Bufotes Rafinesque, 1815

Class: Amphibia > Order: Anura > Family: Bufonidae > Genus: Bufotes
12 species

Bufo Laurenti, 1768, Spec. Med. Exhib. Synops. Rept.: 25. Type species: Bufo vulgaris Laurenti, 1768 (= Bufo bufo) by subsequent designation of Tschudi, 1838, Classif. Batr.: 50, although this was considered invalid by reason of ambiguity by Dubois, 1984, Mem. Mus. Natl. Hist. Nat. Paris, A—Zool., 131: 19, Dubois and Bour, 2010, Zootaxa, 2447: 12–13, and Welter-Schultes and Klug, 2011, Zootaxa, 2814: 55; designated as Bufo viridis Laurenti, 1768, by subsequent designation of Fitzinger, 1843, Syst. Rept.: 32. See discussions by Dubois and Bour, 2010, Zootaxa, 2447: 12–13, and, and 22, and Dubois and Bour, 2012, Zootaxa, 3221: 37–47, who considered Bufo Garsault, 1764, and Bufo Laurenti, 1768, to represent independent nomenclatural acts and Welter-Schultes and Klug, 2011, Zootaxa, 2814: 55, who did not. ICZN action is required to resolve this issue.

BuffoLacépède, 1788, Hist. Nat. Quadrup. Ovip. Serpens, 16mo ed., 2: 329, 460; Lacépède, 1788, Hist. Nat. Quadrup. Ovip. Serpens, Quarto ed., 1: Table following page 618. Incorrect subsequent spelling of Bufo Laurenti, 1768. But see comments by Dubois and Bour, 2010, Zootaxa, 2447: 22.

Batrachus Rafinesque, 1814, Specchio Sci., 2, 2: 103. Substitute name for Bufo Laurenti, 1768. Junior homonym of Batrachus Schaeffer 1760 (a fish).

Bufotes Rafinesque, 1815, Analyse Nat.: 78. Substitute name for "Bufo Daud." (= Bufo Laurenti, 1768). See comment under Bufonidae.

Pseudepidalea 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: 365. Type species: Bufo viridis Laurenti, 1768, by original designation. Synonymy by Dubois and Bour, 2010, Zootaxa, 2447: 22.

Calliopersa Safaei-Mahroo and Ghaffari, 2020, Compl. Guide Amph. Iran: 16. Type species: Bufo surdus Boulenger, 1891. Zoobank publication registration: 90E7552C-444D-4BFE-B0DC-FFA695A4AA55. See comment. 

English Names

Middle Asiatic Toads (Terbish, Munkhbayar, and Munkhbaatar, 2013, Guide Amph. Rept. Mongolia: 16). 

Green Toads (BufotesSafaei-Mahroo, Ghaffari, and Niamir, 2023, Zootaxa, 5279: 13).

Persian Melodious Toads (CalliopersaSafaei-Mahroo, Ghaffari, and Niamir, 2023, Zootaxa, 5279: 17)..

Distribution

Korea, northern and far western China, Baikal region of Russia and extreme western Mongolia, and Korea east through Kyrgyzstan and Afghanistan, Kashmir (India and Pakistan), to Iran, Iraq, and western Europe, the Levant, and Mediterranean North Africa.

