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Natrix natrix (LINNAEUS, 1758)

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Higher TaxaColubridae (Natricinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesNatrix natrix moreotica (BEDRIAGA 1882)
Natrix natrix natrix (LINNAEUS 1758)
Natrix natrix scutata (PALLAS 1771)
Natrix natrix syriaca (HECHT 1930)
Natrix natrix vulgaris LAURENTI 1768 
Common NamesE: Common Grass Snake
G: Gemeine Ringelnatter
Croatian: bjelouška
Russian: Обыкновенный уж
Chinese: 水游蛇
Italian: natrice dei Balcani 
SynonymColuber Natrix LINNAEUS 1758: 220
Natrix vulgaris LAURENTI 1768 (non Coluber vulgaris RAZOUMOWSKY 1789)
Coluber scutatus PALLAS 1771
Coluber torquatus LACÉPÈDE 1789
Coluber persa PALLAS 1814
Tropidonotus natrix — BOIE 1826
Tropidonotus natrix — EICHWALD 1831
Tropidonotus ater EICHWALD 1831
Tropidonotus Natrix var. minax FITZINGER 1832
Tropidonotus Natrix var. murorum FITZINGER 1832
Coluber natrix var. dalmatina SCHINZ 1833
Coluber Natrix — GRAY 1852: 509
Tropidonotus natrix — DUMÉRIL, BIBRON & DUMÉRIL 1854: 555
Tropidonotus natrix var. picturata JAN 1863 (non SCHLEGEL 1837: 314)
Tropidonotus natrix var. bilineata JAN 1864
Tropidonotus natrix var. subbilineata JAN 1864
Coluber sparsus SCHREIBER 1875
Tropidonotus natrix var. lineata NINNI 1880
Tropidonotus natrix var. moreoticus BEDRIAGA 1881: 289 (140)
Tropidonotus natrix var. bilineata — PERACCA 1890
Tropidonotus natrix — BOULENGER 1893: 219
Natrix natrix — STEJNEGER 1907
Tropidonotus natrix — BOULENGER 1913
Tropidonotus natrix bithynius HECHT 1930
Tropidonotus natrix britannicus HECHT 1930
Tropidonotus natrix cephallonicus HECHT 1930
Tropidonotus natrix cypriacus HECHT 1930
Tropidonotus natrix dystiensis HECHT 1930
Tropidonotus natrix syrae HECHT 1930
Natrix natrix schweizeri MÜLLER 1932
Natrix natrix schweizeri — LOTZE 1972
Natrix natrix gotlandica NILSON & ANDRÉN 1981: 366
Natrix natrix fusca CATTANEO 1990
Natrix natrix — ENGELMANN et al 1993
Natrix natrix — SCHLEICH, KÄSTLE & KABISCH 1996: 510
Natrix natrix — GRUBER 2009
Natrix natrix schweizeri — KLESIUS 2013
Natrix natrix — WALLACH et al. 2014: 477
Natrix natrix fusca — KINDLER et al. 2017
Natrix natrix gotlandica — KINDLER et al. 2017

Natrix natrix moreotica (BEDRIAGA 1882)
Tropidonotus natrix moreoticus BEDRIAGA 1882: 140
Tropidonotus natrix cypriacus HECHT 1930
Natrix natrix cypriaca — BLOSAT 1998
Natrix natrix cypriaca — BAIER & WIEDL 2010
Natrix natrix cypriaca — KINDLER et al. 2017
Natrix natrix cypriaca — ZOTOS et al. 2021
Natrix natrix mtDNA lineage 7 — FRITZ & SCHMIDTLER 2020: 659
Natrix natrix moreotica — ASZTALOS et al. 2021

