Abstract
With 36 species and at least nine potentially independent lineages (not formally described yet) occurring mostly in theAndes and adjacent Patagonia and Puna plateau areas, Phymaturus lizards represent one of the most endemic vertebrategroups of the arid southwestern region of South America. Phylogenetic relationships among species of Phymaturus areinferred using mainly a morphological data set of 206 characters. Also available sequences of mitochondrial DNA for sev-en terminals were added for a total evidence analysis. Most information is included in the discrete characters block; mostcharacters involve color pattern, osteology and squamation. Continuous characters were taken from body proportions,squamation and skeletons. Among morphological data, binary polymorphic characters were analyzed applying the scaledcoding criteria. Continuous characters were entered in the analysis using standardized ranges, a method that allows a sim-ple optimization to estimate distances/costs avoiding the arbitrary coding as discrete characters. For our parsimony anal-yses we chose the implied weights method, which underweights homoplastic characters. Several runs were madeanalyzing all the information combined and also separating morphological from molecular datasets. Binary polymor-phisms were also analyzed as missing data. All characters affected by sexual dimorphism were analyzed separating thesexes; female information was more congruent with the total evidence analysis. Characters involving continuous and poly-morphic information are relevant for searching and building phylogenetic hypotheses in Phymaturus. There exists signif-icant congruence between the molecular information analyzed in this study and previous published analyses. Within bothmain clades of Phymaturus, northern subgroups are those more recently originated during the genus diversification. Spe-cies belonging to the puna subclade of the palluma group are arranged in two natural groups, one distributed in the north(Catamarca and La Rioja provinces), and the other in the south, La Rioja and San Juan provinces. Within the patagonicusgroup, the majority of the species are arranged in a south-central Chubut clade, eastern-central Chubut clade, central Rio Negro clade and a Payunia clade.References
Aguilera, G., & Mirande, J.M. (2005) A new species of Jenynsia (Cyprinodontiformes: Anablepidae) from northwestern Argentina and its phylogenetic relationships. Zootaxa, 1096, 29–39.
Avila, L.J., Pérez, C.H.F., Pérez, D.R. & Morando, M. (2011) Two new mountain lizard species of the Phymaturus genus (Squamata: Iguania) from northwestern Patagonia, Argentina. Zootaxa, 2924: 1–21.
Barbour, T. (1921) On a small collection of reptiles from Argentina. Proceedings of the Biological Society of Washington, 34, 139–141.
Boretto, J.M., Ibargüengoytía, N.R., Acosta, J.C., Blanco, G.M., Villavicencio, J. & Marinero, J.A. (2007) Reproductive biology and sexual dimorphism of a high-altitude population of the viviparous lizard Phymaturus punae from the Andes in Argentina. Amphibia–Reptilia, 28, 427–432.
Burt, C.E. & Burt, M.D. (1931) South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, 61, 227–395.
Cei, J.M. (1980) New iguanid lizards from the Famatina mountains of western Argentina. Instituto Biologia Animal, Universidad Nacional de Cuyo, Mendoza, Argentina. Journal of Herpetology, 14, 57–64.
Cei, J.M. & Videla, F. (2003) A new Phymaturus species from volcanic cordilleran mountains of the south-western Mendoza province, Argentina (Liolaemidae, Iguania, Lacertilia, Reptilia). Bolletino del Museo Regionale di Scienze Naturale, 20, 291–314.
Chancellor, G. & Van Wyhe, J. (2009) Charles Darwin’s notebooks from the Voyage of the Beagle. Cambridge: Cambridge University Press. 656 pp.
Chiszar, D., Gingery, T., Gingery, B. & Smith, H.M. (1999) Phymaturus patagonicus (Argentine chuckwalla). Facultative parthenogenesis. Herpetological Review, 30, 98.
Corbalán, V., Scolaro, A. & Debandi, G. (2009) A new species of the genus Phymaturus of the flagellifer group from Central-Western Mendoza, Argentina (Reptilia: Iguania: Liolaemidae). Zootaxa, 2021, 42–56.
Cruz, F.B., Belver, L., Acosta, J.C., Villavicencio, H.J., Blanco, G. & Cánovas, M.G. (2009) Thermal biology of Phymaturus lizards: evolutionary constraints or lack of environmental variation?. Zoology, 112, 425–432.
De Queiroz, K. (1982) The scleral ossicles of sceloporine iguanids: a reexamination with comments on their phylogenetic significance. Herpetologica, 38, 302–311.
