Brachycephalus Fitzinger, 1826

English Names

Saddleback Toads (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 38).

Three-toed Toadlets (Halliday and Adler, 2002, New Encyclop. Rept. Amph.: 85).

Flea Toads (Napoli, Caramaschi, Cruz, and Dias, 2011, Zootaxa, 2739: 33).

Izecksohn's Toads (Psyllophryne [no longer recognized]: Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 38).

Distribution

Southeastern Brazil.

Comment

See comments on the status of the former nominal genera Brachycephalus and Psyllophryne by Pombal, Wistuba, and Bornschein, 1998, J. Herpetol., 32: 70–74, as well as by Kaplan, 2002, Amphibia-Reptilia, 23: 227. See comments under Adelophryne and Euparkerella. Alves, Ribeiro, Haddad, and Reis, 2006, Herpetologica, 62: 221–233, provided considerable information on the external morphology and osteology of the species of Brachycephalus. Silva, Campos, and Sebben, 2007, Zootaxa, 1422: 59–68, commented on the auditory region and the monophyly of the genus. Heinicke, Duellman, and Hedges, 2007, Proc. Natl. Acad. Sci. USA, 104: 10092–10097, provided additional evidence for Brachycephalus being imbedded within the former Eleutherodactylinae. See comment under Brachycephalidae. Pombal, 2010, Bol. Mus. Nac., Rio de Janeiro, N.S., Zool., 526: 1–12, redelimited the species of Brachycephalus. Clemente-Carvalho, Klaczko, Perez, Alves, Haddad, and Reis, 2011, Mol. Phylogenet. Evol., 61: 77–89, reported on molecular phylogenetics and phenotypic diversity. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, confirmed the sister taxon relationship of Brachycephalus and Ischnocnema. Padial, Grant, and Frost, 2014, Zootaxa, 3825: 50–51, summarized the literature of the systematics of the group and corroborated the results of Clemente-Carvalho, Klaczko, Perez, Alves, Haddad, and Reis, 2011, Mol. Phylogenet. Evol., 61: 79–89, that two major clades exist in the genus: 1) a southern clade formed by locally endemic species restricted to the high and humid Atlantic Forest of Paraná (Brachycephalus brunneus, Brachycephalus ferruginus, Brachycephalus izecksohni, Brachycephalus pernix, and Brachycephalus pombali) and the northern clade, allopatric to the first, distributed from farther north in Paraná to Espírito Santo (Brachycephalus alipioi, Brachycephalus didactylus, Brachycephalus ephippium, Brachycephalus garbeanus, Brachycephalus hermogenesi, Brachycephalus nodoterga, Brachycephalus pitanga, Brachycephalus toby, and Brachycephalus vertebralis), with the two species of the former Psyllophryne placed separately in the second clade. Ribeiro, Bornschein, Belmonte-Lopes, Firkowski, Morato, and Pie, 2015, PeerJ, 3 (e1011): 1–35, formalized the two arguably monophyletic groups as the a) Brachycephalus ephippium group (found in the states of Rio de Janeiro, Sao Paulo, and Espirito Santo), b) Brachycephalus pernix group (Parana and Santa Catarina), and c) the Brachycephalus didactylus group for former Psyllophryne. Condez, Monteiro, and Haddad, 2017, Zootaxa, 4290: 395–400, criticized the Brachycephalus species delimitation with reference to the papers of Pie and Ribeiro, 2015, PeerJ, 3(e1179): 1–9; Ribeiro, Bornschein, Belmonte-Lopes, Firkowski, Morato, and Pie, 2015, PeerJ, 3 (e1011): 1–36, and Bornschein, Ribeiro, Blackburn, Stanley, and Pie, 2016, PeerJ, 4(e2629): 1–19, and suggested that given the errors in these papers that several species required reassessment. Pie, Ribeiro, and Bornschein, 2017, Zootaxa, 4350: 587–589, responded to these criticisms and suggested that according to the criteria of Condez et al. (2017), virtually no Brachycephalus species should be valid (including their own), noting published and ongoing molecular work (Firkowski, Bornschein, Ribeiro, and Pie, 2016, Mol. Phylogenet. Evol., 100 : 345–360, iwho reported on comparative species delimitation methods) in support of their position. Pie, Faircloth, Ribeiro, Bornschein, and McCormack, 2018, Biol. J. Linn. Soc., 125: 72–82, reported on phylogenetics and biogeography of the species. Condez, Haddad, and Zamudio, 2020, Biol. J. Linn. Soc., 129: 664–686, reported on the biogeography and evolution of the species. Lyra, Monteiro, Rancilhac, Irisarri, Künzel, Sanchez, Condez, Rojas-Padilla, Solé, Toledo, Haddad, and Vences, 2021, Toxins, 13 (816): 1–16, discussed the molecular phylogeny within the genus with special reference to the "pumpkin toadlet" morphotype and probable toxicity. Folly, Vrcibradic, Siqueira, Rocha, Machado, Lopes, and Pombal, 2022, Ichthyol. & Herpetol., 110: 585–601, redelimited the species groups, rejecting the former Brachycephalus didactylus group as nonmonophyletic, leaving several species unassigned to group (e.g., Brachycephalus hermogenesi, Brachycephalus pulex, Brachycephalus sulfuratus, Brachycephalus didactylus, and Brachycephalus clarissae), and retaining only three species groups: (1) the Brachycephalus ephippium group (Brachycephalus rotenbergae, Brachycephalus margaritatus, Brachycephalus ibitinga, Brachycephalus ephippium, and Brachycephalus garbeanus); (2) the Brachycephalus vertebralis group (Brachycephalus crispus, Brachycephalus garbeanus, Brachycephalus ephippium, Brachycephalus ibitinga, Brachycephalus margaritatus, and Brachycephalus rotenbergae), and the (3) Brachycephalus pernix group (Brachycephalus ferrugineus, Brachycephalus pernix, Brachycephalus pombali, Brachycephalus quirirensis, Brachycephalus actaeus, Brachycephalus auroguttatus, Brachycephalus mariaeterezae, Brachycephalus verrucosus, Brachycephalus brunneus, Brachycephalus coloratus, Brachycephalus curupira, vizecksohni, Brachycephalus leopardus, Brachycephalus tridactylus, Brachycephalus fuscolineatus, Brachycephalus mirissimus, Brachycephalus boticaro, and Brachycephalus olivaceus). Folly, Luna-Dias, Miguel, Ferreira, Machado, Lopes, and Pombal, 2023, Acta Zool., Stockholm, 104: 71–105, reviewed the osteology of the Brachycephalus ephippium group. Bornschein, Ribeiro, and Pie, 2023, Phyllomedusa, 20: 109–115, discussed the taxonomic history and its ambiguities of the systematics of the genus. 

Contained taxa (41 sp.):

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