Theodoxus fluviatilis

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Theodoxus fluviatilis
1854 engraving of Theodoxus fluviatilis
Four shells of Theodoxus fluviatilis
Conservation status
NE[4]
Scientific classification
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
(unranked): clade Neritimorpha

clade Cycloneritimorpha

Superfamily: Neritoidea
Family: Neritidae
Subfamily: Neritininae
Tribe: Theodoxini
Genus: Theodoxus
Subgenus: Theodoxus[5]
Species: T. fluviatilis
Binomial name
Theodoxus fluviatilis
(Linnaeus, 1758)[6]
Synonyms[7][8][9][10]
  • Nerita fluviatilis Linnaeus, 1758
  • Theodoxus velox Anistratenko, 1999
  • Theodoxus brauneri Lindholm, 1908[10] - or as separate species[11][5])
  • Theodoxus brauneri f. lacrymans Lindholm, 1908
  • Theodoxus brauneri f. alboguttata Lindholm, 1908
  • Theodoxus brauneri f. pulherrima Lindholm, 1908
  • Theodoxus lutetianus Montfort, 1810
  • Theodoxus dniestroviensis Put, 1972

Theodoxus fluviatilis, common name the river nerite, is a species of small freshwater and brackish water snail with a gill and an operculum, an aquatic gastropod mollusk in the family Neritidae, the nerites.

Theodoxus fluviatilis is the type species of the genus Theodoxus.[10] Anistratenko (2005)[10] designated the lectotype for Theodoxus fluviatilis.[10]

Subspecies

Several subspecies are (inconsistently) recognized by various authors:

  • Theodoxus fluviatilis fluviatilis (Linnaeus, 1789)[12][13][5] - it has been described from freshwater[14]
  • Theodoxus fluviatilis fluviatilis f. fontinalis Brard, 1815[12] or as a synonym of Theodoxus fluviatilis[5]
  • Theodoxus fluviatilis littoralis (Linnaeus, 1789)[12] - it has been described from brackish water,[14] or as a synonym of Theodoxus fluviatilis[5]
  • Theodoxus fluviatilis sardous (Menke, 1830)[5]
  • Theodoxus fluviatilis subthermalis Issel, 1865[5] - or Theodoxus subthermalis (Bourguignat in Issel, 1865)[7]
  • Theodoxus fluviatilis thermalis (Dupuy, 1851)[5]
    Theodoxus fluviatilis thermalis Syntype MHNT
  • Theodoxus fluviatilis transversetaeniatus A. J. Wagner, 1928[5]
  • Theodoxus fluviatilis dalmaticus Sow. - in Lake Ohrid[15]
  • Theodoxus fluviatilis euxinus (Clessin, 1885) has been considered as a subspecies[13] - see Theodoxus euxinus (Clessin, 1886)[5]

Bunje (2005)[9] does not consider Theodoxus velox Anistratenko, 1999[16] to be a distinct species from Theodoxus fluviatilis.[9]

Distribution

The distribution of this species is European.[17] Its distribution is scattered in Europe and in Western Asia[10] except the Alps and regions immediately north of the Alps.[8] It does not live in Norway.[8][14] Theodoxus fluviatilis has the most widespread distribution of all species in the genus Theodoxus.[9]

Western Europe:

Central Europe:

Northern Europe:

Eastern Europe:

Two views of a shell of Theodoxus fluviatilis from Krokees, Greece.[1]

Southern Europe:

Asia:

Bunje (2005)[9] hypothetized that the ancestral range of Theodoxus fluviatilis was the Ponto-Pannonian region (southern Ukraine, Romania and Hungary).[9] Firstly it colonized northern Italy, Greece and Turkey; in the second phase it colonized Spain, France and Germany.[9] Finally in the Holocene it colonized the British Isles, Sweden and the Baltic Sea.[9]

Shells of Theodoxus fluviatilis have also been found in the following sites:

Description

One row of teeth in the radula of Theodoxus fluviatilis

Theodoxus fluviatilis was originally described (under the name Nerita fluviatilis) by Carl Linnaeus in 1758. Linnaeus's original text (the type description) was very short, and reads as follows:[6]

Nerita fluviatilis, n. 632: testa rugosa, labiis edentilis. Habitat in Europa cataractis.