Díaz Gómez, J.M. (2008) Historical biogeography of Phymaturus (Iguania: Liolaemidae) from Andean and patagonian South America. Zoologica Scripta, 37, 1–7.
Espinoza, R.E., Wiens, J.J. & Tracy, C.R. (2004) Recurrent evolution of herbivory in small, cold-climate lizards: breaking the ecophysiological rules of reptilian herbivory. Proceedings of the National Academy of Sciences, 101, 16819–16824.
Etheridge, R.E. (1995) Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142, 1–34.
Etheridge, R.E. (2000) A review of lizards of the Liolaemus wiegmannii group (Squamata, Iguania, Tropiduridae), and a history of morphological change in the sand-dwelling species. Herpetological Monographs, 14, 293–352.
Farris, J.S. (1969) A successive approximations approach to character weighting. Systematic Zoology, 18, 374–385.
Farris, J.S. (1970) Methods for computing Wagner trees. Systematics Zoology, 19, 83–92.
Farris, J.S. (1983) The logical basis of phylogenetic analysis. In: Platnick, N.I., Funk, V.A. (Eds), Advances in Cladistics II. Columbia University Press, New York, pp. 7–36.
Goloboff, P.A. (1993) Estimating character weights during tree search. Cladistics, 9, 83–91.
Goloboff, P.A. (1995) Parsimony and weighting: a reply to Turner and Zandee. Cladistics, 11, 91–114.
Goloboff, P.A. (1997) Self-weighted optimization: tree searches and character state reconstructions under implied transformation costs. Cladistics, 13, 225–245.
Goloboff, P., Farris, J. & Nixon, K. (2003a) T.N.T.: Tree Analysis Using New Technology. Program and documentation, available from the authors at www.zmuc.dk/public/phylogeny.
Goloboff, P.A., Farris, J.S., Källersjö, M., Oxelman, B., Ramírez, M.J. & Szumik, C.A. (2003b) Improvements to resampling measures of group support. Cladistics, 19, 324–332.
Goloboff, P., Mattoni, C.I. & Quinteros, A.S. (2006) Continuous characters analyzed as such. Cladistics, 22, 589–601.
Goloboff, P.A., Carpenter, J.M., Arias, J.S. & Miranda Esquivel, D.R. (2008) Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics, 24, 758–773.
Grosso, J.R., Cardozo, D.E., Baldo, D. & Lobo, F. (2010) Análisis citogenético en 7 especies del grupo de Phymaturus palluma (Liolaemidae, Phymaturus). Resúmenes del XI Congreso Argentino de Herpetología (2010), 85–86.
Habit, E.M. & Ortiz, J. (1996) Ciclo reproductivo de Phymaturus flagellifer (Reptilia: Tropiduridae). Boletín de la Sociedad de Biología de Concepción, 67, 7–14.
Higgins, D.G. & Sharp, P.M. (1988) Clustal: A package for performing multiple sequence alignement on a microcomputer. Gene, 73, 237–244.
Ibargüengoytía, N.R. (2005) Field, selected body temperature and thermal tolerance of the syntopic lizards Phymaturus patagonicus and Liolaemus elongatus (Iguania: Liolaemidae). Journal of Arid Environments, 62, 435–448.
Kim, J. (1993).Improving the accuracy of phylogenetic estimation by combining different methods. Systematic Biology, 42, 331–340.
Laspiur, A. & Acosta, J.C. (2007) Sexual dimorphism of Liolaemus cuyanus Cei & Scolaro, 1980 (Iguania: Liolaemidae) in a population of San Juan, Argentina. Revista peruana de biología. 14, 047–050.
Laurent, R.F. (1986) Descripciones de nuevos Iguanidae del género Liolaemus. Acta Zoologica Lilloana, 38, 87–105.
Lobo, F. (2005) Las relaciones filogenéticas dentro del grupo chiliensis (Iguania: Liolaemidae: Liolaemus): sumando nuevos caracteres y taxones. Acta Zoológica Lilloana, 49, 67–89.
Lobo, F. & Abdala, C.S. (2007) Descripción de una nueva especie de Phymaturus del grupo de P. palluma de la provincia de Mendoza, Argentina. Cuadernos de Herpetología, 21, 103–113.
Lobo, F. & Espinoza, R.E. (2004) Two new Liolaemus from the Puna region of Argentina and Chile: further resolution of purported reproductive bimodality in Liolaemus alticolor (Iguania: Liolaemidae). Copeia, 850–866.
