Amphibian Species of the World 6.0, an Online Reference

Bufonidae Gray, 1825

English Names

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).

Distribution

Cosmopolitan except for the Australian, Madagascan, and Oceanic regions, although introduced widely.

Comment

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, 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.

Contained taxa (578 sp.):

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