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Abstract:

The frog clade composed of the alsodid genera Alsodes+Eupsophus is the most species-rich of the Patagonian endemic frog clades, including nearly 31 of the slightly more than 50 species of that region. The biology of this group of frogs is poorly known, its taxonomy quite complex (particularly Alsodes), and its diversity in chromosome number striking when compared with other frogs (collectively, there are species having 2n=22, 2n=26, 2n=28, 2n=30 or 2n=34). We present a phylogenetic analysis of this Patagonian frog clade based on mitochondrial and nuclear gene sequences. We sequenced five mitochondrial genes (cytochrome b, cytochrome oxidase I, 12S, 16S, NADH dehydrogenase subunit 1) with three intervening tRNAs, and fragments of three nuclear genes (seven in absentia homolog 1, rhodopsin exon 1, RAG-1), for a maximum of 6510bp for multiple specimens from 26 of the 31 species. We recovered Eupsophus as polyphyletic, with E. antartandicus, E. sylvaticus, and E. taeniatus in Batrachylidae, in accordance with most previous hypotheses. Based on this result, we transfer E. antartandicus and E. taeniatus back to Batrachyla, and E. sylvaticus to Hylorina (resurrected from the synonymy of Eupsophus), remediating the paraphyly of Eupsophus. Our results strongly corroborate the monophyly of Alsodes+Eupsophus (sensu stricto), the individual monophyly of these genera, and the monophyly of the species groups of Eupsophus. They also show the non-monophyly of all non-monotypic species groups of Alsodes proposed in the past. Our results expose several taxonomic problems particularly in Alsodes, and to a lesser extent in Eupsophus. This phylogenetic context suggests a rich evolutionary history of karyotypic diversification in the clade, in part corroborating previous hypotheses. In Alsodes, we predict three independent transformations of chromosome number from the plesiomorphic 2n=26. All these, strikingly, involve increments or reductions of pairs of haploid chromosomes. Finally, the phylogenetic pattern recovered for Alsodes and Eupsophus suggests a trans-Andean origin and diversification of the group, with multiple, independent ingressions over cis-Andean regions. © The Willi Hennig Society 2012.

Registro:

Documento: Artículo
Título:Phylogenetic relationships of a Patagonian frog radiation, the Alsodes+Eupsophus clade (Anura: Alsodidae), with comments on the supposed paraphyly of Eupsophus
Autor:Blotto, B.L.; Nuñez, J.J.; Basso, N.G.; Úbeda, C.A.; Wheeler, W.C.; Faivovich, J.
Filiación:División Herpetología, Museo Argentino de Ciencias Naturales Bernardino Rivadavia-CONICET, Ciudad Autónoma de Buenos Aires, Ángel Gallardo 470, C1405DJR, Argentina
Instituto de Zoología, Universidad Austral de Chile, Casilla 567, Valdivia, Chile
Centro Nacional Patagónico (CENPAT), Blvd. Brown 2915, U-9120-ACF Puerto Madryn, Argentina
Centro Regional Universitario Bariloche, Universidad Nacional del Comahue, Quintral 1250, R-8400-FRF San Carlos de Bariloche, Río Negro, Argentina
Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th St, New York, NY 10024, United States
Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Argentina
Palabras clave:adaptive radiation; chromosome; cladistics; endemic species; frog; geographical distribution; phylogeny; pollution exposure; taxonomy; Andes; Patagonia
Año:2013
Volumen:29
Número:2
Página de inicio:113
Página de fin:131
DOI: http://dx.doi.org/10.1111/j.1096-0031.2012.00417.x
Título revista:Cladistics
