''Brusinaella'' Andrusov, 1912
* † ''Theodoxus petasatus'' (Seninski, 1905)
; Subgenus ''Calvertia'' Bourguignat, 1880

* † ''Theodoxus acuticarinatus'' (Fuchs, 1870)
* † '' Theodoxus amblygonioides'' Wenz, 1930
* † ''Theodoxus amethystinus'' (Brusina, 1874)
* † ''Theodoxus anconae'' (Capellini, 1880)
* † '' Theodoxus angularis'' Gozhik, 2002
* † ''Theodoxus anonymus'' Anistratenko & Gozhik, 1995
* † ''Theodoxus atticus'' Kühn, 1963
* † ''Theodoxus barakovici'' (Brusina, 1902)
* † ''Theodoxus bessarabicus'' (Sinzov, 1896)
* † ''Theodoxus brusinai'' (Pavlović, 1931)
* † ''Theodoxus capellinii'' (Pantanelli, 1876)
* † ''Theodoxus capillaceus'' (Brusina, 1874)
* † ''Theodoxus crescens'' (Fuchs, 1870)
* † ''Theodoxus doderleini'' (d'Ancona, 1869)
* † ''Theodoxus dumortieri'' (Fontannes, 1878)
* † ''Theodoxus fasciatus'' Gozhik in Gozhik & Datsenko, 2007
* † ''Theodoxus gnezdai'' (Brusina, 1884)
* † ''Theodoxus grateloupianus'' (Férussac, 1823)
* † ''Theodoxus hoernesanus'' (Semper, 1867)
* † ''Theodoxus ingulenzis'' Gozhik in Gozhik & Datsenko, 2007
* † ''Theodoxus kalodictya'' (Andrusov, 1909)
* † ''Theodoxus koslinskyi'' (Porumbaru, 1881)
* † ''Theodoxus lamellatus'' (Brusina, 1892)
* † ''Theodoxus lineatus'' (Sinzov, 1896)
* † ''Theodoxus lorkovici'' (Brusina, 1878)
* † ''Theodoxus macedonicus'' Wenz, 1943
* † ''Theodoxus maculosus'' Gozhik in Gozhik & Datsenko, 2007
* † ''Theodoxus mayeri'' (Semper, 1867)
* † ''Theodoxus militaris'' (Neumayr, 1869)
* † ''Theodoxus miljkovici'' (Brusina, 1902)
* † '' Theodoxus millepunctatus'' (Brusina, 1902)
* † ''Theodoxus moeschi'' Locard, 1893
* † ''Theodoxus morulus'' Kühn, 1963
* † ''Theodoxus mutinensis'' (d'Ancona, 1869)
* † ''Theodoxus neumayri'' (Burgerstein, 1877)
* † ''Theodoxus nitens'' Gozhik, 2002
* † ''Theodoxus nivosus'' (Brusina, 1874)
* † ''Theodoxus novorossicus'' (Sinzov, 1896)
* † '' Theodoxus obtusangulaeformis'' Gozhik, 2002
* † ''Theodoxus oxytropida'' (Andrusov, 1909)
* † ''Theodoxus pavlovici'' Milošević, 1983
* † '' Theodoxus philippianus'' Locard, 1883
* † ''Theodoxus pilari'' (Brusina, 1884)
* † '' Theodoxus platystoma'' (Brusina, 1874)
* † ''Theodoxus pseudodanubialis'' (Sinzov, 1896)
* † '' Theodoxus pseudofluviatilis'' Locard, 1893
* † ''Theodoxus pseudograteloupanus'' (Sinzov, 1884)
* † ''Theodoxus radmanesti'' (Fuchs, 1870)
* † ''Theodoxus radovanovici'' (Brusina, 1893)
* † ''Theodoxus reiseri'' (Brusina, 1902)
* † ''Theodoxus rugosus'' (Pavlović, 1931)
* † ''Theodoxus sagittiferus'' (Brusina, 1874)
* † ''Theodoxus schachmaticus'' (Andrusov, 1909)
* † ''Theodoxus scoliogramma'' (Brusina, 1884)
* † ''Theodoxus semidentatus'' (Sandberger, 1875)
* † ''Theodoxus serrulatus'' (Brusina, 1892)
* † ''Theodoxus slavonicus'' (Brusina, 1878)
* † '' Theodoxus sophievkaensis'' Gozhik in Gozhik & Datsenko, 2007
* † ''Theodoxus stanae'' (Brusina, 1893)
* † ''Theodoxus stefanescui'' (Fontannes, 1887)
* † ''Theodoxus stefanescuiformis'' Gozhik in Gozhik & Datsenko, 2007
* † ''Theodoxus subquadrofasciatus'' Gozhik, 2002
* † ''Theodoxus subslavonicus'' Gozhik, 2002
* † ''Theodoxus sundicus'' (Andrusov, 1909)
* † ''Theodoxus suskalovici'' (Pavlović, 1903)
* † ''Theodoxus sycophantus'' (Brusina, 1878)
* † ''Theodoxus tropidophorus'' (Brusina, 1884)
* † ''Theodoxus turbinatus'' (Fuchs, 1870)
* † ''Theodoxus veljetinensis'' (Pavlović, 1903)
* † ''Theodoxus venustus'' (Brusina, 1897)
* † ''Theodoxus xanthozona'' (Brusina, 1884)
* † '' Theodoxus zagradovkaensis'' Gozhik, 2002
* † ''Theodoxus zivkovici'' (Pavlović, 1903)

