Skip to main content Skip to main navigation menu Skip to site footer
Article
Published: 2020-05-19

Unraveling the species diversity and relationships in the Leptodactylus mystaceus complex (Anura: Leptodactylidae), with the description of three new Brazilian species

Centro de Ciências Exatas e da Natureza, Universidade Federal da Paraíba (UFPB), João Pessoa, Paraíba, Brazil
Centro de Ciências Exatas e da Natureza, Universidade Federal da Paraíba (UFPB), João Pessoa, Paraíba, Brazil
4Instituto de Ciências Biológicas e da Saúde, Universidade Federal de Viçosa (UFV), Florestal, Minas Gerais, Brazil 5
Instituto de Ciências Biológicas e da Saúde, Universidade Federal de Viçosa (UFV), Florestal, Minas Gerais, Brazil
Departamento de Botânica e Zoologia, Universidade Federal do Rio Grande do Norte (UFRN), Natal, Rio Grande do Norte, Brazil
8Departamento de Engenharia Florestal, Universidade de Brasília (UnB), Brasília, Distrito Federal, Brazil.
Departamento de Biodiversidade e Centro de Aquicultura, Instituto de Biociências, Universidade Estadual Paulista (UNESP), Rio Claro, São Paulo, Brazil.
Instituto de Ciências Exatas e Naturais do Pontal, Universidade Federal de Uberlândia (UFU), Ituiutaba, Brazil
9Departamento de Biodiversidade e Centro de Aquicultura, Instituto de Biociências, Universidade Estadual Paulista (UNESP), Rio Claro, São Paulo, Brazil. haddad1000@gmail.com; https://orcid.org/0000-0003-1057-5660
Amphibia Amazonia Atlantic Forest Bioacoustics Biodiversity Cerrado Cryptic diversity Dry Diagonal

Abstract

Members of the Leptodactylus mystaceus species complex are widely distributed in forests and open formations of South America east of the Andes. Species of the complex are morphologically similar or indistinguishable among each other, but acoustic data have been the cornerstone for species discrimination across their geographic ranges. In this paper, we re-examine the monophyly, species diversity, and relationships in the L. mystaceus complex on the basis of morphology, coloration, acoustics, and DNA sequences. Morphological and color patterns originally used to the allocation of species to the L. mystaceus complex are also reassessed. Our results revealed three new species from the Brazilian Atlantic Forest and Cerrado, which are named and described herein, based mainly on acoustic and molecular data. Populations assigned to the lineage widely distributed across the South American Dry Diagonal (DD), reported in this study as L. cf. mystaceus, is likely paraphyletic with respect to the nominal species (Amazonian lineage), but additional data are still needed to address the taxonomic status of the DD lineage.

 

References

  1. Andrade, F.S., Silva, L.A., Koroiva, R., Fadel, R.M. & Santana, D.J. (2019) A new species of Pseudopaludicola Miranda-Ribeiro, 1926 (Anura: Leptodactylidae: Leiuperinae) from an Amazonia-Cerrado transitional zone, state of Tocantins, Brazil. Journal of Herpetology, 53, 68–80.

    https://doi.org/10.1670/18-125

    Ahl, E. (1936) Zweif neue Froscharten der Gattung Leptodactylus aus Südamerika. Veröffentlichungen aus dem Deutschen Kolonial – und Übersee-Museum in Bremen, 1, 389–392.

    Audacity Team (2017) Audacity(R): free audio editor and recorder. Version 2.2.1. Available from: https://www.audacityteam.org (accessed 5 May 2018)

    Barrio, A. (1965a) El género Physalaemus (Anura, Leptodactylidae) en la Argentina. Physis, 25, 421–444.

    Barrio, A. (1965b) Afinidades del canto nupcial de las especies cavícolas del género Leptodactylus (Anura—Leptodactylidae). Physis, 70, 401–410.

    Bilate, M., Wogel, H., Weber, L.N. & Abrunhosa, P.A. (2006) Vocalizações e girino de Leptodactylus spixi Heyer, 1983 (Amphibia, Anura, Leptodactylidae). Arquivos do Museu Nacional, 64, 235–245.

