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Type: Article
Published: 2018-01-25
Page range: 511–536
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Diversity of Tropidurus (Squamata: Tropiduridae) in Paraguay—an integrative taxonomic approach based on morphological and molecular genetic evidence

Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Senckenberganlage 25, 60325 Frankfurt a.M., Germany Goethe-University, Institute for Ecology, Evolution & Diversity, Biologicum, Building C, Max-von-Laue-Str. 13, 60438 Frankfurt am Main, Germany Instituto de Investigación Biológica del Paraguay, Del Escudo 1607, 1425 Asunción, Paraguay
Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Senckenberganlage 25, 60325 Frankfurt a.M., Germany Goethe-University, Institute for Ecology, Evolution & Diversity, Biologicum, Building C, Max-von-Laue-Str. 13, 60438 Frankfurt am Main, Germany
Reptilia Iguanoidea 16S COI PRLR South America ring species

Abstract

Tropidurus is a Neotropical genus of iguanoid lizards characterized by a conspicuously enlarged interparietal plate, the presence of gular folds, presence of infradigital keels, and the absence of femoral pores. Currently, 29 species are recognized within the genus, seven of which are present in Paraguay: T. etheridgei, T. torquatus, T. guarani, T. lagunablanca, T. spinulosus, T. tarara, and T. teyumirim. We generated genetic data based on two DNA mitochondrial markers (16S and COI) and one nuclear (PRLR) marker for all the seven Paraguayan species with the goal to identify the taxonomic relationships among taxa based on the intra- and interspecific genetic variation and the construction of molecular clusters. ML and BI analyses match in the recognition of two main clusters: groups torquatus and spinulosus, and within the torquatus group the differentiation between T. catalanensis and T. etheridgei is highly supported. Nevertheless, there is a complete lack of congruence between mitochondrial and nuclear genes in the topology within the spinulosus group. Tropidurus guarani and T. spinulosus are more differentiated from the remaining species of the spinulosus group with genetic p-distances from 4.0 to 6.0. Low distances were found between T. lagunablanca and T. tarara (1.0–1.1%), and slightly higher, among T. teyumirim, T. lagunablanca, and T. tarara (2.0–2.6% respectively). From a morphological perspective, species of the Tropidurus torquatus group are easily distinguished; but we found strong overlaps of scalation characters in the spinulosus group. We interpret the low genetic distances documented among the nominal taxa Tropidurus lagunablanca, T. tarara, and T. teyumirim as evidence for conspecificity. This hypothesis is supported by the lack of morphological characters that would diagnose any of the three taxa. Similarly, we found low genetic distances among populations assigned to the nominal taxa T. guarani and T. spinulosus, including samples from near the type locality of the former, and therefore we recognize only two species of the T. spinulosus complex in Paraguay: T. spinulosus and T. lagunablanca.

 

References

  1. Alcaide, M., Scordato, E.S.C., Price, T.D. & Irwin, D.E. (2014) Genomic divergence in a ring species complex. Nature, 511, 83–85.
    https://doi.org/10.1038/nature13285

    Álvarez, B.B., Cei, J.M. & Scolaro, J.A. (1994) A new subspecies of Tropidurus spinulosus (Cope 1862) from the subtropical wet mesic Paraguayan region (Reptilia Squamata Tropiduridae). Tropical Zoology, 7, 161–179.
    https://doi.org/10.1080/03946975.1994.10539249

    Álvarez, B.B., Lions, M.L., Aguirre, R., Céspedez, J. & Hernando, A. (1995) Herpetofauna del área de influencia del embalse de la represa Yacyretá (Argentina-Paraguay). Facena, 11, 57–73. Available from: http://exa.unne.edu.ar/biologia/herpetologia/public_html/PDF/Herpetofauna%20de%20Yacyreta.pdf (Accessed 25 Jan. 2018)

    Álvarez, B.B., Aguuirre, R.H., Céspedez, J.A., Hernando, A.B. & Tedesco, M.E. (2002) Atlas de los Anfibios y Reptiles de las Provincias de Corrientes, Chaco y Formosa, Argentina. I: Anuros, Cecilidos, Saurios, Anfisbenidos y Serpientes. Universidad Nacional del Nordeste, Corrientes, 160 pp.

