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A new species of Andean frog of the genus Bryophryne from southern Peru Anura: Craugastoridae) and its phylogenetic position, with notes on the diversity of the genus



We describe a new species of terrestrial frog of the genus Bryophryne (Anura: Craugastoridae) from the wet puna and elfin forests of the Amazonian versant of the Andes. The new species seems to be restricted to high altitude environments at elevations between 3506–3651 m in the area now protected by Megantoni National Sanctuary and Manu National Park (Distrito de Echarate, Provincia La Convención, Departamento Cusco, Peru). The new species is characterized by lacking vomerine processes of vomers, by having tympanic annulus and tympanic membrane not evident through the skin, smooth dorsal skin with scattered warts, conspicuous dorsolateral, middorsal, and occipital folds, warty flanks, areolate skin on ventral surfaces of the body, and by lacking finger and toe fringes and basal web on feet. In life, specimens have bright and highly variable dorsal coloration that ranges from olive-green to red with variable combinations of red or orange marks (red or orange in the green form and olive-green in the red form). Molecular phylogenetic analyses of mitochondrial and nuclear DNA place the new species within the genus Bryophryne and as sister group of B. cophites. Bryophryne bustamantei, also sequenced for this study, is found as the sister group of the clade formed by B. cophites and the new species. Bryophryne is found as sister group of Psychrophrynella in maximum likelihood analyses and as the sister group of a large clade of holoadenines in parsimony analyses. The genus Bryophryne now contains nine species, all of them distributed along the Cordillera Oriental of the Peruvian Andes, southeast of the Apurimac River valley.




Andes, Brachycephaloidea, wet puna, Psychrophrynella, Phrynopus, taxonomy, terraranas

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Bazinet, A.L., Zwickl, D.J., Michael, P. & Cummings, P. (2014) A Gateway for Phylogenetic Analysis Powered by Grid Computing Featuring GARLI 2.0. Systematic Biology, 63, 812–818.

Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences (USA), 97, 6585–6590.

Canedo, C. & Haddad, C.F.B. (2012) Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 65, 610–620.

Chaparro, J.C., De la Riva, I., Padial, J.M., Ochoa, J.A. & Lehr, E. (2007) A new species of Phrynopus from Departamento Cusco, southern Peru (Anura: Brachycephalidae). Zootaxa, 1618, 61–68.

Crawford, A.J. (2003) Huge populations and old species of Costa Rican and Panamanian dirt frogs inferred from mitochondrial and nuclear gene sequences. Molecular Ecology, 12, 2525–2540.

De la Riva, I. (2007) Bolivian frogs of the genus Phrynopus, with the description of twelve new species (Anura: Brachycephalidae). Herpetological Monographs, 21, 242–278.

De la Riva, I. & Burrowes, P.A. (2014) A new species of Psychrophrynella (Anura: Craugastoridae) from the Cordillera Real, Department La Paz, Bolivia. Zootaxa, 3887 (4), 459–470.

Duellman, W.E. & Lehr, E. (2009) Terrestrial–Breeding Frogs (Strabomantidae) in Peru. Natur und Tier, Verlag GmbH, 382 pp.

Farris, J.S., Albert, V.A., Källersjö, M., Lipscomb, D. & Kluge, A.G. (1996) Parsimony jackknifing outperforms neighbor–joining. Cladistics, 12, 99–124.

Frost, D.R. (2015) Amphibian Species of the World: an Online Reference. Version 6.0 (Date of access). Electronic Database. American Museum of Natural History, New York, USA. Available from: (accessed 30 June 2015)

Goloboff, P.A. (1996) Methods for faster parsimony analysis. Cladistics, 12, 199–220.

Goloboff, P.A. (1999) Analyzing large data sets in reasonable times, solutions for composite optima. Cladistics, 15, 415–428.

Goloboff, P.J., Farris, S. & Nixon, K. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.

Hedges, S.B., Duellman, W.E. & Heinicke, M.P. (2008) New World direct developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1–182.

Heinicke, M.P., Duellman, W.E. & Hedges, S.B. (2007) Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. Proceedings of the National Academy of Sciences USA, 24, 10092–10097.

Hillis, D. M., Moritz, C. & Mable, B.K. (1996) Molecular Systematics. 2nd Edition. Sinauer Associates Inc., Sunderland, Massachusetts, 655 pp.

Jungfer, K.H., Faivovich, J., Padial, J.M., Castroviejo-Fisher, S., Lyra, M., Berneck, B., Iglesias, P., Kok, P.J.R., MacCulloch, R.D., Trefaut-Rodrigues, M., Verdade, V.K., Torres-Gastello, C.P., Chaparro, J.C., Valdujo, P.H., Reichle, S., Moravec, J., Gvoždík, V., Gagliardi-Urrutia, Ernst, R., De la Riva, I., Means, D.B., Lima, A.P., Señaris, J.C., Wheeler, W.C. & Haddad, C.F.B. (2013) Systematics of spiny-backed treefrogs (Hylidae: Osteocephalus): an Amazonian puzzle. Zoologica Scripta, 42, 351–380.