Título revista abreviado:Cladistics
ISSN:07483007
CODEN:CLADE
Registro:https://bibliotecadigital.exactas.uba.ar/collection/paper/document/paper_07483007_v29_n2_p113_Blotto

Referencias:

  • Barrio, A., Insuetophrynus acarpicus, un nuevo leptodactílido firmisternio sudamericano (Amphibia, Anura) (1970) Physis, 30, pp. 331-341. , Buenos Aires
  • Barrio, A., Rinaldi de Chieri, P., Contribución al esclarecimiento de la posición taxofilética de algunos batracios patagónicos de la familia Leptodactylidae mediante el análisis cariotípico (1971) Physis, 30, pp. 673-685. , Buenos Aires
  • Basso, N.G., Úbeda, C.A., Geographic distribution. Eupsophus vertebralis (1999) Herpetol. Rev., 30, p. 172
  • Burton, T.C., Variation in the hand and superficial throat musculature of Neotropical leptodactylid frogs (1998) Herpetologica, 54, pp. 53-72
  • Burton, T.C., Pointing the way: the distribution and evolution of some characters of the finger muscles of frogs (1998) Am. Mus. Novit., 3229, pp. 1-13
  • Cárdenas-Rojas, D.R., Veloso, A., de Sá, R.O., The tadpole of Eupsophus queulensis (Anura, Cycloramphidae) (2007) Alytes, 25, pp. 45-54
  • Cei, J.M., Remarks on some Neotropical amphibians of the genus Alsodes from southern Argentina (1976) Atti. Soc. Ital. Sci. nat. Museo civ. nat. Milano, 117, pp. 159-164
  • Cei, J.M., Amphibians of Argentina (1980) Monitore Zool. Ital. (NS), 2, pp. 1-609
  • Corbalán, V., Debandi, G., Martínez, F., Alsodes pehuenche (Anura: Cycloramphidae): past, present and future (2010) Cuad. Herpetol., 24, pp. 17-23
  • Correa, C.L., Iturra, P., Méndez, M.A., Phylogenetic relationships of Chilean leptodactylids: a molecular approach based on mitochondrial genes 12S and 16S (2006) Rev. Chil. Hist. Nat., 79, pp. 435-450
  • Correa, C.L., Sallaberry, M., Iturra, P., Collado, G., Méndez, M.A., Amphibia, Anura, Cycloramphidae, Alsodes montanus: new record and geographic distribution map (2008) Check List, 4, pp. 467-471
  • Cuevas, C., A new species of the genus Alsodes (Anura: Neobatrachia) from the Nothofagus forest, coastal range, southern Chile, identified by its karyotype (2008) Zootaxa, 177, pp. 43-53
  • Cuevas, C.C., Formas, J.R., Heteromorphic sex chromosomes in Eupsophus insularis (Amphibia: Anura: Leptodactylidae) (1996) Chromosome Res., 4, pp. 467-470
  • Cuevas, C.C., Formas, J.R., A new species of Alsodes (Amphibia, Anura, Leptodactylidae) from Central Chile (2001) Amphib-reptil., 22, pp. 187-198
  • Cuevas, C.C., Formas, J.R., Cytogenetic analysis of four species of the genus Alsodes (Anura: Leptodactylidae) with comments about the karyological evolution of the genus (2003) Hereditas, 138, pp. 138-147
  • Cuevas, C., Formas, R., A new frog of the genus Alsodes (Leptodactylidae) from Tolhuaca Nacional Park, Andes Range, southern Chile (2005) Amphib-reptil., 26, pp. 39-48
  • Cuevas, C., Formas, R., Novel diploid number and chromosomal formula in a South American frog of the genus Alsodes (Amphibia, Leptodactylidae) (2005) Ital. J. Zool., 72, pp. 87-91
  • Darst, C.R., Cannatella, D.C., Novel relationships among hyloid frogs inferred from 12S and 16S mitochondrial DNA sequences (2004) Mol. Phylogenet. Evol., 31, pp. 462-475
  • De Laet, J., Parsimony and the problem of inapplicables in sequence data (2005) Parsimony, Phylogeny, and Genomics, pp. 81-116. , Albert, V.A. (Ed.), Oxford University Press, London -
  • Faivovich, J., Haddad, C.F.B., Garcia, P.C.A., Frost, D.R., Campbell, J.A., Wheeler, W.C., Systematic review of the frog family Hylidae, with special reference to Hylinae: Phylogenetic analysis and taxonomic revision (2005) Bull. Amer. Mus. Nat. Hist., 294, pp. 1-240
