Bufo spinosus Daudin, 1803

Class: Amphibia > Order: Anura > Family: Bufonidae > Genus: Bufo > Species: Bufo spinosus

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 spinosusRafinesque, 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 sèches 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).

Bufo vulgaris var. spinosus  Schreiber, 1912, Herpetol. Eur., Ed. 2: 212.

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 spinosusMertens, 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 spinosusVan 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 spinosusDubois and Bour, 2010, Zootaxa, 2447: 24.

Common Names

Giant Toad (Schneider and Sinsch, 2004, Z. Feldherpetol., 11: 187 [as Bufo bufo spinosus]).

Spiny Toad (Speybroeck, Beukema, Bok, and Van Der Voort, 2016, Field Guide Amph. Rept. Brit. Eur.: 149). 

Distribution

Iberian Peninsula and southwestern France (north to about Caen and about Lyon; an isolated population on the Isle of Jersey (United Kingdom) into extreme northwestern Italy (Liguria); northwestern Africa in the northern mountains of Morocco, Algeria, and Tunisia; expected in Monaco.

Geographic Occurrence

Natural Resident: Algeria, Andorra, France, Gibralter, Italy, Jersey, Morocco, Portugal, Spain, Tunisia

Likely/Controversially Present: Monaco

Comment

Recuero, Canestrelli, Vörös, Szabó, Poyarkov, Arntzen, Crnobrnja-Isailovic, Kidov, Cogǎlniceanu, Caputo, Nascetti, and Martínez-Solano, 2012, Mol. Phylogenet. Evol., 62: 71-86, considered Bufo spinosus a species distinct from Bufo bufo, redelimited its range, noted that African populations are notably distinct from European Bufo spinosus, 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., 6: 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. Amat Orriols and Roig Fernàndez, 2003, Butll. Soc. Catalana Herpetol., 16:42–72, discussed and mapped the species (as Bufo bufo) in Andorra. 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). Garcia-Porta, Litvinchuk, Crochet, Romano, Géniez, 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 bufoAmor, Kalboussi, and Said, 2013, Basic & Appl. Herpetol., 27: 85–100, discussed the range in Tunisia and its conservation status. Reques, Pleguezuelos, Busack, and de Pous, 2013, Basic & Appl. Herpetol., 27: 23–50, discussed the range and conservation status in Morocco. Mateo, Géniez, and Pether, 2013, Basic & Appl. Herpetol., 27: 51–83, provided a range map, comments on the range and population status in Algeria. Beukema, de Pous, Donaire-Barroso, Bogaerts, Garcia-Porta, Escoriza, Arribas, El Mouden, and Carranza, 2013, Zootaxa, 3661: 41–42, provided an account for Morocco (as Bufo bufo ssp.) and without discussion discarded the evidence of Bufo spinosus being diagnosable from Bufo bufoArntzen, Recuero, Canestrelli, and Martínez-Solano, 2013, Mol. Phylogenet. Evol., 69: 1203–1208, discussed the phylogenetics and species taxonomy of this and near relatives and concluded that Bufo spinosus is a distinct species. Arntzen, McAtear, Ziermann, Ohler, van Alphen, and Martínez-Solano, 2013, Contrib. Zool., Amsterdam, 82: 147–169, reported on the contact zone of Bufo bufo and Bufo spinosus. Arntzen, Wilkinson, Butôt, and Martínez-Solano, 2014, Herpetol. J., 24: 209–216, reported on the population on Jersey, United Kingdom. Mediani, Brito, and Fahd, 2015, Basic & Appl. Herpetol., 29: 81–107, provided a dot map for northern Morocco. Nöllert, Grosse, Buschendorf, and Geiger, 2016, Mertensiella, 24: 7–32, summarized and discussed the taxonomic history of this species and its recognition within the Bufo bufo group and provided a range map. Trujillo, Gutiérrez-Rodríguez, Arntzen, and Martínez-Solano, 2017, Contrib. Zool., Amsterdam, 86: 1–10, reported on a hybridizing population in northwestern France with Bufo bufo with no evidence of restriction to gene flow. Arntzen, McAtear, Butôt, and Martínez-Solano, 2018, Amphibia-Reptilia, 39: 41–50, reported on the hybrid zone between Bufo bufo and Bufo spinosus from the Atlant coast near Caen, France, to the Mediterranean coast near Savona, Italy. Speybroeck, Beukema, Bok, and Van Der Voort, 2016, Field Guide Amph. Rept. Brit. Eur.: 149–151, provided a brief account and range map. Skorinov, Bolshakova, Donaire, Pasynkova, and Litvinchuk, 2018, Russ. J. Herpetol., 25: 253–258, reported on the karyotype.  See detailed account for North Africa by Escoriza and Ben Hassine, 2019, Amph. N. Afr.: 150–159. Channing and Rödel, 2019, Field Guide Frogs & Other Amph. Afr.: 56–57, provided a brief account, photograph, and range map for the African component of the range. See Dufresnes, 2019, Amph. Eur., N. Afr., & Middle East: 73, for brief summary of identifying morphology and biology, a range map, as well as a photograph. Sanna, 2019, Acta Herpetol., Firenze, 14: 117–122, reported on ontogenetic and interspecific variation in skull morphology of this species and Bufo bufo. Mabrouki, Taybi, Skalli, and Sánchez-Vialas, 2019, Basic & Appl. Herpetol., 33: 19–32, commented on and mapped the species in the Mouloua River Basin and adjacent areas in eastern Morocco. Arntzen, Canestrelli, and Martínez-Solano, 2020, Contrib. Zool., Amsterdam, 89: 270–281, reported on the hybrid zone between this species and Bufo bufo, found from the English Channel in central Normandy southwest to the Meditterannean in Liguria in Italy, examining environmental correlates with the location and degree of hybridization and placement of pure populations. Ivanović, Cvijanovíc, Vučić, and Arntzen, 2023, Organisms Divers. Evol., 23: 209–219, reported on skull morphology and cranial kinesis of Bufo bufo and  Bufo spinosus.

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