    Anisimova, M., Gil, M., Dufayard, J.F., Dessimoz, C. & Gascuel, O. (2011) Survey of branch support methods demonstrates accuracy, power, and robustness of fast Likelihood-based approximation schemes. Systematic Biology, 60, 685–699.
    https://doi.org/10.1093/sysbio/syr041

    Aquino, A.L., Scott, N. & Motte, M. (1996) Lista de los anfibios y reptiles del Museo Nacional de Historia Natural del Paraguay (marzo, 1980–setiembre, 1995). In: Romero, O. (Ed.), Colecciones de Fauna y Flora del Museo Nacional de Historia Natural del Paraguay. Museo Nacional de Historia Natural del Paraguay, San Lorenzo, pp. 331–400.

    Aquino-Shuster, A.L., Motte, M. & Sequera, G. (1991) Relación del indígena Chamacoco con la herpetofauna del Alto Paraguay. Boletín del Museo Nacional de Historia Natural del Paraguay, 10, 11–22. [http://www.seam.gov.py/content/bolet%C3%ADn-del-museo-nacional-de-historia-natural-del-paraguay-199110]

    Avila, L.J. & Carrizo, G.R. (2003) Lista comentada y distribución geográfica de la herpetofauna de la provincia de San Luis, Argentina. Acta Zoologica Lilloana, 47, 93–115.

    Batista, A., Veseley, M., Mebert, K., Lotzkat, S. & Köhler, G. (2015) A new species of Dactyloa from eastern Panama, with comments on other Dactyloa species present in the region. Zootaxa, 4039 (1), 57–84.
    https://doi.org/10.11646/zootaxa.4039.1.2

    Bertoni, A. de W. (1914) Fauna paraguaya: Catálogos sistemáticos de los vertebrados del Paraguay. Descripción Física y Económica del Paraguay, 59, 1–83.

    Bertoni, A. de W. (1939) Catálogos sistemáticos de los vertebrados de Paraguay. Revista de la Sociedad Científica del Paraguay, 4, 3–60.

    Boettger, O. (1885) Liste von Reptilien und Batrachiern aus Paraguay. Zeitschrift für Naturwissenschaft, 58, 213–248. Available from: http://www.biodiversitylibrary.org/page/31401231#page/245/mode/1up (Accessed 25 Jan. 2018)

    Boulenger, G.A. (1885) Catalogue of the lizards in the British Museum (Natural History). 2nd edition. Vol. 2. Taylor and Francis, London, 497 pp. Available from: http://www.biodiversitylibrary.org/item/61183#page/7/mode/1up (Accessed 25 Jan. 2018)

    Boulenger, G.A. (1894) List of reptiles and batrachians collected by Dr. J. Bohls near Asunción, Paraguay. Annals and Magazine of Natural History, 13, 342–348.
    https://doi.org/10.1080/00222939408677709

    Boulenger, G.A. (1898) A list of reptiles, batrachians, and fishes collected by Cav. Guido Boggiani in the northern Chaco. Annali del Museo Civico di Storia Naturale di Genova, 19, 125–127.

    Brown, W.M., George, M. & Wilson, A.C. (1979) Rapid evolution of animal DNA. Proceedings of the National Academy of Sciences, 76, 1967–1971.
    https://doi.org/10.1080/00222939408677709

    Burt, C.E. & Burt, M.D. (1933) A preliminary checklist of the lizards of South America. Transactions of the Academy of Science of St. Louis, 28, 1–104.

    Cabral, H. & Weiler, A. (2014) Lista comentada de los reptiles de la Colección Zoológica de la Facultad de Ciencias Exactas y Naturales de Asunción, Paraguay. Cuadernos de Herpetología, 28, 19–28.

    Cacciali, P., Scott, N., Aquino, A.L., Fitzgerald, L.A. & Smith, P. (2016) The reptiles of Paraguay: literature, distribution, and an annotated taxonomic checklist. Special Publications of the Museum of Southwestern Biology, 11, 1–373.

    Cain, A.J. (1954) Animal Species and their Evolution. Hutchinson University Library, London, 210 pp.

    Carreira, S., Meneghel, M. & Achaval, F. (2005) Reptiles de Uruguay. Universidad de la República, Montevideo, 639 pp.

