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.
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. 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.
Ecology
These animals live on stones, and often also under stones, in up to 5âÂÂ6 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 30âÂÂ70 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 <small>Andrusov, 1912</small>
Subgenus Calvertia <small>Bourguignat, 1880</small>
- â Theodoxus acuticarinatus <small>(Fuchs, 1870)</small>
- â Theodoxus amblygonioides <small>Wenz, 1930</small>
- â Theodoxus amethystinus <small>(Brusina, 1874)</small>
- â Theodoxus anconae <small>(Capellini, 1880)</small>
- â Theodoxus angularis <small>Gozhik, 2002</small>
- â Theodoxus anonymus <small>Anistratenko & Gozhik, 1995</small>
- â Theodoxus atticus <small>Kühn, 1963</small>
- â Theodoxus barakovici <small>(Brusina, 1902)</small>
- â Theodoxus bessarabicus <small>(Sinzov, 1896)</small>
- â Theodoxus brusinai <small>(PavloviÃÂ, 1931)</small>
- â Theodoxus capellinii <small>(Pantanelli, 1876)</small>
- â Theodoxus capillaceus <small>(Brusina, 1874)</small>
- â Theodoxus crescens <small>(Fuchs, 1870)</small>
- â Theodoxus doderleini <small>(d'Ancona, 1869)</small>
- â Theodoxus dumortieri <small>(Fontannes, 1878)</small>
- â Theodoxus fasciatus <small>Gozhik in Gozhik & Datsenko, 2007</small>
- â Theodoxus gnezdai <small>(Brusina, 1884)</small>
- â Theodoxus grateloupianus <small>(Férussac, 1823)</small>
- â Theodoxus hoernesanus <small>(Semper, 1867)</small>
- â Theodoxus ingulenzis <small>Gozhik in Gozhik & Datsenko, 2007</small>
- â Theodoxus kalodictya <small>(Andrusov, 1909)</small>
- â Theodoxus koslinskyi <small>(Porumbaru, 1881)</small>
- â Theodoxus lamellatus <small>(Brusina, 1892)</small>
- â Theodoxus lineatus <small>(Sinzov, 1896)</small>
- â Theodoxus lorkovici <small>(Brusina, 1878)</small>
- â Theodoxus macedonicus <small>Wenz, 1943</small>
- â Theodoxus maculosus <small>Gozhik in Gozhik & Datsenko, 2007</small>
- â Theodoxus mayeri <small>(Semper, 1867)</small>
- â Theodoxus militaris <small>(Neumayr, 1869)</small>
- â Theodoxus miljkovici <small>(Brusina, 1902)</small>
- â Theodoxus millepunctatus <small>(Brusina, 1902)</small>
- â Theodoxus moeschi <small>Locard, 1893</small>
- â Theodoxus morulus <small>Kühn, 1963</small>
- â Theodoxus mutinensis <small>(d'Ancona, 1869)</small>
- â Theodoxus neumayri <small>(Burgerstein, 1877)</small>
- â Theodoxus nitens <small>Gozhik, 2002</small>
- â Theodoxus nivosus <small>(Brusina, 1874)</small>
- â Theodoxus novorossicus <small>(Sinzov, 1896)</small>
- â Theodoxus obtusangulaeformis <small>Gozhik, 2002</small>
- â Theodoxus oxytropida <small>(Andrusov, 1909)</small>
- â Theodoxus pavlovici <small>MiloÃ
¡eviÃÂ, 1983</small>
- â Theodoxus philippianus <small>Locard, 1883</small>
- â Theodoxus pilari <small>(Brusina, 1884)</small>
- â Theodoxus platystoma <small>(Brusina, 1874)</small>
- â Theodoxus pseudodanubialis <small>(Sinzov, 1896)</small>
- â Theodoxus pseudofluviatilis <small>Locard, 1893</small>
- â Theodoxus pseudograteloupanus <small>(Sinzov, 1884)</small>
- â Theodoxus radmanesti <small>(Fuchs, 1870)</small>
- â Theodoxus radovanovici <small>(Brusina, 1893)</small>
- â Theodoxus reiseri <small>(Brusina, 1902)</small>
- â Theodoxus rugosus <small>(PavloviÃÂ, 1931)</small>