Lobo, F. & Quinteros, S. (2005a) A morphology-based phylogeny of Phymaturus (Iguania: Liolaemidae) with the description of four new species from Argentina. Papéis Avulsos de Zoologia, 45,143–177.
Lobo, F. & Quinteros, S. (2005b) Taxonomic studies of the genus Phymaturus (Iguania: Liolaemidae): Redescription of Phymaturus patagonicus Koslowsky 1898, and Revalidation and Redescription of Phymaturus spurcus Barbour 1921. Journal of Herpetology, 39, 533–540.
Lobo, F., Abdala, F. & Scrocchi, G.J. (1995) Desarrollo del esqueleto de Liolaemus scapularis (Iguania: Tropiduridae). Bolletino del Museo Regionale di Scienze Naturale, Torino, 13, 77–104.
Lobo, F., Espinoza, R. & Quinteros, S. (2010a) A critical review and systematic discussion of recent classification proposals for liolaemid lizards. Zootaxa, 2549, 1–30.
Lobo, F., Abdala, C. & Valdecantos, S. (2010b) Taxonomic studies of the genus Phymaturus (Iguania: Liolaemidae): description of four new species. South American Journal of Herpetology, 5, 102–126.
Lobo, F., Espinoza, R.E., Sanabria, E. & Quiroga, L. (2012a) A New Phymaturus (Iguania: Liolaemidae) from the Southern Extreme of the Argentine Puna. Copeia (in press).
Lobo, F., Cruz, F.B. & Abdala, C.S. (2012b) Multiple lines of evidence show that Phymaturus agilis Scolaro, Ibargüengoytía, and Pincheira-Donoso, 2008 is a junior synonym of Phymaturus spectabilis Lobo & Quinteros, 2005. Cuadernos de Herpetologia (in press).
Mirande, J.M. (2009) Weighted parsimony phylogeny of the family Characidae (Teleostei: Characiformes). Cladistics, 25, 574–613.
Mooi, R.D. & Gill, A.C. (2010) A Transitioning State or Harmful Mutation in Systematic Ichthyology?. A Reply to Chakrabarty. Copeia, 3, 516–519.
Morando, M. (2004) Sistemática y filogenia de grupos de especies de los géneros Phymaturus y Liolaemus (Squamata: Tropiduridae: Liolaeminae). Unpublished Ph.D. Dissertation, Universidad Nacional de Tucumán, Argentina. 246 pp.
Morrone, J.J. (1994) Distributional patterns of species of Rhytirrhinini (Coleoptera: Curculionidae) and the historical relationships of the andean provinces. Global Ecology and Biogeography Letters, 4,188–194.
Morrone, J.J. (1996) The biogeographical Andean subregion: a proposal exemplified by arthropod taxa (Arachnida, Crustacea, and Hexapoda). Neotropica, 42,104–114.
Núñez, H., Veloso, A., Espejo, P., Veloso, C., Cortés, A. & Araya, S. (2010) Nuevas especies de Phymaturus (grupo “palluma”) para la zona cordillerana central de Chile (Reptilia, Sauria, Liolaemidae). Boletín del Museo Nacional de Historia Natural, Chile 59, 41–74.
Pereyra, E.A. (1992) Sistemática y relaciones evolutivas de las especies de Phymaturus Gravenhorst, 1838 (Sauria-Liolaeminae). Unpublished Magister Dissertation, Universidad de Chile.
Pincheira-Donoso, D. (2004) Una nueva especie del genero Phymaturus (Iguania: Tropiduridae: Liolaemini) del centrosur de Chile. Multequina, 13, 57–70.
Pincheira-Donoso, D., Scolaro, J.A. & Sura, P. (2008) A monographic catalogue on the systematics and phylogeny of the South American iguanian lizard family Liolaemidae (Squamata, Iguania). Zootaxa, 1800, 1–85.
Ramírez, M.J. (2003) The spider subfamily Amaurobioidinae (Aranae, Anyphaenidae): a phylogenetic revision at the generic level. Bulletin of the American Museum.of Natural History. 277, 1–262.
Rocha, C.F.D. (1999) Home range of the tropidurid lizard Liolaemus lutzae: sexual and body size differences. Revista Brasilera de Biología, 59, 125–130.
Roig-Juñent, S., Domínguez, M.C., Flores, G.E. & Mattoni, C. (2006) Biogeographic history of South American arid lands: A view from its arthropods using TASS analysis. Journal of Arid Environments, 66, 404–420.
Schulte, II, J.A., Macey, J.R., Espinoza, R.E. & Larson, A. (2000) Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal. Biological Journal of the Linnean Society, 69, 75–102.
