Abstract
The genus Sceloporus is one of the largest genus of lizards in North and Central America, with 22 species groups. Among these, the torquatus group has a notably wide geographic distribution with populations occurring from southern United States to Guatemala. In spite of the taxonomical work done with the group, some problems remain unsolved. We therefore obtained the phylogeny of the torquatus group, based on 925 bp of the ribosomal 16S gene, 912 bp of the ribosomal 12S gene, and 893 bp of the ND4 gene, for a total of 54 specimens of 25 taxa. The genes were analyzed, both separately and combined, by means of maximum parsimony and Bayesian inference analyses. The subspecies of S. serrifer did not form a monophyletic group. The sequence data refuted the morphological evidence that suggested that S. s. plioporus and S. cyanogenys are closely related to S. s. serrifer and to S. s. prezygus. Regardless, these last two were recovered as sister taxa. Moreover, evidence was found that S. ornatus does not form a monophyletic group, and that S. ornatus ornatus and S. oberon are a single species, despite their marked differences in coloration and scutelation. In addition, the non-monophyly of S. mucronatus was confirmed and the phylogenetic relationships of its different species were determined. At the same time, the subspecies of S. dugesii were recovered as a monophyletic group, refuting the nonmonophyly of this taxon suggested in the phylogenetic hypothesis of the entire genus.References
Arévalo, E., Davis, S.K. & Sites, J.W., Jr. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of Sceloporus grammicus complex (Phrynosomatidae) in central México. Systematic Biology, 43, 387–481.
Baird, S.F. (1859) Description of new genera and species of North American lizards in the museum of the Smithsonian Institution. Proceedings of the Academy of Natural Sciences of Philadelphia, 10, 253–256.
Bell, E.L., Smith, H.M., & Chizar, D. (2003) An Annoted list of the species-group names applied to the lizard genus Sceloporus. Acta Zoologica Mexicana, n.s., 103–174.
Barrett, M., Donoghue, M.J., & Sober, E. (1991) Against consensus. Systematic Zoology, 40, 486–493.
Benabib, M., Kjer, K.M. & Sites, J.W., Jr. (1997) Mitochondrial DNA sequence-based phylogeny and the evolution of viviparity in the Sceloporus scalaris group (Reptilia: Squamata). Evolution, 51, 1262–1275.
Brandley, M. C. & De Queiroz, K. (2004) Phylogeny, ecomorphological evolution, and historical biogeography of the Anolis cristatellus series. Herpetological Monographs, 18, 90–126.
Cope, E. D. (1875) Check list of North American Batrachia and Reptilia with a systematic list of the higher groups and an essay on geographic distribution based on the specimens in United States National Museum. Bulletin of United States Natural Museum, 1, 1–104.
Cope, E. D. (1885) A contribution to the herpetology of Mexico. I. The collection of the Comisión Científica. IV. Cozumel Island. VI. A synopsis of the Mexican species of the genus Sceloporus. Wieg. Proceedings of American Philosophical Society, 22, 379–404.
Chrapliwy, P. S. (1964) Taxonomy and distribution of jarrovii complex of lizards of the torquatus group, genus Sceloporus. Ph.D. dissertation. University of Illinois, Urban, IL.
Dugès, A. A. D. (1877) Una nueva especie de saurio. La Naturaleza, 4, 29–34.
Felsenstein, J. (1985) Confidents limits on phylogenies: an approach using bootstrap. Evolution, 39, 783–781.
Fetzner, J.W. Jr. (1999) Extracting High-Quality DNA from shed reptile skins: A simplified method. Biotechniques, 26, 1052–1054.
Filatov D.A. (2002) ProSeq: A software for preparation and evolutionary analysis of DNA sequence data sets. Molecular Ecology Notes, 2, 621–624.
Forstner, M.R., Davis, S. K. & Arévalo, E. (1995) Support for the hypothesis of anguimorph ancestry for the suborder Serpentes from phylogenetic analysis of mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 4, 93–102.
