Bufonidae Gray, 1825
Bufonina Gray, 1825, Ann. Philos., London, Ser. 2, 10: 214. Type genus: Bufo Laurenti, 1768.
Bufonoidea — Fitzinger, 1826, Neue Class. Rept.: 37 (explicit family); Schinz, 1833, Naturgesch. Abbild Rept.: 228; Wied-Neuwied, 1865, Nova Acta Phys. Med. Acad. Caesar Leopold Carol., Halle, 32: 121; Laurent, 1967, Acta Zool. Lilloana, 22: 207; Lynch, 1973, in Vial (ed.), Evol. Biol. Anurans: 162; Duellman, 1975, Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 42: 5.
Bufonidea — Fitzinger, 1827, Isis von Oken, 20: 264.
Bufonini — Bonaparte, 1835, Iconograph. Fauna Ital., 2 (Fasc. 14): unnumbered (although "124" in left hand lower corner).
Bufonidae — Bell, 1839, Hist. Brit. Rept.: 105.
Bufoidae — Swainson, 1839, Nat. Hist. Fishes Amph. Rept., 2: 88.
Bufoniformes — Duméril and Bibron, 1841, Erp. Gen., 8: plate opposite page 53, 640. Explicit family-group name.
Atelopoda Fitzinger, 1843, Syst. Rept.: 32. Type genus: Atelopus Duméril and Bibron, 1841. Synonymy by Duellman and Lynch, 1969, Herpetologica, 25: 239; McDiarmid, 1971, Sci. Bull. Nat. Hist. Mus. Los Angeles Co., 12: 52; and Trueb, 1971, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co., 216: 1-40.
Bufonia — Gravenhorst, 1845, Das Thierreich: 43.
Adenomidae Cope, 1861 "1860", Proc. Acad. Nat. Sci. Philadelphia, 12: 371. Type genus: Adenomus Cope, 1861. Synonymy ; by implication of synonymy of Adenomus with Bufo) by Cope, 1862, Proc. Acad. Nat. Sci. Philadelphia, 14: 358; Günther, 1864, Rept. Brit. India: 421; and Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 281.
Bufonides — Bruch, 1862, Würzb. Naturwiss. Z., 3: 221.
Bufonida — Haeckel, 1866, Gen. Morphol. Organ., 2: cxxxii.
Phryniscina — Mivart, 1869, Proc. Zool. Soc. London, 1869: 288. Explicit subfamily.
Dendrophryniscina Jiménez de la Espada, 1870, J. Sci. Math. Phys. Nat., Lisboa, 3: 65. Type genus: Dendrophryniscus Jiménez de la Espada, 1870. Treatment as a subfamily of Bufonidae by Gadow, 1901, Amphibia and Reptiles: 139; subsequently ignored.
Dendrophryniscidae — Jiménez de la Espada, 1870, J. Sci. Math. Phys. Nat., Lisboa, 3: 65.
Bufonida — Knauer, 1878, Naturgesch. Lurche: 109; Bayer, 1885 "1884", Abh. K. Böhm. Ges. Wiss., Prague, 12: 18.
Bufoniidae — Boulenger, 1893, Zool. Rec., 29: 39.
Bufonidi — Acloque, 1900, Fauna de France, 1: 489.
Dendrophryniscinae — Gadow, 1901, Amphibia and Reptiles: 139.
Bufoninae — Fejérváry, 1917, Földtani Közlöny, Budapest, 47: 148.
Bufavidae Fejérváry, 1920, Ann. Hist. Nat. Mus. Natl. Hungarici, 18: 30. Type genus: Bufavus Portis, 1885. Fossil taxon. Synonymy by implication of Sanchíz, 1998, Handb. Palaeoherpetol., 4: 125.
Bufonidae — Miranda-Ribeiro, 1926, Arq. Mus. Nac., Rio de Janeiro, 27: 19.
Tornierobatidae Miranda-Ribeiro, 1926, Arq. Mus. Nac., Rio de Janeiro, 27: 19. Type genus: Tornierobates Miranda-Ribeiro, 1926. Synonymy by Noble, 1926, Am. Mus. Novit., 212: 15.
Dendrophryniscidae — Miranda-Ribeiro, 1926, Arq. Mus. Nac., Rio de Janeiro, 27: 19.