- â Theodoxus sagittiferus <small>(Brusina, 1874)</small>
- â Theodoxus schachmaticus <small>(Andrusov, 1909)</small>
- â Theodoxus scoliogramma <small>(Brusina, 1884)</small>
- â Theodoxus semidentatus <small>(Sandberger, 1875)</small>
- â Theodoxus serrulatus <small>(Brusina, 1892)</small>
- â Theodoxus slavonicus <small>(Brusina, 1878)</small>
- â Theodoxus sophievkaensis <small>Gozhik in Gozhik & Datsenko, 2007</small>
- â Theodoxus stanae <small>(Brusina, 1893)</small>
- â Theodoxus stefanescui <small>(Fontannes, 1887)</small>
- â Theodoxus stefanescuiformis <small>Gozhik in Gozhik & Datsenko, 2007</small>
- â Theodoxus subquadrofasciatus <small>Gozhik, 2002</small>
- â Theodoxus subslavonicus <small>Gozhik, 2002</small>
- â Theodoxus sundicus <small>(Andrusov, 1909)</small>
- â Theodoxus suskalovici <small>(PavloviÃÂ, 1903)</small>
- â Theodoxus sycophantus <small>(Brusina, 1878)</small>
- â Theodoxus tropidophorus <small>(Brusina, 1884)</small>
- â Theodoxus turbinatus <small>(Fuchs, 1870)</small>
- â Theodoxus veljetinensis <small>(PavloviÃÂ, 1903)</small>
- â Theodoxus venustus <small>(Brusina, 1897)</small>
- â Theodoxus xanthozona <small>(Brusina, 1884)</small>
- â Theodoxus zagradovkaensis <small>Gozhik, 2002</small>
- â Theodoxus zivkovici <small>(PavloviÃÂ, 1903)</small>
Subgenus Neritonyx <small>Andrusov, 1912</small>
Subgenus Ninniopsis <small>Tomlin, 1930</small>
Subgenus Neritaea <small>Roth, 1855</small>
- Theodoxus anatolicus <small>(Récluz, 1841)</small>
- â Theodoxus groyanus <small>(Férussac, 1823)</small>
- Theodoxus jordani <small>(G. B. Sowerby I, 1836)</small>
- â Theodoxus micans <small>(Gaudry & Fischer in Gaudry, 1867)</small>
- Theodoxus niloticus <small>(Reeve, 1856)</small> - synonym: Theodoxus africanus (Reeve, 1856)
- â Theodoxus subdoricus <small>Schütt, 1976</small>
- Theodoxus subterrelictus <small>Schütt, 1963</small>
- Theodoxus peloponensis <small>(Récluz, 1841)</small>
- Theodoxus varius <small>(Menke, 1828)</small>
Subgenus Theodoxus <small>Montfort, 1810</small>
- Theodoxus altenai <small>Schütt, 1965</small>
- Theodoxus baeticus <small>(Lamarck, 1822)</small>
- â Theodoxus becenensis <small>(CobÃÂlcescu, 1883)</small>
- â Theodoxus brenneri <small>(Handmann, 1882)</small>
- â Theodoxus carasiensis <small>Jekelius, 1944</small>
- â Theodoxus constantiae <small>Stefanescu, 1896</small>
- â Theodoxus culceri <small>(Porumbaru, 1881)</small>
- â Theodoxus cyrtocelis <small>(Krauss, 1852)</small>
- Theodoxus danubialis <small>(Pfeiffer, 1828)</small>
- â Theodoxus denisluensis <small>(Oppenheim, 1919)</small>
- â Theodoxus deperditus <small>(Almera, 1894)</small>
- â Theodoxus elongatulus <small>(Philippi, 1844)</small>
- â Theodoxus eugenii <small>Jekelius, 1944</small>
- Theodoxus euxinus <small>(Clessin, 1886)</small>
- Theodoxus fluviatilis <small>(Linnaeus, 1758)</small> - type species. synonyms: Theodoxus brauneri <small>(Lindholm, 1908)</small> (but as separate species per); Theodoxus brauneri f. lacrymans <small>Lindholm, 1908</small>; Theodoxus brauneri f. alboguttata <small>Lindholm, 1908</small>; Theodoxus brauneri f. pulherrima <small>Lindholm, 1908;</small> Theodoxus lutetianus <small>Montfort, 1810</small>. Bunje (2005) does not consider Theodoxus velox <small>Anistratenko, 1999</small> to be a distinct species from Theodoxus fluviatilis.