Fox, S. F. (1975) Natural selection on morphological phenotypes of the lizard Uta stansburiana. Evolution, 29, 95–107.
Hernández-Gallegos, O., Rodríguez-Romero, F. & Casas-Andreu, G. (2003) Sceloporus torquatus melanogaster. Herpetological Review, 34, 385.
Huelsenbeck, J. P. & Ronquist, F. (2001) MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Kluge, A. G. & Wolf, A. J. (1993) Cladistics: What´s in a word?. Cladistics, 9, 183–199.
Larget, B. & Simon, D. L. (1999) Markov chain Monte Carlo algorithms for the Bayesian analysis of phylogenetic tress. Molecular Biology and Evolution, 16, 750–759.
Larsen, K.R. & Tanner, W. W. (1974) Numeric analysis of the lizard genus Sceloporus with special reference to cranial osteology. Great Basin Naturalist, 35, 1–20.
Larsen, K.R. & Tanner, W. W. (1975) Evolution of the sceloporine lizards (Iguanidae). Great Basin Naturalist, 35, 1–20.
Lee, J. C. (1996) The Amphibians and Reptiles of the Yucatan Peninsula. Cornell University Press, E.U.
Lee, J. C. (2000) A Field Guide to the Amphibians and Reptiles of the Maya World, the Lowlands of México, Northern Guatemala, and Belize. Cornell University Press, Ithaca, New York, 402 pp.
Leviton, A. E., Gibbs, R. H., Heal, E. & Dawson, C. E. (1985) Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, 1985, 802–832.
Lewis, P.O. (2001) Phylogenetics systematics turns over a new leaf. Trends in Ecology and Evolution, 16, 30–37.
Lyons-Weiler, J., Hoelzer, G. A. & Tausch, R. J. (1998) Optimal outgroup analysis. Biological Journal of the Linnean Society, 64, 493–511.
Martin, P. S. (1952) A new subspecies of the iguanid lizard Sceloporus serrifer from Tamaulipas, Mexico. Occasional Papers of the Museum of Zoology University of Michigan, 534, 1–7.
Nylander, J. A. A. (2001) Taxon sampling in phylogenetic analysis: Problems and strategies reviewed. Introductory Research Essay No. 1. Department of Systematic Zoology, Evolutionary Biology Centre, Uppsala University.
Olson , R. E. (1987) Taxonomic revision of the lizards Sceloporus serrifer and cyanogenys of the Gulf Coastal Plain. Bulletin of the Maryland Herpetological Society, 23, 158–167.
Ortí, G. & Meyer, A. (1997) The radiation of characiform fishes and the limits of resolution of mitochondrial ribosomal DNA sequences. Systematic Biology, 46, 75–100.
Posada, D. & Buckley, T. R. (2004) Model selection and model averaging in phylogenetics: advantages of Akaike Information Criterion and Bayesian approaches over likelihood ratio tests. Systematic Biology, 53, 793–808.
Posada, D., & Crandall, K. A. (1998) Modeltest: Testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Rambaut, A., & Drummond, A. (2005) Tracer v1.2.1 2003- MCMC Trace File Analyser University of Oxford. Available from http://evolve.zoo.ox.ac.uk/beast/
Reeder, T.W. (1995) Phylogenetic relationship among phrynosomatid lizards as inferred from mitochondrial ribosomal DNA sequences: substitutional bias and information content of transitions relative to transversions. Molecular Phylogenetics and Evolution, 4, 203–222.
Reeder, T.W. (2003) A phylogeny of the Australian Sphenomorphus group (Scincidae: Squamata) and the phylogenetic placement of the crocodile skinks (Tribolonotus): Bayesian approaches to assessing congruence and obtaining confidence in maximum likelihood inferred relationships. Molecular Phylogenetics and Evolution, 27, 384–397.