Comment

Bufotes is the Bufo viridis group of previous authors (e.g., Martin, 1972, in Blair (ed.), Evol. Genus Bufo: 63; Inger, 1972, in Blair (ed.), Evol. Genus Bufo: 107, excluding Epidalea calamita). The literature of these species is so intertwined with that of Bufo, that the Bufonidae record should be consulted for access to relevant regional literature. 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: 365, named Pseudepidalea on the basis of its presumed phylogenetic distance from Epidalea (Epidalea calamita), although the monophyly of Epidalea + Pseudepidalea has been widely assumed by previous authors, it has not been adequately tested. Borkin, Rosanov, and Litvinchuk, 2001 "2000", Russ. J. Herpetol., 7: 171–180, discussed geographic distribution of the various diploid and triploid populations and provide a new classification, now largely superseded. Batista, Carranza, Carretero, and Harris, 2006, Butll. Soc. Catalana Herpetol., 17: 24–33, noted three mtDNA lineages in nominal Bufo viridis (now Bufotes species complex), one in North Africa and Sicilty, another in the rest of Europe, and one in Sardinia and Mallorca. Stöck, Moritz, Hickerson, Frynta, Dujsebayeva, Eremchenko, Macey, Papenfuss, and Wake, 2006, Mol. Phylogenet. Evol., 41: 663–689, reported on the phylogenetics and biogeography of the nominal Bufo viridis subgroup (= Pseudepidalea of Frost et al., 2006, but including Epidalea calamita). The taxonomic history is confused and confusing and names used in various faunal works are frequently misleading. Stöck, Günther, and Böhme, 2001, Zool. Abh. Staatl. Mus. Tierkd. Dresden, 51: 295–302, revised partly the nomenclature of the Asian taxa of this species including data on name-bearing types (providing also translations of non-English diagnoses and type descriptions), on type localities, nomenclatural and systematic histories, karyotypes, bioacoustics, distribution, proposed current taxonomic status, and a tentative identification key. This was later expanded upon and refined by Stöck, Moritz, Hickerson, Frynta, Dujsebayeva, Eremchenko, Macey, Papenfuss, and Wake, 2006, Mol. Phylogenet. Evol., 41: 663–689—the place to start to understand this group. Smith and Chiszar, 2006, Herpetol. Conserv. Biol., 1: 6–8, implied that this taxon should be considered a subgenus of Bufo; see comment under Bufonidae. Van Bocxlaer, Biju, Loader, and Bossuyt, 2009, BMC Evol. Biol., 9 (e131): 1–10, and Van Bocxlaer, Loader, Roelants, Biju, Menegon, and Bossuyt, 2010, Science, 327: 679–682, on the basis of molecular evidence suggested that Pseudepidelea is far from Epidalea, likely monophyletic, with the exception of "Pseudepidalea" brongersmai, which may be distantly related to this group. Speybroeck, Beukema, and Crochet, 2010, Zootaxa, 2492: 6–7, regarded Epidalea and Pseudepidalea to be junior synonyms of Bufo on the basis of plesiomorphic retention of hybridization ability. See comment under Bufo for discussion of nomenclatural issues surrounding the use of this name. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, confirmed the monophyly of this taxon (although this is difficult to see because the authors explicitly adopted a non-monophyletic and out-dated taxonomy), provided a tree for their exemplar species, and suggested that it is the sister taxon of Schismaderma. Guignard, Büchi, Gétaz, Betto-Colliard, and Stöck, 2012, Biol. J. Linn. Soc., 105: 584–590, discussed the effect of genome size on call properties of toads in this group. Dubois and Bour, 2010, Zootaxa, 2447: 1–52, regarded Bufotes Rafinesque, 1815, as the oldest available generic name for this group, but also placed it as a subgenus of Bufo Garsault, 1764, which is inconsistent with our current understanding of relationships within Bufonidae as suggested by Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, who inferred this taxon as being the sister taxon of Schismaderma. Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept.: 290, considered Bufotes as subgenus of Bufo, cherry-picking their citation to literature (excluding any reference to  Van Bocxlaer, Biju, Loader, and Bossuyt, 2009, BMC Evol. Biol., 9 (e131): 1–10, Van Bocxlaer, Loader, Roelants, Biju, Menegon, and Bossuyt, 2010, Science, 327: 679–682, or Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, which provided results not congruent with the story that Fouquette and Dubois wanted to tell) to avoid recognizing that treating this genus as a subgenus of Bufo also requires under current understanding of phylogeny all Old-World bufonids, such as Sabahphrynus, Nectophryne, and Ansonia to be treated as subgenera of Bufo as well.  Terbish, Munkhbayar, and Munkhbaatar, 2013, Guide Amph. Rept. Mongolia: 16–17, provided a brief account, photograph, and spot map for Mongolia. Fei and Ye, 2016, Amph. China, 1: 788–802, reviewed the species of ChinaDufresnes, Lymberakis, Kornilios, Savary, and Perrin, 2018, BMC Evol. Biol., 18 (67): 1–12, reported on phylogenetics, phylogeography, and hybrid zones between Bufotes variabilis, Bufotes siculus, Bufotes viridis, and Bufotes balearicus in the Aegean region. Betto-Colliard, Hofmann, Sermier, Perrin, and Stöck, 2018, Proc. R. Soc. London, Ser. B, Biol. Sci., 285 (20172667): 1–8, discussed genetic divergence and asymmetric parental genome contributions in hybrid speciation within the genus. Dufresnes, Mazepa, Jablonski, Oliveira, Wenseleers, Shabanov, Auer, Ernst, Koch, Ramírez-Chaves, Mulder, Simonovo, Tiutenko, Kryvokhyzhar, Wennekes, Zinenko, Korshunov, Al-Johany, Peregontsev, Masroor, Betto-Colliard, Denoël, Borkin, Skorinov, Pasynkova, Mazanaeva, Rosanov, Dubey, and Litvinchuk, 2019, Mol. Phylogenet. Evol., 141: 1–25, provided a substantial phylogeographic study and revision of the genus on the basis of molecular and phenotypic evidence. Recognition of Calliopersa (including both Bufotes luristanica and Bufotes surdus) renders Bufotes paraphyletic, at least if the tree of Dufresnes, Mazepa, Jablonski, Oliveira, Wenseleers, Shabanov, Auer, Ernst, Koch, Ramírez-Chaves, Mulder, Simonovo, Tiutenko, Kryvokhyzhar, Wennekes, Zinenko, Korshunov, Al-Johany, Peregontsev, Masroor, Betto-Colliard, Denoël, Borkin, Skorinov, Pasynkova, Mazanaeva, Rosanov, Dubey, and Litvinchuk, 2019, Mol. Phylogenet. Evol., 141: 12, is correct. Further studies may redefine this taxon to allow Bufotes monophyletic but until then I hesitate to recognize it (DRF). Che, Jiang, Yan, and Zhang, 2020, Amph. Rept. Tibet: 141, provided molecular tree that suggested that Bufotes is paraphyletic with respect to Epidalea (which is the older generic name) suggesting (to at least DRF) that we will likely see either Bufotes placed into the synonymy of Epidalea, or a three-genus  solution of Bufotes, Calliopersa, and Epidalea. The arrangement with Epidalea and Bufotes (including Calliopersa and Bufotes as subgenera) is inconsistent with the phylogeny as currently understood (DRF). 

Contained taxa (12 sp.):

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