- â Theodoxus gregarius <small>(Thomä, 1845)</small>
- Theodoxus heldreichi <small>(Martens, 1879)</small>
- â Theodoxus hisingeri <small>(Bellardi & Michelotti, 1841)</small>
- Theodoxus hispalensis <small>(Martens, 1879)</small>
- â Theodoxus jekeliusi <small>JuriÃ
¡iÃÂ-PolÃ
¡ak, 1979</small>
- â Theodoxus leobersdorfensis <small>(Handmann, 1887)</small>
- Theodoxus meridionalis <small>(Philippi, 1836)</small>
- â Theodoxus milessii <small>Papp, 1979</small>
- â Theodoxus moosbrunnensis <small>Papp, 1953</small>
- â Theodoxus morellii <small>(Bellardi & Michelotti, 1841)</small>
- Theodoxus pallasi <small>Lindholm, 1924</small> - synonym: Theodoxus lituratus <small>(Eichwald, 1838)</small>
- â Theodoxus pilidei <small>(Tournouër, 1879)</small>
- Theodoxus prevostianus <small>(Pfeiffer, 1828)</small>
- â Theodoxus punctatolineatus <small>(Sinzov, 1896)</small>
- Theodoxus saulcyi <small>(Bourguignat, 1852)</small>
- â Theodoxus scamandrius <small>(Calvert & Neumayr, 1880)</small>
- â Theodoxus semiplicatus <small>(Neumayr in Herbich & Neumayr, 1875)</small>
- â Theodoxus simplex <small>(Fuchs, 1877)</small>
- â Theodoxus subglobosus <small>(Eichwald, 1853)</small>
- Theodoxus subthermalis <small>(Bourguignat in Issel, 1865)</small> or Theodoxus fluviatilis subthermalis <small>Issel, 1865</small>
- Theodoxus transversalis <small>(Pfeiffer, 1828)</small>
- â Theodoxus trifasciatus <small>(Grateloup, 1839)</small>
- â Theodoxus turislavicus <small>Jekelius, 1944</small>
- Theodoxus valentinus <small>(Graells, 1846)</small>
- â Theodoxus zografi <small>(Brusina, 1902)</small>
Subgenus ?
