Pelophylax ridibundus (Pallas, 1771)

Class: Amphibia > Order: Anura > Family: Ranidae > Genus: Pelophylax > Species: Pelophylax ridibundus

Rana esculenta Linnaeus, 1758, Syst. Nat., Ed. 10, 1: 212. Type(s): By indication including the frogs illustrated by Roesel von Rosenhof, 1758, Hist. Nat. Ran. Nost.: 51, pl. 13, 14, and frontispiece; specimen not know to exist according to Lönnberg, 1896, Bih. K. Svenska Vetensk. Akad. Handl., 22: 1–45, and Andersson, 1900, Bih. K. Svenska Vetensk. Akad. Handl., 26: 1–29. The only surviving type specimen is actually a specimen of Rana arvalis according to Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 143, and who first noted that the the specimen illustrated as Rana viridis aquaticae Roesel von Rosenhof, 1758, Hist. Nat. Ran. Nost.: pl. 13, should be considered a syntype and so designated the specimen illustrated as the lectotype of the taxon. Type locality: "In Europae fontibus"; restricted to "Mitteldeutschland" (= central Germany) by Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 19; restricted to "Nürnberg", Germany, by Mertens and Müller, 1940, Abh. Senckenb. Naturforsch. Ges., 451: 18, and by lectotype designation of Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 143–144. A Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana viridis Linnaeus, 1761, Fauna Svec.: 102. Reported by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 160, as an unavailable name published as a synonym of Rana esculenta. The name actually reported by Linnaeus is non-binomial, Rana viridis aquatica attributed to Roesel von Rosenhof, 1758, Hist. Nat. Ran. Nost.: 51, pl. 13. Synonymy by Günther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 12, and Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 160. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana ridibunda Pallas, 1771, Reise Vers. Prov. Russ. Reich, 1: 458. Type(s): Not stated. Type locality: "versus mare Caspium; Volgae et Iaico" (= opposite to the Caspian Sea, Volga, and on the Yaik [= Ural] River), Russia and Kazakhstan; restricted to "Gurjew [= Guryev, Kazakhstan], Nord Küste des Kaspischen Meeres" by Mertens and Müller, 1928, Abh. Senckenb. Naturforsch. Ges., 41: 20; to legitimize this restriction Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 139, suggested that a neotype designation be made from this locality.

Rana vulgaris Lacépède, 1788, Hist. Nat. Quadrup. Ovip. Serpens, 16mo ed., 2: 457. Lacépède, 1788, Hist. Nat. Quadrup. Ovip. Serpens, Quarto ed., 1: Table following page 618 and referencing account starting on page 503. Replacement name for Rana esculenta Linnaeus, 1758. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen. Rejected as published in a nonbinominal work by Opinion 2104, Anonymous, 2005, Bull. Zool. Nomencl., 62: 55.

Bufo ridibundusBonnaterre, 1789, Tab. Encyclop. Method. Trois Reg. Nat., Erp.: 15; Schneider, 1799, Hist. Amph. Nat.: 226.

Rana gigas Gmelin, 1789, Syst. Nat., Ed. 13, 1(3): 1053. Types: Not stated or known to exist; Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 150, suggested that a neotype was needed to stabilize nomenclature. Type locality: "Persia" = Iran. Synonymy with Rana ridibunda by Eichwald, 1842, Nouv. Mem. Soc. Imp. Nat. Moscou, 7: 159 (as Rana cachinnans); Schreiber, 1875, Herpetol. Eur.: 119; Mertens and Wermuth, 1960, Amph. Rept. Europas: 57. See discussion by Smith, Schneider, and Smith, 1977, J. Herpetol., 11: 423–425. Preoccupied by Rana gigas Wallbaum, 1784 (= Bufo marinus).

Rana persica Schneider, 1799, Hist. Amph. Nat.: 149. Substitute name for Rana gigas Gmelin, 1789. NHMW 19885.1 designated neotype by Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 31. Neotype from Rasht, Gilan Province, Iran. 

Rana caucasica Pallas, 1814 "1831", Zoograph. Rosso-Asiat. Sist. Omn. Animal.: 7. Types: Not stated or known to exist. Type locality: Caucasus, along the itinerary of Güldenstadt's travels, according to Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 154. A junior synonym of Rana cachinnans under first revisor principle (Camerano, 1882 "1881", C. R. Assoc. Franç. Avanc. Sci., Paris, 10: 687). Synonymy (with Rana esculenta) by Schreiber, 1875, Herpetol. Eur.: 119, and Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 38. Boulenger, 1898, Tailless Batr. Eur., 2: 270, allied this form with what is now Rana ridibunda. Considered to have been based on specimens of both Rana ridibunda and Bufo verrucosissimus by Kuzmin, 1999, Amph. Former Soviet Union: 238. Synonymy with Pelophylax ridibunda by Litvinchuk, Borkin, Skorinov, and Rosanov, 2008, Russ. J. Herpetol., 15: 20. 

Rana bufoides Güldenstadt In Pallas, 1814 "1831", Zoograph. Rosso-Asiat. Sist. Omn. Animal.: 15. Unavailable name published as a synonym, according to Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 154. Synonymy, with Rana caucasica, by Pallas, 1814 "1831", Zoograph. Rosso-Asiat. Sist. Omn. Animal.: 15.

Rana cachinnans Pallas, 1814 "1831", Zoograph. Rosso-Asiat. Sist. Omn. Animal.: 15. Types: Not stated or known to exist; adult figured in Pallas, 1842, Icon. Zoograph. Rosso-Asiat.: pl. 1, fig 2, designated lectotype by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 150. Type locality: Around the Caspian Sea, on the lower Volga and in all the small rivers and ponds of the Chersones Tauricus [= Crimea]; restricted by lectotype designation to "Krymskij Poluostrov (Crimea), Ukraine". Synonymy with Rana ridibunda by Eichwald, 1842, Nouv. Mem. Soc. Imp. Nat. Moscou, 7: 159 (as Rana cachinnans). Synonymy (with Rana esculenta) by Günther, 1864, Rept. Brit. India: 408; Schreiber, 1875, Herpetol. Eur.: 118; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 38. Boulenger, 1898, Tailless Batr. Eur., 2: 270, allied this form with what is now Rana ridibunda.

Ranaria esculentaRafinesque, 1814, Specchio Sci., 2, 2: 102. Based on misidentified specimens from Sicily.

Rana alpina Fitzinger, 1826, Neue Class. Rept.: 64. Type(s): Not stated although presumably in NHMW. Type locality: "Alpen R. Ex Europa, Asutria et Styria". Nomen nudum. Preoccupied by Rana alpina Laurenti. Synonymy with XXX by XXX. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana hispanica Fitzinger, 1826, Neue Class. Rept.: 64. Type(s): Not designated, presumably originally NHMW. Type locality: "Spanische R. Ex. Euopa, Hispania". Nomen nudum.

Rana maritima Risso, 1826, Hist. Nat. Principales Product. Eur. Merid., 3: 92. Types: Not stated or known to exist; Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 155, suggested that a neotype designation was needed to avoid nomenclaturally threatening Rana perezi and Rana lessonae. Type locality: "Marais des bords de la mer" = marshes near the Mediterranean sea in the department of Alpes Maritimes, France. Synonymy by Bonaparte, 1838, Iconograph. Fauna Ital., 2 (Fasc. 22): 123; Bonaparte, 1840, Mem. Accad. Sci. Torino, Ser. 2, 2: 443, Duméril and Bibron, 1841, Erp. Gen., 6: 345; Günther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 12; Schreiber, 1875, Herpetol. Eur.: 119; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 38. See comment. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana bosci Bory de Saint-Vincent, 1828, Résumé Herpetol. Hist. Nat. Rept.: 266. Types: Not stated or known to exist. Type locality: "marais de la Camargue, à l'embouchure du Rhône" [= Marshes of Camargue, Bouches-du-Rhône, France]. Placed on the Official Index of Rejected and Invalid Specific Names in Zoology by Anonymous, 1962, Bull. Zool. Nomencl., 19: 280. Synonymy with Rana maritima by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 156. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana viridis Eichwald, 1830, Naturhist. Skizze Lithuanien: 234. Types: Not stated or known to exist. Type localities: Volhynien, Podolien [southern Ukraine]. Nomen nudum. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 156. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Bufo caucasicusEichwald, 1831, Zool. Special.: 167. Based on misidentifications.

Rana esculenta var. tigrina Eichwald, 1831, Zool. Special.: 166. Types: Not stated or known to exist. Type locality: Persia [= Iran]. Synonymy with Rana ridibunda by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 156. Preoccupied by Rana tigrina Merrem, 1820 (= Rana tigerina Linnaeus, 1758). Synonymy with Pelophylax persicus by Safaei-Mahroo, Ghaffari, and Niamir, 2023, Zootaxa, 5279: 30.

Rana dentex Krynicki, 1837, Bull. Soc. Imp. Nat. Moscou, 10: 63. Syntypes: Presumably lost; lectotype designated by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 157, as the specimen figured in pl. 2 and described on page 65 of the original paper by Krynicki. Vedmederya, Zinenko, and Barabanov, 2009, Russ. J. Herpetol., 16: 205, noted two surviving syntypes and designated one, MNKNU 26237 as lectotype by Vedmederya, Zinenko, and Barabanov, 2009, Russ. J. Herpetol., 16: 205. Type localities: "Caucasus in amne Podkumok [= Pyatigorsk]. . . Adj. Kaleniczenkow—in lacu arundine tecto . . . in circuitu Stauropolis" = River Podkumok near 'Kaleniczenkow'—reed-covered lake near Stavropol, Russia; restricted to river Podkumok by Mertens and Wermuth, 1960, Amph. Rept. Europas: 57, and by lectotype designation. Synonymy with Rana esculenta tigrina Eichwald by Eichwald, 1842, Nouv. Mem. Soc. Imp. Nat. Moscou, 7: 157. Synonymy (with Rana esculenta) by Schreiber, 1875, Herpetol. Eur.: 119, and Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 38. Boulenger, 1891, Proc. Zool. Soc. London, 1891: 376, allied this form with what is now Rana ridibunda. Synonymy Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 157.