    Carvalho, A.L.G. (2013) On the distribution and conservation of the South American lizard genus Tropidurus Wied-Neuwied, 1825 (Squamata: Tropiduridae). Zootaxa, 3640 (1), 42–56.
    https://doi.org/10.11646/zootaxa.3640.1.3

    Carvalho, A.L.G. (2016) Three new species of the Tropidurus spinulosus group (Squamata: Tropiduridae) from Eastern Paraguay. American Museum Novitates, 3853, 1–44.
    https://doi.org/10.1206/3853.1

    Carvalho, A.L.G., Sena, M.A., Peloso, P.L.V., Machado, F.A., Montesinos, R., Silva, H.R., Campbell, G. & Rodrigues, M.T. (2016) A new Tropidurus (Tropiduridae) from the semiarid Brazilian Caatinga: evidence for conflicting signal between mitochondrial and nuclear loci affecting the phylogenetic reconstruction of South American collared lizards. American Museum Noviates, 3852, 1–66.
    http://hdl.handle.net/2246/6637

    Cei, J.M. (1982) A new species of Tropidurus (Sauria, Iguanidae) from the arid Chacoan and western regions of Argentina. Occasional Papers of the Museum of Natural History University of Kansas, 97, 1–10

    Cei, J.M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Museo Regionale Sci. Naturale Torino, Monografie, 14, 1–949.

    Céspedez, J.A., Lions, M.L., Álvarez, B.B. & Schaefer, E.F. (2001) Inventario de anfibios y reptiles del Parque Nacional Chaco, Argentina. Natura Neotropicalis, 32, 163–169.

    Cochran, D.M. (1961) Type specimens of reptiles and amphibians in the United States National Museum. United States National Museum Bulletin, 220, 1–291.
    https://doi.org/10.5962/bhl.part.26967

    Cope, E.D. (1862) Catalogue of the reptiles obtained during the exploration of the Parana, Paraguay, Vermejo and Uruguay rivers, by Capt. Thos. J. Page, U. S. N.; and of those procured by Lieut. N. Michler, U. S. Top. Eng., commander of the expedition conducting the survey of the Atrato River. Proceedings of the Academy of Natural Science of Philadelphia, 14, 346–359. Available from: http://www.biodiversitylibrary.org/page/1951818#page/358/mode/1up (Accessed 25 Jan. 2018)

    Cruz, F.B., Teisaire, E. & Nieto, L. (1997) Reproductive biology of the lizard Tropidurus spinulosus in the Chaco of Salta, Argentina. Studies on Neotropical Fauna and Environment, 32, 28–32.
    https://doi.org/10.1076/snfe.32.1.28.13465

    Duré Rodas, A. (1995) Estudio de reptiles y anfibios. Biota, 2, 20–24.

    Elter, O. (1981) Cataloghi. V - La collezione erpetologica del Museo di Zoologia dell’Università di Torino. Museo Regionale di Scienze Naturali. Torino, 116 pp.

    Frost, D.R. (1992) Phylogenetic analysis and taxonomy of the Tropidurus group of lizards (Iguania: Tropiduridae). American Museum Novitates, 3033, 1–68.

    Frost, D.R. & Hills, D.M. (1990) Species in concept and practice: herpetological applications. Herpetologica, 46, 87–104.

    Frost, D.R., Crafts, H.M., Fitzgerald, L.A. & Titus, T.A. (1998) Geographic variation, species recognition, and molecular evolution of Cytochrome Oxidase I in the Tropidurus spinulosus complex (Iguania: Tropiduridae). Copeia, 1998, 839–851.
    https://doi.org/10.2307/1447331

    Frost, D.R., Rodrigues, M.T., Grant, T. & Titus, T.A. (2001) Phylogenetics of the lizard genus Tropidurus (Squamata: Tropiduridae: Tropidurinae): direct optimization, descriptive efficiency, and sensitivity analysis of congruence between molecular data and morphology. Molecular Phylogenetics and Evolution, 21, 352–371.
    https://doi.org/10.1006/mpev.2001.1015

    Gallardo, J.M. (1969) Las especies de saurios (Reptilia) de la Provincia de Santa Fé, Argentina, y consideraciones sobre su ecología y zoogeografía. Neotropica, 15, 73–81.