- â Theodoxus abnormis <small>(Jenkins, 1864)</small>
- â Theodoxus almelae <small>(Revilla, 1958)</small>
- Theodoxus astrachanicus <small>Starobogatov, 1994</small>
- â Theodoxus banaticus <small>Jekelius, 1944</small>
- â Theodoxus bohotinensis <small>(Simionescu & Barbu, 1940)</small>
- â Theodoxus bolivari <small>(Royo Gómez, 1922)</small>
- â Theodoxus bukowskii <small>(Oppenheim, 1919)</small>
- Theodoxus cariosus <small>(Wood, 1828)</small>
- Theodoxus coronatus <small>(Leach, 1815)</small>
- â Theodoxus crenulatus <small>(Klein, 1853)</small>
- â Theodoxus cunici <small>(Brusina, 1892)</small>
- Theodoxus danasteri <small>(Lindholm, 1908)</small>
- â Theodoxus doricus <small>(Neumayr, 1880)</small>
- Theodoxus euxinus <small>(Clessin, 1886)</small>
- Theodoxus gloeri <small>OdabaÃ
Âi & Arslan, 2015</small>
- Theodoxus gurur <small>Sands & Glöer, 2020</small>
- â Theodoxus hellenicus <small>(Bukowski, 1896)</small>
- â Theodoxus imbricatus <small>(Brusina, 1878)</small>
- â Theodoxus intracarpaticus <small>Jekelius, 1944</small>
- â Theodoxus licherdopoli <small>(Stefanescu, 1896)</small>
- Theodoxus luteofasciatus <small>(Miller, 1879)</small>
- Theodoxus macri <small>(Sowerby, 1849)</small>
- Theodoxus maresi <small>(Bourguignat, 1864)</small>
- Theodoxus marteli <small>(Pallary, 1920)</small>
- Theodoxus milachevitchi <small>Golikov & Starobogatov, 1966</small>
- â Theodoxus mutinensis <small>(d'Ancona, 1869)</small>
- Theodoxus numidicus <small>(Récluz, 1841)</small>
- Theodoxus pallasi <small>Lindholm, 1924</small>
- Theodoxus pallidus <small>Dunker, 1861</small>
- â Theodoxus pappi <small>Sauerzopf, 1952</small>
- â Theodoxus paradisii <small>(Magrograssi, 1928)</small>
- â Theodoxus patrae <small>Esu & Girotti, 2015</small>
- â Theodoxus percarinatus <small>(Oppenheim, 1919)</small>
- Theodoxus pilidei <small>(Tournouêr, 1879)</small>
- â Theodoxus politioanei <small>Jekelius, 1944</small>
- â Theodoxus politus <small>Jekelius, 1944</small>
- Theodoxus poppei <small>Ahuir, 2020</small>
- â Theodoxus postcrenulatus <small>Papp, 1953</small>
- â Theodoxus prozlatarici <small>Jekelius, 1944</small>
- â Theodoxus pseudodacicus <small>Neubauer, Harzhauser, Georgopoulou, Mandic & Kroh, 2014</small>
- â Theodoxus quadrifasciatus <small>(Bielz, 1864)</small>
- â Theodoxus reticulatus <small>PanÃÂ, 1990</small>
- â Theodoxus rhodiensis <small>(Tournouër in Fischer, 1877)</small>
- Theodoxus sarmaticus <small>(Lindholm, 1901)</small>
- Theodoxus schultzii <small>(Grimm, 1877)</small>
- â Theodoxus scriptus <small>(Stefanescu, 1896)</small>
- â Theodoxus serratiliniformis <small>Geyer, 1914</small>
- â Theodoxus sinjanus <small>(Brusina, 1876)</small>
- â Theodoxus soceni <small>Jekelius, 1944</small>
- â Theodoxus sphaeroidalis <small>(Revilla, 1958)</small>
- â Theodoxus spratti <small>(Jenkins, 1864)</small>
- â Theodoxus stoicai <small>Neubauer, Harzhauser, Georgopoulou, Mandic & Kroh, 2014</small>
- Theodoxus subthermalis <small>Bourguignat in Issel, 1865</small>
- â Theodoxus susuzianus <small>Harzhauser, Neubauer & HoÃ
Âgör, 2018</small>
- â Theodoxus timisensis <small>Jekelius, 1944</small>
- â Theodoxus trilophosensis <small>Rust, 1997</small>
- Theodoxus velascoi <small>(Graëlls, 1846)</small>
- Theodoxus velox <small>V. Anistratenko in O. Anistratenko, Starobogatov & V. Anistratenko, 1999</small>