Scolaro, J. (2010) Redescripción del Neotipo de Phymaturus palluma: un aporte a la delimitación de su tierra típica (Reptilia, Sauria, Liolaemidae). Boletín del Museo Nacional de Historia Natural, Chile, 59, 29–39.
Scolaro, J. & Cei, J.M. (2003) Una excepcional nueva especie de Phymaturus de la precordillera de Chubut, Argentina (Liolaemidae, Iguania, Lacertilia, Reptilia). Facena, 19, 107–112.
Scolaro, J.A. & Ibargüengoytía, N.R. (2007) A new species of Phymaturus from rocky outcrops in the central steppe of Rio Negro province, Patagonia Argentina (Reptilia: Iguania: Liolaemidae). Zootaxa, 1524, 47–55.
Scolaro, J.A. & Ibargüengoytía, N.R. (2008) A new fragment for the understanding of the puzzling evolutive process of the Phymaturus genus: a new species of the patagonicus group from Patagonia, Argentina (Reptilia: Iguania: Liolaemidae). Zootaxa, 1936, 38–50.
Scolaro, J.A. & Pincheira-Donoso, D. (2010) Lizards at the end of the world: two new species of Phymaturus of the patagonicus clade (Squamata, Liolaemidae) revealed in southern Patagonia of Argentina. Zootaxa, 2393, 17–32.
Scolaro, J.A. & Tappari, O.F. (2009) Una nueva especie del género Phymaturus del “grupo patagonicus” en los afloramientos rocosos del sudoeste de la provincia de Río Negro, Patagonia Argentina (Reptilia: Iguania: Liolaemidae). Naturalia Patagónica, 4, 81–94.
Scolaro, J.A., Ibargüengoytía, N.R. & Pincheira-Donoso, D. (2008) When starvation challenges the tradition of niche conservatism: On a new species of the saxicolous genus Phymaturus from Patagonia Argentina with pseudoarboreal foraging behavior (Iguania, Liolaemidae). Zootaxa, 1786, 48–60.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX-Windows interface: Flexible strategies for multiple sequence alignement aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882.
Thompson, J.D., Higgins, D.G. & Gibson, T.J. (1994) The ClustalW: Improving the sensitivity of progressive multiple sequence alignement through sequence weighting, position, specific gap penalties and weight matrix choice. Nucleic Acids Research, 22, 4673–4680.
Torres Carvajal, O. (2007). Phylogeny and biogeography of a large radiation of Andean lizards (Iguania, Stenocercus). Zoologica Scripta, 36, 311–326.
Valdecantos, S. & Lobo, F. (2007) Dimorfismo sexual en Liolaemus multicolor y L. irregularis (Iguania: Liolaemidae). Revista Española de Herpetología, 21, 55–69.
Vanhooydonck. B., Cruz, F.B., Abdala, C.S., Moreno Azócar, D.L., Bonino, M.F. & Herrel, A. (2010) Sex-specific evolution of bite performance in Liolaemus lizards (Iguania: Liolaemidae): the battle of the sexes. Biological Journal of the Linnean Society, 101, 461–475.
Verrastro, L. (2004) Sexual dimorphism in Liolaemus occipitalis (Iguania, Tropiduridae). Iheringia, Série Zoologia, 94, 45–48.
Vidal, M.A., Ortiz, J.C., Ramirez, C.C. & Lamborot, M. (2005) Intraspecific variation in morphology and sexual dimorphism in Liolaemus tenuis (Tropiduridae). Amphibia-Reptilia, 26, 343–351.
Vidal, M.A., Habit, E., Victoriano, P., González-Gajardo, A. & Ortiz, J.C. (2010) Thermoregulation and activity pattern of the high-mountain lizard Phymaturus palluma (Tropiduridae) in Chile. Zoologia, 27, 13–18.
Videla, F. (1982) Observaciones etológicas preliminares sobre Phymaturus palluma (Reptilia, Iguanidae) del roquedal precordillerano mendocino. Boletín del Museo de Ciencias Naturales y Antropología. J. C. Moyano, Mendoza, Argentina, 3, 57–62.
Wassersug, R.J. (1976) A procedure for differential staining of cartilage and bone in whole formalin fixed vertebrates. Stain Technology, 51, 131–134.
Wiens, J.J. (2000) Coding morphological variation within species and higher taxa for phylogenetic analysis. In J. J. Wiens (Ed.) Phylogenetic Analysis of Morphological Data (pp. 115–145). Washington D.C.: Smithsonian Institution Press.