Bufo cachinnansHohenacker, 1837, Bull. Soc. Imp. Nat. Moscou, 10: 145.

Rana hispanica Bonaparte, 1839, Iconograph. Fauna Ital., 2 (Fasc. 24): pl. 81 (description by indication). Holotype: Specimen figured in pl. 81, fig. 5 of the original publication according to Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 157. Type locality: Not stated; restricted to Sicily, Italy, by Mertens and Wermuth, 1960, Amph. Rept. Europas: 55, this restriction supported by discussion in  Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 158. Based on a hybridogen of Rana bergeri x Rana ridibunda.

Rana taurica Bonaparte, 1840, Mem. Accad. Sci. Torino, Ser. 2, 2: 443. Types: Not designated or known to exist. Type locality: Not designated; given as Tauricus Chersonesus (= Krymskij Poluostrov, Crimea, Ukraine) by XXX. Unavailable name, published as a synonym of Rana esculenta Linnaeus. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 160.

Rana fluviatilis Bonaparte, 1840, Mem. Accad. Sci. Torino, Ser. 2, 2: 443. Types: Not stated; Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 159, suggested designating a neotype of this species that is a specimen of Rana hispanica, which Bonaparte thought might actually apply to this form. Type locality: Not designated; given as Rivers of Lombardia, Italy, by XXX. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 158. Presumably based on a hybridogen of Rana bergeri x Rana ridibunda.

Rana viridis Duméril and Bibron, 1841, Erp. Gen., 6: 343. Types: animals figured by Roesel von Rosenhof, 1758, Hist. Nat. Ran. Nost., as Rana viridis among many other illustrations and descriptions cited by Duméril and Bibron, 1841, Erp. Gen., 6: 343–45; lectotype designated by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 160, as the adult female figured under the name Rana viridis aquatica Roesel von Rosenhof, 1758, Hist. Nat. Ran. Nost.pl. 13. Type localities: Numerous by indication; restricted to Nürnberg, Germany, by lectotype designation. Nomen oblitum. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 156, who noted that this description provided nomenclatural status for Rana viridis Eichwald, 1830, a nomen nudum. Name coined as a junior synonym and homonym of Rana viridis Linnaeus, 1761. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta tigrina Eichwald, 1842, Nouv. Mem. Soc. Imp. Nat. Moscou, 7: 157. Substitute name for Rana dentex Drynicki, 1837.

Pelophylax esculentusFitzinger, 1843, Syst. Rept.: 31.

Rana esculenta var. lineata Massalongo, 1854, Mem. Accad. Agric. Commerc. Arti Verona, 29: 427. Types: Not stated or known to exist. Type locality: "provincia Veronese", Italy. Preoccupied by Rana lineata Schneider, 1799, Hist. Amph. Nat.: 138 (= Leptodactylus lineatus). Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 161. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. vulgaris Massalongo, 1854, Mem. Accad. Agric. Commerc. Arti Verona, 29: 427. Types: Not stated or known to exist. Type locality: "provincia Veronese", Italy. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 161. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. roseo-virescens Massalongo, 1854, Mem. Accad. Agric. Commerc. Arti Verona, 29: 427. Types: Not stated or known to exist. Type locality: "provincia Veronese", Italy. Synonymy by Lessona, 1877, Atti R. Accad. Lincei, Ser. 3, 1: 1060; Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 161. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. marmorata Massalongo, 1854, Mem. Accad. Agric. Commerc. Arti Verona, 29: 427. Types: Not stated or known to exist. Type locality: "provincia Veronese", Italy. Senior homonym of Rana marmorata Hallowell, 1861 (= Pelophylax nigromaculatus). Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 161. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Pelophylax hispanicusFitzinger, 1861 "1860", Sitzungsber. Akad. Wiss. Wien, Phys. Math. Naturwiss. Kl., 42: 414.

Rana esculenta var. hispanicaSteindachner, 1867, Reise Österreichischen Fregatte Novara, Zool., Amph.: 16; Boettger, 1883, Rept. Amph. Marocco: 46. [Based on misidentified specimens from Morocco]; Boscá, 1880, Bull. Soc. Zool. France, 5: 260.

Rana agrorum Gistel, 1868, Die Lurche Europas: 165. Types: Not stated or known to exist. Type locality: Not stated; designated as Germany by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 164. Synonymy Mertens, 1936, Senckenb. Biol., 18: 76; Mertens and Wermuth, 1960, Amph. Rept. Europas: 55. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. cachinnansKessler, 1872, Trudy Sankt Peterburgskogo Obschesva Estestvoispytatelei, 3: 6; Camerano, 1882 "1881", C. R. Assoc. Franç. Avanc. Sci., Paris, 10: 680; Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 207.

Rana esculenta viridisLataste, 1880, Bull. Soc. Zool. France, 5: 61.

Rana esculenta var. ridibundaBoettger, 1880, Ber. Senckenb. Naturforsch. Ges., 1879–80: 213; Boulenger, 1885, Proc. Zool. Soc. London, 1885: 666; Boulenger, 1898, Tailless Batr. Eur., 2: 270.

Rana esculenta var. bedriagae Camerano, 1882 "1881", C. R. Assoc. Franç. Avanc. Sci., Paris, 10: 688. Syntypes: MZUT, but not reported recently. Type locality: "Damas [= Damascus]", Syria. Synonymy by Tarkhnishvili, Todua, and Iankoshvili, 2024, Caucasiana, 3: 1. 

Rana esculenta var. viridisBedriaga, 1883, Arch. Naturgesch., 49: 256; Camerano, 1884 "1883", Mem. Accad. Sci. Torino, Ser. 2, 35: 245 (based on Italian specimens, presumably of Rana ridibunda).

Rana fortis Boulenger, 1884, Zoologist, 3rd Ser., 1884: 220. Syntypes: Including BMNH 1947.2.31.35–51 (formerly 1884.11.20.15–24) by museum records, MCZ 6822–30 (according to Barbour and Loveridge, 1929, Bull. Mus. Comp. Zool., 69: 327) and MNHNP 84.254–256 (according to Guibé, 1950 "1948", Cat. Types Amph. Mus. Natl. Hist. Nat.: 36); MNHNP 1884.255 designated lectotype by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 170. Type locality: "North Germany"; restricted to "Berlin, Germany" by Boulenger, 1885, Proc. Zool. Soc. London, 1885: 666. Synonymy by Boulenger, 1885, Zool. Rec., 21: 16; Boulenger, 1885, Proc. Zool. Soc. London, 1885: 666; Boulenger, 1891, Proc. Zool. Soc. London, 1891: 376; Barbour and Loveridge, 1929, Bull. Mus. Comp. Zool., 69: 327; Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 170.

Rana (Baliopygus) ridibundaSchulze, 1891, Jahresber. Abhandl. Naturwiss. Ver. Magdeburg, 1890: 177.

Rana (Baliopygus) viridisSchulze, 1891, Jahresber. Abhandl. Naturwiss. Ver. Magdeburg, 1890: 177.

Rana esculenta var. alpinaBoettger, 1892, Kat. Batr. Samml. Mus. Senckenb. Naturforsch. Ges.: 7.

Rana esculenta var. maxima Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 207. Types: Not designated. Type locality: "Hungaria" (as cited by Schreiber, 1875, Herpetol. Eur.: 118) for his Rana esculenta var. m.

Rana esculenta var. nigromaculata Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 208. Types: Not designated or known to exist. Type locality: Not designated although implied to be in Bohemia. Based on Rana esculenta var. a of Schreiber, 1875, Herpetol. Eur.: 208. Preoccupied by Rana nigromaculata Hallowell as noted by Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 208.

Rana esculenta var. concolor Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 208. Types: Not designated. Type locality: "Croatia" (as cited by Schreiber, 1875, Herpetol. Eur.: 118) for his Rana esculenta var. e on which this taxon is based. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. immaculata  Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 208. Types: Not designated. Type locality: Not designated although implied to be Bohemia. Based on Rana esculenta var. f of Schreiber, 1875, Herpetol. Eur.: 118. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. grisescens  Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 208. Types: Not designated. Type locality: Not designated although implied to be Bohemia. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. paulopunctata Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 209. Types: Not designated. Type locality: "Chersones taur." (from Schreiber, 1875, Herpetol. Eur.: 119). Based on Rana esculenta var. r of Schreiber, 1875, Herpetol. Eur.: 119. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. rubiginosa  Prazák, 1898, Zool. Jahrb., Jena, Abt. Syst., 22: 209. Types: Not designated. Type locality: Not designated although implied to be in Bohemia. Based on Rana esculenta var. t of Schreiber, 1875, Herpetol. Eur.: 119. Presumably a Pelophylax lessonae X Pelophylax ridibundus hybridogen.

Rana esculenta var. susana Boulenger, 1905, Ann. Mag. Nat. Hist., Ser. 7, 16: 552. Syntypes: BMNH (2 specimens), by original designation, these being BMNH 1905.10.14.53–54 (now renumbered 1947.2.28.97–98) according to museum records. Type locality: "not far from the ruins of Susa, at an altitude of about 400 feet . .. . [one of two specimens from] Kerkbah River", Iran. Status rejected by Boulenger, 1918, Ann. Mag. Nat. Hist., Ser. 9, 2: 247, although placement was ambiguous. 

Rana florinskii Kashchenko and Sipacev, 1913, Annu. Mus. Zool. Acad. Imp. Sci. St. Petersbourg, 18: 232–236. Types: Not designated. Type locality: Near the summer quay on Bolshoe Dikoe Lake, between the rivers Malaya Kirgizka and Bolshaya Kirgizka, 6.4 km from Tomsk City, Russia. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 172, who noted subsequently (Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 207), that the type might represent an escapee rather than a natural record or occurrence.

Rana (Rana) esculentaBoulenger, 1920, Rec. Indian Mus., 20: 9.