Natrix natrix scutata (PALLAS 1771)
Coluber scutatus PALLAS 1771: 379
Coluber persa PALLAS 1814: 41
Coluber minutus PALLAS 1814
Tropidonotus ater EICHWALD 1831
Tropidonotus Natrix var. colchica NORDMANN in DEMIDOFF 1842
Tropidonotus Natrix var. nigra NORDMANN in DEMIDOFF 1842
Natrix vibakari continentalis NIKOLSKY 1925
Tropidonotus natrix bucharensis HECHT 1930
Tropidonotus natrix schirvanae HECHT 1930
Natrix natrix persa — FORCART 1950
Natrix megalocephala ORLOV & TUNIYEV 1987
Natrix megalocephala — ENGELMANN et al 1993
Natrix megalocephala — ORLOV & TUNIYEV 1999
Natrix natrix scutata — BISCHOFF 2002
Natrix natrix persa — SCHLÜTER 2003
Natrix natrix persa — RUTSCHKE et al. 2005
Natrix megalocephala — ANANJEVA et al. 2006
Natrix megalocephala — GRUBER 2009
Natrix natrix persa — AFRASIAB 2011
Natrix megalocephala — TUNIYEV et al. 2014
Natrix megalocephala — WALLACH et al. 2014: 477
Natrix natrix persa — KINDLER et al. 2017
Natrix megalocephala — ÇAKMAK et al. 2018
Natrix megalocephala — TUNIYEV et al. 2019
Natrix natrix persa — SCHMID 2019
Natrix natrix persa — GEBHART 2020
Natrix natrix persa — FRITZ & SCHMIDTLER 2020: 658
Natrix natrix scutata — FRITZ & SCHMIDTLER 2020: 658
Natrix natrix scutata — MILKO et al. 2021
Natrix natrix scutata — ASZTALOS et al. 2021
Natrix natrix scutata — BORKIN et al. 2022

Natrix natrix syriaca (HECHT 1930)
Tropidonotus natrix syriacus HECHT 1930
Natrix natrix syriaca — KINDLER et al. 2017
Natrix natrix syriaca — FRITZ & SCHMIDTLER 2020: 658
Natrix natrix syriaca — ASZTALOS et al. 2021 (incertae sedis)

Natrix natrix vulgaris LAURENTI 1768
Natrix vulgaris LAURENTI 1768
Natrix natrix vulgaris — FRITZ & SCHMIDTLER 2020: 658
Natrix natrix vulgaris — ASZTALOS et al. 2021
Natrix natrix vulgaris — HASSL 2023 
DistributionNorway, Sweden, Finland, France (Corsica), Belgium, Netherlands, Luxembourg, Germany, Poland, Czech Republic (formerly Czechoslovakia), Denmark, Austria, Switzerland, Italy, Hungary, Romania, Croatia (including some adriatic islands), Slovenia, Slovakia, Bosnia and Hercegovina, Montenegro, Macedonia, Serbia, Albania, Bulgaria, Greece (Limnos, Lesbos, Paros, Antiparos, Despotiko, Chios, Samos, Samothraki, Andros, Corfu, Milos: Kimolos, Karpathos), Cyprus,
N Iran, Iraq, Syria, NW China (Xinjiang)

Russia (incl. Siberia), Estonia, Latvia, Lithuania, Belarus, Moldova, Ukraine, Armenia, Republic of Georgia, Azerbaijan, Kazakhstan, Turkmenistan, Kyrgyzstan, NW Mongolia; elevations 0-3000 m.

moreotica: Greece. Type locality: “im Norden des Peloponnes.”

natrix: Norway, Sweden, Denmark, Germany, etc. Type locality: Fada mill pond, approximately two kilometres southeast of the Berga­-Tuna Estate, Nyköping, Sweden. Type locality: Europe

scutata: Finland, Russia (east of the Dnjepr), West Caucacus, Ukraine, Latvia, Estonia, Poland, Belarus, Turkey; Type locality: lower Ural River near Atyrau, Kazakhstan.

syriaca: N Syria. Type locality: Zincirli, İslahiye/Gaziantep, Turkey.

vulgaris: Poland, S Germany, Austria, Czech republic, Hungary, Slovakia, Romania, Bulgaria; Type locality: Oberedlitz, Ge­meinde Thaya, Niederösterreich (Lower Austria), Austria.