    Gallardo, J.M., Tio Vallejo, M. & Miranda, M.E. (1985) Estudio sobre la distribución de los saurios de Santiago del Estero, República Argentina (Reptilia: Sauria). Historia Natural, 5, 97–103.

    Giraudo, A.R. & Contreras, A.O. (1994) Lista preliminar de los reptiles registrados en el Departmento de Ñeembucú, Paraguay. Boletín de la Asociación Herpetológica Argentina, 10, 1–4.

    Glez-Peña D., Gómez-Blanco. D., Reboiro-Jato, M., Fdez-Riverola, F. & Posada, D. (2010) ALTER: program-oriented conversion of DNA and protein alignments. Nucleic Acid Research, 38, 14–18.
    https://doi.org/10.1093/nar/gkq321

    Gudynas, E. & Skuk, G. (1983) A new species of the iguanid lizard genus Tropidurus from temperate South America (Lacertilia: Iguanidae). Contribuciones en Biología del Centro Educativo Don Orione, 10, 1–10.

    Harvey, M.B. & Gutberlet, R.L. (1998) Lizards of the genus Tropidurus (Iguania: Tropiduridae) from the Serranía de Huanchaca, Bolivia: new species, natural history, and a key to the genus. Herpetologica, 54, 493–520.

    Hellmich, W. (1960) Die Sauria des Gran Chaco und seiner Randgebiete. Bayerische Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, 101, 1–131.

    Huelsenbeck, J.P., Ronquist, F. (2001) Mrbayes: bayesian inference of phylogeny. Bioinformatics, 17, 754–755.
    https://doi.org/10.1093/bioinformatics/17.8.754

    Irwin, D.E., Irwin, J.H. & Price, T.D. (2001) Ring species as bridges between microevolution and speciation. Genetica, 112–113, 223–243.
    https://doi.org/10.1023/A:1013319217703

    Irwin, D.E., Alcaide, M., Delmore, K.E., Irwin, J.H. & Owens, G.L. (2016) Recurrent selection explains parallel evolution of genomic regions of high relative but low absolute differentiation in a ring species. Molecular Ecology, 25, 4488–4507. https://doi.org/10.1111/mec.13792

    Ivanova, N.V., Dewaard, J.R. & Hebert, P.D. (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Notes, 6, 998–1002.
    https://doi.org/10.1111/j.1471-8286.2006.01428.x

    Kacoliris, F.P., Berkunsky, I. & Williams, J. (2006) Herpetofauna of the Argentinean Impenetrable Great Chaco. Phyllomedusa, 5, 149–157. https://doi.org/10.11606/issn.2316-9079.v5i2p149-157

    Kalyaanamoorthy, S., Minh, B.Q.,Wong, T.K.F., Haeseler, A. & Jermiin, L.S. (2016) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589.
    https://doi.org/10.1038/nmeth.4285

    Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.
    https://doi.org/10.1093/molbev/mst010

    Katoh, K. & Toh, H. (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics, 9, 286–298.
    https://doi.org/10.1093/bib/bbn013

    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

    Köhler, G., Batista, A., Veseley, M., Ponce, M., Carrizo, A. & Lotzkat, S. (2012) Evidence for the recognition of two species of Anolis formerly referred to as A. tropidogaster (Squamata: Dactyloidae). Zootaxa, 3348, 1–23.

    Köhler, G., Townsend, J.H. & Petersen, C.B. (2016) A taxonomic revision of the Norops tropidonotus complex (Squamata, Dactyloidae), with the resurrection of N. spilorhipis (Álvarez del Toro and Smith, 1956) and the description of two new species. Mesoamerican Herpetology, 3, 8–41.

    Koslowsky, J. (1898) Enumeración sistemática y distribución geográfica de los reptiles argentinos. Revista del Museo de La Plata, 8, 161–200.

    Kuntz, T.S. & Borges-Martins, M. (2013) A new microendemic species of Tropidurus (Squamata: Tropiduridae) from southern Brazil and revalidation of Tropidurus catalanensis Gudynas & Skuk, 1983. Zootaxa, 3681 (4), 413–439.
    https://doi.org/10.11646/zootaxa.3681.4.6

    Lions, M.L. & Álvarez, B.B. (1998) Desarrollo del esqueleto de Tropidurus etheridgei (Iguania: Tropiduridae). Revista Española de Herpetología, 12, 7–18.