The shell is depressed, strongly calcified and with three whorls.[18][32] It is basically whitish or yellowish with dark reddish or violetish reticulate pattern, very variably arranged (adapted to environmental factors), sometimes partly in bands, occasionally evenly dark.[8] It is very variable in colors and in color patterns (it has great polymorphism in colors).[10] In the Mediterranean region there are slightly different forms.[8] The shell may be corroded.[14] Operculum is light reddish with red margin,[8] with broad rib and without apophysis.[8]

The width of the shell is 5-9 mm,[8] up to 11 mm[22] or up to 13 mm.[8] The height of the shell is 4-6.5 mm,[8] or up to 7 mm.[22]

The maximum width of the shell of freshwater population is 13.1 mm.[14] The maximum height of the shell of freshwater population is 9.3 mm.[14] The maximum weight of the shell is 343 mg.[14]

The maximum width of the shell of brackish water population is 9.3 mm.[14] The maximum height of the shell of brackish water population is 5.8 mm.[14] The maximum weight of the shell is 124 mg.[14]

The animal is light yellow with head black.[8] Tentacles are greyish and long.[8] Eyes are big and black, foot is whitish.[8]

Drawing of the reproductive system of a male by Lehmann (1873)[3] shows testis (on the left), prostate, vas deferens and penis (on the right).
Drawing of the reproductive system as firstly described (correctly) by Gustave Gilson (1896)[2] of a female shows:
1 - ovarium
2 - oviduct
3 - uterus
4 - diverticulum
5 - connection between bursa copulatrix and uterus
6 - receptaculum seminis
7 - bursa copulatrix
8 - vagina.

Ecology

This small snail inhabits the central and lower parts of rivers (up to 13 m deep),[8] also in brackish water[8] in tidal rivers of the estuary.[18] Sometimes it lives in lakes on unvegetated bottom.[18] It rarely lives in springs (rheocrenes), in ground water and in caves.[18] For example there was pH 7.8-8.9 in lakes in Åland Islands with Theodoxus fluviatilis.[25]

The ability of Theodoxus fluviatilis to live in freshwater and also in brackish water is a representation of phenotypic plasticity of this species.[9] It can live up to 60 m depth in coastal waters.[14] Brackish water populations can live in salinity up to 15 ‰ in the Baltic Sea[12][23] or up to 18 ‰ in the Baltic Sea and in the Black Sea.[9] Populations from brackish water tolerates higher salinity than populations from freshwater.[14] Brackish water populations have much higher accumulation of ninhydrin-positive substances in the foot.[33]

This species requires a stony substrate.[8] It lives on pebbles, sometimes on boulders and rarely on dead wood.[18] It tolerates mild organic pollution, low oxygen content (down to below 2 mg/liter) but does not tolerate long periods of droughts or ice.[8] It lives in mesotrophic waters, sometimes in oligotrophic.[18] Theodoxus fluviatilis has large phenotypic plasticity and it was found to be living on stones and on the dead wood in freshwater environment; while it lives on stones and on Fucus vesiculosus, Potamogeton spp. and Zostera marina in brackish water in the Baltic Sea.[14] It can also occur on aggregates of Mytilus.[14]Theodoxus fluviatilis together with the isopod Saduria entomon has been found to be a dominant part of the fauna biomass in the central and northern Baltic Sea.[34] Brackish water populations can reach densities up to 200-1000 snails per m².[14] Theodoxus fluviatilis dalmaticus in Lake Ohrid can reach population densities up to 6412 snails per m².[15]

It feeds on algae cover[8] and diatoms on stones.[14]

Sexes are separate (dioecious) and cross-fertilization occurs.[18] There is unique structure of the flagellum of spermatozoon: the flagellum is divided into two parts.[35] Eggs are laid in an egg mass[18] from mid-April to October.[8] There are less than 10 eggs.[18] Only one egg develops and other eggs serve as nutrition for the embryo.[36]

0.5-1 mm sized juveniles hatch after 30 days (in 25°C) or after 65 days (in 20°C).[12][8] Sexual maturity is in less than 1 year.[18] The life span is 2-3 years.[8]

Parasites of Theodoxus fluviatilis include:

This species is threatened mainly by river engineering, and water pollution in densely populated regions.[8]

References

This article incorporates public domain text from references[6][8]