-------- Currently invalid subspecies: --------

calabra: Italy (Calabria); Type locality: 10 km NNE di Serro San Bruno, provincia di Catanzaro.

cypriaca: Cyprus; Type locality: Cyprus

gotlandica: Sweden (Gotland Island); Type locality: Muske mire, South Gotland, Sweden.

fusca: Greece (W Cyclades); Type locality: Kea, W Cyclades.

megalocephala (invalid): Soviet Union, S Russia (Caucasus), Azerbaijan, Republic of Georgia, Russia, Turkey; Type locality: Pitsunda, Republic of Georgia

persa: Italy, Slovakia, Bulgaria, Bosnia-Herzegovina, Greece (Lesbos, Rhodos), Turkey, N Iran, Syria, Croatia (Krk etc.); Type locality: Persia.

schweizeri: Greece; Type locality: Μήλος (Milos), Cycla­des, Greece.

siculus: Italy (Sicily); Type locality: Sicily.  
Reproductionoviparous. N. tessellata hybridizes with N. natrix and N. maura (Mebert 2011, Trapp 2017). 
TypesNeotype: NRM 8260, designated by Norén & Åhlander 2020. Original type lost, was NRM = NHRM (Stockholm) (Andersson 1899: 14).
Holotype: ZMB 19636 [algericus]
Holotype: ZMB 31425 [bithynius]
Holotype: ZMB 33060 [britannicus]
Holotype: MZUF (Museo Zoologico Univ. Firenze), male [calabra]
Holotype: ZMB 18422 [cephallonicus]
Holotype: ZMB 16122 [cypriaca]
Holotype: ZMB 13534 [dystiensis]
Holotype: private collection (A. Cattaneo), NN/KEA 10 (2), male; 20.V.1988; A. Cattaneo leg. [fusca]
Holotype: GNM Ba.Re.Su. 1031 (Göteborg Natural History Museum) [gotlandica]
Holotype: ZISP 11846, female [megalocephala]
Lectotype: MNHN-RA 3468 [persa]
Holotype: ZMB 1938 [schirvanae]
Syntypes: lost, previously ZSM (SLM) 1869 (female), ZSM (SLM) 1870a-b (female, male) (all lost), from "Insel Milos", collected by H. Schweizer, 05.1927 and 1932. See also Franzen & Glaw 2007 [schweizeri]
Holotype: unlocated, suspected to be in ZISP but not found there fide I. Doronin, pers. comm., 19 May 2019 [scutata]
Holotype: ZMB 13533 [syrae]
Holotype: ZMB 13768 [syriaca]
Neotype: NMW 36405 [vulgaris] 
DiagnosisSize: medium-sized to large, males are smaller than the females, 70-90 cm on average; females may be much larger and especially more thickset , measuring on average 80-120 cm but to 140 cm ; there are doubtful records of specimens of 180-200 cm. (modified after Geniez 2018)

Head: The head is clearly distinct from the body, and is quite oval in young animals , becoming thicker , wider and more triangular in large individuals , especially females. The medium-sized eyes have a round pupil and a grey, yellowish, orangish or red iris, depending on subspecies. (modified after Geniez 2018)

Scalation: The dorsal scales are clearly keeled, arranged in 19 rows at mid-body. 1 preocular is higher than it is wide, 3-4 postoculars (sometimes 2), generaIly 7 supralabial plates on each side, 150- 200 ventral plates, subcordals divided and arranged in pairs. (modified after Geniez 2018)

Coloration: generally greyish with a white, yellow or sometimes orangish collar composed of 2 more or less attached wide transverse marks and bordered at the rear by 2 black triangular marks, sometimes joined on the nape. Quite variable in colour, more often than not slate or mouse-grey, but also greenish, yellowish-brown , orange or brick-red. There are normally well-separated black spots or bars on the back and flanks that occur in a regular, alternating manner. There are black vertical bars on the sides of the labial plates, characteristic of this species. Some populations have 2 pale lines along the length of the body (e.g., within subspecies persa). Melanistic individuals occur rarely in some populations, quite commonly in others (e.g., Natrix natrix persa in Turkey). The belly is whitish-grey or yellowish with black marks normally in a checkered pattern, sometimes with a large black band with pale edges, sometimes entirely black. (modified after Geniez 2018)

