Rhinella Fitzinger, 1826

Class: Amphibia > Order: Anura > Family: Bufonidae > Genus: Rhinella
93 species

Oxyrhynchus Spix, 1824, Animal. Nova Spec. Nov. Test. Ran. Brasil.: 49. Type species: not designated. Junior homonym of Oxyrhynchus Leach, 1818 (fish). Spix's original ranking of the name is unclear, either as a genus or a subgenus, although since the time of Cuvier, 1829, Regne Animal., Ed. 2, 2: 112 (p. 84 in the 1831 edition) the original usage has been considered to have been as a subgenus although (at least according to Cuvier) not distinguishable from Bufo. Synonymy with Bufo by Cuvier, 1829, Regne Animal., Ed. 2, 2: 112, and Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 281.

Rhinella Fitzinger, 1826, Neue Class. Rept.: 39. Type species: Bufo (Oxyrhynchus) proboscideus Spix, 1824, by monotypy. Synonymy with Bufo by Cuvier, 1829, Regne Animal., Ed. 2, 2: 112 (treatment as a subgenus of Bufo).

Chaunus Wagler, 1828, Isis von Oken, 21: 744. Type species: Chaunus marmoratus Wagler, 1828 (= Bufo granulosus Spix), by monotypy. Synonymy with Rhinella by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211.

Otilophes Cuvier, 1829, Regne Animal., Ed. 2, 2: 112. Type species: Rana margaritifera Laurenti, 1768, by original designation. Coined as a subgenus of Bufo as "Otilophes Cuv."; not formed in Latin (as Otilophis) until Cuvier, 1831, Animal Kingdom (M'Murtrie), 2: 84. Synonymy with Bufo by Duméril and Bibron, 1841, Erp. Gen., 8: 662; Boulenger, 1881 "1880", Proc. Zool. Soc. London, 1880: 545-574. Synonymy with Rhinella by implication of 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.

Otilophis — Cuvier, 1831, Animal Kingdom (M'Murtrie), 2: 84. Incorrect subsequent spelling. Accepted by Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept.: 316, as the original spelling. 

Phryniscus Wiegmann, 1834, in Meyen (ed.), Reise in die Erde K. Preuss. Seehandl., 3(Zool.): 514. Subsequently published by Wiegmann, 1834, Nova Acta Phys. Med. Acad. Caesar Leopold Carol., Halle, 17: 264. Type species: Phryniscus nigricans Wiegmann, 1834, by monotypy. Synonymy (with Atelopus) by Günther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 43. Synonymy with Bufo by Boulenger, 1894, Ann. Mag. Nat. Hist., Ser. 6, 14: 374 (not Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2, who retained Phyniscus as a synonym of Atelopus). The name Phryniscus was published twice: first in 1834 (Wiegmann, 1834, in Meyen (ed.), Reise in die Erde K. Preuss. Seehandl., 3(Zool.): 514), and then in early 1835 (Wiegmann, 1834, Nova Acta Phys. Med. Acad. Caesar Leopold Carol., Halle, 17: 185–268). For additional discussion see Lavilla, 1997, Cuad. Herpetol., 11: 75–80, Anonymous, 1910, Cat. Books Mss Maps Brit. Mus., Vol. 3, and Bauer and Adler, 2001, Arch. Nat. Hist., London, 28: 313–326. Synonymy with Chaunus by implication of 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. Synonymy with Rhinella by implication by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211.

OsilophusTschudi, 1838, Classif. Batr.: 52, 89. Incorrect subsequent spelling of Otilophes or Otilophis Cuvier.

Otolophus Fitzinger, 1843, Syst. Rept.: 32. Type species: Bufo margaritifer Daudin, 1802, by original designation. Synonymy with Bufo by Kellogg, 1932, Bull. U.S. Natl. Mus., 160: 28. Possibly an incorrect subsequent spelling of Otilophes or Otilophis Cuvier.. Synonymy with Rhinella by implication of 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.

