Phrynopus Peters, 1873

Common Names

Andes Frogs (Frank and Ramus, 1995, Compl. Guide Scient. Common Names Amph. Rept. World: 82).

Distribution

Upper humid montane forests and supra-treeline grasslands in the Cordillera Oriental in central Peru, and one locality in the Cordillera Occidental in Peru.

Comment

Lynch, 1975, Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 35: 1–51, provided synonymies and discussion of the relationships of and within this genus. Species groups noted in comments with the species records follow his revision. According to Lynch, Phrynopus is most closely related to some species of Eleutherodactylus. Cannatella, 1984, Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 113: 1–16, reinvestigated the phylogenetic reconstructions of Lynch. Even though differences were noted, Cannatella refrained from redefining species groups because of the topological instability among the alternative trees generated. See comment under Phyllonastes. Lynch, 1994, Rev. Acad. Colomb. Cienc. Exact. Fis. Nat., 19: 195–203, suggested that Phrynopus might be a subset of Eleutherodactylus, most closely related to species such as Eleutherodactylus nicefori. Duellman, 2000, Herpetologica, 56: 273–285, reviewed the species of northern Peru. Lehr, Fritzsch, and Müller, 2005, Zool. Scripta, 34: 593–603, provided mtDNA sequence evidence that Phyllonastes is nested within Phrynopus, and that Phrynopus is deeply polyphyletic; they excluded "Phrynopus" carpish (to Microhylidae) and Phrynopus simonsii to Eleutherodactylus to render a monophyletic Phrynopus. (They did not actually accomplish this, however, as their results show Phrynopus as paraphyletic with respect to Phyllonastes—DRF.) 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: 125, unambiguously placed Phrynopus within a paraphyletic Eleutherodactylus. See comments by Lehr, 2006, Herpetologica, 62: 343, regarding the difficulty of assigning species among Eleutherodactylus, Phyllonastes, and Phrynopus, and his suggestion that Phrynopus will ultimately be restricted to species south of the Huancabamba Depression in Peru. Heinicke, Duellman, and Hedges, 2007, Proc. Natl. Acad. Sci. USA, 104: 10092–10097, suggested that Phrynopus is polyphyletic, with Phrynopus cophites being closer to Barycholos, Phrynopus bracki the sister taxon of Pristimantis, and "Phrynopus" brunneus and "Phrynopus" peraccai being most closely related to "Eleutherodactylus" dolops. Hedges, Duellman, and Heinicke, 2008, Zootaxa, 1737: 1–182, redelimited and restricted Phrynopus to a putatively monophyletic group, with other species being assigned to other genera. Trueb and Lehr, 2008, Phyllomedusa, 7: 11–24, discussed the comparative osteology within Phrynopus. See Duellman and Lehr, 2009, Terrest.-breeding Frogs in Peru: 104–126, for brief accounts of the Peruvian species. Pyron and Wiens, 2011, Mol. Phylogenet. Evol., 61: 543–583, confirmed the monophyly of this taxon and provided a tree of their molecular exemplar species. Padial, Grant, and Frost, 2014, Zootaxa, 3825: 53, transferred this genus into Holoadeninae.  Lehr and Rodríguez, 2017, Salamandra, 53: 327–338, provided a comparison of the species from the Cordillera de Carpish in central Peru. von May, Lehr, and Rabosky, 2018, PeerJ, 6(e4313): 1–27, reported on molecular phylogenetics of the group and found evidence of monophyly. Venegas, Barboza, De la Riva, and Padial, 2018, Zootaxa, 4446: 511, provided a map of the species. Ospina-Sarria and Grant, 2022, Zool. J. Linn. Soc., 195: 976–994, reported on morphology of pelivic and thigh musculature and digital discs of the Brachycephaloidea, and identified an unambiguous synapomorphy of Phrynopus. 

Contained taxa (37 sp.):

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