    Bokermann, W.C.A. (1969) Uma nova espécie de Leptodactylus de Mato Grosso (Anura, Leptodactylidae). Revista Brasileira de Biologia, 29, 13–16.

    Boulenger, G.A. (1894) List of reptiles and batrachians collected by Dr. J. Bohls near Asuncion, Paraguay. Annals and Magazine of Natural History, 13, 342–348.

    https://doi.org/10.1080/00222939408677709

    Boulenger, G.A. (1902) Descriptions of new batrachians and reptiles from north western Ecuador. Annals and Magazine of Natural History, 9, 51–57.

    https://doi.org/10.1080/00222930208678538

    Boulenger, G.A. (1918) Descriptions of new South American batrachians. Annals and Magazine of Natural History, 2, 427–433.

    https://doi.org/10.1080/00222931808673782

    Brandão, R.A., Álvares, G.F.R. & de Sá, R.O. (2013) The advertisement call of the poorly known Leptodactylus tapiti (Anura, Leptodactylidae). Zootaxa, 3616, 284–286.

    https://doi.org/10.11646/zootaxa.3616.3.6

    Brocchi, P. (1877) Sur quelques batraciens raniformes et bufoniformes de l’Amérique Centrale. Bulletin de la Société Philomathique de Paris, Series 7, 1, 175–197.

    Bruford, M.W., Hanotte, O., Brookfield, J.F.Y. & Burke, T. (1992) Single and multilocus DNA fingerprinting. In: Hoelzel, A.R. (Ed.) Molecular Genetic Analysis of Populations: A Practical Approach. I.R.L. Press, Oxford, pp. 225–269.

    Burmeister, H. (1861) Reise durch die La Plata-Staaten mit besonderer Rücksicht auf die Physische Beschaffenheit und den Culturzustand der Argentinische Republik. Ausgefuhrt in den Jahren 1857, 1858, 1859 un 1860. Vol. 2. H. M. Schmidt, Halle, 538 pp.

    Caramaschi, U., Feio, R.N. & São Pedro, V.A. (2008) A new species of Leptodactylus Fitzinger (Anura, Leptodactylidae) from Serra do Brigadeiro, state of Minas Gerais, southeastern Brazil. Zootaxa, 1861 (1), 44–54.

    https://doi.org/10.11646/zootaxa.1861.1.4

    Carvalho, T.R. & Ron, S.R. (2011) Advertisement call of Leptodactylus labrosus Jiménez de la Espada, 1875 (Anura, Leptodactylidae): an unusual advertisement call within the L. fuscus group. Herpetology Notes, 4, 325–326.

    Carvalho, T.R., Giaretta, A.A., Angulo, A., Haddad, C.F.B. & Peloso, P.L.V. (2019) A new Amazonian species of Adenomera (Anura: Leptodactylidae) from the Brazilian state of Pará: a tody-tyrant voice in a frog. American Museum Novitates, 3919, 1–21.

    https://doi.org/10.1206/3919.1

    Carvalho, T.R., Leite, F.S.F. & Pezzuti, T.L. (2013) A new species of Leptodactylus Fitzinger (Anura, Leptodactylidae, Leptodactylinae) from montane rock fields of the Chapada Diamantina, northeastern Brazil. Zootaxa, 3701 (3), 349–364.

    https://doi.org/10.11646/zootaxa.3701.3.5

    Cassini, C.S., Orrico, V.G.D., Dias, I.R., Solé, M. & Haddad, C.F.B. (2013) Phenotypic variation of Leptodactylus cupreus Caramaschi, São-Pedro & Feio 2008 (Anura, Leptodactylidae). Zootaxa, 3616 (1), 73–84.

    https://doi.org/10.11646/zootaxa.3616.1.6

    Cei, J.M. (1950) Leptodactylus chaquensis n. sp. y el valor sistematico real de la especie Linneana Leptodactylus ocellatus en la Argentina. Acta Zoologica Lilloana, 9, 395–423.

    Cope, E.D. (1862) On some new and little known American Anura. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 151–159.