;Subgenus ''Neritonyx'' Andrusov, 1912
* † ''Theodoxus unguiculatus'' (Seninski, 1905)

;Subgenus ''Ninniopsis'' Tomlin, 1930
* † ''Theodoxus colchicus'' (Andrusov, 1912)

;Subgenus ''Neritaea'' Roth, 1855
* ''Theodoxus anatolicus'' (Récluz, 1841)"''Neritaea''"
Fauna Europaea, last update 27 January 2011, accessed 12 April 2011.
* † ''Theodoxus groyanus'' (Férussac, 1823)
* ''Theodoxus jordani'' (Sowerby)
* † ''Theodoxus micans'' (Gaudry & Fischer in Gaudry, 1867)
* '' Theodoxus niloticus'' (Reeve, 1856) - synonym: ''Theodoxus africanus'' (Reeve, 1856)
* † ''Theodoxus subdoricus'' Schütt, 1976
* ''Theodoxus subterrelictus'' Schütt, 1963
* ''Theodoxus peloponensis'' (Récluz, 1841)
* ''Theodoxus varius'' (Menke, 1828)

;Subgenus ''Theodoxus'' Montfort, 1810

* ''Theodoxus altenai'' Schütt, 1965"Species in genus ''Theodoxus ''"
(n=20). AnimalBase, accessed 11 April 2011. Schütt H. (1965). "Zur Systematik und Ökologie türkischer Süßwasserprosobranchier". ''Zoologische Mededelingen'' 41(3): 43-72, Plate 1
abstract PDF