Rana mangischlakensis Ahl, 1925, Zool. Anz., 65: 19. Holotype: Not stated, but presumably ZMB, unnumbered in the original publication. Type locality: "Mangischlak", Kazakhstan. Thought to be related to Rana tibetana and Rana feae in the original publication (now Paa). Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 172.

Rana ridibunda ridibundaMertens, 1925, Abh. Senckenb. Naturforsch. Ges., 39: 55; Mertens and Wermuth, 1960, Amph. Rept. Europas: 57.

Rana ridibunda kurt-mülleri Gayda, 1940 "1939", Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano, 79: 267. Holotype: ZMB, according to the original publication. Type locality: "Arsen presso Petrelli" = Erzen river near Petrela, Albania.

Rana esculenta var. hungarica Toutain, 1966, Bull. Soc. Linn. Normandie, Ser. 10, 7: 180–181. Nomen nudum. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 207.

Rana kl. esculentaDubois and Günther, 1982, Zool. Jahrb., Jena, Abt. Syst., 109: 290–305.

Rana (Rana) ridibundaDubois, 1987 "1986", Alytes, 5: 41, by implication.

Rana (Rana) bedriagae — Dubois, 1987 "1986", Alytes, 5: 41, by implication.

Rana (Rana) kl. esculentaDubois, 1987 "1986", Alytes, 5: 41.

Rana ridibunda caralitana Arikan, 1988, Istanbul Univ. Fen Fak. Biyoloji Der., 53: 82. Holotype: ZDEU 18.1982.11, by original designation. Type locality: "Beysehir Lake", Konya, Turkey.Synonymy with Pelophylax bedriagae by Sinsch, Werding, and Kaya, 2023, Animals, 13 (1725): 1.

Pelophylax ridibundaFei, Ye, and Huang, 1990, Key to Chinese Amph.: 135; Ye, Fei, and Hu, 1993, Rare and Economic Amph. China: 231; Fei, 1999, Atlas Amph. China: 164.

Rana (Pelophylax) ridibundaDubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 332.

Rana (Pelophylax) kl. esculentaDubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 332.

Rana (Pelophylax) bedriagae — Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 332.

Rana balcanica Schneider and Sinsch, 1992, Z. Zool. Syst. Evolutionsforsch., 30: 297–315. Type(s): Not designated; holotype subsequently noted as ZFMK 52825 by Schneider, Sinsch, and Sofianidou, 1993, Z. Zool. Syst. Evolutionsforsch., 31: 47–63; Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 179, designated as lectotype "the adult male which produced the mating call represented in oscillogram of fig. 2c of Schnieder and Sinsch (1992)." This second lectotype designation is apparently unnecessary and invalid. Type locality: "includes Greece except for Thrace . . . has also been found at Lake Scutari in Yugoslavia [on Montenegro–Albanian border]"; designated as "ca. 2 km west of Gefyra (near Thessaloniki, Macedonia) in a pond directly on the eastern bank of the Axios River", Greece, by Schneider, Sinsch, and Sofianidou, 1993, Z. Zool. Syst. Evolutionsforsch., 31: 60. Synonymy by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 175. The argument by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 179–180, that Rana balcanica Schneider and Sinsch, 1992, and Rana balcanica Schneider, Sinsch, and Sofianidou, 1993, represent different nominal taxa is here rejected.

Rana levantina Schneider and Sinsch, 1992, Z. Zool. Syst. Evolutionsforsch., 30: 297–315. Type(s): Not designated; holotype subsequently noted as ZFMK 52836 by Schneider, Sinsch, and Nevo, 1992, Zool. Anz., 228: 97–106. Type locality: Not designated; subsequently noted as "Birket Ata, 5 km south of Hadera, Israel" by Schneider, Sinsch, and Nevo, 1992, Zool. Anz., 228: 97–106. Synonymy with Rana bedriagae by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 179, without discussion (following unpublished dissertation of P. Beerli, 1994), and by Schneider and Sinsch, 1999, Z. Zool. Syst. Evolutionsforsch., 37: 57.

Rana terentievi Mezhzherin, 1992, Dopov. Akad. Nauk Ukr., 5: 152. Holotype: ZIK 25441, by original designation; considered ZIL Amph A3 by Pisanets, 2001, Cat. Types Specimens Ukran. Acad. Sci., 1: 92 (apparently renumbered—DRF). Type locality: Obi-Garm, Tajikistan. 

Rana cerigensis Beerli, Hotz, Tunner, Heppich, and Uzzell, 1994, Not. Nat., Philadelphia, 470: 4. Holotype: MHNG 2543.92, by original designation. Type locality: Olympus, Greece—"Olympos . . . , northern Karpathos, Greece (35°44′N/27°10′E), 300 m". Synonymy by Dufresnes, Monod-Broca, Bellati, Canestrelli, Ambu, Wielstra, Dubey, Crochet, Denoël, and Jablonski, 2024, Global Change Biol., 30 (e17180): 7, Table S1. 

Rana (Pelophylax) kl. grafi Crochet, Dubois, Ohler, and Tunner, 1995, Bull. Mus. Natl. Hist. Nat. Paris, Sect. A, Zool., 17: 17. Holotype: MNHNP 1993.3374, by original designation. Type locality: "Gorges de l'Aiguillon, between Lussan (44° 09′ N, 4° 22′ E) and La Bastide, Gard, France". A Pelophylax perezi X Pelophylax ridibundus hybridogen.

Rana (Pelophylax) kl. maritima — Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 155.

Rana (Pelophylax) susana — Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 171. Provisional recognition as a distinct species on the basis of the distinctiveness of the descriptions of green frogs from northern and southern Iran.

Rana (Pelophylax) kl. hispanica — Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 157.

Rana (Pelophylax) kurtmuelleri — Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 179.

Rana (Pelophylax) terentievi — Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 179.

Rana (Pelophylax) cerigensis — Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 180.

Rana bedriagae caralitana — Jdeidi, Bilgin, and Kence, 2001, Turkish J. Zool., 25: 153-158.

Rana caralitana — Jdeidi, Bilgin, and Kence, 2001, J. Morphol., 248: 245.

Hylarana ridibundaChen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, by implication.

Hylarana kurtmuelleri — Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, by implication.

Hylarana terentievi — Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, by implication.

Hylarana cerigensis — Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, by implication.

Hylarana caralitana — Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, by implication.

Hylarana bedriagae — Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, by implication.

Pelophylax caralitanus — Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 369; Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1-13; by implication.

Pelophylax ridibundusFrost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 369; Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1–13; by implication.

Pelophylax kurtmuelleri — Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 369; Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1–13; by implication.

Pelophylax terentievi — Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 369; Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1–13, by implication.

Pelophylax cerigensis — Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 297: 369; Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1-13; by implication.

Pelophylax bedriagae bedriagae — Bülbül, Matsui, Kutrup, and Eto, 2011, Zool. Sci., Tokyo, 28: 936.

Pelophylax bedriagae caralitanus — Bülbül, Matsui, Kutrup, and Eto, 2011, Zool. Sci., Tokyo, 28: 935.

Pelophylax bedriagae caralitanus — Bülbül, Matsui, Kutrup, and Eto, 2011, Zool. Sci., Tokyo, 28: 935.

Pelophylax cypriensis Plötner, Baier, Akın, Mazepa, Schreiber, Beerli, Litvinchuk, Bilgin, Borkin, and Uzzell, 2012, Zoosyst. Evol., Berlin, 88: 276. Holotype: ZISP 10531, by original designation. Type locality: "Cyprus, southeast of Troodos (34.91611111° N, 32.90027778° E; 1383 m elevation".

Pelophylax gigas — Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 31. 

Pelophylax persicus — Safaei-Mahroo, Ghaffari, and Niamir, 2023, Zootaxa, 5279: 30.

Pelophylax ridibundus kurtmuelleri — Sindaco and Razzetti, 2021, Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano, 8: 40, but citing Speybroeck, Beukema, and Crochet, 2010, Zootaxa, 2492, for the arrangement. 

Pelophylax ridibundus cypriensis — Dufresnes, Monod-Broca, Bellati, Canestrelli, Ambu, Wielstra, Dubey, Crochet, Denoël, and Jablonski, 2024, Global Change Biol., 30 (e17180): 7, Table S1. 

Pelophylax ridibundus cerigensis — Dufresnes, Monod-Broca, Bellati, Canestrelli, Ambu, Wielstra, Dubey, Crochet, Denoël, and Jablonski, 2024, Global Change Biol., 30 (e17180): 7, Table S1.

Common Names

Laughing Frog (Shaw, 1802, Gen. Zool., 3(1): 114; Hellmich, 1962, Rept. Amph. Eur.: 74; Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 127).

German River Frog (Boulenger, 1885, Proc. Zool. Soc. London, 1885: 666).

Marsh Frog (Hellmich, 1962, Rept. Amph. Eur.: 74; Arnold and Burton, 1978, Field Guide Rept. Amph. Brit. Eur.: 85; Stumpel-Rienks, 1992, Ergänzungsband Handbuch Rept. Amph. Eur., Trivialnamen der Herpetofauna Eur.: 59; Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 109; Kuzmin, 1999, Amph. Former Soviet Union: 362; Arnold, 2002, Rept. Amph. Eur., Ed. 2: 89).

Eurasian Marsh Frog (Arakelyan, Danielyan, Corti, Sindaco, and Leviton, 2011, Herpetofauna of Armenia: 44).

Lake Frog (Arnold and Burton, 1978, Field Guide Rept. Amph. Brit. Eur.: 85; Stumpel-Rienks, 1992, Ergänzungsband Handbuch Rept. Amph. Eur., Trivialnamen der Herpetofauna Eur.: 59; Günther, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 154).

Central Asian Pond Frog (Fei, 1999, Atlas Amph. China: 164).

Green Frog (Rana esculenta [a hybrid with Rana lessonae]: Shaw, 1802, Gen. Zool., 3(1): 103; Gray, 1831, in Cuvier, Animal Kingdom (Griffith), 9—Appendix: 100; Cuvier, 1831, Animal Kingdom (M'Murtrie), 2: 78; Wood, 1863, Illust. Nat. Hist., 3: 159; Stumpel-Rienks, 1992, Ergänzungsband Handbuch Rept. Amph. Eur., Trivialnamen der Herpetofauna Eur.: 56).