    Costa-Reis, R. & Genovez, P.F. (2013) Território sagrado: exílio, diáspora e reconquista Krenak no vale do Rio Doce, Resplendor, MG. Boletim Goiano de Geografia, 33, 1–15.

    https://doi.org/10.5216/bgg.v33i1.23628

    Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772.

    https://doi.org/10.1038/nmeth.2109

    de Sá, R.O., Grant, T., Camargo, A., Heyer, W.R., Ponssa, M.L. & Stanley, E. (2014) Systematics of the Neotropical genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the relevance of non-molecular evidence, and species accounts. South American Journal of Herpetology, 9, S1–S128.

    https://doi.org/10.2994/SAJH-D-13-00022.1

    de Sá, R.O., Heyer, W.R. & Camargo, A. (2005) Are Leptodactylus didymus and L. mystaceus phylogenetically sibling species (Amphibia, Anura, Leptodactylidae)? Herpetologica Petropolitana, Russian Journal of Herpetology, 12, 90–92.

    Dória, T.A.F., Canedo, C.C. & Napoli, M.F. (2018) Processes influencing anuran coexistence on a local scale: A phylogenetic and ecological analysis in a restinga environment. South American Journal of Herpetology, 13, 183–201.

    https://doi.org/10.2994/SAJH-D-17-00044.1

    Duellman, W.E. (1970) The Hylid Frogs of Middle America. Monographs of the Museum of Natural History of the University of Kansas, Lawrence, 753 pp.

    Duméril, A.M.C. & Bibron, G. (1840) Plate. In: D’Orbigny, A. & Bibron, G. (Eds.), Voyage dans l’Amérique Méridionale (Le Brésil, La République Orientale de l’Uruguay, La République Argentine, La Patagonie, La Republique du Chili, La Republique de Bolivia, La République du Pérou), exécuté pendant las annees 1826, 1827, 1828, 1829, 1830, 1831, 1832 et 1833. Vol. 5. Part 1. M. le Ministre de l’Instruction Publique, Paris and Strasbourg, pl. 13.

    Ferreira, R.B., Mônico, A.T., Silva, E.T., Lirio, F.C.F., Zocca, C., Mageski, M.M., Tonini, J.F.R., Beard, K.H., Duca, C. & Silva-Soares, T. (2019) Amphibians of Santa Teresa, Brazil: the hotspot further evaluated. ZooKeys, 857, 139–162.

    https://doi.org/10.3897/zookeys.857.30302

    Giaretta, A.A. & Costa, H.C.M. (2007) A redescription of Leptodactylus jolyi Sazima and Bokermann (Anura, Leptodactylidae) and the recognition of a new closely related species. Zootaxa, 1608 (1), 1–10.

    https://doi.org/10.11646/zootaxa.1608.1.1

    Gridi-Papp, M. (2007) SoundRuler: Acoustic Analysis for Research and Teaching. Available from: http://soundruler.sourceforge.net (accessed 5 May 2018)

    Gualberto, R.D. (2017) O lugar contado: Narrativas em torno à Chapada dos Negros em Arraias—TO. Dissertação (Mestrado em Estudos de Cultura e Território)—Universidade Federal do Tocantins, Programa de Pós-Graduação em Estudos de Cultura e Território, Araguaína, 125 pp.

    Guimarães, C.S., Assis, C.L., Thomassen, H., Leite, F.S.F. & Feio, R.N. (2019) Anfíbios do Parque Estadual do Rio Doce, Minas Gerais. Universidade Federal de Viçosa, Departamento de Biologia Animal, Museu de Zoologia João Moojen, Viçosa, Minas Gerais, 57 pp.

    Günther, A.C.L.G. (1859) On the reptiles from St. Croix, West Indies, collected by Messrs. A. and E. Newton. Annals and Magazine of Natural History, 3, 209–217.

    https://doi.org/10.1080/00222935908697110

    Heyer, W.R. (1969) The adaptive ecology of the species groups of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Evolution, 23, 421–428.

    https://doi.org/10.1111/j.1558-5646.1969.tb03525.x

    Heyer, W.R. (1978) Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Science Bulletin, Natural History Museum of Los Angeles County, 29, 1–85.