Eurhina Fitzinger, 1843, Syst. Rept.: 32. Type species: Oxyrhynchus proboscideus Spix, 1824, by original designation. Synonymy with Bufo by implication of Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 315, who treated the type species as a member of Bufo. Synonymy with Rhinella by implication of 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.

Chilophryne Fitzinger, 1843, Syst. Rept.: 32. Type species: Bufo dorbignyi Duméril and Bibron, 1841, by original designation. Synonymy by Keferstein, 1867, Nachr. Ges. Wiss. Göttingen, 18: 352 (as a subgenus of Bufo); Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 281. Synonymy with Chaunus by implication of 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. Synonymy with Chaunus ; by implication by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211.

Docidophryne Fitzinger, 1843, Syst. Rept.: 32. Type species: Bufo agua Daudin, 1802, by original designation. Synonymy with Bufo (sensu lato) by Tschudi, 1845, Arch. Naturgesch., 11: 168. Synonymy (with Phrynoidis) by Cope, 1862, Proc. Acad. Nat. Sci. Philadelphia, 14: 358. Synonymy with Chaunus by implication of 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. Synonymy with Chaunus ; by implication by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211.

Trachycara Tschudi, 1845, Arch. Naturgesch., 11: 169. Type species: Trachycara fusca Tschudi, 1845, by monotypy. Synonymy with Bufo by Peters, 1873, Monatsber. Preuss. Akad. Wiss. Berlin, 1873: 624. Synonymy with Rhinella by implication of 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.

OtilophusGünther, 1859 "1858", Cat. Batr. Sal. Coll. Brit. Mus.: 69; Boulenger, 1882, Cat. Batr. Sal. Coll. Brit. Mus., Ed. 2: 281. Incorrect subsequent spelling of Otilophis Cuvier, 1829. Not Otolophus Fitzinger, 1843.

Aruncus Philippi, 1902, Supl. Batr. Chil. Descr. Hist. Fis. Polit. Chile: 4. Type species: Aruncus valdivianus Philippi, 1902, by monotypy. Considered a nomen dubium by Nieden, 1923, Das Tierreich, 46: 146. Tentative synonymy with Bufo (sensu lato) by Cei, 1958, Invest. Zool. Chilen., 4: 266; Gorham, 1974, Checklist World Amph.: 146. Synonymy with Chaunus by implication of 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. Synonymy with Rhinella (by implication) by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211.

Stenodactylus Philippi, 1902, Supl. Batr. Chil. Descr. Hist. Fis. Polit. Chile: 40. Type species: Stenodactylus ventralis Philippi, 1902, by monotypy. Considered a nomen dubium by Nieden, 1923, Das Tierreich, 46: 146. Synonymy by Philippi, 1902, Supl. Batr. Chil. Descr. Hist. Fis. Polit. Chile: 158. Tentative synonymy with Bufo (sensu lato) by Gorham, 1974, Checklist World Amph.: 146. Synonymy with Chaunus by implication of 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. Synonymy with Rhinella (by implication) by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211.

OtylophusCei, 1953, Arch. Zool. Ital. Torino, 38: 511. Incorrect subsequent spelling.

Rhamphophryne Trueb, 1971, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co., 216: 6. Type species: Rhamphophryne acrolopha Trueb, 1971, by original designation. Synonymy with Rhinella by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211, by implication.

Atelophryniscus McCranie, Wilson, and Williams, 1989, Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 129: 2. Type species: Atelophryniscus chrysophorus McCranie, Wilson, and Williams, 1989, by original designation. Synonymy with the "Bufo" veraguensis group by Pramuk and Lehr, 2005, J. Herpetol., 39: 610, and therefore associated with Chaunus in the sense of 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. Formal synonymy with Rhinella by Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211, by implication.