Atelopodidae — Parker, 1934, Monogr. Frogs Fam. Microhylidae: 8; Davis, 1935, Field Mus. Nat. Hist. Publ., Zool. Ser., 20: 91; Gallardo, 1961, 1° Reunion Trab. Comun. Cienc. Nat. Geograf., Univ. Nac. Litoral, Santa Fe, Argentina: 205.
Atelopodinae — Davis, 1935, Field Mus. Nat. Hist. Publ., Zool. Ser., 20: 91.
Nectophrynoidini — Dubois, 1982, Monit. Zool. Ital., N.S., Suppl., 16: 50. Type genus: Nectophrynoides Noble, 1926.
Tornierobatinae — Dubois, 1983, Bull. Mens. Soc. Linn. Lyon, 52: 273.
Adenominae — Dubois, 1983, Bull. Mens. Soc. Linn. Lyon, 52: 273.
Tornierobatini — Dubois, 1987 "1986", Alytes, 5: 25.
Nectophrynini — Dubois, 1987 "1986", Alytes, 5: 27.
Stephopaedini Dubois, 1987 "1986", Alytes, 5: 27. Type genus: Stephopaedes Channing, 1978.
Nomina inquirenda - Name(s) unassigned to a living or extinct population
Rana marginata Linnaeus, 1758, Syst. Nat., Ed. 10, 1: 212. Types: Not designated; although Laurenti, 1768, Spec. Med. Exhib. Synops. Rept.: 30, notes specimens in "Museo regio suecico", presumably available to Linnaeus. Daudin, 1802 "An. XI", Hist. Nat. Rain. Gren. Crap., Quarto: 54, regarded this taxon to be synonymous with his Rana typhonia (= Leptodactylus fuscus). (See Daudin, 1802 "An. XI", Hist. Nat. Rain. Gren. Crap., Quarto for access to NHRM type.) Type locality: "Indiis". Andersson considered one specimen from the Linnaean collection to be Linnaeus' Rana marginata, and synonymous with Bufo molitor Tschudi and Bufo andianus Cope. However, this name has not been used subsequently and sufficient ambiguities exist in Andersson's discussion to render this name a nomen dubium. Lönnberg, 1896, Bih. K. Svenska Vetensk. Akad. Handl., 22: 35, considered the taxon unidentifiable. * Bufo marginatus—Andersson, 1900, Bih. K. Svenska Vetensk. Akad. Handl., 26: 25.
Bufo siamensis Schneider, 1799, Hist. Amph. Nat.: 231. Type(s): By indication frog(s) mentioned in "Turpin Histoire civile et naturelle de Siam I page 256". Type locality: "Siam" (= Thailand).
Bufo arboreus Schneider, 1799, Hist. Amph. Nat.: 231. Type(s): By indication including frog(s) mentioned in "Britanni Itinerarium exceptum in Promtuario physico Lichtenbergii Vol. III. P. III. page 77." Type locality: "Americae eiusdem septentrionalis".
Bufo panamensis Daudin in Sonnini de Manoncourt and Latreille, 1801 "An. X", Hist. Nat. Rept., 2: 129. Type(s): MNHNP 742, according to Dubois and Bour, 2010, Zootaxa, 2447: 15. Type locality: "isthme de Panama".
Palaeophrynos Tschudi, 1838, Classif. Batr.: 52. Type species: Palaeophrynos gessneri Tschudi, 1838. Fossil taxon. Synonymy with Bufo (sensu lato) by Sanchíz, 1998, Handb. Palaeoherpetol., 4: 12, and Dubois and Bour, 2010, Zootaxa, 2447: 24. * Palaeophryne—Fitzinger, 1843, Syst. Rept.: 32. Incorrect subsequent spelling of Palaeophrynos Tschudi, 1838.
Toads (Fowler, 1907, Annu. Rep. N.J. State Mus. for 1906: 85; Conant, Cagle, Goin, Lowe, Neill, Netting, Schmidt, Shaw, Stebbins, and Bogert, 1956, Copeia, 1956: 176; Channing, 2001, Amph. Cent. S. Afr.: v).
True Toads (Ananjeva, Borkin, Darevsky, and Orlov, 1988, Dict. Amph. Rept. Five Languages: 35; Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 38; Anders, 2002, in Schleich and Kästle (eds.), Amph. Rept. Nepal: 144).
Cosmopolitan except for the Australian, Madagascan, and Oceanic regions, although introduced widely.
