Bufo spinosus Daudin, 1803
Bufo spinosus Daudin, 1803 (An. XI), Hist. Nat. Gen. Part. Rept., 8: 199. Types: MNHNP. Type locality: "en France dans les pays montagneux, . . . aux environs de Brives et de Bordeaux", southern France.
Batrachus spinosus — Rafinesque, 1814, Specchio Sci., 2, 2: 102.
Bufo vinearum Lesson, 1841, Act. Soc. Linn. Bordeaux, Ser. 3, 12: 61. Types: Not stated or known to exist. Type locality: "dans les vignes des terres hautes et sčches de l'arrondissement de Sant-Jean-d'Angély". Synonymy by Boulenger, 1898, Tailless Batr. Eur., 2: 214, with Bufo vulgaris. Placed here provisionally on the basis of geography.
Platosphus gervais de l'Isle, 1877, J. Zool., Paris, 6: 473. Holotype: MNHNP (uncatalogued), incomplete articulated specimen. Type locality: "Dufort", France. [Lower Pleistocene fossil.] Synonymy by Sanchíz, 1998, Handb. Palaeoherpetol., 4: 122.
Bufo rubeta var. robustior Lataste In Boscá, 1880, Bull. Soc. Zool. France, 5: 256. Holotype: Deposition of specimen not stated, but presumably in Lataste's personal collection which was deposited ultimately in the BMNH—DRF. Type locality: "environs de Ciudad-Real", Spain. Synonymy by implication of Bufo rubeta being a synonym of Bufo bufo and specifically by Gorham, 1974, Checklist World Amph.: 78, and on the basis of geography (DRF).
Pelobates Wilsoni Boscá, 1919, Bol. R. Soc. Esp. Hist. Nat., 19: 89. Holotype: MNCN, by original designation. Type locality: " Estación Alpina de de Cercedilla (Madrid), a mas de 1.500 metros sobre el nivel del mar . . . . vertiente segogiana [of the Sierra Guadarrama]". Synonymy with Bufo bufo by Boulenger, 1919, Bol. R. Soc. Esp. Hist. Nat., 19: 293-294.
Bufo bufo spinosus — Mertens, 1925, Abh. Senckenb. Naturforsch. Ges., 39: 46. Mertens, 1955, Senckenb. Biol., 36: 288; Pasteur and Bons, 1959, Trav. Inst. Scient. Chérifien, Rabat, Ser. Zool., 17: 141; Schneider and Sofianidou, 1985, Zool. Anz., 214: 209-221.
Bufo bufo gredosicola Müller and Hellmich, 1935, Zool. Anz., 112: 54. Holotype: ZSM (SLM 2049), lost according to Glaw and Franzen, 2006, Spixiana, München, 29: 159. Type locality: "Laguna Grande de Gredos, 2027 m", central Spain.
Bufo spinosus — Van Bocxlaer, Biju, Loader, and Bossuyt, 2009, BMC Evol. Biol., 9 (e131): 3.
Bufo (Bufo) bufo gredosicola — Dubois and Bour, 2010, Zootaxa, 2447: 24.
Bufo (Bufo) bufo spinosus — Dubois and Bour, 2010, Zootaxa, 2447: 24.
English Names
Giant Toad (Schneider and Sinsch, 2004, Z. Feldherpetol., 11: 187 [as Bufo bufo spinosus]).
Distribution
Iberian Peninsula and southwestern France; northwestern Africa in the northern mountains of Morocco, Algeria, and Tunisia.
Comment
Recuero, Canestrelli, Vörös, Szabó, Poyarkov, Arntzen, Crnobrnja-Isailovic, Kidov, Cogalniceanu, Caputo, Nascetti, and Martínez-Solano, 2012, Mol. Phylogenet. Evol., 62: 71-86, considered Bufo spinosus a species distinct from Bufo bufo, redelimited its range and diagnosis, and suggested on the basis of molecular evidence that it is the sister taxon of Bufo bufo + Bufo verrucosissimus. Prior to this study, the status of populations in southern Europe and the eastern Mediterranean had been the source of much confusion regarding the status of Bufo spinosus. Kutrup, Yilmaz, Canakci, Belduz, and Doglio, 2006, Amphibia-Reptilia, 27: 268-273, discussed geographic variation in 16S rRNA and rejected the Turkish populations of Bufo bufo spinosus as a distinct subspecies or species, suggesting instead that it should be placed in the synonymy of Bufo bufo verrucosissimus (thereby implying that they regarded Bufo verrucosissimus as part of Bufo bufo). Barbadillo-Escrivá, 1987, Guia INCAFO Anf. Rept. Peninsula Iberica: 150-154, provided an account for the Iberian populations as Bufo bufo. Llorente, Montori, Santos, and Carretero, 1995, Atlas Amf. Rept. Catalunya Andorra: 66-68, provided an account as Bufo bufo and detailed range map for Catalunya and Andorra. Schneider and Sinsch, 2004, Z. Feldherpetol., 11: 187-201, considered Bufo spinosus to be a species distinct from Bufo bufo, although the distribution of both taxa was uncertain and Nöllert and Nöllert, 1992, Die Amph. Eur.: 286-292, noted considerable hybridization with diagnostic Bufo bufo in Portugal, central and southern France, the southern Alps and the northern part of the Balkans. Previously Bufo spinosus had been considered either a synonym or subspecies of Bufo bufo by Bonaparte, 1835, Iconograph. Fauna Ital., 2 (Fasc. 14): unnumbered; Tschudi, 1838, Classif. Batr.: 88; Duméril and Bibron, 1841, Erp. Gen., 8: 672 (as Bufo vulgaris); Boulenger, 1881 "1880", Proc. Zool. Soc. London, 1880: 545-574; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 303. See Salvador, 1996, Smithson. Herpetol. Inform. Serv., 109, and Schneider, 1974, Salamandra, 10: 55-60, for accounts for the North African population (as Bufo bufo). Lizana, 1997, in Pleguezuelos (ed.), Dist. Biogeogr. Anf. Rep. Esp. Portugal: 152-154, provided a brief account, photograph, and detailed map for Iberia. Rivera, Escoriza, Maluquer-Margalef, Arribas, and Carranza, 2011, Amf. Rept. Catalunya: 75-77, provided a brief account for northeastern Spain and adjacent France, as Bufo bufo. Martínez-Solano and González, 2008, Biol. J. Linn. Soc., 95: 824-839, noted a relative deep genetic divergence between Iberian and North African populations of Bufo bufo (all of which are now considered Bufo spinosus). Recuero, Canestrelli, Vörös, Szabó, Poyarkov, Arntzen, Crnobrnja-Isailovic, Kidov, Cogalniceanu, Caputo, Nascetti, and Martínez-Solano, 2012, Mol. Phylogenet. Evol., 62: 71-86, on the basis of mt and nuDNA concludd that Bufo spinosus is the sister taxon of Bufo verrucosissimus, and that African populations are notably distinct from European Bufo spinosus. Garcia-Porta, Litvinchuk, Crochet, Romano, Geniez, Lo-Valvo, Lymberakis, and Carranza, 2012, Mol. Phylogenet. Evol., 63: 113-130, subsequently largely confirmed these results, including noting the deep divergences among the African populations and from the European populations, and regarded Bufo spinosus as a subspecies of Bufo bufo.
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