Common Water-frog (Rana esculenta [a hybridogen with Rana lessonae]: Gadow, 1901, Amphibia and Reptiles: 263).

Edible Frog (Rana esculenta [a hybrid with Rana lessonae]: Lacépède, 1802, Nat. Hist. Ovip. Quadruped. (Kerr transl.): 197; Wood, 1863, Illust. Nat. Hist., 3: 159; Boulenger, 1891, Proc. Zool. Soc. London, 1891: 374; Flower, 1933, Proc. Zool. Soc. London, 1933: 844; Arnold and Burton, 1978, Field Guide Rept. Amph. Brit. Eur.: 87; Stumpel-Rienks, 1992, Ergänzungsband Handbuch Rept. Amph. Eur., Trivialnamen der Herpetofauna Eur.: 55; Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 107; Kuzmin, 1999, Amph. Former Soviet Union: 384; Arnold, 2002, Rept. Amph. Eur., Ed. 2: 92).

Graf's Frog ("Rana grafi" [hybridogenetic hemiclone]: original publication).

Boulenger's Green Frog (Rana fortisDubois and Ohler, 1996 "1994", Zool. Polon., 39: 170).

Graf's Hybrid Frog ("Rana grafi" [hybridogenetic hemiclone]: Arnold, 2002, Rept. Amph. Eur., Ed. 2: 93); Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 115 (Pelophylax perezi x Pelophylax ridibundus)  

Eurasian Marsh Frog (Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 105). 

Giant Marsh Frog (Pelophylax gigas [status uncertain]: Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 31).

Balkan Frog (Pelophylax kurtmuelleriSofianidou, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 130).

Greek Marsh Frog (Pelophylax kurtmuelleriArnold, 2002, Rept. Amph. Eur., Ed. 2: 96).

Bedriaga's Frog (Rana bedriagaeFrank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 106).

Hadera Frog (Rana levantina [no longer recognized]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 108).

Levant Water Frog (Rana bedriagae:Arnold, 2002, Rept. Amph. Eur., Ed. 2: 97).

Levant Green Frog (Rana bedriagae:Baha El Din, 2006, Guide Rept. Amph. Egypt: 42).

Levantine Green Frog (Pelophylax bedriagae:Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 384).

Bedriaga's Marsh Frog (Pelophylax bedriagae:Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 27).

Beyşehir frog (Pelophylax caralitanusArısoy and Başkale, 2019, Herpetozoa, Wien, 32: 195). 

Cyprus Water Frog (Pelophylax cypriensisBaier, Sparrow, and Wiedl, 2013, Amph. Rept. Cyprus, 2nd ed.: 52; Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 110).

Terentjev's Frog (Rana terentieviKuzmin, 1999, Amph. Former Soviet Union: 389).

Central Asian Pond Frog (Rana terentieviFei, Ye, and Jiang, 2010, Colored Atlas of Chinese Amph.: 289).

Karpathos Water Frog (Rana cerigensisArnold, 2002, Rept. Amph. Eur., Ed. 2: 96).

Karpathos Frog (Pelophylax cerigensisDufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 112).

Distribution

Central Europe east of northeastern France, north to southern shore of Baltic Sea (and extreme southern Finland), south to extreme northeastern Spain, Italy and the Balkans including eastern Greece; east to ca. 81° E in Asiatic Russia and Uzbekistan, south to northern and eastern Iran (see comment) to the Turkmenistan border and Afghanistan border. Isolated populations in the high Asir and Ha'il regions of of western and north-central Saudi Arabia, and oases of eastern Saudi Arabia; introduced into England and Italy; introduced  in the Kamchatka Peninsula of Russia.   

Upper Nile (from approximately the vicinity of Luxor) and the Delta of Egypt through Israel to western, southwestern, and southern Türkiye; Limnos I. (Greece) in the eastern Aegean; another section of the range is found in eastern Iraq to Iran (see comment); isolated record in south-central Bulgaria; Introduced into Belgium, Italy (Sardinia), Switzerland, and Malta; introduced hybrid population (with Pelophylax ridibundus) in Petropavlovsk–Kamchatsky and in the Paratunka River valley of the Kamchatka Peninsula of Russia; questionably in Bulgaria.

Eastern Iran east throughout Afghanistan, to southern Tajikistan (south of the Gissar Ridge) and adjacent northwestern Xinjiang, China; expected in extreme southeastern Azerbaijan. Possible records in southeastern Pakistan. 

Karpathos Island and Rhodos Island, Greece, with mainland population requiring genetic confirmation.

Geographic Occurrence

Natural Resident: Afghanistan, Albania, Armenia, Austria, Azerbaijan, Bahrain, Belarus, Bosnia and Herzegovina, Bulgaria, Croatia, Cyprus, Czech Republic, Denmark, Estonia, Finland, France, Georgia, Germany, Greece, Hungary, Iran, Iraq, Israel, Italy, Jordan, Kazakhstan, Kosovo, Kyrgyzstan, Latvia, Lebanon, Liechtenstein, Lithuania, Luxembourg, Moldova, Montenegro, Netherlands, North Macedonia, Palestine, State of, Poland, Romania, Russia, Saudi Arabia, Serbia, Slovakia, Slovenia, Switzerland, Syria, Tajikistan, Türkiye, Turkmenistan, Ukraine, Uzbekistan

Likely/Controversially Present: China, People's Republic of, Pakistan

Introduced: Belgium, China, People's Republic of, Italy, Malta, Spain, Switzerland, United Kingdom

Comment

Dufresnes, Monod-Broca, Bellati, Canestrelli, Ambu, Wielstra, Dubey, Crochet, Denoël, and Jablonski, 2024, Global Change Biol., 30 (e17180): 1–30, provided a substantial revision (and biogeography_ which considered several nominal species to be subspecies as well as identifying several nu- and mtDNA lineages that may require taxonomic recognition. A quotation (slightly modified to fit style) from their Table S1 summarizes the situation thusly: The numerous lineages previously assigned or associated (“cf.”) to Pelophylax ridibundus, Pelophylax kurtmuelleri, Pelophylax bedriagae, Pelophylax cypriensis, Pelophylax cerigensis or Pelophylax caralitanus all belong to a fairly young diversification (<3.5 My, their Fig. 10) characterized by extensive admixture across the most divergent lineages in northern Greece (Hotz, Beerli, Uzzell, Guex, Pruvost, Schreiber, and Plötner, 2013, J. Heredity, 104: 57–71), which altogether preliminary suggests a single species with high phylogeographic diversity (see also Speybroeck, Beukema, Dufresnes, Fritz, Jablonski, Lymberakis, Martínez-Solano, Razzetti, Vamberger, Vences, Vörös, and Crochet, 2020, Amphibia-Reptilia, 41: 139–189). Lineages with available names are thus here treated as subspecies of Pelophylax ridibundus, and the remaining lineages are listed as unnamed subspecies (Pelophylax r. cf. ridibundus). If future investigations confirm their distinctiveness and leads to new taxonomic descriptions, the nominal taxa Pelophylax ridibundus may face a nomenclatural availability issue, as its type locality (N of the Caspian Sea in W-Kazakhstan) corresponds to hybrid populations (Plötner, Köhler, Uzzell, Beerli, Schreiber, Guex, and Hotz, 2009, Mol. Phylogenet. Evol., 53: Fig. 4) with Central Asian lineages, and the name fortis (described from N-Germany) may be used instead (e.g., Dufresnes and Mazepa, 2020, Encycloped. Life Sci., 1: 718–726). Jablonski, Wynn, Masroor, Papenfuss, Litvinchuk, and Mazepa, 2021, Herpetozoa, Wien, 34: 141–147, discussed specimens and records from southwestern Pakistan which may be assignable to this species. Papežík, Sciberras, Benovics, Sciberras, Deidun, and Mikulíček, 2024, Biol. Invasions, 26: 1045–1059, discussed detected mtDNA markers for "Pelophylax cf. bedriagae" or "Pelophylax caralitanus" from collections from Malta, suggesting human introduction. Hosseinian Yousefkani, Shafiei Bafti, Rezazadeh Mashizi, and Moeinadini, 2024, North-West. J. Zool., Romania, 20(e242501): 85–89, discussed the possible effects of climate change on range in Iran,  provided a current dot map and projected climate change map. 