- Theodoxus vespertinus <small>(Sowerby II, 1849)</small>
- Theodoxus wesselinghi <small>Sands & Glöer, 2020</small>
- Theodoxus wilkei <small>Sands & Glöer, 2020</small>
- â Theodoxus zlatarici <small>(Brusina, 1902)</small>
Synonyms:
- â Theodoxus burdigalensis <small>(d'Orbigny, 1852)</small>: synonym of â Theodoxus (Theodoxus) trifasciatus <small>(Grateloup, 1839)</small>
- Theodoxus cariosus <small>(Wood, 1828)</small> and Theodoxus cariosa are synonyms of Neritina cariosa <small>(Wood, 1828)</small>
- â Theodoxus dacicus <small>Jekelius, 1944</small>: synonym of â Theodoxus (Theodoxus) leobersdorfensis dacicus <small>Jekelius, 1944</small>
- Theodoxus doriae <small>Issel, 1865</small>: synonym of Theodoxus fluviatilis <small>(Linnaeus, 1758)</small>: synonym of Theodoxus (Theodoxus) fluviatilis <small>(Linnaeus, 1758)</small>
- Theodoxus euphraticus <small>(Mousson, 1874)</small>: synonym of Neritina euphratica <small>Mousson, 1874</small>
- â Theodoxus intracarpatica <small>Jekelius, 1944</small>: synonym of â Theodoxus intracarpaticus <small>Jekelius, 1944</small>
- Theodoxus lutetianus <small>Montfort, 1810</small>: synonym of Theodoxus fluviatilis <small>(Linnaeus, 1758)</small>: synonym of Theodoxus (Theodoxus) fluviatilis <small>(Linnaeus, 1758)</small>
- â Theodoxus mariae <small>(Handmann, 1887)</small>: synonym of â Neritina mariae <small>Handmann, 1887</small>
- â Theodoxus martensi <small>(Brusina, 1884)</small>: synonym of â Ninnia martensi <small>(Brusina, 1884)</small>
- Theodoxus neglectus <small>(Pease, 1861)</small>: synonym of Neritina neglecta <small>(Pease, 1861)</small>
- â Theodoxus oslavanensis <small>Rzehak, 1893</small>: synonym of â Theodoxus (Theodoxus) cyrtocelis austriacus <small>(Rzehak, 1893)</small>
- Theodoxus oualaniense <small>(Lesson, 1831)</small>: synonym of Clithon oualaniense <small>(Lesson, 1831)</small>
- â Theodoxus perlongus <small>Jekelius, 1944</small>: synonym of â Theodoxus (Theodoxus) zografi perlongus <small>Jekelius, 1944</small>
- â Theodoxus petralbensis <small>Jekelius, 1944</small>: synonym of â Theodoxus (Theodoxus) zografi petralbensis <small>Jekelius, 1944</small>
- â Theodoxus pictus <small>(Férussac, 1823)</small>: synonym of â Agapilia picta <small>(Férussac, 1823)</small>
- Theodoxus reclivatus <small>Say, 1822</small>: synonym of Neritina usnea <small>(Röding, 1798)</small>
- â Theodoxus rumanus <small>(Stefanescu, 1896)</small>: synonym of â Theodoxus (Calvertia) stefanescui <small>(Fontannes, 1887)</small>
- â Theodoxus scamandri <small>(Calvert & Neumayr, 1880)</small>: synonym of â Theodoxus (Theodoxus) scamandrius <small>(Calvert & Neumayr, 1880)</small>
- Theodoxus vespertinus <small>(G. B. Sowerby II, 1849)</small>: synonym of Neritina vespertinus <small>(G. B. Sowerby II, 1849)</small>
- â Theodoxus vetranici <small>(Brusina, 1902)</small>: synonym of â Neritina vetranici <small>(Brusina, 1902)</small>
- Theodoxus virginea <small>(Linnaeus, 1758)</small>: synonym of Neritina virginea <small>(Linnaeus, 1758)</small>
- â Theodoxus (Vittoclithon) pictus bizonalis <small>(Grateloup, 1839)</small>: synonym of â Agapilia picta bizonalis <small>(Grateloup, 1839)</small>
References
This article incorporates public domain text from the reference
- Eichhorst T.E. (2016). Neritidae of the world. Volume 2. Harxheim: Conchbooks. pp. 696âÂÂ1366
External links