    Heyer, W.R. (1979) Systematics of the pentadactylus species group of the frog genus Leptodactylus (Amphibia: Leptodactylidae). Smithsonian Contributions to Zoology, 301, 1–43.

    https://doi.org/10.5479/si.00810282.301

    Heyer, W.R. (1983) Clarification of the names Rana mystacea Spix, 1824, Leptodactylus amazonicus Heyer, 1978, and a description of a new species, Leptodactylus spixi (Amphibia: Leptodactylidae). Proceedings of the Biological Society of Washington, 96, 270–272.

    Heyer, W.R. (1994) Variation within the Leptodactylus podicipinus–wagneri complex of frogs (Amphibia: Leptodactylidae). Smithsonian Contributions to Zoology, 546, 1–124.

    https://doi.org/10.5479/si.00810282.546.i

    Heyer, W.R. (2000) The lectotype of Cystignathus hylodes Reinhardt and Lütken. Journal of Herpetology, 34, 150–153.

    https://doi.org/10.2307/1565254

    Heyer, W.R. (2005) Variation and taxonomic clarification of the large species of the Leptodactylus pentadactylus species group (Amphibia: Leptodactylidae) from Middle America, northern South America, and Amazonia. Arquivos de Zoologia, 37, 269–348.

    https://doi.org/10.11606/issn.2176-7793.v37i3p269-348

    Heyer, W.R. & Crombie, I. (2005) Leptodactylus lauramiriamae, a distinctive new species of frog (Amphibia: Anura: Leptodactylidae) from Rondônia, Brazil. Proceedings of the Biological Society of Washington, 118, 590–595.

    https://doi.org/10.2988/0006-324X(2005)118[590:LLADNS]2.0.CO;2

    Heyer, W.R. & de Sá, R.O. (2011) Variation, systematics, and relationships of the Leptodactylus bolivianus complex (Amphibia: Anura: Leptodactylidae). Smithsonian Contributions to Zoology, 635, 1–58.

    https://doi.org/10.5479/si.00810282.635.1

    Heyer, W.R. & Juncá, F.A. (2003) Leptodactylus caatingae, a new species of frog from eastern Brazil (Amphibia: Anura: Leptodactylidae). Proceedings of the Biological Society of Washington, 116, 317–329.

    Heyer, W.R. & Scott Jr., N.J. (2006) The advertisement call of Leptodactylus laticeps (Amphibia, Anura, Leptodactylidae): Predatory aural luring? Herpetological Natural History, 9, 189–194.

    Heyer, W.R. & Thompson, A.S. (2000) Leptodactylus rugosus Noble. Catalogue of American Amphibians and Reptiles, 705, 1–5.

    Heyer, W.R., García-Lopez, J.M. & Cardoso, A.J. (1996) Advertisement call variation in the Leptodactylus mystaceus species complex (Amphibia: Leptodactylidae) with a description of a new sibling species. Amphibia-Reptilia, 17, 7–31.

    https://doi.org/10.1163/156853896X00252

    Heyer, W.R., Heyer, M.M. & de Sá, R.O. (2006) Leptodactylus fragilis. Catalogue of American Amphibians and Reptiles, 830, 1–26.

    Heyer, W.R., Heyer, M.M. & de Sá, R.O. (2010) Leptodactylus syphax Bokermann. Catalogue of American Amphibians and Reptiles, 868, 1–9.

    Heyer, W.R., Rand A.S., Cruz, C.A.G., Peixoto, O.L. & Nelson, C.E. (1990) Frogs of Boracéia. Arquivos de Zoologia, 31, 231–410.

    Izecksohn, E. (1976) Uma nova espécie de Leptodactylus do Estado do Rio de Janeiro, Brasil (Amphibia, Anura, Leptodactylidae). Revista Brasileira de Biologia, 36, 527–530.