Comment under Pelophylax ridibundus prior to 17 March 2024: Unnamed taxa similar to Pelophylax ridibundus revealed in peninsular Italy by Uzzell and Hotz, 1979, Mitt. Zool. Mus. Berlin, 55: 13–27; in Greece and Serbia and Montenegro by Hotz and Uzzell, 1982, Proc. Acad. Nat. Sci. Philadelphia, 134: 50–79, and by Tunner and Heppich, 1982, Z. Zool. Syst. Evolutionsforsch., 20: 209–223. Status of North African frogs is uncertain. See Pelophylax saharicus. See Ma, 1979, Nat. Hist., Beijing, 1979: 37, for Chinese record. Records for the southwestern Balkans, Levant, and Egypt refer to Pelophylax epeiroticus, Pelophylax balcanicus, Pelophylax bedriagae, and Pelophylax levantinus. See account by Ye, Fei, and Hu, 1993, Rare and Economic Amph. China: 231. Nöllert and Nöllert, 1992, Die Amph. Eur.: 341–346, provided an account and polygon map. See also Mezhzherin, 1997, Zool. Zh., 76: 933–939, for discussion of morphological and biochemical variation in population in Kazakhstan. See Balletto, Cherchi, and Gasperetti, 1985, Fauna Saudi Arabia, 7: 368, for account of Arabian populations. Briggs and Ault, 1985, Herpetol. Rev., 16: 72–75, detailed the range in southwestern Saudi Arabia. See accounts by Kuzmin, 1999, Amph. Former Soviet Union: 362–377, Kuzmin, 2013, Amph. Former Soviet Union, Ed. 2: 231–244, (these two for the former USSR) and Fei, 1999, Atlas Amph. China: 164–165. Başoğlu and Özeti, 1973, Türkiye Amphibileri: 97–99, provided an account for the Turkish populations. See comment under Pelophylax bedriagae. Günther, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 154–155, discussed relevant biology and range. Arnold, 2002, Rept. Amph. Eur., Ed. 2: 89–90, provided a brief account, figure, and map, as did Günther in Engelmann, Fritzsche, Günther, and Obst, 1993, Lurche Kriechtiere Eur.: 181–182. Özeti and Yilmaz, 1994, Türkiye Amfibileri: 145–148, provided an account for Turkish populations (Rana ridibunda ridibunda and Rana ridibunda caralitana [now Pelophylax caralitanus]). Wycherley, Doran, and Beebee, 2002, Biol. J. Linn. Soc., 77: 355–365, reported on geographic varation in male advertisement call characteristics. Sartaeva, 2000, Selevinia, 2000: 38–42, provided a brief report on morphometric variation in Kazakhstan. Kyriakopoulou-Sklavounou, Xeros, Charilaou, and Tsiora, 2003, Ann. Soc. R. Zool. Belg., 133: 191–193, on the basis of allele frequencis suggested that Rana ridibunda from Crete is more closely related to Rana balcanica than to Greek populations of Rana ridibunda, suggesting that the latter is a composite. Pikulik, 1996, Zemnavodnyia Pauzuny: 85–88, provided an account for the Belarus population. Lever, 2003, Naturalized Rept. Amph. World: 222–224, discussed the introductions in England, Belgium, France, Italy, Russia, and Kyrghyzstan. Casola, Marracci, Bucci, Ragghianti, Mancino, Hotz, Uzzell, and Guex, 2004, J. Zool. Syst. Evol. Res., 42: 234–244, provided a molecular tree of western Palearctic water frogs in the topology Rana saharica (Rana cretensis (Rana perezi (Rana shqiperica (Rana epeirotica ("Rana lessonae" + Rana ridibunda))))). Kuzmin and Maslova, 2003, Adv. Amph. Res. Former Soviet Union, 8: 340–342, provided an account for Far East Russia. Because each individual of "Pelophylax esculentus" is a hybrid "Pelophylax esculentus" must be regarded as a kind of frog, not a species, Subgenus Rana, a pattern-class in the Pelophylax ridibundus-Pelophylax lessonae hybridogenetic complex. "Pelophylax esculentus" had a hybrid origin from Pelophylax lessonae and Pelophylax ridibundus, according to Berger, 1968, Acta Zool. Cracov., 13: 301–324. See Dubois, 1982, Alytes, 1: 42–49, for an outline of the complex. See also Berger, 1983, Experientia, 39: 127–130, and Spolsky and Uzzell, 1986, Mol. Biol. Evol., 3: 44–56. Many older records apply to Rana lessonae. Unidentified members of this complex have been introduced on Sardinia, according to Auclair, Auclair, and Brugiere, 1983, Alytes, 2: 53–55. See account by Kuzmin, 1999, Amph. Former Soviet Union: 3384–389. Günther, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 138–139, discussed range and relevant literature. Rana maritima was removed from the synonymy of Rana esculenta, by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 155, where it had been placed by Bonaparte, 1840, Mem. Accad. Sci. Torino, Ser. 2, 2: 443, Duméril and Bibron, 1841, Erp. Gen., 6: 345; Günther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 12; Schreiber, 1875, Herpetol. Eur.: 119; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 38, on the assumption that this was the correct name for the taxon subsequently named Rana grafi. Dubois, 1998, Amphibia-Reptilia, 19: 21, suggested that the name Rana maritima might apply to Rana bergeri or Rana hispanica. See comment under Rana grafi. Arnold, 2002, Rept. Amph. Eur., Ed. 2: 92, provided a brief account, figure, and map. Wycherley, Doran, and Beebee, 2002, Biol. J. Linn. Soc., 77: 355–365, reported on geographic varation in male advertisement call characteristics. Borkin, Korshunov, Lada, Litvinchuk, Rosanov, Shabanov, and Zinenko, 2004, Russ. J. Herpetol., 11: 194–213, reported on hybridization with Pelophylax lessonae, and the hybridogen Pelophylax "esculenta" in eastern Ukraine. Brief account for Europe provided by Günther in Engelmann, Fritzsche, Günther, and Obst, 1993, Lurche Kriechtiere Eur.: 173–175. Gislén and Kauri, 1959, Acta Vert., Stockholm, 1: 295–300, reported on Swedish populations.Pelophylax kl. grafi is a hybridogenetic hemilineage associated with Rana perezi and Rana ridibunda is "Rana grafi". Because each individual frog is a hybrid "Pelophylax grafi" cannot be considered a lineage of frogs, but a kind of frog. Discovered by Graf, Karch, and Moreillon, 1977, Experientia, 33: 1582–1584, and Graf and Polls-Pelaz, 1989, in Dawley and Bogart (eds.), Evol. Ecol. Unisexual Vert.: 289–302, "Rana grafi" was formerly referred to in the literature as Graf's Frog (e. g., Dubois, 1992, XXX),and briefly the name Rana maritima was used for this taxon (Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 155). Subsequently, Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 206, on the basis of additional evidence suggested that application of the name Rana maritima remained unsettled, and the name Rana grafi was coined for Graf's frog. Arnold, 2002, Rept. Amph. Eur., Ed. 2: 93–94, provided a brief account, figure, and map as did Günther in Engelmann, Fritzsche, Günther, and Obst, 1993, Lurche Kriechtiere Eur.: 179–181 (as "Grafscher Hybridfrosch"). The hybridogen now called Pelophylax kl. hispanicus was noted by Uzzell and Hotz, 1979, Mitt. Zool. Mus. Berlin, 55: 15, as a distinct hybridogenetic population. Hispanicus was removed from the synonymy of Rana perezi by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 157, where it had been placed by XXX. While noting that it may still be found to be a junior synonym of Rana lessonae, Dubois and Ohler attached the name Rana hispanica to the hybridogenetic population formerly known as the "southern hybrid" by Uzzell and Hotz, 1979, Mitt. Zool. Mus. Berlin, 55: 15. Records for this species from Corsica are based on specimens of Rana bergeri (J. Speybroeck, personal commun.). Baloutch and Kami, 1995, Amph. Iran: 152–157, provided an account for Iran. Ogielska, Kierzkowski, and Rybacki, 2005, Canad. J. Zool., 82: 1894–1901, discussed genome composition in the Pelophylax lessonae-Pelophylax ridibundus-Pelophylax "esculentus" hybridogenetic complex (as Rana). See comment under Pelophylax kurtmuelleri. Valakos, Pafilis, Sotiropoulos, Lymberakis, Maragou, and Foufopoulos, 2008, Amph. Rept. Greece: 118–119, discussed the species in Greece. Holsbeek, Mergeay, Hotz, Plötner, Volckaert, and De Meester, 2008, Mol. Ecol., 17: 5023-5035, noted introduced populations in Belgium. See account of the hybridogenetic system Pelophylax kl. esculentus in Italy, with which this species interacts, by Capula, Sacchi, and Razzetti, 2007, in Lanza et al. (eds.), Fauna d'Italia, 42 (Amph.): 386–392. See account for Pelophylax ridibundus in Italy by Capula, 2007, in Lanza et al. (eds.), Fauna d'Italia, 42 (Amph.): 402–404. Fei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 3: 1071–1075, noted that records from Xinjiang, China, are referable to Pelophylax terentievi. See comments by Speybroeck, Beukema, and Crochet, 2010, Zootaxa, 2492: 8, regarding the potential for cryptic species in nominal Pelophylax ridibundus in Europe. Bergers and Luijten, 2009, in Creemers and van Delft (eds.), Amph. Rept. Nederland: 242–247, provided a detail summary of the biology in the Netherlands, and Dorenbosch, 2009, in Creemers and van Delft (eds.), Amph. Rept. Nederland: : 236–241, provided a detailed account for the same area for its hybridogenetic associate "Rana esculenta". Rivera, Escoriza, Maluquer-Margalef, Arribas, and Carranza, 2011, Amf. Rept. Catalunya: 96–98, provided a brief account for northeastern Spain and adjacent France. Stojanov, Tzankov, and Naumov, 2011, Die Amph. Rept. Bulgariens: 239–246, provided a fairly detailed account and range map for Bulgaria (and for its hybridogenetic associate Pelophylax "esculentus" on page 233–238). Arakelyan, Danielyan, Corti, Sindaco, and Leviton, 2011, Herpetofauna of Armenia: 44–45, provided a brief account for Armenia and environs. Cogǎlniceanu, Székely, Samoilă, Iosif, Tudor, Plăiaşu, Stănescu, and Rozylowicz, 2013, ZooKeys, 296: 35–57, provided a dot map for Romania of this species and its hybridogenetic parasite Pelophylax kl. esculentus. Behbahani, Nokhbatolfoghahai, and Esmaeili, 2014, Iranian J. Anim. Biosyst., 10: 11–28, found significant interpopulational variation among sites in Fars Province, Iran, in life history and developmental patterns. Mikulíček, Kautman, Kautman, and Pruvost, 2014, J. Zool. Syst. Evol. Res., 53: 124–132, reported on hybridogenetic system with Pelophylax lessonae in western Slovakia (producing Pelophylax kl. esculenta). Safaei-Mahroo, Ghaffari, Fahimi, Broomand, Yazdanian, Najafi-Majd, Hosseinian Yousefkani, Rezazadeh, Hosseinzadeh, Nasrabadi, Mashayekhi, Motesharei, Naderi, and Kazemi, 2015, Asian Herpetol. Res., 6: 257–290, reported on distribution and conservation status in Iran. Karaica, Buj, Čavlović, and Stanković, 2016, Salamandra, 52: 161–170, reported on comparative morphology and ecology of the Pelophylax ridibundus/Pelophylax lessonae/Pelophylax "esculentus" complex in Croatia. MtDNA analysis suggests that the specimens assigned to this nominal species in western Iran actually represent and unnamed species, according to Pesarakloo, Rastegar-Pouyani, Rastegar-Pouyani, Kami, Najibzadeh, Khosravani, and Oraie, 2017, MtDNA, Part A, 28: 392–398. Speybroeck, Beukema, Bok, and Van Der Voort, 2016, Field Guide Amph. Rept. Brit. Eur.: 185–187, provided a brief account for this species and its hybridogenetic associate, Pelophylax kl. grafi, and range map. Litvinchuk, Borkin, Litvinchuk, and Rosanov, 2015, Russ. J. Herpetol., 22: 188–196, reported on the hybridogenetic Pelophylax esculentus complex in Kaliningrad Oblast', Russia. Herczeg, Vörös, Christiansen, Benovics, and Mikulíček, 2016, J. Zool. Syst. Evol. Res., 55: 129–137, reported on the Pelophylax lessonae/Pelophylax ridibundus/Pelophylax kl. esculentus hybridogenetic system in eastern Hungary. Beşir and Gül, 2019, Herpetol. Notes, 12: 45–51, provided a dot map for Rize Province, northeastern Türkiye. Lyapkov, Ermakov, and Titov, 2017, Zool. Zh., 9: 1384–1391, discussed an introduced hybrid population (with Pelophylax bedriagae) in the Kamchatka Peninsula of Russia. Martin, Guillemin, Nivet-Mazerolles, Landsmann, Dubos, and Eudeline, 2017, Amph. Rept. Conserv., 11: 93–107, discussed the range in central Uzbekistan. Alshammari and Ibnrahim, 2018, Adv. Bioresearch, Agra, 9: 165–177, reported on habitat and localities in the Ha'il Region of north-central Saudi Arabia. Al-Qahtani and Al-Johany, 2018, Saudi J. Biol. Sci., 25: 1380–1386, discussed and mapped this species in southwestern Saudi Arabia. Mulder, 2019, Amph. Rept. Conserv., 13: 162–172, reported the species from the Mount Ararat region of extreme eastern Türkiye. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 105, for brief summary of identifying morphology and biology, a range map, as well as a photograph. On p. 113, he provided an account of the hybridogenetic associate Pelophylax kl. esculentus (Pelophylax lessonae x Pelophylax ridibundus), and on p. 115 an account for another hybridogenetic associate, Pelophylax kl. grafi (Pelophylax perezi x Pelophylax ridibundus). Ermakov, Ivanov, Titov, Svinin, and Litvinchuk, 2019, Russ. J. Herpetol., 26: 367–370, reported on a multiplex PCR method for identifying Pelophylax ridibundus, Pelophylax bedriagae, and Pelophylax lessonae as well as the hybridogenetic associates Pelophylax kl. esculentus. Key to the species (as Pelophylax gigas), synonymy, distribution (including map), and access to literature provided for Iran by Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 1–331. Yaşar, Çiçek, Mulder, and Tok, 2021, North-West. J. Zool., Romania, 17(e201512): 232–275, discussed and grid-mapped the range in Türkiye. Papežík, Kubala, Jablonski, Doležálková-Kaštánková, Choleva, Benovics, and Mikulíček, 2021, Salamandra, 57: 105–123, discussed the difficulty of identifying this species in the Balkans from Pelophylax kurtmuelleri, Pelophylax shqipericus, and Pelophylax epeiroticus, and within a context of molecular markers provided the best characters for separating the species. Miura, Vershinin, Vershinina, Lebedinskii, Trofimov, Sitnikov, and Ito, 2021, Genes, 12 (244): 1–13, reported on intrapopulation variation in genome elimination in the hybridogenetic Pelophylax "esculentus" system. Gasimova, 2021, Biharean Biol., 15: 112–116, discussed the range in Azerbaijan, and provided a dot map. Mohaymeni, Nikcheh, Kami, and Mohammadi, 2022, J. Wildl. Biodivers., Arak, 6: 54–71, reported on morphological and morphometric variation in Pelophylax spp. of Iran, suggesting that the western populations be referred to Pelophylax cf. bedriagae and agreeing with Pesarakloo et al. (2017, above) that eastern populations in Iran represent an unnamed species. Mollov, Natchev, Koynova, Kambourov, Rashkov, Dimitrov, Todorov, Petrova, Vladov, and Uzunov, 2022, Ecol. Balkanica, Spec. Edit. 5: 43–74, discussed the presence of this species in protected areas of southeastern Bulgaria. Nekrasova and Marushchak, 2023, Biodiversity Data J., 11(e99036): 1–15, modeled the distribution in Ukraine. 