Original description (moreoticus): “... schwarz gefärbt und mit zwei am Nacken anfangenden und sich über die ganze Länge des Rückens hinziehenden, schmalen, gelben Binden versehen.” (Bedriaga 1882: 140) 
CommentType species: Coluber Natrix LINNAEUS 1758 is the type species of the genus Natrix LAURENTI 1768. Note that the generic nomen Serpens Garsault, 1764 is a senior doxisonym of Natrix Laurenti, 1768 (DUBOIS & BOUR 2010). Kuhl in Boie 1827: 518 designated Coluber natrix Linnaeus as the type species of Tropidonotus which, however, is pre-occupied by Tropidonotus BOIE 1826: 205 who did not designate any type species although he included T. tigrinus and T. vibakari (fide Williams & Wallach 1989: 152). Stejneger (1907) placed Tropidonotus in the genus Natrix, where it remained until Malnate (1960) divided Natrix sensu lato into six genera, resurrecting Rhabdophis Fitzinger, 1843.

Distribution: see maps for N. natrix and N. helvetica and their various clades in Kindler et al. 2018. Populations west of the Rhine, including the UK, are now considered as N. helvetica. Populations from Spain and Portugal are now assigned to N. astreptophora (Pokrant et al. 2016). For a map of subspecies see ASZTALOS et al. 2021: Fig. 10. Not on Sardinia where it is replaced by N. helvetica cetti (Corti et al. 2022).

Synonymy partly after KHALIKOV & ANANJEVA (pers. comm.) and Fritz & Schmidtler 2020. Kaiser et al. 2013 considered the generic names Guystebbinsus Hoser 2012, Jackyhosernatrix Hoser 2012 invalid and rejected their use instead of Natrix. Natrix megalocephala is synonymous with Natrix natrix scutata (VENCHI & SINDACO 2006, BÖHME 2009: 61) or Natrix natrix (Kindler et al. 2013). Tropidonotus natrix var. picturata JAN 1863 as well as N. n. moreoticus BEDRIAGA 1881 are most likely mutations of Natrix natrix (Müller & Mertens 1932). Asztalos et al. 2022 synonymized cypriaca with moreotica and persa with scutata.

Habitat: terrestrial, usually near freshwater; Böse (1897) reported sightings of this species in the open ocean, 1 km and more away from the shore.

Subspecies: Only subspecies listed under subspecies are currently recognized. THORPE (1975) recognized only 4 subspecies: natrix, cetti, corsa, and helvetica (with N. n. natrix to include cypriaca, fusca, gotlandica, persa, schweizeri, scutata, and syriaca and helvetica to include helvetica, astreptophora, lanzai, and sicula). SOCHUREK (1979) suggested to recognize N.n. algirus. Natrix natrix helvetica was sometimes considered as valid species. See Orlov & Tuniyev (1987) for differences between N. n persa, N. n. scutata and Natrix megalocephala. Kindler et al. 2013, 2017 found many mismatches between morphologically defined taxa and mitochondrial clades (see e.g. Fig. S1 in Kindler et al. 2017). Baran 1976 synonymized syriaca with persa. Pokrant et al. (2016) suggested that Natrix (n.) astreptophora (Seoane, 1884) should be recognized as a distinct species.

Hybridization: In the French Pyrenees region, N. n. astreptophora and N. n. helvetica hybridize occasionally. N. n. astreptophora differs consistently from all other grass snakes by its reddish iris coloration and in having significantly fewer ventral scales and another skull morphology. Natrix natrix helvetica “hybridizes extensively with N. n. natrix” (FRITZ et al. 2012). N. natrix also hybridizes with N. tessellata, at least in Turkey (Asztalos et al. 2022).

Genome: the genome of this and the 4 other species of Natrix were sequenced by Schöneberg et al. 2023. 
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