    Jansen, M., Bloch, R., Schulze, A. & Pfenninger, M. (2011) Integrative inventory of Bolivia’s lowland anurans reveals hidden diversity. Zoologica Scripta, 40, 567–583.

    https://doi.org/10.1111/j.1463-6409.2011.00498.x

    Jiménez de la Espada, M. (1875) Vertebrados del Viaje al Pacífico Verificado de 1862 a 1865 por una Comisión de Naturalistas Enviada por el Gobierno Español. Batrachios. A. Miguel Ginesta, Madrid, 208 pp.

    https://doi.org/10.5962/bhl.title.5769

    Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30, 3059–3066.

    https://doi.org/10.1093/nar/gkf436

    Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.

    https://doi.org/10.1093/bioinformatics/bts199

    Ligges, U., Krey, S., Mersmann, O. & Schnackenberg, S. (2017) tuneR: analysis of music and speech. Available from https://CRAN.R-project.org/package=tuneR (accessed 20 September 2018)

    Lutz, A. (1926) Observações sobre batrachios brasileiros/Observations on Brazilian batrachians. Memórias do Instituto Oswaldo Cruz, 19, 139–174.

    https://doi.org/10.1590/S0074-02761926000200001

    Matlab (2004) The language of technical computing. Version 6.5.2. MathWorks, Inc., Natick. [program]

    Nascimento, J., Lima, J.D., Suárez, P., Baldo, D., Andrade, G.V., Pierson, T.W., Fitzpatrick, B.M., Haddad, C.F.B., Recco-Pimentel, S.M. & Lourenço, L.B. (2019) Extensive cryptic diversity within the Physalaemus cuvieri-Physalaemus ephippifer species complex (Amphibia, Anura) revealed by cytogenetic, mitochondrial, and genomic markers. Frontiers in Genetics, 10, e719.

    https://doi.org/10.3389/fgene.2019.00719

    Palumbi, S.R. (1996) Nucleic acids II: The polymerase chain reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular Systematics. Sinauer, Sunderland, pp. 205–247.

    Pereira-Ribeiro, J., Ferreguetti, A.C., Linause, T.M., Cozer, J.S., Bergallo, H.G. & Rocha, C.F.D. (2019) Diversity and distribution of anurans from Mata das Flores State Park, Espírito Santo, southeastern Brazil. Oecologia Australis, 23, 292–300.

    https://doi.org/10.4257/oeco.2019.2302.08

    Pons, J., Barraclough, T.G., Gomez-Zurita, J., Cardoso, A., Duran, D.P., Hazell, S., Kamoun, S., Sumlin, W.D. & Vogler, A.P. (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology, 55, 595–609.

    https://doi.org/10.1080/10635150600852011

    Ponssa, M.L. (2008) Cladistic analysis and osteological descriptions of the frog species in the Leptodactylus fuscus species group (Anura, Leptodactylidae). Journal of Zoological Systematics and Evolutionary Research, 46, 249–266.

    https://doi.org/10.1111/j.1439-0469.2008.00460.x

    Puillandre, N., Lambert, A., Brouillet, S. & Achaz, G. (2011) ABGD, automatic barcode gap discovery for primary species delimitation. Molecular Ecology, 21, 1864–1877.

    https://doi.org/10.1111/j.1365-294X.2011.05239.x

    R Core Team (2018) R: a language and environment for statistical computing, version 3.5.0. R Foundation for Statistical Computing, Vienna, Austria. Available from http://www.r-project.org (accessed 5 May 2018)

    Rambaut, A. (2014) FigTree: a graphical viewer of phylogenetic trees. Version 1.4.2. Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed 5 May 2018)

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer. Version 1.7. Available from: http://beast.community/tracer (accessed 5 May 2018)

    Reid, N. & Carstens, B. (2012) Phylogenetic estimation error can decrease the accuracy of species delimitation: a Bayesian implementation of the general mixed Yule-coalescent model. BMC Evolutionary Biology, 12, 1–11.

    https://doi.org/10.1186/1471-2148-12-196

    Reinhardt, J.T. & Lütken, C.F. (1862) Bidrag til Kundskab om Brasiliens Padder og Krybyr. Förste Afdeling: Padderne og Öglerne. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjøbenhavn, 2, 143–242.