Comment under Pelophylax kurtmuelleri prior to 17 March 2024: Related and most similar to Pelophylax ridibundus (as Rana) according to the original publication. In the Section Pelophylax, subgenus Pelophylax; by implication. Sinsch and Eblenkamp, 1994, Z. Zool. Syst. Evolutionsforsch., 32: 35–43, discussed relationships (of Rana balcanica) with Rana ridibunda and Rana levantina. A more extensive description of Rana balcanica appeared (Schneider, Sinsch, and Sofianidou, 1993, Z. Zool. Syst. Evolutionsforsch., 31: 47–63) after the inadvertent earlier naming of this species. See Schneider and Haxhiu, 1994, Zool. Jahrb., Jena, Abt. Syst., 121: 248–262, for discussion of Rana balcanica (and Rana epeirotica) in Albania. See Sofianidou, Schneider, and Sinsch, 1994, Alytes, 12: 93–108, for comparative electrophoretic study. See comment under Rana epeiroticaSofianidou, 1996, Israel J. Zool., 42: 149–157, reported on an electrophoretic study of a hybrid zone between Pelophylax epeiroticus (as Rana epeirotica) and Pelophylax kurtmuelleri (as Rana balcanica). Sofianidou, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 130–131, discussed (as Rana balcanica) range and relevant literature. Arnold, 2002, Rept. Amph. Eur., Ed. 2: 96, provided a brief account, figure, and map. Lever, 2003, Naturalized Rept. Amph. World: 202–203, reported the species as introduced in Denmark (as Rana balcanica). Gavrilovic, Cvetkovic, Džukić, and Petkovski, 1999, in Legakis (ed.), Contr. Zoogeogr. Ecol. E. Mediterr. Region, 1: 205–210, reported on differences from Pelophylax ridibundus (as Rana balcanica). Lymberakis, Poulakakis, Manthalou, Tsigenopoulos, Magoulas, and Mylonas, 2007, Mol. Phylogenet. Evol., 44: 115–125, questioned on the basis of molecular evidence the distinctiveness of Pelophylax kurtmuelleri from Pelophylax ridibundusCrochet and Dubois, 2004, In Gasc et al. (eds)., Atlas Amph. Rept. Europe, Ed. 2: 499, discussed the ambiguity of data separating this taxon from Pelophylax ridibundusBressi, 2007, Atti Mus. Civ. Stor. Nat. Trieste, 53: 3–10, reported this species in Slovenia. See brief account for Greece by Valakos, Pafilis, Sotiropoulos, Lymberakis, Maragou, and Foufopoulos, 2008, Amph. Rept. Greece: 114–115. Sacchi, Razzetti, and Capula, 2007, in Lanza et al. (eds.), Fauna d'Italia, 42 (Amph.): 392–396, and Razzetti, Pellitteri-Rosa, Sacchi, Tiso, and Bernini, 2010, Atti VIII Congr. Naz. Soc. Herpetol. Ital. 2010: 341–347, discussed the introduced Italian population. Bressi, 2007, Atti Mus. Civ. Stor. Nat. Trieste, 53: 3–10, reported the species in Slovenia. Razzetti, Pellitteri-Rosa, Sacchi, Tiso, and Bernini, 2010, Atti VIII Congr. Naz. Soc. Herpetol. Ital. 2010: 341–347, discussed the expansion of the introduced range from Liguria into Piemonte and Lombardia, Italy. Dubey, Leuenberger, and Perrin, 2014, Biol. J. Linn. Soc., 112: 442–449, noted an introduced population in Switzerland. Kolenda, Pietras-Lebioda, Hofman, Ogielska, and Pabijan, 2017, Amphibia-Reptilia, 38: 187–196, present genetic evidence for the possible introduction into southwestern Poland from the border areas of Greece, Albania, and North Macedonia. Szabolcs, Mizsei, Jablonski, Vági, Mester, Végvári, and Lengyel, 2017, Amphibia-Reptilia, 38: 435–448, provided a dot map and discussion of range in Albania. Vucić, Jelić, Klobučar, Prkljačić, and Jelić, 2018, Salamandra, 54: 147–157, reported on hybridization levels with Pelophylax shqipericus in Lake Skadar, Montenegro. Bellati, Bassu, Nulchis, and Corti, 2019, BioInvasions Rec., 8: 8–25, reported an introduced population on Sardinia, Italy. Bisconti, Martino, Chiocchio, Siclari, and Canestrelli, 2019, BioInvasions Rec., 8: 26–33, reported an introduced population in Aspromonte National Park, Calabria, southern Italy. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 106, for brief summary of identifying morphology and biology, a range map, as well as a photograph. Papežík, Kubala, Jablonski, Doležálková-Kaštánková, Choleva, Benovics, and Mikulíček, 2021, Salamandra, 57: 105–123, discussed the difficulty of identifying this species in the Balkans with respect to Pelophylax shqipericus, Pelophylax epeiroticus, and Pelophylax ridibundus, and within that context molecular markers provided the best characters for separating the species. Corti, Biaggini, Nulchis, Cogoni, Cossu, Frau, Mulargia, Lunghi, and Bassu, 2022, Acta Herpetol., Firenze, 17: 125–133, reported on the range in Sardinia. Related and most similar to Pelophylax ridibundus (as Rana) according to the original publication. In the Section Pelophylax, subgenus Pelophylax; by implication. Sinsch and Eblenkamp, 1994, Z. Zool. Syst. Evolutionsforsch., 32: 35–43, discussed relationships (of Rana balcanica) with Rana ridibunda and Rana levantina. A more extensive description of Rana balcanica appeared (Schneider, Sinsch, and Sofianidou, 1993, Z. Zool. Syst. Evolutionsforsch., 31: 47–63) after the inadvertent earlier naming of this species. See Schneider and Haxhiu, 1994, Zool. Jahrb., Jena, Abt. Syst., 121: 248–262, for discussion of Rana balcanica (and Rana epeirotica) in Albania. See Sofianidou, Schneider, and Sinsch, 1994, Alytes, 12: 93–108, for comparative electrophoretic study. See comment under Rana epeiroticaSofianidou, 1996, Israel J. Zool., 42: 149–157, reported on an electrophoretic study of a hybrid zone between Pelophylax epeiroticus (as Rana epeirotica) and Pelophylax kurtmuelleri (as Rana balcanica). Sofianidou, 1997, in Gasc et al. (eds.), Atlas Amph. Rept. Eur.: 130–131, discussed (as Rana balcanica) range and relevant literature. Arnold, 2002, Rept. Amph. Eur., Ed. 2: 96, provided a brief account, figure, and map. Lever, 2003, Naturalized Rept. Amph. World: 202–203, reported the species as introduced in Denmark (as Rana balcanica). Gavrilovic, Cvetkovic, Džukić, and Petkovski, 1999, in Legakis (ed.), Contr. Zoogeogr. Ecol. E. Mediterr. Region, 1: 205–210, reported on differences from Pelophylax ridibundus (as Rana balcanica). Lymberakis, Poulakakis, Manthalou, Tsigenopoulos, Magoulas, and Mylonas, 2007, Mol. Phylogenet. Evol., 44: 115–125, questioned on the basis of molecular evidence the distinctiveness of Pelophylax kurtmuelleri from Pelophylax ridibundusCrochet and Dubois, 2004, In Gasc et al. (eds)., Atlas Amph. Rept. Europe, Ed. 2: 499, discussed the ambiguity of data separating this taxon from Pelophylax ridibundusBressi, 2007, Atti Mus. Civ. Stor. Nat. Trieste, 53: 3–10, reported this species in Slovenia. See brief account for Greece by Valakos, Pafilis, Sotiropoulos, Lymberakis, Maragou, and Foufopoulos, 2008, Amph. Rept. Greece: 114–115. Sacchi, Razzetti, and Capula, 2007, in Lanza et al. (eds.), Fauna d'Italia, 42 (Amph.): 392–396, and Razzetti, Pellitteri-Rosa, Sacchi, Tiso, and Bernini, 2010, Atti VIII Congr. Naz. Soc. Herpetol. Ital. 2010: 341–347, discussed the introduced Italian population. Bressi, 2007, Atti Mus. Civ. Stor. Nat. Trieste, 53: 3–10, reported the species in Slovenia. Razzetti, Pellitteri-Rosa, Sacchi, Tiso, and Bernini, 2010, Atti VIII Congr. Naz. Soc. Herpetol. Ital. 2010: 341–347, discussed the expansion of the introduced range from Liguria into Piemonte and Lombardia, Italy. Dubey, Leuenberger, and Perrin, 2014, Biol. J. Linn. Soc., 112: 442–449, noted an introduced population in Switzerland. Kolenda, Pietras-Lebioda, Hofman, Ogielska, and Pabijan, 2017, Amphibia-Reptilia, 38: 187–196, present genetic evidence for the possible introduction into southwestern Poland from the border areas of Greece, Albania, and North Macedonia. Szabolcs, Mizsei, Jablonski, Vági, Mester, Végvári, and Lengyel, 2017, Amphibia-Reptilia, 38: 435–448, provided a dot map and discussion of range in Albania. Vucić, Jelić, Klobučar, Prkljačić, and Jelić, 2018, Salamandra, 54: 147–157, reported on hybridization levels with Pelophylax shqipericus in Lake Skadar, Montenegro. Bellati, Bassu, Nulchis, and Corti, 2019, BioInvasions Rec., 8: 8–25, reported an introduced population on Sardinia, Italy. Bisconti, Martino, Chiocchio, Siclari, and Canestrelli, 2019, BioInvasions Rec., 8: 26–33, reported an introduced population in Aspromonte National Park, Calabria, southern Italy. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 106, for brief summary of identifying morphology and biology, a range map, as well as a photograph. Papežík, Kubala, Jablonski, Doležálková-Kaštánková, Choleva, Benovics, and Mikulíček, 2021, Salamandra, 57: 105–123, discussed the difficulty of identifying this species in the Balkans with respect to Pelophylax shqipericus, Pelophylax epeiroticus, and Pelophylax ridibundus, and within that context molecular markers provided the best characters for separating the species. Corti, Biaggini, Nulchis, Cogoni, Cossu, Frau, Mulargia, Lunghi, and Bassu, 2022, Acta Herpetol., Firenze, 17: 125–133, reported on the range in Sardinia. Foufopoulos, Roussos, Kalogiannis, Kalb, Strachinis, and Brock, 2024, Herpetozoa, Wien, 27: 231–256, discussed the range and natural history on the Sporades Islands in the Aegean Sea of Greece. 

