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Wikipedia / 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.

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 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:

  1. Subgenus Brusinaella Andrusov, 1912

  2. † Theodoxus petasatus (Seninski, 1905)

  3. Subgenus Calvertia Bourguignat, 1880

  4. † Theodoxus acuticarinatus (Fuchs, 1870)

  5. † Theodoxus amblygonioides Wenz, 1930

  6. † Theodoxus amethystinus (Brusina, 1874)

  7. † Theodoxus anconae (Capellini, 1880)

  8. † Theodoxus angularis Gozhik, 2002

  9. † Theodoxus anonymus Anistratenko & Gozhik, 1995

  10. † Theodoxus atticus Kühn, 1963

  11. † Theodoxus barakovici (Brusina, 1902)

  12. † Theodoxus bessarabicus (Sinzov, 1896)

  13. † Theodoxus brusinai (Pavlović, 1931)

  14. † Theodoxus capellinii (Pantanelli, 1876)

  15. † Theodoxus capillaceus (Brusina, 1874)

  16. † Theodoxus crescens (Fuchs, 1870)

  17. † Theodoxus doderleini (d'Ancona, 1869)

  18. † Theodoxus dumortieri (Fontannes, 1878)

  19. † Theodoxus fasciatus Gozhik in Gozhik & Datsenko, 2007

  20. † Theodoxus gnezdai (Brusina, 1884)

  21. † Theodoxus grateloupianus (Férussac, 1823)

  22. † Theodoxus hoernesanus (Semper, 1867)

  23. † Theodoxus ingulenzis Gozhik in Gozhik & Datsenko, 2007

  24. † Theodoxus kalodictya (Andrusov, 1909)

  25. † Theodoxus koslinskyi (Porumbaru, 1881)

  26. † Theodoxus lamellatus (Brusina, 1892)

  27. † Theodoxus lineatus (Sinzov, 1896)

  28. † Theodoxus lorkovici (Brusina, 1878)

  29. † Theodoxus macedonicus Wenz, 1943

  30. † Theodoxus maculosus Gozhik in Gozhik & Datsenko, 2007

  31. † Theodoxus mayeri (Semper, 1867)

  32. † Theodoxus militaris (Neumayr, 1869)

  33. † Theodoxus miljkovici (Brusina, 1902)

  34. † Theodoxus millepunctatus (Brusina, 1902)

  35. † Theodoxus moeschi Locard, 1893

  36. † Theodoxus morulus Kühn, 1963

  37. † Theodoxus mutinensis (d'Ancona, 1869)

  38. † Theodoxus neumayri (Burgerstein, 1877)

  39. † Theodoxus nitens Gozhik, 2002

  40. † Theodoxus nivosus (Brusina, 1874)

  41. † Theodoxus novorossicus (Sinzov, 1896)

  42. † Theodoxus obtusangulaeformis Gozhik, 2002

  43. † Theodoxus oxytropida (Andrusov, 1909)

  44. † Theodoxus pavlovici Milošević, 1983

  45. † Theodoxus philippianus Locard, 1883

  46. † Theodoxus pilari (Brusina, 1884)

  47. † Theodoxus platystoma (Brusina, 1874)

  48. † Theodoxus pseudodanubialis (Sinzov, 1896)

  49. † Theodoxus pseudofluviatilis Locard, 1893

  50. † Theodoxus pseudograteloupanus (Sinzov, 1884)

  51. † Theodoxus radmanesti (Fuchs, 1870)

  52. † Theodoxus radovanovici (Brusina, 1893)

  53. † Theodoxus reiseri (Brusina, 1902)

  54. † Theodoxus rugosus (Pavlović, 1931)

  55. † Theodoxus sagittiferus (Brusina, 1874)

  56. † Theodoxus schachmaticus (Andrusov, 1909)

  57. † Theodoxus scoliogramma (Brusina, 1884)

  58. † Theodoxus semidentatus (Sandberger, 1875)

  59. † Theodoxus serrulatus (Brusina, 1892)

  60. † Theodoxus slavonicus (Brusina, 1878)

  61. † Theodoxus sophievkaensis Gozhik in Gozhik & Datsenko, 2007

  62. † Theodoxus stanae (Brusina, 1893)