    Lopez, J.V., Cilver, M., Stephens, J.C., Johnson, W.E. & O’Brien, S.J. (1997) Rates of nuclear and cytoplasmic mitochondrial DNA sequence divergence in mammals. Molecular Biology and Evolution, 14, 277–286.
    https://doi.org/10.1093/oxfordjournals.molbev.a025763

    Lotzkat, S., Hertz, A., Bienentreu, J.F. & Köhler, G. (2013) Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus. Zootaxa, 3626, 1–54.
    https://doi.org/10.11646/zootaxa.3626.1.1

    Mayr, E. & Ashlock, P.K. (1991) Principles of Systematic Zoology. McGraw-Hill Inc. New York, 475 pp.

    Meyer, C.P. (2003) Molecular systematics of cowries (Gastropoda: Cypraeidae) and diversification patterns in the tropics. Biological Journal of the Linnean Society, 79, 401–459.
    https://doi.org/10.1046/j.1095-8312.2003.00197.x

    Minh, B.Q. Minh Anh, T.N. & Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30, 1188–1195.
    https://doi.org/10.1093/molbev/mst024

    Monahan, W.B., Pereira, R.J. & Wake, D.B. (2012) Ring distributions leading to species formation: a global topographic analysis of geographic barriers associated with ring species. BMC Biology, 10, 20.
    https://doi.org/10.1186/1741-7007-10-20

    Motte, M., Cacciali, P. Aquino, A.L. & Yanosky, A. (2004) Anfibios y reptiles de los humedales del Paraguay. In: Salas Dueñas, D.A., Mereles, F. & Yanosky, A.A. (Eds.), Los Humedales de Paraguay. Comité Nacional de Humedales del Paraguay. Asunción, pp. 167–174.

    Motte, M., Núñez, K., Cacciali, P., Brusquetti, F., Scott, N. & Aquino, A.L. (2009) Categorización del state de conservación de los anfibios y reptiles de Paraguay. Cuadernos de Herpetología, 23, 5–18.

    Noonan, B.P. & Yoder, A.D. (2009) Anonymous nuclear markers for Malagasy plated lizards (Zonosaurus). Molecular Ecology Resources, 9, 402–404.
    https://doi.org/10.1111/j.1755-0998.2008.02250.x

    Oliveira, D.P., Carvalho, V.T. & Hrbek, T. (2016) Cryptic diversity in the lizard genus Plica (Squamata): phylogenetic diversity and Amazonian biogeography. Zoologica Scripta, 45, 630–641.
    https://doi.org/10.1111/zsc.12172

    Palumbi, J.H. (1991) The simple fool’s guide to PCR. University of Hawaii, Honolulu, 94 pp.

    Pelegrin, N. (2007) Presence of a polydactylous Tropidurus etheridgei (Squamata: Iguanidae: Tropiduridae) in the Dry Chaco of Córdoba Province, Argentina. Cuadernos de Herpetología, 21, 115–116.

    Pelegrin, N. & Leynaud, G.C. (2006) Reptlie fauna of the Chancaní Reserve (Arid Chaco, Argentina): species list and conservation status. Herpetozoa, 19, 85–86.

    Peracca, M.G. (1895) Viaggio del dott. Alfredo Borelli nella Republica Argentina e nel Paraguay. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Università di Torino, 10, 1–32.

    Pérez-Iglesias, J.M., Jofré, L.E. & Rueda, M.P. (2017) Primeros registros de la herpetofauna en dos áreas naturales protegidas de la provincia de Santiago del Estero (Argentina). Cuadernos de Herpetología, 31, 49–57.

    Rodrigues, M.T. (1981) Uma nova espécie de Tropidurus do Brasil (Sauria, Iguanidae). Papéis Avulsos de Zoologia, 34, 145–149.

    Rodrigues, M.T. (1984) Uma nova espécie brasileira de Tropidurus com crista dorsal (Saura, Iguanidae). Papéis Avulsos de Zoologia, 35, 169–175.