* ''Theodoxus baeticus'' (Lamarck, 1822)"''Theodoxus ''"
Fauna Europaea, last update 27 January 2011, accessed 12 April 2011.
* † ''Theodoxus becenensis'' (Cobălcescu, 1883)
* † ''Theodoxus brenneri'' (Handmann, 1882)
* † ''Theodoxus carasiensis'' Jekelius, 1944
* † ''Theodoxus constantiae'' Stefanescu, 1896
* † ''Theodoxus culceri'' (Porumbaru, 1881)
* † ''Theodoxus cyrtocelis'' (Krauss, 1852)
* ''Theodoxus danubialis'' (Pfeiffer, 1828)
* † ''Theodoxus denisluensis'' (Oppenheim, 1919)
* † ''Theodoxus deperditus'' (Almera, 1894)
* † ''Theodoxus elongatulus'' (Philippi, 1844)
* † ''Theodoxus eugenii'' Jekelius, 1944
* ''Theodoxus euxinus'' (Clessin, 1886)
* ''Theodoxus fluviatilis'' (Linnaeus, 1758) - type species.Anistratenko V. V. (2005). "Lectotypes for ''Tricolia pullus'', ''Gibbula divaricata'' and ''Theodoxus fluviatilis'' (Mollusca, Gastropoda) revisited". ''Vestnik zoologii'' 39(6): 3-10
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synonyms: ''Theodoxus brauneri'' (Lindholm, 1908) (but as separate species per); ''Theodoxus brauneri'' f. ''lacrymans'' Lindholm, 1908; ''Theodoxus brauneri'' f. '' alboguttata'' Lindholm, 1908; ''Theodoxus brauneri'' f. ''pulherrima'' Lindholm, 1908; ''Theodoxus lutetianus'' Montfort, 1810. Bunje (2005)PDF
does not consider ''Theodoxus velox'' Anistratenko, 1999Anistratenko O. Y., Starobogatov Y. I. & Anistratenko V. V. (1999). "Mollusks of the genus ''Theodoxus '' (Gastropoda, Pectinibranchia, Neritidae) from the Black and the Azov seas basin". ''Vestnik Zoologii'' 33: 11-19. to be a distinct species from ''Theodoxus fluviatilis''.
* † ''Theodoxus gregarius'' (Thomä, 1845)
* ''Theodoxus heldreichi'' (Martens, 1879)
* † ''Theodoxus hisingeri'' (Bellardi & Michelotti, 1841)
* ''Theodoxus hispalensis'' (Martens, 1879)
* † ''Theodoxus jekeliusi'' Jurišić-Polšak, 1979
* † ''Theodoxus leobersdorfensis'' (Handmann, 1887)
* ''Theodoxus meridionalis'' (Philippi, 1836)
* † ''Theodoxus milessii'' Papp, 1979
* † ''Theodoxus moosbrunnensis'' Papp, 1953
* † ''Theodoxus morellii'' (Bellardi & Michelotti, 1841)
* '' Theodoxus pallasi'' Lindholm, 1924 - synonym: ''Theodoxus lituratus'' (Eichwald, 1838)
* † '' Theodoxus pilidei'' (Tournouër, 1879)
* ''Theodoxus prevostianus'' (Pfeiffer, 1828)
* † ''Theodoxus punctatolineatus'' (Sinzov, 1896)
* ''Theodoxus saulcyi'' (Bourguignat, 1852)
* † ''Theodoxus scamandrius'' (Calvert & Neumayr, 1880)
* † ''Theodoxus semiplicatus'' (Neumayr in Herbich & Neumayr, 1875)
* † ''Theodoxus simplex'' (Fuchs, 1877)
* † ''Theodoxus subglobosus'' (Eichwald, 1853)
* ''Theodoxus subthermalis'' (Bourguignat in Issel, 1865) or ''Theodoxus fluviatilis subthermalis'' Issel, 1865
* '' Theodoxus transversalis'' (Pfeiffer, 1828)
* † ''Theodoxus trifasciatus'' (Grateloup, 1839)
* † ''Theodoxus turislavicus'' Jekelius, 1944
* ''Theodoxus valentinus'' (Graells, 1846)
* † ''Theodoxus zografi'' (Brusina, 1902)

;Subgenus ?