Katoh, K., Kuma, K., Toh, H. & Miyata, T. (2005) MAFFT, a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30, 3059–3066.

Lanfear, R., Calcott, B., Ho, S.Y. & Guindon, S. (2012) PartitionFinder, combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29, 1695–1701.

Lehr, E. & Catenazzi, A. (2008) A new species of Bryophryne (Anura: Strabomantidae) from southern Peru. Zootaxa, 1784, 1–10.

Lehr, E. & Catenazzi, A. (2009) Three new species of Bryophryne (Anura: Strabomantidae) from the region of Cusco, Peru. South American Journal of Herpetology, 4, 125–138.

Lehr, E. and Catenazzi, A. (2010) Two new species of Bryophryne (Anura: Strabomantidae) from high elevations in southern Peru (Region of Cusco). Herpetologica, 66, 308–319.

Lehr, E., & Oróz, A. (2012) Two new species of Phrynopus (Anura: Strabomantidae) from the Cordillera de Carpish in central Peru (Departamento de Huánuco). Zootaxa, 3512, 53–63.

Lehr, E., Fritzsch, G. & Müller, A. (2005) Analysis of Andes frogs (Phrynopus, Leptodactylidae, Anura) phylogeny based on 12S and 16S mitochondrial rDNA sequences. Zoologica Scripta, 6, 593–603.

Lehr, E., Moravec, J. & Cusi, J.C. (2012) Two new species of Phrynopus (Anura, Strabomantidae) from high elevations in the Yanachaga-Chemillén National park in Peru (Departamento de Pasco). ZooKeys, 235, 51–71.

Lynch, J.D. (1975). A review of the Andean leptodactylid frog genus Phrynopus. Occasional Papers of the Museum of Natural History, University of Kansas, 35, 1–51.

Mamani, L. & Malqui, S. (2014) A new species of Phrynopus (Anura: Craugastoridae) from the central Peruvian Andes. Zootaxa, 3838 (2), 207–214.

Padial, J. M., Castroviejo-Fisher, S. and De la Riva, I. 2009. The phylogenetic relationships of Yunganastes revisited (Anura: Terrarana). Molecular Phylogenetics and Evolution, 52, 911–915.

Padial, J.M., Chaparro, J.C., Castroviejo-Fisher, S., Guayasamín, J.M., Lehr, E., Delgado, A.J., Vaira, M., Teixeira, M., Aguayo, R. & De la Riva, I. (2012) A revision of species diversity in the Neotropical genus Oreobates (Anura: Strabomantidae), with the description of three new species from the Amazonian slopes of the Andes, and the proposal of candidate species. American Museum Novitates, 3752, 1–55.

Padial, J.M., Grant, T. & Frost, D.R. (2014). Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa, 3825 (1), 1–132.

Pereyra, M.O., Cardozo, D., Baldo, J.L. & Baldo, D. (2014) Description and phylogenetic position of a new species of Oreobates (Anura: Craugastoridae) from northwestern Argentina. Herpetologica, 70, 211–227.

Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2,800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543–583.

Teixeira, M. Jr., Amaro, R.C., Recoder, R.S., de Sena, M.A. & Rodrigues, M.T. (2012) A relict new species of Oreobates (Anura, Strabomantidae) from the seasonally dry tropical forests of Minas Gerais, Brazil, and its implication to the biogeography of the genus and that of South American dry forests. Zootaxa, 3158, 37–52.

Wheeler, W.C. (1996) Optimization alignment, the end of multiple sequence alignment in phylogenetics? Cladistics, 12, 1–10.

Wheeler, W.C. (2003) Implied alignment, a synapomorphy-based multiple sequence alignment method and its use in cladogram search. Cladistics, 19, 261–268.

Wheeler, W.C., Arango, C.P., Grant, T., Janies, D., Varón, A., Aagesen, L., Faivovich, J., D'Haese, C., Smith, W.L. & Giribet, G. (2006) Dynamic Homology and Phylogenetic Systematics, A Unified Approach Using POY. American Museum of Natural History, New York, 365 pp.

Wheeler, W.C., Lucaroni, N., Hong, L., Crowley, L.M. & Varón, A. (2015) POY version 5: phylogenetic analysis using dynamic homologies under multiple optimality criteria. Cladistics, 31, 189–196.

Wiens, J.J., Fetzner, J.W., Parkinson, C.L. & Reeder, T.W. (2005) Hylid frog phylogeny and sampling strategies for speciose clades. Systematic Biology, 54, 719–748.

Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D. Dissertation, The University of Texas at Austin. [unkown pagination]


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