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Pelophylax Fitzinger, 1843
Palmirana Ritgen, 1828, Nova Acta Phys. Med. Acad. Caesar Leopold Carol., Halle, 14: 278. Nomen nudum. See Dubois, 1981, Monit. Zool. Ital., N.S., Suppl., 15: 249-250. Synonymy with Pelophylax by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 183.
Pelophylax Fitzinger, 1843, Syst. Rept.: 31. Type species: Rana esculenta Linneus, 1758, by original designation.
Baliopygus Schulze, 1891, Jahresber. Abhandl. Naturwiss. Ver. Magdeburg, 1890: 177. Type species: Rana ridibunda Pallas, 1771, by subsequent designation of Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 183. The earlier designation of Rana esculenta Linnaeus, 1758, by Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 93, is in error inasmuch as Rana esculenta was not among the species listed by Schulze. Synonymy with Rana by Stejneger, 1907, Bull. U.S. Natl. Mus., 58: 93.
Nomina inquirenda - Name(s) unassigned to a living or extinct population
Hyla ranaeformis Laurenti, 1768, Spec. Med. Exhib. Synops. Rept.: 33. Holotype: By indication including specimens figured by Seba, 1734, Locuplet. Rer. Nat. Thesaur. Descript. Icon. Exp. Univ. Phys. Hist., 2: pl. 13, fig. 2 (var. a), and pl. 70, fig. 4 (var. b). Type locality: "prior in Lemno" (= mainly in Limnos, Greece) and [var. b] "Surinami". Considered by Dubois and Ohler, 1996 "1994", Zool. Polon., 39: 205, to be assignable to some member of Pelophylax, and who formed the combination Rana (Pelophylax) ranaeformis. Schinz, 1822, Thierr. Naturgesch., 2: 167, only mentioned the frog figured in pl. 13, fig. 2 of Seba, 1734, Locuplet. Rer. Nat. Thesaur. Descript. Icon. Exp. Univ. Phys. Hist., 1, thereby implying a lectotype designation of the Limnos specimen.* Calamita ranaeformis—Merrem, 1820, Tent. Syst. Amph.: 170. (DRF finds the association of this nominal taxon with Pelophylax to be questionable, because to assign the frog illustrated by Seba in vol. 2, pl. 13, fig. 2, to Pelophylax requires a number of assumptions regarding the long-lost frog in question; the frog as illustrated lacks webbing between the toes, has expanded toe tips and, five toes on the forefeet.) Nevertheless, even if the assignment to Pelophylax is accepted, the name must be treated as a nomen oblitum.
Hyla gibbosa Lacépède, 1788, Hist. Nat. Quadrup. Ovip. Serpens, Quarto ed., 1: Table following page 618 and referencing account on page 559. Replacement name for Hyla ranaeformis Laurenti, 1768. Rejected as published in a nonbinominal work by Opinion 2104, Anonymous, 2005, Bull. Zool. Nomencl., 62: 55.
Common Names
Green Frogs (Common usage).
Water Frogs (Common usage).
Distribution
North Africa, Europe, Middle East and east through Russia, the Caucasus, and Afghanistan to southern Far East Russia and China to Korea, Japan, Taiwan; Eritrea; introduced on Canary Is.; isolated populations in the high Asir region of of western Saudi Arabia, and oases of eastern Saudi Arabia and Bahrain.
Comment
Dubois, 1982, Alytes, 1: 42–49, discussed the Rana esculenta complex (now Pelophylax). Dubois, 1977, Mem. Soc. Zool. France, 39: 161–284; Hotz and Bruno, 1980, Rend. Accad. Naz. Sci. XL, Mem. Sci. Fis. Natur., Ser. 5, 4: 49–112; Uzzell, 1982, Amphibia-Reptilia, 3: 135–143; and Berger, 1983, Experientia, 39: 127–130, discussed the "Rana esculenta" complex. Günther, 1998, Zool. Abh. Staatl. Mus. Tierkd. Dresden, 50 (Suppl.): 99–108, discussed the species taxonomy of western Palearctic water frogs (Pelophylax ridibundus complex; as Rana in the original). Günther, 1991, Mitt. Zool. Mus. Berlin, 67: 39–53, summarized the distribution, taxonomy, and evolutionary history of the "Rana esculenta complex". Plötner and Ohst, 2001, Mitt. Mus. Naturkd. Berlin, Zool., 77: 5–21, discussed the relationships of the western Palearctic water frogs (Pelophylax saharicus, Pelophylax perezi, Pelophylax lessonae, Pelophylax bergeri, Pelophylax shqipericus, Pelophylax ridibundus, Pelophylax bedriagae, Pelophylax cretensis). Pelophylax was considered a synonym of Rana by Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 7, a subgenus of Rana by Dubois, 1992, Bull. Mens. Soc. Linn. Lyon, 61: 332, and a genus by Fei, Ye, and Huang, 1990, Key to Chinese Amph.: 133–134. Fei, Ye, Huang, Jiang, and Xie, 2005, in Fei et al. (eds.), Illust. Key Chinese Amph.: 