Comment under Pelophylax bedriagae prior to 17 March 2024: Most closely related to Pelophylax perezi (as Rana) according to the original publication (but see below). In the Section Pelophylax, subgenus Pelophylax; by implication. Relationships (as Rana levantina) with Pelophylax ridibundusPelophylax perezi, and Pelophylax balcanicus discussed by Sinsch and Eblenkamp, 1994, Z. Zool. Syst. Evolutionsforsch., 32: 35–43. Plötner, 1998, J. Zool. Syst. Evol. Res., 36: 191–201, suggested that Pelophylax bedriagae is the sister taxon of Pelophylax ridibundus. See also Schneider, Sinsch, and Nevo, 1992, Zool. Anz., 228: 97–106, for a formal description of Pelophylax levantinusPelophylax bedriagae removed from the synonymy of Pelophylax ridibundus (as Rana), by Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 332, for populations in Israel formerly referred to Rana ridibunda; on the basis of call data supplied by Schneider and Sofianidou, 1985, Zool. Anz., 214: 309–319 (who considered the Israel population an unnamed subspecies of Rana ridibunda). This use of Rana levantina accepted by Beerli, Hotz, Tunner, Heppich, and Uzzell, 1994, Not. Nat., Philadelphia, 470: 1–9, for Anatolian water frogs. Schneider, 1997, Zool. Middle East, 15: 51–66, reported on the advertisement call in the vicinity of Damascus, Syria. See Plötner, 1998, J. Zool. Syst. Evol. Res., 36: 191–201, and Plötner and Ohst, 2001, Mitt. Mus. Naturkd. Berlin, Zool., 77: 5–21, for discussion of relationships. See study of Sinsch and Schneider, 1999, J. Zool. Syst. Evol. Res., 37: 67–75, for multivariate comparison of Pelophylax bedriagae and Pelophylax ridibundusPlötner, Ohst, Böhme, and Schreiber, 2002 "2001", Amphibia-Reptilia, 22: 397–412, provided evidence that nominal Rana bedriagae is composed of several cryptic species (one on Cyprus [now Pelophylax cypriensis], two in Syria, and three in Anatolian Türkiye [including what is now Pelophylax caralitanus]), interleaved with various cryptic species currently confused with Pelophylax ridibundus: one in Greece, and two in Kazakhstan (one of which may be Pelophylax ridibundus sensu stricto). Disi, Modry, Necas, and Rifai, 2001, Amph. Rept. Hashemite Kingdom Jordan: 102–104, provided an account for the Jordanian population. Arnold, 2002, Rept. Amph. Eur., Ed. 2: 97, provided a brief account, figure, and map. Sinsch, Schneider, Kaya, and Arikan, 2003 "2002", Herpetol. J., 12: 141–153, discussed morphometrics of Turkish frogs and discerned two allopatric morphs, the one restricted to Ulubey, Türkiye, is clearly an unnamed species. Schneider, 2001, Bonn. Zool. Beitr., 50: 35–48, delimited the range in southern Türkiye. Schneider, 2001, Herpetofauna, Weinstadt, 23: 24–33, provided a detailed account of the species. Hraoui-Bloquet, Sadek, Sindaco, and Venchi, 2002, Zool. Middle East, 27: 36, provided definitive records for Lebanon. Sciberras and Schembri, 2006, Herpetol. Rev., 37: 102, provided as record for Malta which they considered to represent an introducted population. Baha El Din, 2006, Guide Rept. Amph. Egypt: 41–43, provided an account for Egypt. See comment under Pelophylax cerigensis. See brief account for Greece by Valakos, Pafilis, Sotiropoulos, Lymberakis, Maragou, and Foufopoulos, 2008, Amph. Rept. Greece: 98–99, although the presence of this species in that country is disputed by more recent literature. Holsbeek, Mergeay, Hotz, Plötner, Volckaert, and De Meester, 2008, Mol. Ecol., 17: 5023–5035, noted mtDNA introgression of this species in introduced populations of Pelophylax ridibundus in Belgium. Bülbül, Matsui, Kutrup, and Eto, 2011, Zool. Sci., Tokyo, 28: 930–936, reported on mtDNA trees and suggested that pending resolution of the presence or scale of hybridization between Pelophyax bedriagae and Pelophylax caralitanus that they be considered subspecies, although previous morphological work had considered them distinct species; these authors also noted an unnamed species that is likely the sister taxon of Pelophylax caralitanus + Pelophylax bedriagae in south-central Türkiye and likely into adjacent Syria. See comments on distribution and population status in Egypt by Ibrahim, 2013, Basic & Appl. Herpetol., 27: 110–111. Dubey, Leuenberger, and Perrin, 2014, Biol. J. Linn. Soc., 112: 442–449, noted an introduced population in Switzerland. Strachinis and Roussos, 2016, Herpetol. Notes, 9: 237–248, reported the species from Limnos I., northern Aegean, Greece.  MtDNA analysis suggests that the species extends through Iraq to western Iran, according to Pesarakloo, Rastegar-Pouyani, Rastegar-Pouyani, Kami, Najibzadeh, Khosravani, and Oraie, 2017, MtDNA, Part A, 28: 392–398. Speybroeck, Beukema, Bok, and Van Der Voort, 2016, Field Guide Amph. Rept. Brit. Eur.: 192–194, provided a brief account (of Pelophylax cf bedriagae) and range map. Pesarakloo, Najibzadeh, Rastegar-Pouyani, and Rastegar-Pouyani, 2018, Biologia, Bratislava, 73: 673–681, reported on bioacoustics and morphometrics of the western Iran population, concluding that these populations are Pelophylax bedriagae, not Pelophylax ridibundaToli, Siarabi, Bounas, Pafilis, Lymberakis, and Sotiropoulos, 2018, Acta Herpetol., Firenze, 13: 117–123, presented evidence that this nominal species may contain at least two lineages, one in the Middle East and Cyprus [now Pelophylax cypriensis] and the the east Aegean Sea area which is more closely related to Pelophylax cerigensisLyapkov, Ermakov, and Titov, 2017, Zool. Zh., 9: 1384–1391, discussed an introduced hybrid population (with Pelophylax ridibundus) in the Kamchatka Peninsula of Russia. Bellati, Bassu, Nulchis, and Corti, 2019, BioInvasions Rec., 8: 8–25, reported two different introduced cryptic taxa of nominal Pelophylax bedriagae on Sardinia, Italy.  See detailed account for Egypt by Escoriza and Ben Hassine, 2019, Amph. N. Afr.: 285–289. Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 384–385, provided a brief account, photograph, and range map for the African component of the range. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 109, for brief summary of identifying morphology and biology, a range map, as well as a photograph. Pesarakloo and Najibzadeh, 2019, J. Anim. Diversity, 1: 11–19, reported on larval development and morphology. Ermakov, Ivanov, Titov, Svinin, and Litvinchuk, 2019, Russ. J. Herpetol., 26: 367–370, reported on a multiplex PCR method for identifying Pelophylax ridibundus, Pelophylax bedriagae, and Pelophylax lessonae as well as the hybridogenetic associates Pelophylax kl. esculentusLukanov, Popgeorgiev, and Tzankov, 2018, Acta Scient. Nat., 5: 54–63, reported the species on the basis of acoustic and morphological characters from south-central Bulgaria. Ivanov, Ruchin, Fayzulin, Chikhlyaev, Litvinchuk, Kirillov, Svinin, and Ermakov, 2019, Nat. Conserv. Res., 4: 125–128, reported natural transfer of mtDNA from Pelophylax cf. bedriagae into Pelophylax lessonae in the Republic of Mordovia, Russia. Cyprus records now apply to Pelophylax cypriensisYildiz, 2020, Amph. Rept. Conserv., 14 (2: e242): 147, reported the species from Kilis Province, southern Türkiye. Key to the species, synonymy, distribution (including map), and access to literature provided by Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 1–331. Mohaymeni, Nikcheh, Kami, and Mohammadi, 2022, J. Wildl. Biodivers., Arak, 6: 54–71, reported on morphological and morphometric variation in Pelophylax spp. of Iran, agreeing substantially with Pesarakloo et al.(2018, above) that the western populations be referred to Pelophylax cf. bedriagae and agreeing with Pesarakloo et al. (2017 above) that eastern populations in Iran represent and unnamed species. Mollov, Natchev, Koynova, Kambourov, Rashkov, Dimitrov, Todorov, Petrova, Vladov, and Uzunov, 2022, Ecol. Balkanica, Spec. Edit. 5: 43–74, discussed the presence of this species in protected areas of southeastern Bulgaria. Corti, Biaggini, Nulchis, Cogoni, Cossu, Frau, Mulargia, Lunghi, and Bassu, 2022, Acta Herpetol., Firenze, 17: 125–133, reported on the range in Sardinia. Khaloei, Fathinia, and Shafaeipour, 2023, Curr. Herpetol., Kyoto, 42: 55–63, reported on skeletochronology in southern Iran. 