  • Farris, J.S., The logical basis of phylogenetic analysis (1983) Advances in Cladistics: Proceedings of the Third Meeting of the Willi Hennig Society, pp. 7-36. , Platnick, N.I., Funk, V.A. (Eds), Columbia University Press, New York -
  • Farris, J.S., Albert, V.A., Källersjo, M., Lipscomb, D., Kluge, A.G., Parsimony jackknifing outperforms neighbour-joining (1996) Cladistics, 12, pp. 99-124
  • Fernández de la Reguera, P.A., Identifying species in the Chilean frogs by principal component analysis (1987) Herpetologica, 43, pp. 173-177
  • Formas, J.R., The karyotypes of the Chilean frogs Eupsophus emiliopugini and E. vertebralis (Amphibia: Anura: Leptodactylidae) (1991) Proc. Biol. Soc. Wash., 104, pp. 7-11
  • Formas, J.R., El cariotipo de la rana chilena Eupsophus contulmoensis (Anura, Leptodactylidae), con comentarios sobre la evolución cariológica del género Eupsophus (1992) Bol. Soc. Biol. Concepc., 63, pp. 77-82
  • Formas, J.R., Brieva, L., Advertisement calls and relationships of Chilean frogs Eupsophus contulmoensis and E. insularis (Amphibia: Anura: Leptodactylidae) (1994) Proc. Biol. Soc. Wash., 107, pp. 391-397
  • Formas, J.R., Vera, M.A., Reproductive patterns of Eupsophus roseus and E. vittatus (1980) J. Herpetol., 14, pp. 11-14
  • Formas, J.R., Vera, M.I., Karyological relationships among frogs of the genus Alsodes, with description of the karyotypes of A. vanzolinii and A. verrucosus (1983) Copeia, 1983, pp. 1101-1107
  • Formas, J.R., Pugín, E., Jorquera, B., La identidad de batracio chileno Heminectes rufus Philippi, 1902 (1975) Physis, 34, pp. 147-157. , Buenos Aires
  • Formas, J.R., Vera, M.I., Lacrampe, S., Allozymic and morphological differentiation in the South American frogs genus Eupsophus (1983) Comp. Biochem. Physiol., B, 75, pp. 475-478
  • Formas, J.R., Cuevas, C., Nuñez, J., A new species of Alsodes (Amphibia: Anura: Leptodactylidae) from southern Chile (1998) Proc. Biol. Soc. Wash., 111, pp. 521-530
  • Formas, J.R., Cuevas, C.C., Brieva, L.M., A new species of Alsodes (Anura: Leptodactylidae) from Cerro Mirador, Cordillera Pelada, southern Chile (2002) Proc. Biol. Soc. Wash., 115, pp. 708-719
  • Formas, J.R., Nuñez, J.N., Cuevas, C.C., Identidad de la rana austral chilena Eupsophus coppingeri (Amphibia, Anura, Neobatrachia): evidencias morfológicas, cromosómicas y moleculares (2008) Rev. Chil. Hist. Nat., 81, pp. 3-20
  • Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M., Gemmell, N.J., Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses (2007) PLoS ONE, 2, pp. e1109
  • Frost, D.R., (2011), http://research.amnh.org/vz/herpetology/amphibia/, Amphibian Species of the World: An Online Reference. Version 5.5 (31 January 2011). Electronic database accessible at American Museum of Natural History, New York, USA; Frost, D.R., Grant, T., Faivovich, J., Bain, R., Haas, A., Haddad, C.F.B., de Sá, R.O., Wheeler, W.C., The amphibian tree of life (2006) Bull. Amer. Mus. Nat. Hist., 297, pp. 1-370
  • Gallardo, J.M., A propósito de los Telmatobiinae (Anura, Leptodactylidae) Patagónicos (1970) Neotropica, 16, pp. 73-85
  • Goloboff, P.A., Analyzing large data sets in reasonable times: solutions for composite optima (1999) Cladistics, 15, pp. 415-428
  • Goloboff, P.A., Parsimony, likelihood, and simplicity (2003) Cladistics, 19, pp. 91-103