  63. † Theodoxus stefanescui (Fontannes, 1887)

  64. † Theodoxus stefanescuiformis Gozhik in Gozhik & Datsenko, 2007

  65. † Theodoxus subquadrofasciatus Gozhik, 2002

  66. † Theodoxus subslavonicus Gozhik, 2002

  67. † Theodoxus sundicus (Andrusov, 1909)

  68. † Theodoxus suskalovici (Pavlović, 1903)

  69. † Theodoxus sycophantus (Brusina, 1878)

  70. † Theodoxus tropidophorus (Brusina, 1884)

  71. † Theodoxus turbinatus (Fuchs, 1870)

  72. † Theodoxus veljetinensis (Pavlović, 1903)

  73. † Theodoxus venustus (Brusina, 1897)

  74. † Theodoxus xanthozona (Brusina, 1884)

  75. † Theodoxus zagradovkaensis Gozhik, 2002

  76. † Theodoxus zivkovici (Pavlović, 1903)

  77. Subgenus Neritonyx Andrusov, 1912

  78. † Theodoxus unguiculatus (Seninski, 1905)

  79. Subgenus Ninniopsis Tomlin, 1930

  80. † Theodoxus colchicus (Andrusov, 1912)

  81. Subgenus Neritaea Roth, 1855

  82. Theodoxus anatolicus (Récluz, 1841)

  83. † Theodoxus groyanus (Férussac, 1823)

  84. Theodoxus jordani (G. B. Sowerby I, 1836)

  85. † Theodoxus micans (Gaudry & Fischer in Gaudry, 1867)

  86. Theodoxus niloticus (Reeve, 1856) - synonym: Theodoxus africanus (Reeve, 1856)

  87. † Theodoxus subdoricus Schütt, 1976

  88. Theodoxus subterrelictus Schütt, 1963

  89. Theodoxus peloponensis (Récluz, 1841)

  90. Theodoxus varius (Menke, 1828)

  91. Subgenus Theodoxus Montfort, 1810

  92. Theodoxus altenai Schütt, 1965

  93. Theodoxus baeticus (Lamarck, 1822)

  94. † Theodoxus becenensis (Cobălcescu, 1883)

  95. † Theodoxus brenneri (Handmann, 1882)

  96. † Theodoxus carasiensis Jekelius, 1944

  97. † Theodoxus constantiae Stefanescu, 1896

  98. † Theodoxus culceri (Porumbaru, 1881)

  99. † Theodoxus cyrtocelis (Krauss, 1852)

  100. Theodoxus danubialis (Pfeiffer, 1828)

  101. † Theodoxus denisluensis (Oppenheim, 1919)

  102. † Theodoxus deperditus (Almera, 1894)

  103. † Theodoxus elongatulus (Philippi, 1844)

  104. † Theodoxus eugenii Jekelius, 1944

  105. Theodoxus euxinus (Clessin, 1886)

  106. Theodoxus fluviatilis (Linnaeus, 1758) - type species. 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) does not consider Theodoxus velox Anistratenko, 1999 to be a distinct species from Theodoxus fluviatilis.

  107. † Theodoxus gregarius (Thomä, 1845)

  108. Theodoxus heldreichi (Martens, 1879)

  109. † Theodoxus hisingeri (Bellardi & Michelotti, 1841)

  110. Theodoxus hispalensis (Martens, 1879)

  111. † Theodoxus jekeliusi Jurišić-Polšak, 1979

  112. † Theodoxus leobersdorfensis (Handmann, 1887)

  113. Theodoxus meridionalis (Philippi, 1836)

  114. † Theodoxus milessii Papp, 1979

  115. † Theodoxus moosbrunnensis Papp, 1953

  116. † Theodoxus morellii (Bellardi & Michelotti, 1841)

  117. Theodoxus pallasi Lindholm, 1924 - synonym: Theodoxus lituratus (Eichwald, 1838)

  118. † Theodoxus pilidei (Tournouër, 1879)

  119. Theodoxus prevostianus (Pfeiffer, 1828)

  120. † Theodoxus punctatolineatus (Sinzov, 1896)

  121. Theodoxus saulcyi (Bourguignat, 1852)

  122. † Theodoxus scamandrius (Calvert & Neumayr, 1880)