Nomina inquirenda - Name(s) unassigned to a living or extinct population

Bufo semilunatus Schneider, 1799, Hist. Amph. Nat.: 215. Type(s): "Museum Blochianum" (= ZMB). Type locality: "Surinamo". *Rana semilunataShaw, 1802, Gen. Zool., 3(1): 174 (Crescent Toad). See Daudin, 1803 "An. XI", Hist. Nat. Gen. Part. Rept., 8: 208. Presumably a member of Rhinella.

Bufo lacunatus Wiegmann, 1833, Isis von Oken, 26: 656. Types: Originally in ZMB, but not located. Type locality: Not stated. Provisionally allocated to a member of the Bufo marinus group by Dubois and Ohler, 1999, J. South Asian Nat. Hist., 4: 170.

English Names

South American Toads (Frost, McDiarmid, and Mendelson, 2008, in Crother (ed.), Herpetol. Circ., 37: 11; Frost, McDiarmid, Mendelson, and Green, 2012, in Crother (ed.), Herpetol. Circ., 39: 21; Frost, Lemmon, McDiarmid, and Mendelson, 2017, in Crother (ed.), Herpetol. Circ., 43: 19).

Beaked Toads (Rhamphophryne [no longer recognized]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 45).

Rio Viejo Toads (Atelophryniscus [no longer recognized]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 39).

Distribution

Lower Rio Grande Valley region of southern Texas (USA) and southern Sonora (Mexico) south through tropical Mexico and Central America to southern South America; one species (Rhinella marina) introduced widely (Antilles, Hawaii, Fiji, Philippines, Taiwan, Ryukyu Is. (Japan), New Guinea, Australia, and many Pacific islands).