  • Goloboff, P.A., Pol, D., Parsimony and Bayesian phylogenetics (2005) Parsimony, Phylogeny, and Genomics, pp. 148-159. , Albert, V.A. (Ed), Oxford University Press, London -
  • Goloboff, P.A., Farris, J.S., Nixon, K.C., TNT, a free program for phylogenetic analysis (2008) Cladistics, 24, pp. 774-786
  • Grant, T., Frost, D.R., Caldwell, J.P., Gagliardo, R., Haddad, C.F.B., Kok, P.J.R., Means, D.B., Wheeler, W., Phylogenetic systematic of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae) (2006) Bull. Am. Mus. Nat. Hist., 299, pp. 1-262
  • Green, D.M., Sessions, S.K., Karyology and Cytogenetics (2007) Amphibian Biology, pp. 2756-2841. , Heatwole, H., Tyler, M. (Eds), Surrey Beatty and Sons, Chipping Norton, Australia -
  • Hall, T.A., (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis, 41, pp. 95-98. , program for Windows 95/98/NT. Nucl. Acids Symp. Ser
  • Heusser, C.J., Late-glacial-Holocene climate of Lake District of Chile (1984) Quat. Res., 22, pp. 77-90
  • Heyer, W.R., Crombie, R.I., Natural history notes on Craspedoglossa stejnegeri and Thoropa petropolitana (Amphibia: Salientia: Leptodactylidae) (1979) J. Wash. Acad. Sci., 69, pp. 17-20
  • Huelsenbeck, J.P., Ronquist, F., MrBayes: Bayesian inference of phylogeny (2001) Bioinformatics, 17, pp. 754-755
  • Ibarra-Vidal, H., Ortiz, J.C., Torres-Pérez, F., Eupsophus septentrionalis n. sp., nueva especie de Leptodactylidae (Amphibia) de Chile central (2004) Bol. Soc. Biol. Concepc., 75, pp. 91-102
  • Iturra, P., Veloso, A., Further evidence for early sex chromosome differentiation of Anuran species (1989) Genetica, 78, pp. 25-31
  • Jorquera, B., Pugín, E., Garrido, O., Goicoechea, O., Formas, R., Procedimiento de desarrollo en dos especies del género Rhinoderma (1981) Medio Ambiente, 5, pp. 58-71
  • Kaplan, M., Evaluation and redefinition of the states of anuran pectoral girdle architecture (2004) Herpetologica, 60, pp. 84-97
  • Lavilla, E.O., Condrocráneo y esqueleto visceral en larvas de Cycloramphus stejnegeri (Leptodactylidae) (1991) Amphib-reptil., 12, pp. 33-38
  • Lavilla, E.O., Estructura del condrocráneo y esqueleto visceral de larvas de Alsodes barrioi (Anura: Leptodactylidae) (1992) Acta Zool. Lill., 42, pp. 13-18
  • Lutz, B., Biologia e taxonomia de Zachaenus parvulus (1944) Bol. Mus. Nac., Rio de Janeiro, NS, Zool., 17, pp. 1-66
  • Lynch, J.D., Evolutionary relationships, osteology, and zoogeography of leptodactyloid frogs (1971) Misc. Publ. Mus. Nat. Hist. Univ. Kans., 53, pp. 1-238
  • Lynch, J.D., A re-assessment of the telmatobiine leptodactylid frogs of Patagonia (1978) Occ. Pap. Mus. Nat. Hist. Univ. Kansas, 72, pp. 1-57
  • Méndez, M.A., Torres-Pérez, F., Correa, C., Soto, E.R., Nuñez, J.J., Veloso, A., Armesto, J., Genetic divergence in the endangered frog Insuetophrynus acarpicus (Anura: Leptodactylidae) (2006) Herpetol. J., 16, pp. 93-96
  • Nicoli, L., On the putative presence of Eupsophus (Anura: Cycloramphidae) in central Patagonia during the Oligocene (2012) Herpetologica, 68, pp. 236-243
  • Nixon, K.C., The parsimony ratchet, a new method for rapid parsimony analysis (1999) Cladistics, 15, pp. 407-414
  • Nixon, K.C., (2002) WinClada, vers. 1.00.8, , Published by the author, Ithaca, NY
  • Nuñez, J.J., Úbeda, C.A., The tadpole of Eupsophus nahuelbutensis (Anura: Neobatrachia): external morphology, chondrocranium, and comments on its natural history (2009) Zootaxa, 2126, pp. 58-68
  • Nuñez, J.J., Rabanal, F.E., Formas, J.R., Description of a new species of Eupsophus (Amphibia: Neobatrachia) from the Valdivian Coastal range, Southern Chile: an integrative taxonomic approach (2012) Zootaxa, 3305, pp. 53-68