  1. 1.0 1.1 "Picture summary of Theodoxus-fluviatilis_04.jpg". AnimalBase, last modified 20 June 2008, accessed 13 April 2011.
  2. Gilson G. (1896) "The female organs of Neritina fluviatilis". Proceedings of the Malacological Society of London 2: 81-83.
  3. Lehmann R. (1873). Die lebenden Schnecken und Muscheln der Umgegend Stettins und in Pommern mit besonderer Berücksichtigung ihres anatomischen Baues. R. Friedländer & Sohn, Berlin. page 261. Plate 19, figure 94.
  4. 2006 IUCN Red List of Threatened Species. <www.iucnredlist.org>. Cited 14 June 2007.
  5. 5.0 5.1 5.2 5.3 5.4 5.5 5.6 5.7 5.8 5.9 "Theodoxus (Theodoxus) fluviatilis". Fauna Europaea, last update 27 January 2011, accessed 12 April 2011.
  6. 6.0 6.1 6.2 Linnaeus C. (1758). Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 10th edition. Vermes. Testacea: 700-781. Holmiae. (Salvius). page 777.
  7. 7.0 7.1 Kantor 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.
  8. 8.0 8.1 8.2 8.3 8.4 8.5 8.6 8.7 8.8 8.9 8.10 8.11 8.12 8.13 8.14 8.15 8.16 8.17 8.18 8.19 8.20 8.21 8.22 8.23 8.24 "Species summary for Theodoxus fluviatilis". AnimalBase, last modified 21 September 2009, accessed 11 April 2011.
  9. 9.0 9.1 9.2 9.3 9.4 9.5 9.6 9.7 9.8 9.9 9.10 Bunje, P. M. E. (2005). "Pan-European phylogeography of the aquatic snail Theodoxus fluviatilis (Gastropoda: Neritidae)". Molecular Ecology 14 (14): 4323–4340. doi:10.1111/j.1365-294X.2005.02703.x. PMID 16313596.  PDF.
  10. 10.0 10.1 10.2 10.3 10.4 10.5 10.6 Anistratenko V. V. (2005). "Lectotypes for Tricolia pullus, Gibbula divaricata and Theodoxus fluviatilis (Mollusca, Gastropoda) revisited". Vestnik zoologii 39(6): 3-10. PDF .
  11. "Species in genus Theodoxus" (n=20). AnimalBase, accessed 11 April 2011.
  12. 12.0 12.1 12.2 12.3 12.4 Glöer P. (2002). Die Süßwassergastropoden Nord- und Mitteleuropas. Die Tierwelt Deutschlands, ConchBooks, Hackenheim, 326 pp., ISBN 3-925919-60-0.
  13. 13.0 13.1 13.2 Yildirim M. Z., Koca S. B. & Kebapçi U. (2006). "Supplement to the Prosobranchia (Mollusca: Gastropoda) Fauna of Fresh and Brackish Waters of Turkey". Turkish Journal of Zoology 30: 197-204. PDF
  14. 14.0 14.1 14.2 14.3 14.4 14.5 14.6 14.7 14.8 14.9 14.10 14.11 14.12 14.13 14.14 14.15 14.16 14.17 14.18 14.19 14.20 14.21 14.22 14.23 14.24 14.25 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.PDF.
  15. 15.0 15.1 15.2 Smiljkov, S.; Budzakoska-Gjoreska, B.; Sapkarev, J.; Trajanovski, S. (2007). "Dominant species of the gastropod fauna from the littoral region in Lake Ohrid of R. Macedonia". Prilozi / Makedonska akademija na naukite i umetnostite, Oddelenie za bioloski i medicinski nauki = Contributions / Macedonian Academy of Sciences and Arts, Section of Biological and Medical Sciences 28 (1): 137–144. PMID 17921924. . PDF.
  16. Anistratenko 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.
  17. (Slovak) Lisický M. J. (1991). Mollusca Slovenska [The Slovak molluscs]. VEDA vydavateľstvo Slovenskej akadémie vied, Bratislava, 344 pp.
  18. 18.0 18.1 18.2 18.3 18.4 18.5 18.6 18.7 18.8 18.9 18.10 18.11 18.12 Falkner G., Obrdlík P., Castella E. & Speight M. C. D. (2001). Shelled Gastropoda of Western Europe. München: Friedrich-Held-Gesellschaft, 267 pp.
  19. Theodoxus fluviatilis, accessed 25 September 2008
  20. (German) Schultz H. & Schultz O. (2001). "Erstnachweis der Gemeinen Kahnschnecke, Theodoxus fluviatilis (LINNAEUS, 1758) in Österreich (Gastropoda: Neritidae)". Annalen des Naturhistorischen Museums in Wien 103B: 231-241. PDF.