Comment

The literature of Rhinella is deeply intertwined with that of Bufo (sensu lato) and any reference to "Bufo" in South America refers either to this genus, Nannophryne or Rhaebo. Literature and work to 1972 on Bufo (sensu lato, including Rhinella) were summarized in Blair, 1972, Evol. Genus Bufo. Pramuk and Kadivar, 2003, Herpetologica, 59: 277–280, provided a key to Bufo species (now Chaunus, Incilius, Rhaebo, or Rhinella) from Ecuador and Peru. Barrio-Amorós, 1999 "1998", Acta Biol. Venezuelica, 18: 7–10, reported on literature and distribution for the species in Venezuela. Graybeal, 1997, Zool. J. Linn. Soc., 119: 297–338, discussed phylogeny in the group (as part of Bufo). Duellman and Schulte, 1992, Copeia, 1992: 162–172, reviewed the similarity-defined groups of former Bufo in South America. See comment under Andinophryne atelopoides. Caramaschi and Pombal, 2006, Pap. Avulsos Zool., São Paulo, 46: 257, discussed morphological synapomorphies for Rhinella. Pramuk and Kadivar, 2003, Herpetologica, 59: 277–280, provided a key to Bufo species (now Incilius, Rhaebo, or Rhinella) from Ecuador and Peru. Pauly, Hillis, and Cannatella, 2004, Evolution, 58: 2517–2535, provided a tree of many exemplars as part of a study of New World "Bufo". Barrio-Amorós, 1999 "1998", Acta Biol. Venezuelica, 18: 7–10, reported on literature and distribution for the species in Venezuela. Graybeal, 1997, Zool. J. Linn. Soc., 119: 297–338, discussed phylogeny in the group (as part of Bufo). Duellman and Schulte, 1992, Copeia, 1992: 162–172, reviewed the similarity-defined groups of "Bufo" in South America and provided a key to the then Bufo veraguensis group. Cei, 1980, Monit. Zool. Ital., N.S., Monogr., 2: 157–213, discussed the species found in Argentina (as Bufo). Maciel, Schwartz, Colli, Castro, Fontes, and Schwartz, 2006, Biochem. Syst. Ecol., 34: 457–466, reported on phylogenetics of the Rhinella crucifer group (as Bufo) on the basis of indolealkylamines and protein skin secretions. Thomé, Zamudio, Giovanelli, Haddad, Baldissera, and Alexandrino, 2010, Mol. Phylogenet. Evol., 55: 1018–1031, also reported on the molecular phylogenetics of the group as well as post-Pliocene biogeography. Pramuk, 2006, Zool. J. Linn. Soc., 146: 407–452, based on the study of nuDNA, mtDNA, and morphology suggested that former Chaunus is paraphyletic with respect to Rhinella (sensu stricto) and Rhamphophryne, and this unit is the sister taxon of Anaxyrus + Incilius. See Trueb, 1971, Contrib. Sci. Nat. Hist. Mus. Los Angeles Co., 216: 6, for discussion and review of former Rhamphophryne, as well as synonymies for all of the species, except for Rhamphophryne proboscidea. Graybeal and Cannatella, 1995, Herpetologica, 51: 125, noted that there is no unambiguous evidence in support of the monophyly of former Rhamphophryne. 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: 215, suggested that Rhamphophryne is either imbedded within Rhinella (sensu stricto), or its sister taxon, and that Crepidophryne is likely allied with Rhamphophryne. Pramuk, 2006, Zool. J. Linn. Soc., 146: 407–452, who did not study Crepidophryne, found Rhamphophryne to be most closely related to Rhinella chavin, among her exemplars. Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 211, placed Chaunus and Rhamphophryne into the synonymy of Rhinella, but did not address Crepidophryne. They also demonstrated that the Rhinella spinulosus group is not monophyletic. Kwet, Di-Bernardo, and Maneyro, 2006, Iheringia, Zool., 96: 479–485, provided a key to the Rhinella marina group (as the Chaunus marinus group). Fouquet, Gaucher, Blanc, and Vélez-Rodriguez, 2007, Zootaxa, 1663: 17–32, discussed the Rhinella margaritifera group. Pramuk, Robertson, Sites, and Noonan, 2008, Global Ecol. Biogeograph., 17: 72–83, found Rhinella (sensu lato) to be the sister taxon of Anaxyrus + Incilius. Savage and Bolaños, 2009, Zootaxa, 2005: 4, on the basis of results of Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 203–212, regarded Rhinella (composed of the Rhinella margaritifer group and Rhinella veraguensis group) to form the sister taxon of Chaunus (composed of the Rhinella marinus, Rhinella crucifer, Rhinella granulosus, and Rhinella spinulosus groups). The problem is that, regardless of the results of Chaparro, Pramuk, and Gluesenkamp, 2007, Herpetologica, 63: 203–212, Pramuk, Robertson, Sites, and Noonan, 2008, Global Ecol. Biogeograph., 17: 72–83, on the basis of larger by somewhat different dataset found Chaunus (sensu Savage and Bolaños) to be paraphyletic on a and this needs to be clarified prior to partition. Moreover the hypothesized monophyly