* † ''Theodoxus abnormis'' (Jenkins, 1864)
* † ''Theodoxus almelae'' (Revilla, 1958)
* ''Theodoxus astrachanicus'' Starobogatov, 1994Kantor Yu I., Vinarski M. V., Schileyko A. A. & Sysoev A. V. (published online on March 2, 2010). "Catalogue of the continental mollusks of Russia and adjacent territories"
Version 2.3.1.
/ref>
* † ''Theodoxus banaticus'' Jekelius, 1944
* † '' Theodoxus bohotinensis'' (Simionescu & Barbu, 1940)
* † ''Theodoxus bolivari'' (Royo Gómez, 1922)
* † ''Theodoxus bukowskii'' (Oppenheim, 1919)
* ''Theodoxus cariosus'' (Wood, 1828)
* ''Theodoxus coronatus'' (Leach, 1815)
* † ''Theodoxus crenulatus'' (Klein, 1853)
* † ''Theodoxus cunici'' (Brusina, 1892)
* ''Theodoxus danasteri'' (Lindholm, 1908)
* † ''Theodoxus doricus'' (Neumayr, 1880)
* ''Theodoxus euxinus'' (Clessin, 1886)
* '' Theodoxus gloeri'' Odabaşi & Arslan, 2015
* ''Theodoxus gurur'' Sands & Glöer, 2020
* † ''Theodoxus hellenicus'' (Bukowski, 1896)
* † ''Theodoxus imbricatus'' (Brusina, 1878)
* † ''Theodoxus intracarpaticus'' Jekelius, 1944
* ''Theodoxus karasuna''
* † ''Theodoxus licherdopoli'' (Stefanescu, 1896)
* ''Theodoxus luteofasciatus'' (Miller, 1879)
* ''Theodoxus macri'' (Sowerby, 1849)
* ''Theodoxus maresi'' (Bourguignat, 1864)
* ''Theodoxus marteli'' (Pallary, 1920)
* ''Theodoxus milachevitchi'' Golikov & Starobogatov, 1966
* ''Theodoxus mutinensis''
* ''Theodoxus numidicus'' (Récluz, 1841)
* '' Theodoxus olivaceus''
* '' Theodoxus pallasi'' Lindholm, 1924
* ''Theodoxus pallidus'' Dunker, 1861
* † '' Theodoxus pappi'' Sauerzopf, 1952
* † ''Theodoxus paradisii'' (Magrograssi, 1928)
* † ''Theodoxus patrae'' Esu & Girotti, 2015
* † ''Theodoxus percarinatus'' (Oppenheim, 1919)
* '' Theodoxus pilidei'' (Tournouêr, 1879)
* † ''Theodoxus politioanei'' Jekelius, 1944
* † ''Theodoxus politus'' Jekelius, 1944
* ''Theodoxus poppei'' Ahuir, 2020
* † ''Theodoxus postcrenulatus'' Papp, 1953
* † ''Theodoxus prozlatarici'' Jekelius, 1944
* † ''Theodoxus pseudodacicus'' Neubauer, Harzhauser, Georgopoulou, Mandic & Kroh, 2014 †
* † Theodoxus quadrifasciatus (Bielz, 1864) †

* † ''Theodoxus reticulatus'' Pană, 1990

* † ''Theodoxus rhodiensis'' (Tournouër in Fischer, 1877)
* ''Theodoxus sarmaticus'' (Lindholm, 1901)
* ''Theodoxus schultzii'' (Grimm, 1877)Zettler M. L. (2007). "A redescription of ''Theodoxus schultzii'' (Grimm, 1877), an endemic neritid gastropod of the Caspian Sea". ''Journal of Conchology'' 39(3): 245-252
abstract

* † ''Theodoxus scriptus'' (Stefanescu, 1896)
* † ''Theodoxus serratiliniformis'' Geyer, 1914Glöer P. (2002). ''Die Süßwassergastropoden Nord- und Mitteleuropas''. Die Tierwelt Deutschlands, ConchBooks, Hackenheim, 326 pp., .
* † ''Theodoxus sinjanus'' (Brusina, 1876)
* † ''Theodoxus soceni'' Jekelius, 1944
* † '' Theodoxus sphaeroidalis'' (Revilla, 1958)
* † ''Theodoxus spratti'' (Jenkins, 1864)
* † ''Theodoxus stoicai'' Neubauer, Harzhauser, Georgopoulou, Mandic & Kroh, 2014
* ''Theodoxus subthermalis'' Bourguignat in Issel, 1865
* † ''Theodoxus susuzianus'' Harzhauser, Neubauer & Hoşgör, 2018
* † '' Theodoxus timisensis'' Jekelius, 1944
* † ''Theodoxus trilophosensis'' Rust, 1997
* ''Theodoxus velascoi'' (Graëlls, 1846)
* ''Theodoxus velox'' V. Anistratenko in O. Anistratenko, Starobogatov & V. Anistratenko, 1999
* ''Theodoxus vespertinus'' (Sowerby II, 1849)
* ''Theodoxus wesselinghi'' Sands & Glöer, 2020
* ''Theodoxus wilkei'' Sands & Glöer, 2020
* † ''Theodoxus zlatarici'' Brusina, 1902)