    Rödder, D., Teixeira, R.L., Ferreira, R.B., Dantas, R.B., Pertel, W. & Guarniere, G.J. (2007) Anuran hotspots: the municipality of Santa Teresa, Espírito Santo, southeastern Brazil. Salamandra, 43, 91–110.

    Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an online reference. American Society of Ichthyologists and Herpetologists, Washington, DC. Available from http://www.asih.org/ (accessed 16 August 2016)

    Santana, D.J., São Pedro, V.A., Hote, P.S., Roberti, H.M., Sant’Anna, A.C., Figueiredo-de-Andrade, C.A. & Feio, R.N. (2010) Anurans in the region of the High Muriaé River, state of Minas Gerais, Brazil. Herpetology Notes, 3, 1–10.

    Santos, M.T.T., Oliveira, S.H., Carvalho, T.R., Zaidan, B.F., Silva, N.R., Berneck, B.M. & Garcia, P.C.A. (2019) A new species of Paratelmatobius (Anura: Leptodactylidae: Paratelmatobiinae) from the Atlantic Forest of southern Brazil. Zootaxa, 4648 (3), 473–493.

    https://doi.org/10.11646/zootaxa.4648.3.4

    Sazima, I. & Bokermann, W.C.A. (1978) Cinco novas espécies de Leptodactylus do centro e sudeste Brasileiro (Amphibia, Anura, Leptodactylidae). Revista Brasileira de Biologia, 38, 899–912.

    Schneider, J.G. (1799) Historia Amphibiorum naturalis et literariae. Impressus Ieanae [Jena], Friederici Frommanni, 264 pp.

    https://doi.org/10.5962/bhl.title.78757

    Schneider, R.G., Cardozo, D.E., Brusquetti, F., Kolenc, F., Borteiro, C., Haddad, C., Basso, N.G. & Baldo, D. (2019) A new frog of the Leptodactylus fuscus species group (Anura: Leptodactylidae), endemic from the South American Gran Chaco. PeerJ, 7, 1–29.

    https://doi.org/10.7717/peerj.7869

    Silva, L.A., Carvalho, P.S., Pereira, E.A., Fadel, R.M., Dantas, S.P., Brandão, R.A., Santana, D.J. (2020) Richness, diversity patterns, and taxonomic notes of amphibians from the Tocantins state. Biota Neotropica, 20, 1–22.

    https://doi.org/10.1590/1676-0611-bn-2019-0838

    Silva-Soares, T. & Scherrer, P.V. (2013) Amphibians of Parque Estadual do Forno Grande, State of Espírito Santo, Southeastern Brazil: Species composition and conservation. North-Western Journal of Zoology, 9, 113–120.

    Spix, J.B. (1824) Animalia nova sive species novae Testudineum et Ranarum, quas in itenere per Brasiliam, annis 1817–1820. F. S. Hübschmann, Munich, 53 pp.

    https://doi.org/10.5962/bhl.title.3665

    Suchard, M.A., Lemey, P., Baele, G., Ayres, D.L., Drummond, A.J. & Rambaut, A. (2018) Bayesian phylogenetic and phylodynamic data integration using BEAST 1.10. Virus Evolution, 4, 1–5.

    https://doi.org/10.1093/ve/vey016

    Sueur, J., Aubin, T. & Simonis, C. (2008) Seewave: a free modular tool for sound analysis and synthesis. Bioacoustics, 18, 213–226.

    https://doi.org/10.1080/09524622.2008.9753600

    Tamura, K. & Nei, M. (1993) Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and Evolution, 10, 512–526.

    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: molecular evolutionary genetics analysis, version 6.0. Molecular Biology and Evolution, 30, 2725–2729.

    https://doi.org/10.1093/molbev/mst197

    Werneck, F.P. (2011) The diversification of eastern South American open vegetation biomes: Historical biogeography and perspectives. Quaternary Science Reviews, 30, 1630–1648.

    https://doi.org/10.1016/j.quascirev.2011.03.009