of Chaunus in the sense of Savage and Bolaños (2009) does not survive the analysis of Van Bocxlaer et al., 2010. Smith and Chiszar, 2006, Herpetol. Conserv. Biol., 1: 6–8, implied that this taxon should be considered a subgenus of Bufo; see comment under Bufonidae. Van Bocxlaer, Biju, Loader, and Bossuyt, 2009, BMC Evol. Biol., 9 (e131): 1–10, on the basis of molecular evidence suggested that Rhinella is the sister taxon of the Old World bufonid radiation and not in a monophyletic group with Incilius and Anaxyrus. Should this be corroborated by additional study, the discussion will be over of whether and New World component of former "Bufo" can be treated as Bufo. Narvaes and Rodrigues, 2009, Arq. Zool., São Paulo, 40: 1–73, revised the Rhinella granulosa group and provided a key and accounts for the species. Pauly, Hillis, and Cannatella, 2009, Herpetologica, 65: 115–128, on the basis of a largely undisclosed analysis (and in ignorance of the Van Bocxlaer et al., 2009, paper) considered Anaxyrus, Incilius, and Rhinella to be subgenera within a redelimited Bufo to form a single inclusive genus, Bufo. This was rejected as without a sufficient evidential basis by Frost, McDiarmid, and Mendelson, 2009, Herpetologica, 65: 136–153, compared with other analyses based on substantially more evidence. Van Bocxlaer, Loader, Roelants, Biju, Menegon, and Bossuyt, 2010, Science, 327: 679–682, suggested on the bais of molecular evidence that Rhinella is more closely related to Old World taxa than to other New World taxa. Vallinoto, Sequeira, Sodré, Bernardi, Sampaio, and Schneider, 2010, Zool. Scripta, 39: 128–140, reported on the molecular phylogenetics of the Rhinella marina group and suggested that it may be paraphyletic with respect to the Rhinella crucifer group. Maciel, Garcia, Colli, and Schwartz, 2010, Mol. Phylogenet. Evol., 57: 787–797, assumed monophyly of the Rhinella marina group on the basis of morphology presented by Pramuk (2006) and provided a molecular phylogenetic analysis of the species. Guerra, Baldo, Rosset, Borteiro, and Kolenc, 2011, Zootaxa, 3092: 26–42, reported on advertisement calls of the members of the Rhinella granulosa group. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, in their study of Genbank sequences, confirmed the monophyly of this taxon (although this is difficult to see on their tree because the authors explicitly adopted a non-monophyletic and out-dated taxonomy; e.g., recognizing Rhamphophryne and a polyphyletic "Bufo"), provided a tree for their exemplar species, and suggested that it is the sister taxon of what other authors refer to as Anaxyrus + Incilius. Mueses-Cisneros and Moreno-Quintero, 2012, Herpetotropicos, Mérida, 7: 39–54, noted two unnamed species of the Rhinella margaritifera group in Nariño, southwestern Colombia. Moravec, Lehr, Cusi, Córdova, and Gvoždík, 2014, ZooKeys, 371: 35–56, delimited the Rhinella festae species group. Köhler, 2011, Amph. Cent. Am.: 102–103 and 116, compared the species of Rhinella (as Chaunus, now a junior synonym of Rhinella) of Central America, provided identifications key, and photographs and range maps of the species. Fouquette and Dubois, 2014, Checklist N.A. Amph. Rept.: 290, considered Incilius a subgenus of Bufo, cherry-picking their citation to literature (excluding any reference to  Van Bocxlaer, Biju, Loader, and Bossuyt, 2009, BMC Evol. Biol., 9 (e131): 1–10, Van Bocxlaer, Loader, Roelants, Biju, Menegon, and Bossuyt, 2010, Science, 327: 679–682, or Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, which provided results not congruent with the story that Fouquette and Dubois wanted to tell) to avoid recognizing that treating this genus as a subgenus of Bufo also requires under current understanding of phylogeny all Old-World bufonids, such as SabahphrynusNectophryne, and Ansonia to be treated as subgenera of Bufo as well. Blotto, Pereyra, and Baldo, 2014, J. Herpetol., 48: 434–438, reported on larval morphology of the Rhinella granulosa group and suggested characteristics that may be synapomorphic. Oliveira, Weber, and Napoli, Herpetol. J., 24: 229–236, compared chondrocranial and hyobranchia larval morphology among Rhinella abei, Rhinella crucifer, Rhinella granulosa, Rhinella henseli, Rhinella hoogmoedi, Rhinella icterica, Rhinella jimi, Rhinella ornata, Rhinella pombali, Rhinella pygmaea, and Rhinella schneideri. Pereyra, Baldo, Blotto, Iglesias, Thomé, Haddad, Barrio-Amorós, Ibáñez D., and Faivovich, 2016, Cladistics, 32: 36–53, reported on the molecular phylogenetics of the Rhinella granulosa group. 

Contained taxa (93 sp.):

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