;Synonyms:
* † ''Theodoxus burdigalensis'' (d'Orbigny, 1852) : synonym of † ''Theodoxus (Theodoxus) trifasciatus'' (Grateloup, 1839)
* ''Theodoxus cariosus'' (Wood, 1828) and ''Theodoxus cariosa'' are synonyms of ''Neritina cariosa'' (Wood, 1828)Goodwin D. R. (2006). "The Discovery of ''Neritina'' (''Theodoxus '') ''cariosa'' (Wood, 1828) on the Island of Maui, Hawaii (Gastropoda: Neritidae)". ''Visaya Net'' 11 pp
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* † ''Theodoxus dacicus'' Jekelius, 1944: synonym of † ''Theodoxus (Theodoxus) leobersdorfensis dacicus'' Jekelius, 1944
* ''Theodoxus doriae'' Issel, 1865: synonym of ''Theodoxus fluviatilis'' (Linnaeus, 1758) represented as ''Theodoxus (Theodoxus) fluviatilis'' (Linnaeus, 1758)
* ''Theodoxus euphraticus'' (Mousson, 1874)Mansoorian A. (2001). "Freshwater Gastropod of Khuzestan Province, South-West Iran". ''Iranian Int. J. Sci.'' 2(2): 9 pp
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is a synonym of ''Neritina euphratica'' Mousson, 1874Glöer P. & Pešić V. (2012). "The freshwater snails (Gastropoda) of Iran, with descriptions of two new genera and eight new species". ''ZooKeys'' 219: 11-61, .
* † ''Theodoxus intracarpatica'' Jekelius, 1944 : synonym of † ''Theodoxus intracarpaticus'' Jekelius, 1944
* ''Theodoxus lutetianus'' Montfort, 1810: synonym of ''Theodoxus fluviatilis'' (Linnaeus, 1758) represented as ''Theodoxus (Theodoxus) fluviatilis'' (Linnaeus, 1758)
* † ''Theodoxus mariae'' (Handmann, 1887): synonym of † ''Neritina mariae'' Handmann, 1887
* † ''Theodoxus martensi'' (Brusina, 1884): synonym of † '' Ninnia martensi'' (Brusina, 1884)
* ''Theodoxus neglectus'' is a synonym of ''Neritina neglecta'' (Pease, 1861)
* † ''Theodoxus oslavanensis'' Rzehak, 1893: synonym of † ''Theodoxus (Theodoxus) cyrtocelis austriacus'' (Rzehak, 1893)
* ''Theodoxus oualaniense'' (Lesson, 1831): synonym of Checked: verified by a taxonomic editorClithon oualaniense (Lesson, 1831)
* † ''Theodoxus perlongus'' Jekelius, 1944: synonym of † ''Theodoxus (Theodoxus) zografi perlongus'' Jekelius, 1944
* † ''Theodoxus petralbensis'' Jekelius, 1944: synonym of † ''Theodoxus (Theodoxus) zografi petralbensis'' Jekelius, 1944
* † ''Theodoxus pictus'' (Férussac, 1823): synonym of † '' Agapilia picta'' (Férussac, 1823)
* ''Theodoxus reclivatus'' Say, 1822: synonym of ''Neritina usnea'' (Röding, 1798)
* † ''Theodoxus rumanus'' (Stefanescu, 1896): synonym of † ''Theodoxus (Calvertia) stefanescui'' (Fontannes, 1887)
* † ''Theodoxus scamandri'' (Calvert & Neumayr, 1880): synonym of † ''Theodoxus (Theodoxus) scamandrius'' (Calvert & Neumayr, 1880)
* ''Theodoxus vespertinus'' : synonym of ''Neritina vespertinus'' (Sowerby, 1849)
* † ''Theodoxus vetranici'' (Brusina, 1902): synonym of † ''Neritina vetranici'' (Brusina, 1902)
* ''Theodoxus virginea'' (Linnaeus, 1758): synonym of ''Neritina virginea'' (Linnaeus, 1758)
* † ''Theodoxus (Vittoclithon) pictus bizonalis'' (Grateloup, 1839): synonym of † ''Agapilia picta bizonalis'' (Grateloup, 1839)

References

This article incorporates public domain text from the reference "Genus summary for ''Theodoxus ''"
AnimalBase, last modified 24 February 2009, accessed 11 April 2011.

* Eichhorst T.E. (2016). Neritidae of the world. Volume 2. Harxheim: Conchbooks. Pp. 696-1366

External links

* Bandel K. (2001). "The history of ''Theodoxus '' and ''Neritina'' connected with description and systematic evaluation of related Neritimorpha (Gastropoda)". '' Mittelungen aus dem Geologisch-Palaontologischen Institut Universitat Hamburg'' 85: 65-164
abstract

{{Taxonbar, from=Q148631

Neritidae
Gastropod genera
Taxonomy articles created by Polbot