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Ceratophryidae Tschudi, 1838
Ceratophrydes Tschudi, 1838, Classif. Batr.: 26. Type genus: Ceratophrys Wied-Neuwied, 1824.
Ceratophreidina — Bonaparte, 1850, Conspect. Syst. Herpetol. Amph.: 1 p.
Ceratophreina — Bonaparte, 1850, Conspect. Syst. Herpetol. Amph.: 1 p.
Ceratophrydidae — Cope, 1863, Proc. Acad. Nat. Sci. Philadelphia, 15: 50.
Batrachophryidae Cope, 1875, Bull. U.S. Natl. Mus., 1: 9. Type genus: Batrachophrynus Peters, 1873. Synonymy by implication of the results of Córdova and Descailleaux, 2005, in Lavilla and De la Riva (eds.), Monogr. Herpetol., 7: 187-217.
Telmatobiidae — Miranda-Ribeiro, 1920, Rev. Mus. Paulista, São Paulo, 12: 320.
Ceratophriidae — Waite, 1927, Trans. R. Soc. S. Aust., 51: 328.
Ceratophryinae — Parker, 1933, Trop. Agric., Trinidad, 10: 11.
Ceratophryidae — Parker, 1933, Trop. Agric., Trinidad, 10: 12; 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: 127.
Ceratophrydae — Lutz, 1954, Mem. Oswaldo Cruz, Rio de Janeiro, 52: 156.
Ceratophryninae — Reig, 1960, Acta Trab. Primer Congr. Sudam. Zool., 4: 117.
Ceratophrynidae — Reig and Limeses, 1963, Physis, Buenos Aires, 24: 125.
Stombinae Gallardo, 1965, Pap. Avulsos Zool., São Paulo, 17: 82. Type genus: Stombus Gravenhorst, 1825.
Ceratophrynae — Cei, 1970, Acta Zool. Lilloana, 27: 183.
Ceratophrynini — Laurent and Teran, 1981, Fund. Miguel Lillo, Misc., 71: 7.
Common Horned Frogs (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 71).
Lynch, 1971, Misc. Publ. Mus. Nat. Hist. Univ. Kansas, 53: 1-238, considered Ceratophryinae (in his restricted sense) to be the primitive subfamily from which Telmatobiinae (in his sense as including Eleutherodactylinae [now the bulk of Brachycephaloidea]) was derived (a view that J.D. Lynch in Frost, 1985, Amph. Species World: 235, subsequently rejected). Reig, 1972, in Blair (ed.), Evol. Genus Bufo: 14-36, and Estes and Reig, 1973, in Vial (ed.), Evol. Biol. Anurans: 11-63, considered Ceratophryinae to be a distinct family, Ceratophryidae, which gave rise directly to Bufonidae. Also considered a distinct family by Cei, 1980, Monit. Zool. Ital., N.S., Monogr., 2: 216. Laurent, 1986, in Grassé and Delsol (eds.), Traite de Zool., 14: 687-689, transferred Macrogenioglottus, Odontophrynus, and Proceratophrys (as the tribe Odontophrynini) into this subfamily, with Ceratophrys (including Chacophrys) and Lepidobatrachus being retained in Ceratophrynini. See Maxson and Ruibal, 1988, J. Herpetol., 22: 228-231, for discussion of intra-subfamilial relationships. Haas, 2003, Cladistics, 19: 23-89, suggested that Ceratophryini and Odontophryini might not form a monophyletic group, each being closer to other groups of frogs; the former being closer to hylids and the latter being closer to other leptodactylids and bufonids. 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: 126, showed that Ceratophryini and Odontophryini were not closest relatives, the content of Odontophrynini being transferred to Cycloramphidae. These authors also reformulated the content of Ceratophryidae on phylogenetic content and recognized two subfamilies, Ceratophryinae (Atelognathus, Batrachyla, Ceratophrys, Chacophrys, and Lepidobatrachus) and Telmatobiinae (Telmatobius). Grant, Frost, Caldwell, Gagliardo, Haddad, Kok, Means, Noonan, Schargel, and Wheeler, 2006, Bull. Am. Mus. Nat. Hist., 299: 126, reformulated the subfamilies, recognizing a third: Batrachylinae. Roelants, Gower, Wilkinson, Loader, Biju, Guillaume, Moriau, and Bossuyt, 2007, Proc. Natl. Acad. Sci. USA, 104: 887-892, on the basis of less inclusive sampling, a smaller molecular dataset, and a different analytical method from Grant et al. (2006) found Ceratophryidae to not be demonstrably monophyletic, with Telmatobiinae and Batrachylinae, forming a monophyletic group, and Ceratophryinae possibly being distant from that group. Bossuyt and Roelants, 2009, in Hedges and Kumar (eds.), Timetree of Life: 357-364, treated these units as Telmatobiidae (including Batrachylinae and Telmatobiinae) and Ceratophryidae. Heinicke, Duellman, Trueb, Means, MacCulloch, and Hedges, 2009, Zootaxa, 2211: 1-35, suggested on the basis of considerable molecular evidence, but narrower taxon sampling than Frost et al. (2006) that Ceratophryidae is not monophyletic, with Batrachylus (Batrachylinae, presumably) possibly being outside of an inclusive monophyletic group containing Telmatobiinae and Ceratophryinae. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543-583, on the basis of their study of Genbank sequences, regarded the three subfamilies as distinct families (Ceratophryidae, Batrachylidae, and Telmatobiidae) on the basis of the inferred lack of monophyly of Ceratophryidae (sensu lato), and arrangement followed here. Fabrezi, 2006, J. Zool. Syst. Evol. Res., 44: 153-166, reported on phylogenetics of this taxon. See comment under Batrachylidae. Fabrezi and Quinzio, 2008, Zool. J. Linn. Soc., 154: 752-780, reported on the morphological distinctiveness of this taxon. Blackburn and Wake, 2011, In Zhang (ed.), Zootaxa, 3148: 39-55, commented on the fossil members of the taxon and reviewed the nomenclatural history of the family-group. Cole, Townsend, Reynolds, MacCulloch, and Lathrop, 2013, Proc. Biol. Soc. Washington, 125: 317-578, provided identification keys and accounts for the species in Guyana. Vitt and Caldwell, 2013, Herpetology, 4th Ed., provided a summary of life history, diagnosis, and taxonomy.
Contained taxa (12 sp.):
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