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Pelophylax bedriagae (Camerano, 1882)
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.
Rana (Rana) bedriagae — Dubois, 1987 "1986", Alytes, 5: 41, by implication.
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 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 Polon., 39: 179, without discussion (following unpublished dissertation of P. Beerli, 1994), and by Schneider and Sinsch, 1999, Z. Zool.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 (Pelophylax) bedriagae — Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 332.
Hylarana bedriagae — Chen, Murphy, Lathrop, Ngo, Orlov, Ho, and Somorjai, 2005, Herpetol. J., 15: 237, by implication.
Pelophylax bedriagae — 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.
English Names
Bedriaga's Frog (Rana bedriaga: Frank 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 (Arnold, 2002, Rept. Amph. Eur., Ed. 2: 97).
Levant Green Frog (Baha El Din, 2006, Guide Rept. Amph. Egypt: 42).
Levantine Green Frog (Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 384).
Bedriaga's Marsh Frog (Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 27).
Distribution
Upper Nile (from approximately the vicinity of Luxor) and the Delta of Egypt through Israel to western and southern Turkey; Limnos and Rhodes Is. (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.
Comment
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 ridibundus, Pelophylax 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 levantinus. Pelophylax 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 ridibundus. Plötner, 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 Turkey [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, Turkey, is clearly an unnamed species. Schneider, 2001, Bonn. Zool. Beitr., 50: 35–48, delimited the range in southern Turkey. 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 Turkey 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 ridibunda. Toli, 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 cerigensis. Lyapkov, 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. esculentus. Lukanov, 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. Cyprus records now apply to Pelophylax cypriensis. Yildiz, 2020, Amph. Rept. Conserv., 14 (2: e242): 147, reported the species from Kilis Province, southern Turkey. 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. Corti, Biaggini, Nulchis, Cogoni, Cossu, Frau, Mulargia, Lunghi, and Bassu, 2022, Acta Herpetol., Firenze, 17: 125–133, reported on the range in Sardinia.
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