  123. † Theodoxus semiplicatus (Neumayr in Herbich & Neumayr, 1875)

  124. † Theodoxus simplex (Fuchs, 1877)

  125. † Theodoxus subglobosus (Eichwald, 1853)

  126. Theodoxus subthermalis (Bourguignat in Issel, 1865) or Theodoxus fluviatilis subthermalis Issel, 1865

  127. Theodoxus transversalis (Pfeiffer, 1828)

  128. † Theodoxus trifasciatus (Grateloup, 1839)

  129. † Theodoxus turislavicus Jekelius, 1944

  130. Theodoxus valentinus (Graells, 1846)

  131. † Theodoxus zografi (Brusina, 1902)

  132. Subgenus ?

  133. † Theodoxus abnormis (Jenkins, 1864)

  134. † Theodoxus almelae (Revilla, 1958)

  135. Theodoxus astrachanicus Starobogatov, 1994

  136. † Theodoxus banaticus Jekelius, 1944

  137. † Theodoxus bohotinensis (Simionescu & Barbu, 1940)

  138. † Theodoxus bolivari (Royo Gómez, 1922)

  139. † Theodoxus bukowskii (Oppenheim, 1919)

  140. Theodoxus cariosus (Wood, 1828)

  141. Theodoxus coronatus (Leach, 1815)

  142. † Theodoxus crenulatus (Klein, 1853)

  143. † Theodoxus cunici (Brusina, 1892)

  144. Theodoxus danasteri (Lindholm, 1908)

  145. † Theodoxus doricus (Neumayr, 1880)

  146. Theodoxus euxinus (Clessin, 1886)

  147. Theodoxus gloeri Odabaşi & Arslan, 2015

  148. Theodoxus gurur Sands & Glöer, 2020

  149. † Theodoxus hellenicus (Bukowski, 1896)

  150. † Theodoxus imbricatus (Brusina, 1878)

  151. † Theodoxus intracarpaticus Jekelius, 1944

  152. † Theodoxus licherdopoli (Stefanescu, 1896)

  153. Theodoxus luteofasciatus (Miller, 1879)

  154. Theodoxus macri (Sowerby, 1849)

  155. Theodoxus maresi (Bourguignat, 1864)

  156. Theodoxus marteli (Pallary, 1920)

  157. Theodoxus milachevitchi Golikov & Starobogatov, 1966

  158. † Theodoxus mutinensis (d'Ancona, 1869)

  159. Theodoxus numidicus (Récluz, 1841)

  160. Theodoxus pallasi Lindholm, 1924

  161. Theodoxus pallidus Dunker, 1861

  162. † Theodoxus pappi Sauerzopf, 1952

  163. † Theodoxus paradisii (Magrograssi, 1928)

  164. † Theodoxus patrae Esu & Girotti, 2015

  165. † Theodoxus percarinatus (Oppenheim, 1919)

  166. Theodoxus pilidei (Tournouêr, 1879)

  167. † Theodoxus politioanei Jekelius, 1944

  168. † Theodoxus politus Jekelius, 1944

  169. Theodoxus poppei Ahuir, 2020

  170. † Theodoxus postcrenulatus Papp, 1953

  171. † Theodoxus prozlatarici Jekelius, 1944

  172. † Theodoxus pseudodacicus Neubauer, Harzhauser, Georgopoulou, Mandic & Kroh, 2014

  173. † Theodoxus quadrifasciatus (Bielz, 1864)

  174. † Theodoxus reticulatus Pană, 1990

  175. † Theodoxus rhodiensis (Tournouër in Fischer, 1877)

  176. Theodoxus sarmaticus (Lindholm, 1901)

  177. Theodoxus schultzii (Grimm, 1877)

  178. † Theodoxus scriptus (Stefanescu, 1896)

  179. † Theodoxus serratiliniformis Geyer, 1914

  180. † Theodoxus sinjanus (Brusina, 1876)

  181. † Theodoxus soceni Jekelius, 1944

  182. † Theodoxus sphaeroidalis (Revilla, 1958)

  183. † Theodoxus spratti (Jenkins, 1864)