    Rodrigues, M.T. (1986) Um novo Tropidurus com crista dorsal do Brasil, com comentarios sobre suas relações, distribuição e origem (Sauria: Iguanidae). Papéis Avulsos de Zoologia, 36, 171–179.

    Rodrigues, M.T. (1987) Sistemática, ecologia e zoogeografia dos Tropidurus do grupo torquatus ao sul do Rio Amazonas (Sauria, Iguanidae). Arquivos de Zoologia, 31, 105–230.
    https://doi.org/10.11606/issn.2176-7793.v31i3p105-230

    Rodrigues, M.T. (1988) Tropidurus psammonastes: uma nova especie do grupo torquatus com notas sobre seu cariótipo e distribução (Sauria, Iguanidae). Papéis Avulsos de Zoologia, 36, 307– 313.

    Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes version 3: bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
    https://doi.org/10.1093/bioinformatics/btg180

    Sabaj Pérez, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an online reference. Version 6.5. American Society of Ichthyologists and Herpetologists, Washington, DC. Available from: http://www.asih.org/ (accessed 27 July 2017)

    Schenkel, E. (1901) Achter Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel, 13, 142–199.

    Schouten, G.B. (1929) Notas sobre la oología de algunos saurios del Paraguay y de los países limítrofes. Revista Chilena de Historia Natural, 33, 518–521.

    Scott, N.J. & Lovett, J.W. (1975) A collection of reptiles and amphibians from the Chaco of Paraguay. Occasional Papers of the University of Connecticut, 2, 257–266.

    Stucky, B.J. (2012) SeqTrace: a graphical tool for rapidly processing DNA sequencing chromatograms. Journal of Biomolecular Techniques, 23, 90–93.
    https://doi.org/10.7171/jbt.12-2303-004

    Talbot, J.J. (1978) Ecological notes on the Paraguayan Chaco herpetofauna. Journal of Herpetology, 12, 433–434.
    https://doi.org/10.2307/1563636

    Talbot, J.J. (1979) Una nueva lista sistemática de reptiles del Paraguay. Informes Científicos del Instituto de Ciencias Básicas, 2, 76–94.

    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

    Townsend, T.M., Alegre, R.E., Kelley, S.T., Wiens, J.J. & Reeder, T.W. (2008) Rapid development of multiple nuclear loci for phylogenetic analysis using genomic resources: an example from squamate reptiles. Molecular Phylogenetics and Evolution, 47, 129–42.
    https://doi.org/10.1016/j.ympev.2008.01.008

    Trifinopoulos, J., Nguyen, L.T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44, W232–W235.
    https://doi.org/10.1093/nar/gkw256

    Vold, T. & Buffett, D.A. (Eds.) (2008) Ecological Concepts, Principles and Applications to Conservation. Biodiversity BC, British Columbia, 36 pp. [http://www.biodiversitybc.org/assets/pressReleases/BBCPrinciplesWEB.pdf]

    Werneck, F.P., Leite, R.N., Geurgas, S.R. & Rodrigues, M.T. (2015) Biogeographic history and cryptic diversity of saxicolous Tropiduridae lizards endemic to the semiarid Caatinga. BMC Evolutionary Biology, 15, 94.
    https://doi.org/10.1186/s12862-015-0368-3

    Wied-Neuwied, M. (1820) Reise nach Brasilien in den Jahren 1815 bis 1817. Vol. 1. Heinrich Ludwig Bronner, Frankfurt, 398 pp. [https://archive.org/details/reisenachbrasili01wied]

    Yachdav, G., Wilzbach, S., Rauscher, B., Sheridan, R., Sillitoe, I., Procter, J., Lewis, S.E., Rost, B. & Goldberg, T. (2016) MSAViewer: interactive JavaScript visualization of multiple sequence alignments. Bioinformatics, 32, 3501–3503.
    https://doi.org/10.1093/bioinformatics/btw474

    Ziegler, T., Unger, J., Feiler, A. & Lehr, E. (2002) The first Gran Chaco Expedition of the Museum für Tierkunde Dresden: records of amphibians, reptiles and mammals from the Dry Chaco of Paraguay (Amphibia, Reptilia, Mammalia). Faunistische Abhandlungen Staatliches Museum für Tierkunde Dresden, 23, 219–238.