109–113, provided a key to the Chinese species. Pelophylax is well-known for its hybridigenetic systems, two of which have been named either as species (e.g., Rana esculenta Linnaeus) or more recently as kleptons (Dubois and Günther, 1982, Zool. Jahrb., Jena, Abt. Syst., 109: 290–305; e.g., Rana kl. esculenta). Rather than treat these hemiclonal classes of perpetual F1 hybrids as lineages, the associated Linnean names are inserted in the synonymy of each parental taxon. So far, the hybridigenetic hybrids that have been named in a Linnaean sense are: Rana kl. esculenta Linnaeus, 1758, and Rana kl. grafi Crochet, Dubois, Ohler, and Tunner, 1995. While this method of naming these peculiar systems of hybridization is useful, hybridogenetic kleptons are not taxa, but classes of hybrids formed through one kind of interaction of taxa; see Bogart, 2003, in Sever (ed.), Reprod. Biol. Phylogeny Urodela: 109–134, for discussion. I therefore place the names of these kleptons in the synonymys of both hybridizing species, but endorse this naming convention as a way of pointing to "kinds" of frogs. Note, however, that Christiansen and Reyer, 2009, Evolution, 63: 1754–1768, have documented lineage formation in at least some populations of Pelophylax kl. esculentus. Lymberakis, Poulakakis, Manthalou, Tsigenopoulos, Magoulas, and Mylonas, 2007, Mol. Phylogenet. Evol., 44: 115–125, reported on the phylogenetics of the eastern Mediterranean species. On the basis of results provided by 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: 1–370, Che, Pang, Zhao, Wu, Zhao, and Zhang, 2007, Mol. Phylogenet. Evol., 43: 1–13, and Cai, Che, Pang, Zhao, and Zhang, 2007, Zootaxa, 1531: 49–55, Pelophylax cannot be regarded as a subgenus or synonym of Rana without also placing a large number of nominal genera within Rana as well in order to avoid paraphyly of Rana. These include, but are not limited to: Huia, Meristogenys, Amolops, Clinotarsus, Lithobates, Odorrana, Babina, and Hylarana. Wiens, Sukumaran, Pyron, and Brown, 2009, Evolution, 63: 1217–1231, suggested that Pelophylax is the sister taxon of a group composed of Rana, Lithobates, Hylarana, Glandirana, Odorrana, and Sanguirana, although they did not adopt this taxonomy. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, in their study of Genbank sequences confirmed this arrangement, but continued the application of a nonmonophyletic taxonomy.Fei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 3: 1042–1088, provided keys and accounts for the species of China. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, in their study of Genbank sequences confirmed the monophyly of this taxon (although this is obscured by their adoption of an antiquated and nonmonophyletic taxonomy) and provided a tree of exemplars. Hofman, Pabijan, Osikowski, Litvinchuk, and Szymura, 2016, MtDNA, Part A, 27: 3434–3437, reported on complete mitogenomes for Pelophylax cypriensis, Pelophylax epeiroticus, Pelophylax kurtmuelleri, and Pelophylax shqipericus and provided MrBayesian and maximum-likelihood trees for 10 species of Pelophylax. Hoffmann, Plötner, Pruvost, Christiansen, Röthlisberger, Choleva, Mikulíček, Cogǎlniceanu, Sas-Kovács, Shabanov, Morozov-Leonov, and Reyer, 2015, Mol. Ecol., 24: 4371–4391, reported on genetic diversity and distribution patterns of diploid and polyploid hybrid members of the Pelophylax esculentus complex. Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 384–385, provided brief accounts, photographs, and range maps for the African species. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 98–99, for a discussion of the difficulty of identifying species in this taxon. Cuevas, Patrelle, Ciavatti, Gendre, Sourrouille, Géniez, Doniol-Valcroze, and Crochet, 2022, Salamandra, 58: 218–230, described a molecular technique that allows a comparatively rapid identification of members of the Pelophylax perezi–Pelophylax grafi hybridogenetic system and the Pelophylax lessonae–Pelophylax esculentus hybridogenetic system. Ualiyeva, Ermakov, Litvinchuk, Guo, Ivanov, Xu, Li, Xu, Arifulova, Kaptyonkina, Khromov, Krainyuk, Sarzhanov, and Dujsebayeva, 2022, Diversity, 14 (869): 1–17, reported on mtDNA phylogeography of the Pelophylax ridibundus complex in Kasakhstan and Northwest China.
Contained taxa (13 sp.):
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