  • Nuñez, J.J., Wood, N.K., Rabanal, F.E., Fontanella, F.M., Sites Jr., J.W., Amphibian phylogeography in the Antipodes: Refugia and postglacial colonization explain mitochondrial haplotype distribution in the Patagonian frog Eupsophus calcaratus (Cycloramphidae) (2011) Mol. Phylogenet. Evol., 58, pp. 343-352
  • Padial, J.M., Castroviejo-Fisher, S., Köhler, J., Vilà, C., Chaparro, J.C., de la Riva, I., Deciphering the products of evolution at the species level: the need for an integrative taxonomy (2009) Zool. Scr., 38, pp. 431-447
  • Pol, D., Empirical problems of the hierarchical likelihood ratio test for model selection (2004) Syst. Biol., 53, pp. 949-962
  • Posada, D., jModelTest: Phylogenetic Model Averaging (2008) Mol. Biol. Evol., 25, pp. 1253-1256
  • Posada, D., Buckley, T.R., Model selection and model averaging in phylogenetics: advantages of Akaike information criterion and Bayesian approaches over likelihood ratio tests (2004) Syst. Biol., 53, pp. 793-808
  • Posada, D., Crandall, K.A., Modeltest: testing the model of DNA substitution (1998) Bioinformatics, 14, pp. 817-818
  • Pyron, R.A., Wiens, J.J., A large-scale phylogeny of Amphibia including over 2,800 species, and a revised classification of extant frogs, salamanders, and caecilians (2011) Mol. Phylogenet. Evol., 61, pp. 543-583
  • Rabassa, J., Coronato, A., Martínez, O., Late Cenozoic glaciations in Patagonia and Tierra del Fuego: an updated review (2011) Biol. J. Linn. Soc., 103, pp. 316-335
  • Roelants, K., Gower, D.J., Wilkinson, M., Loader, S.P., Biju, S.D., Guillaume, K., Moriau, L., Bossuyt, F., Global patterns of diversification in the history of modern amphibians (2007) Proc. Natl Acad. Sci. USA, 104, pp. 887-892
  • Schaeffer, B., Anurans from the Early Tertiary of Patagonia (1949) Bull. Amer. Mus. Nat. Hist., 93, pp. 41-68
  • Sérsic, A.N., Cosacov, A., Cocucci, A.A., Johnson, L.A., Pozner, R., Avila, L.J., Sites Jr., J.W., Morando, M., Emerging phylogeographical patterns of plants and terrestrial vertebrates from Patagonia (2011) Biol. J. Linn. Soc., 103, pp. 475-494
  • Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmoougin, F., Higgins, D.G., The Clustal X Windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools (1997) Nucleic Acids Res., 25, pp. 4876-4882
  • Úbeda, C.A., Eupsophus calcaratus (Anura, Leptodactylidae): ampliación de su distribución geográfica y hábitats en Argentina (2000) Cuad. Herpetol., 14, pp. 71-74
  • Úbeda, C.A., Nuñez, J.J., New parental care behaviours in two telmatobiine genera from temperate Patagonian forests: Batrachyla and Eupsophus (Anura: Leptodactylidae) (2006) Amphib-reptil., 27, pp. 441-444
  • Úbeda, C.A., Ramilo, E., Chehébar, C., Vidoz, F., Geographic distribution. Eupsophus emiliopugini (1999) Herpetol. Rev., 30, p. 230
  • Van Bocxlaer, I., Biju, S.D., Loader, S.P., Bossuyt, F., Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot (2009) BMC Evol. Biol., 9, p. 131
  • Varón, A., Vinh, L.S., Bomash, I., Wheeler, W.C., (2009), http://research.amnh.org/scicomp/projects/poy.php, POY 4.1.1. American Museum of Natural History; Varón, A., Vinh, L.S., Wheeler, W.C., POY version 4: phylogenetic analysis using dynamic homologies (2010) Cladistics, 26, pp. 72-85
  • Veloso, A., Díaz, N., Iturra, P., Penna, M., Descripción de una nueva especie de telmatobino del género Alsodes (Amphibia, Leptodactylidae) de la cordillera de Nahuelbuta (sur de Chile) (1981) Medio Ambiente, 5, pp. 72-77