  21. Red List of the molluscs (Mollusca) of the Czech Republic
  22. 22.0 22.1 22.2 22.3 22.4 22.5 (Czech) Horsák M., Juřičková L., Beran L., Čejka T. & Dvořák L. (2010). "Komentovaný seznam měkkýšů zjištěných ve volné přírodě České a Slovenské republiky. [Annotated list of mollusc species recorded outdoors in the Czech and Slovak Republics]". Malacologica Bohemoslovaca Suppl. 1: 1-37. PDF.
  23. 23.0 23.1 Glöer P. & Meier-Brook C. (2003). Süsswassermollusken. DJN, pp. 134, pages 29, 108, ISBN 3-923376-02-2.
  24. Čejka T. & Horsák M. (2002). "First records of Theodoxus fluviatilis and Sphaerium solidum (Mollusca) from Slovakia". Biologia, Bratislava 57(5): 561-562.
  25. 25.0 25.1 25.2 Carlsson R. (2000). "The distribution of te gastropods Theodoxus fluviatilis (L.) and Potamoyrgus antipodarum (Gray) in lakes on the Åland Islands, southwestern Finland". Boreal Environment Research 5: 187-195. PDF.
  26. Nagorskaya L., Moroz M., Laeno T., Veznovetz V., Pillot H. M., Dijkstra K. D. B. & Reemer M. (2002). "Macrofauna in floodplain pools and dead branches of the Pripyat river, Belarus". The Institute of Zoology NAS Belarus, 158 pp., page 56. PDF.
  27. (Russian) Butenko O. (Бутенко О. И.) (2001). "Моллюски рода Theodoxus (Gastropada, Neritidae) в Одесском заливе Черного моря. [Mollusks of Theodoxus genus (Gastropoda, Neritidae) in the Odessa Bay (the Black Sea)]". Ekologiya Morya 58: 27-28. PDF.
  28. Alexandrov B., Boltachev A., Kharchenko T., Lyashenko A., Son M., Tsarenko P. & Zhukinsky V. (2007). "Trends of aquatic alien species invasions in Ukraine". Aquatic Invasions 2(3): 215-242. PDF.
  29. Beran L. (2009). "The first record of Anisus vorticulus (Troschel, 1834) (Gastropoda: Planorbidae) in Croatia?". Malacologica Bohemoslovaca 8: 70. PDF.
  30. Fehér Z. & Eröss Z. P. (2009). "Checklist of the Albanian mollusc fauna". Schriften zur Malakozoologie 25 22-38. PDF.
  31. (Portuguese) Callapez P. (2003). "Moluscos marinhos e fluviais do Paleolítico superior da Gruta do Caldeirão (Tomar, Portugal): evidências de ordem sistemática, paleobiológica e paleobiogeográfica". Revista Portuguesa de Arqueologia 6(1): 5-15. PDF.
  32. Theodoxus fluviatilis. Marine Species Identification Portal, accessed 11 April 2011.
  33. Symanowski, F.; Hildebrandt, J. P. (2009). "Differences in osmotolerance in freshwater and brackish water populations of Theodoxus fluviatilis (Gastropoda: Neritidae) are associated with differential protein expression". Journal of Comparative Physiology B 180 (3): 337–346. doi:10.1007/s00360-009-0435-4. PMID 20012055. 
  34. Kautsky H. (1989). "Quantitative distribution of plant and animal communities of the phytobenthic zone in the Baltic Sea". Askö Laboratory, Stockholm. .
  35. Giusti, F.; Selmi, M. G. (1982). "The morphological peculiarities of the typical spermatozoa of Theodoxus fluviatilis (L.) (Neritoidea) and their implications for motility". Journal of ultrastructure research 78 (2): 166–177. PMID 7086934. 
  36. (Czech) Beran L. (1998). Vodní měkkýši ČR. Vlašim, 113 pp., ISBN 80-902469-4-X. page 45.
  37. (Russian) Chernogorenko, M. I.; Komarovova, T. I.; Kurandina, D. P. (1978). "Life cycle of the trematode, Plagioporus skrjabini Kowal, 1951 (Allocreadiata, Opecoelidae)". Parazitologiia (in Russian) 12 (6): 479–486. PMID 733319. .

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