  184. † Theodoxus stoicai Neubauer, Harzhauser, Georgopoulou, Mandic & Kroh, 2014

  185. Theodoxus subthermalis Bourguignat in Issel, 1865

  186. † Theodoxus susuzianus Harzhauser, Neubauer & Hoşgör, 2018

  187. † Theodoxus timisensis Jekelius, 1944

  188. † Theodoxus trilophosensis Rust, 1997

  189. Theodoxus velascoi (Graëlls, 1846)

  190. Theodoxus velox V. Anistratenko in O. Anistratenko, Starobogatov & V. Anistratenko, 1999

  191. Theodoxus vespertinus (Sowerby II, 1849)

  192. Theodoxus wesselinghi Sands & Glöer, 2020

  193. Theodoxus wilkei Sands & Glöer, 2020

  194. † Theodoxus zlatarici (Brusina, 1902)

  195. Synonyms:

  196. † Theodoxus burdigalensis (d'Orbigny, 1852): synonym of † Theodoxus (Theodoxus) trifasciatus (Grateloup, 1839)

  197. Theodoxus cariosus (Wood, 1828) and Theodoxus cariosa are synonyms of Neritina cariosa (Wood, 1828)

  198. † Theodoxus dacicus Jekelius, 1944: synonym of † Theodoxus (Theodoxus) leobersdorfensis dacicus Jekelius, 1944

  199. Theodoxus doriae Issel, 1865: synonym of Theodoxus fluviatilis (Linnaeus, 1758): synonym of Theodoxus (Theodoxus) fluviatilis (Linnaeus, 1758)

  200. Theodoxus euphraticus (Mousson, 1874): synonym of Neritina euphratica Mousson, 1874

  201. † Theodoxus intracarpatica Jekelius, 1944: synonym of † Theodoxus intracarpaticus Jekelius, 1944

  202. Theodoxus lutetianus Montfort, 1810: synonym of Theodoxus fluviatilis (Linnaeus, 1758): synonym of Theodoxus (Theodoxus) fluviatilis (Linnaeus, 1758)

  203. † Theodoxus mariae (Handmann, 1887): synonym of † Neritina mariae Handmann, 1887

  204. † Theodoxus martensi (Brusina, 1884): synonym of † Ninnia martensi (Brusina, 1884)

  205. Theodoxus neglectus (Pease, 1861): synonym of Neritina neglecta (Pease, 1861)

  206. † Theodoxus oslavanensis Rzehak, 1893: synonym of † Theodoxus (Theodoxus) cyrtocelis austriacus (Rzehak, 1893)

  207. Theodoxus oualaniense (Lesson, 1831): synonym of Clithon oualaniense (Lesson, 1831)

  208. † Theodoxus perlongus Jekelius, 1944: synonym of † Theodoxus (Theodoxus) zografi perlongus Jekelius, 1944

  209. † Theodoxus petralbensis Jekelius, 1944: synonym of † Theodoxus (Theodoxus) zografi petralbensis Jekelius, 1944

  210. † Theodoxus pictus (Férussac, 1823): synonym of † Agapilia picta (Férussac, 1823)

  211. Theodoxus reclivatus Say, 1822: synonym of Neritina usnea (Röding, 1798)

  212. † Theodoxus rumanus (Stefanescu, 1896): synonym of † Theodoxus (Calvertia) stefanescui (Fontannes, 1887)

  213. † Theodoxus scamandri (Calvert & Neumayr, 1880): synonym of † Theodoxus (Theodoxus) scamandrius (Calvert & Neumayr, 1880)

  214. Theodoxus vespertinus (G. B. Sowerby II, 1849): synonym of Neritina vespertinus (G. B. Sowerby II, 1849)

  215. † Theodoxus vetranici (Brusina, 1902): synonym of † Neritina vetranici (Brusina, 1902)

  216. Theodoxus virginea (Linnaeus, 1758): synonym of Neritina virginea (Linnaeus, 1758)

  217. † Theodoxus (Vittoclithon) pictus bizonalis (Grateloup, 1839): synonym of † Agapilia picta bizonalis (Grateloup, 1839)

References

This article incorporates public domain text from the reference

  • Eichhorst T.E. (2016). Neritidae of the world. Volume 2. Harxheim: Conchbooks. pp.696–1366