Saiki, R.K., Gelfand, D.H., Stoffel, S., Scharf, S.J., Higuchi, R., Horn, G.T., Mullis, K.B. & Erlich, H.A. (1998) Primer-directed enzymatic amplification of DNA with a thermostable DNA polymerase. Science, 239, 487–491.
Sanderson, M. J. & Kim, J. (2000) Parametric phylogenetics?. Systematic Biology, 49, 817–829.
Sanderson, M. J. & H. B. Shaffer. 2002. Troubleshooting molecular phylogenetic analyses. Annual Review of Ecology and Systematics, 33, 49–72.
Sites, J. W., Jr., Archie, J. W., Cole, Ch. J. & Flores-Villela, O. (1992) A review of phylogenetic hypotheses for lizards of the genus Sceloporus (Phrynosomatidae): Implications for ecological and evolutionary studies. Bulletin of the American Museum of Natural History, 213, 1–110.
Smith, A. B. (1994) Rooting molecular trees: problems and strategies. Biological Journal of the Linnean Society, 51, 279–292.
Smith, H. M. (1936) Description of new species of lizards of the genus Sceloporus from Mexico. Proceedings of the Biological Society of Washington, 49, 87–96.
Smith, H. M. (1938) The lizard of the torquatus group of the genus Sceloporus Wiegmann 1828. University of Kansas, Scientific Bulletin, 24, 539–693.
Smith, H. M. (1939) The Mexican and Central American lizards of the genus Sceloporus. Field Museum of Natural History, Publications in Zoology Series, 26,1–397.
Smith, H. M. & Brown, B. C. (1941) A new subspecies of Sceloporus jarrovii from Mexico. Zoological Series of the Field Museum of Natural History, 24, 253–257.
Smith, H. M. & Taylor, E. H. (1950) An annotated checklist and key to the reptiles of Mexico exclusive of the snakes. Bulletin of United States Natural Museum, 199, 1–253.
Soulé, M. & Kerfoot, C. (1972) On the climatic determination of scale size in a lizard. Systematic Zoology, 21, 97–105.
Stuart, L. C. (1970) A brief review of the races of Sceloporus serrifer Cope with especial reference to Sceloporus serrifer prezygus. Herpetologica 26, 141–149.
Swofford, D. L. (1998) “PAUP”: Phylogenetic Analysis Using Parsimony (And other Methods), version 4. Sinauer, Sunderland, MA.
Swofford, D. L. & Olsen, G. J. (1990) Phylogeny reconstruction. In: Hillis, D. M. & Moritz, C. (Eds.). Molecular Systematics. Sunderland, Mass. Sinauer Press, pp. 411–501.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The Clustal X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 24, 4876–4882.
Webb, R. G. & Hensely (1959) Notes on reptiles from the Mexican state of Durango. Publications of the Museum, Michigan State University. Biological series,1, 252–258.
Wenzel, J. W. & Sidall M. E. (1999) Noise. Cladistics, 15, 51–64.
Wiens, J. J. (1998a) Combining data sets with different phylogenetic histories. Systematic Biology, 47, 568–581.
Wiens, J. J., & Penkrot, T. A. (2002) Delimiting species using DNA and morphological variation and discordant species limits in spiny lizards (Sceloporus). Systematic Biology, 51, 69–91.
Wiens, J. J., Reeder, T.W. & Nieto-Montes de Oca, A. (1999) Molecular phylogenetics and evolution of sexual dichromatism among populations of the Yarrow´s spiny lizard (Sceloporus jarrovii). Evolution, 53, 1884–1897.
Wiens, J. J., & Reeder, T.W. (1997) Phylogeny of spiny lizards (Sceloporus) based on molecular and morphological evidence. Herpetological Monographs, 11, 1–101.
Wilcox, T.P., Zwickl, D.J., Heath, T.A. & Hillis, D.M. (2002) Phylogenetic relationships of the dwarf boas and comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, 25, 361–371.