  • Veloso, A., Celis-Diez, J.L., Guerrero, P.C., Méndez, M.A., Iturra, P., Simonetti, J.A., Description of a new Eupsophus species (Amphibia, Leptodactylidae) from the remnants of Maulino Forest, Central Chile (2005) Herpetol. J., 15, pp. 159-165
  • Vences, M., Thomas, M., Bonett, R.M., Vieites, D.R., Deciphering amphibian diversity through DNA barcoding: chances and challenges (2005) Philos. Trans. R. Soc. Lond. B Biol. Sci., 360, pp. 1859-1868
  • Vera Candioti, M.F., Úbeda, C., Lavilla, E.O., Morphology and metamorphosis of Eupsophus calcaratus tadpoles (Anura: Leptodactylidae) (2005) J. Morphol., 264, pp. 161-177
  • Vera Candioti, M.F., Nuñez, J.J., Úbeda, C., Development of the nidicolous tadpoles of Eupsophus emiliopugini (Anura: Cycloramphidae) until metamorphosis, with comments on systematic relationships of the species and its endotrophic developmental mode (2011) Acta Zool., 92, pp. 27-45
  • Villagrán, C., Historia de los bosques templados del sur de Chile durante el Tardiglacial y Postglacial (1991) Rev. Chil. Hist. Nat., 64, pp. 447-460
  • Wheeler, W.C., Optimization alignment: the end of multiple sequence alignment in phylogenetics? (1996) Cladistics, 12, pp. 1-9
  • Wheeler, W.C., Optimization alignment: down, up, error, and improvements (2002) Techniques in Molecular Systematics and Evolution, pp. 55-69. , De Salle, R., Giribet, G., Wheeler, W.C. (Eds), Birkhaüser Verlag, Bassel -
  • Wheeler, W.C., Iterative pass optimization of sequence data (2003) Cladistics, 19, pp. 254-260
  • Wheeler, W.C., Implied alignment: a synapomorphy-based multiple-sequence alignment method and its use in cladogram search (2003) Cladistics, 19, pp. 261-268
  • Wheeler, W.C., Aagesen, L., Arango, C., Faivovich, J., Grant, T., D'Haese, C., Janies, D., Giribet, G., (2006) Dynamic Homology and Phylogenetic Systematics: A Unified Approach Using POY, , American Museum of Natural History, New York
  • Wiens, J.J., Fetzner Jr., J.W., Parkinson, C.L., Reeder, T.W., Hylid frog phylogeny and sampling strategies for speciose clades (2005) Syst. Biol., 54, pp. 719-748

Citas:

---------- APA ----------
Blotto, B.L., Nuñez, J.J., Basso, N.G., Úbeda, C.A., Wheeler, W.C. & Faivovich, J. (2013) . Phylogenetic relationships of a Patagonian frog radiation, the Alsodes+Eupsophus clade (Anura: Alsodidae), with comments on the supposed paraphyly of Eupsophus. Cladistics, 29(2), 113-131.
http://dx.doi.org/10.1111/j.1096-0031.2012.00417.x
---------- CHICAGO ----------
Blotto, B.L., Nuñez, J.J., Basso, N.G., Úbeda, C.A., Wheeler, W.C., Faivovich, J. "Phylogenetic relationships of a Patagonian frog radiation, the Alsodes+Eupsophus clade (Anura: Alsodidae), with comments on the supposed paraphyly of Eupsophus" . Cladistics 29, no. 2 (2013) : 113-131.
http://dx.doi.org/10.1111/j.1096-0031.2012.00417.x
---------- MLA ----------
Blotto, B.L., Nuñez, J.J., Basso, N.G., Úbeda, C.A., Wheeler, W.C., Faivovich, J. "Phylogenetic relationships of a Patagonian frog radiation, the Alsodes+Eupsophus clade (Anura: Alsodidae), with comments on the supposed paraphyly of Eupsophus" . Cladistics, vol. 29, no. 2, 2013, pp. 113-131.
http://dx.doi.org/10.1111/j.1096-0031.2012.00417.x
---------- VANCOUVER ----------
Blotto, B.L., Nuñez, J.J., Basso, N.G., Úbeda, C.A., Wheeler, W.C., Faivovich, J. Phylogenetic relationships of a Patagonian frog radiation, the Alsodes+Eupsophus clade (Anura: Alsodidae), with comments on the supposed paraphyly of Eupsophus. Cladistics. 2013;29(2):113-131.
http://dx.doi.org/10.1111/j.1096-0031.2012.00417.x