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, reviewed systematic literature, discussed phylogeny, and provided a taxonomy of bufonids, although because these authors partitioned Bufo (sensu lato), much of the literature of that nominal genus refers to taxa other than Bufo (sensu stricto). McDiarmid, 1971, Sci. Bull. Nat. Hist. Mus. Los Angeles Co., 12: 1–66, discussed the relationships of the genera Atelopus, Dendrophryniscus, Melanophryniscus, and Oreophrynella and reformulated the family to include much of the content of former Atelopodidae. Ruiz-Carranza and Hernández-Camacho, 1976, Caldasia, 11: 93–148, discussed the relationships of the genera Atelopus, Crepidophryne, Dendrophryniscus, Melanophryniscus, Oreophrynella, Osornophryne, and Rhamphophryne. Cruz and Peixoto, 1982, Rev. Brasil. Biol., 42: 627–629, discussed the relationships between Atelopus pernambucensis (now Frostius), Atelopus, and Dendrophryniscus. Pauly, Hillis, and Cannatella, 2004, Evolution, 58: 2517–2535, discussed relationships among New World bufonids. Cannatella, 1986, Herpetologica, 42: 197–205, discussed the relationships of Frostius, Atelopus, Osornophryne, Oreophrynella, Melanophryniscus, and Dendrophryniscus. Dubois, 1987 "1986", Alytes, 5: 25, provisionally recognized five subfamilies: Bufoninae (for Bufo), Atelopodinae (for Atelopus, Crepidophryne, Dendrophryniscus, Melanophryniscus, Oreophrynella, Osornophryne, Peltophryne, and Rhamphophryne), Tornieriobatinae (containing three tribes—Nectophrynini [for Laurentophryne, Nectophryne, Werneria, and Wolterstorffina], Stephopaedini [for Stephopaedes, Mertensophryne, and Schismaderma], and Tornieriobatini [for Capensibufo, Didynamipus, and Nectophrynoides (sensu lato)]), Adenominae (for Ansonia, Bufoides, Leptophryne, Pedostibes, Pelophryne, and Pseudobufo [and without explaining where Adenomus was placed, then in the synonymy of Bufo]), and Allophryninae (now in Centrolenidae). These taxa were recognized on the basis of previously published evidence (e.g., Grandison, 1978, Monit. Zool. Ital., N.S., Suppl., 11: 119–172; Grandison, 1981, Monit. Zool. Ital., N.S., Suppl., 15: 187–215; Wake, 1980, Copeia, 1980: 193–209), although some proposed groups (e.g., Dubois' Atelopodinae) are clearly polyphyletic on the basis of literature published at the time. Accounts and keys to most of the genera (prior to the partition of Bufo) are supplied by Laurent, 1986, in Grassé and Delsol (eds.), Traite de Zool., 14: 594–797. Graybeal and Cannatella, 1995, Herpetologica, 51: 105–133, discussed the evidence for the monophyly of various genera. Graybeal, 1997, Zool. J. Linn. Soc., 119: 297–338, discussed a phylogeny of bufonids and did not recognize Dubois' subfamilies. Unfortunately, Graybeal presented only her molecular evidence and did not disclose the relevant morphological evidence. Channing, 2001, Amph. Cent. S. Afr.: 56–61, provided a key to the species of southern Africa. Manthey and Grossmann, 1997, Amph. Rept. Südostasiens: 25–44, provided accounts and keys to the species of the Sunda Shelf. Cei, 1980, Monit. Zool. Ital., N.S., Monogr., 2: 157–213, discussed the species found in Argentina. Savage, 2002, Amph. Rept. Costa Rica: 185–212, provide accounts and keys to the species of Costa Rica. Liu and Hu, 1961, Tailless Amph. China: 113–126, discussed the Chinese species. Anders, 2002, in Schleich and Kästle (eds.), Amph. Rept. Nepal: 145–162, provided a key and accounts for the species of Nepal. Stöck, Günther, and Böhme, 2001, Zool. Abh. Staatl. Mus. Tierkd. Dresden, 51: 253–319, reviewed the nomenclature and taxonomy of Asian members of the Bufo viridis group (now Bufotes). Clarke, 2001, Afr. J. Herpetol., 50: 19–30, provided a critique of the systematics of African bufonids with discussion of the evidentiary basis of alternative viewpoints. Smith and Chiszar, 2006, Herpetol. Conserv. Biol., 1: 6–8, discussed the taxonomy of former "Bufo" insofar as it affects North American species. Because the implied combinations from that paper were unclear and provisional in text, the intention of the authors was verified by email correspondence (DRF); these authors intended world-wide application of their view, a view that if applied would require that most, if not all bufonid genera other than Melanophryniscus, Osornophryne, Atelopus, Truebella, and Dendrophryniscus, would be treated as subgenera of a hugely enlarged Bufo. Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept., 1(Amph.): 287, are the most recent authors to include the North American genera (Anaxyrus, Incilius, and Rhinella, as well as the extralimital Amietophrynus, Bufo, Bufotes, Capensibufo, Peltophryne, Stephopaedes, Torrentophryne, and Vandijkophrynus) as subgenera of Bufo in an attempt to maintain the North American status quo; like other authors who have suggested this, they ignore the fact that this synonymy requires under our current understanding of phylogeny that all Old World genera also be synonymized with Bufo, something that is unlikely to happen in any non-colonial world. Of course, Fouquette and Dubois (2014) may alternatively be comfortable with a paraphyletic Bufo, something certainly consistent with Dubois' publication record, but this would hardly represent a step forward. Matsui, Yambun Imbun, and Sudin, 2007, Zool. Sci., Tokyo, 24: 1159–1166, provided additional evidence for the polyphyly of "Bufo", with Ingerophrynus being in a clade with Sabahphrynus, Leptophryne, and Didynamipus; Duttaphrynus being likely closely related to Pelophryne, and Phrynoidis being the sister taxon of Pedostibes. Pramuk, Robertson, Sites, and Noonan, 2008, Global Ecol. Biogeograph., 17: 72–83, provided a phylogenetic hypothesis of Bufonidae and applied a monophyletic taxonomy. Díaz and Cádiz, 2008, Guía Taxon. Anf. Cuba: 1–294, provided accounts for the bufonids of Cuba. Fei, Hu, Ye, and Huang, 2009, Fauna Sinica, Amph. 2: 486–590, provided a key and accounts for the Chinese species. Van Bocxlaer, Biju, Loader, and Bossuyt, 2009, BMC Evol. Biol., 9 (e131): 1–10, provided an analysis of bufonid diversity, focused primarily on Asian diversity, that suggested that the New World component of diversity is paraphyletic with respect to the Old World component. This same tree is provided by Van Bocxlaer, Loader, Roelants, Biju, Menegon, and Bossuyt, 2010, Science, 327: 679–682. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, largely confirmed the results of previous authors, although this is obscured by their explicit adoption of an out-dated and non-monophyletic taxonomy. See generic accounts for more information. Powell, Collins, and Hooper, 2011, Key Herpetofauna U.S. & Canada, 2nd Ed.: 38–42, provided a key to the genera and species of North America north of Mexico. Martín, Alonso-Zarazaga, and Sanchíz, 2012, Graellsia, 68: 159–180, discussed the allocation of fossils previously associated with the name Bufo, but referrable to other currently-recognized genera. Cole, Townsend, Reynolds, MacCulloch, and Lathrop, 2013, Proc. Biol. Soc. Washington, 125: 317–578, provided identification keys and accounts for the species in Guyana. Fei, Ye, and Jiang, 2012, Colored Atlas Chinese Amph. Distr.: 246–274, provided accounts, photographs, and range maps for Chinese taxa. Oliver-López, Woolrich-Piña, and Lemos-Espinal, 2009, Fam. Bufonidae Mex., reviewed the species of Mexico. Vitt and Caldwell, 2013, Herpetology, 4th Ed., provided a summary of life history, diagnosis, and taxonomy. Brandvain, Pauly, May, and Turelli, 2014, Genetics, 197: 743–747, constrained the paraphyletic "Bufo" by excluding all genera that render it paraphyletic, then reported on relative branch lengths of within the tree and tested whether these were correlated with F1 viability. Portik and Papenfuss, 2015, BMC Evol. Biol., 15 (152): 1–19, provided a molecular phylogeny of bufonids that largely corroborated the current generic taxonomy, but suggested that Pedostibes as currently understood is nonmonophyletic and by excluding Incilius bocourti avoided a strong test of its monophyly.
Contained taxa (588 sp.):
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