Comment under Pelophylax caralitanus prior to 17 March 2024: Pelophylax caralitanus was removed from the synonymy of Pelophylax bedriagae by Jdeidi, Bilgin, and Kence, 2001, J. Morphol., 248: 246, where it had been placed by Böhme and Wiedl, 1994, Zool. Middle East, 10: 39; Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 178; Schneider and Sinsch, 1999, Z. Zool. Syst. Evolutionsforsch., 37: 57. Budak, Tok, and Ayaz, 2000, Turkish J. Zool., 24: 135–137, considered Rana ridibunda caralitana as part of Pelophylax ridibundus (as Rana) but did not note the revisionary literature placing this taxon in the synonymy of Pelophylax bedriagaeJdeidi, Bilgin, and Kence, 2001, Turkish J. Zool., 25: 153–158, suggested that Rana bedriagae caralitana may be a distinct species. Tosunoglu, Ayaz, and Göçmen, 2005, Anadolu Univ. Bil. Teknol. Dergisi, 6: 55–59, discussed the morphometric distinctiveness of Pelophylax ridibundus caralitanus, but were agnostic on whether caralitana was conspecific with bedriagae or ridibundusBülbül, Matsui, Kutrup, and Eto, 2011, Zool. Sci., Tokyo, 28: 930–936, reported on mtDNA trees and suggested that pending resolution of the presence or scale of hybridization between Pelophyax bedriagae and Pedlophylax caralitanus that they be considered subspecies, although previous morphological work had considered them distinct species; these authors also noted an unnamed species that is likely the sister taxon of Pelophylax caralitanus + Pelophylax bedriagae in south-central Türkiye and likely into adjacent Syria. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 111, for brief summary of identifying morphology and biology, a range map, as well as a photograph. Yaşar, Çiçek, Mulder, and Tok, 2021, North-West. J. Zool., Romania, 17(e201512): 232–275, discussed and grid-mapped the range in Türkiye. Kıraç, Gidiş, Mert, and Başkale, 2022, Amph. Rept. Conserv., 16 (e303): 76–85, reported on the alarming and pending habitat diminution due to climate change. Yıldırım, Ilgaz, Kumlutaş, Candan, Najafi-Majd, Başkale, and Kaya, 2023, Anat. Rec., 306: 378–385, provided a comparative osteology. Sinsch, Werding, and Kaya, 2023, Animals, 13 (1725): 1–13, discussed vocalization and the existing evidence and replaced Pelophylax caralitanus into the synonymy of Pelophylax bedriagae. On the basis of mtDNA analysis Toli, Bounas, Christopoulos, Pafilis, and Sotiropoulos, 2023, Amphibia-Reptilia, 44: 277–287, transferred the records of this species from Rhodes, Greece, to Pelophylax cerigensis

Comment under Pelophylax cypriensis prior to 17 March 2024: Confused with Pelophylax bedriagae prior to its naming according to Plötner, Baier, Akın, Mazepa, Schreiber, Beerli, Litvinchuk, Bilgin, Borkin, and Uzzell, 2012, Zoosyst. Evol., Berlin, 88: 261–283, who noted that the population in Cyprus represented an unnamed and genetically distinctive species, which they named Pelophylax cypriensis, more closely related to Pelophylax ridibundus or Pelophylax cretensis than to populations of Pelophylax bedriagae. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 110, for brief summary of identifying morphology and biology, a range map, as well as a photograph.

Comment under Pelophylax persicus prior to 17 March 2024: See comments in Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 179. See accounts by Kuzmin, 1999, Amph. Former Soviet Union: 389–391, and Kuzmin, 2013, Amph. Former Soviet Union, Ed. 2: 255, for the former USSR. See comment under Pelophylax ridibundus. In the Pelophylax nigromaculatus group of Fei, Ye, Huang, Jiang, and Xie, 2005, in Fei et al. (eds.), Illust. Key Chinese Amph.: 111. Fei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 3: 1071–1075, provided an account, figures, and map for China and included it in their Pelophylax nigromaculatus group. Fei, Ye, and Jiang, 2010, Colored Atlas of Chinese Amph.: 289, provided a brief account including photographs. Fei, Ye, and Jiang, 2012, Colored Atlas Chinese Amph. Distr.: 323, provided an account, photographs, and a range map. See localities and range map for Afghanistan by Wagner, Bauer, Leviton, Wilms, and Böhme, 2016, Proc. California Acad. Sci., Ser. 4, 63: 457–565. Safaei-Mahroo, Ghaffari, and Niamir, 2023, Zootaxa, 5279: 1–112, provided for Iran an identification key including this species, photographs, habitat and conservation threats, a review of the literature, discussion of nomenclature and systematics, as well as dot and modeled distribution maps. 

Comment under Pelophylax cerigensis prior to 17 March 2024: In the "western Palearctic water frog group" according to the original publication. Lymberakis, Poulakakis, Manthalou, Tsigenopoulos, Magoulas, and Mylonas, 2007, Mol. Phylogenet. Evol., 44: 115–125, questioned on the basis of molecular evidence the distinctiveness of Pelophylax cerigensis from Pelophylax bedriagae. See brief account by Valakos, Pafilis, Sotiropoulos, Lymberakis, Maragou, and Foufopoulos, 2008, Amph. Rept. Greece: 102–103. See photograph, map, description of geographic range and habitat, and conservation status (as Rana cerigensis) in Stuart, Hoffmann, Chanson, Cox, Berridge, Ramani, and Young, 2008, Threatened Amph. World: 501. Toli, Siarabi, Bounas, Pafilis, Lymberakis, and Sotiropoulos, 2018, Acta Herpetol., Firenze, 13: 117–123, discussed the molecular phylogenetics of the species. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 112, for brief summary of identifying morphology and biology, a range map, as well as a photograph. On the basis of mtDNA analysis Toli, Bounas, Christopoulos, Pafilis, and Sotiropoulos, 2023, Amphibia-Reptilia, 44: 277–287, transferred the records of Pelophylax bedriagae from Rhodes, Greece, to Pelophylax cerigensis

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