Gene and species trees of a Neotropical group of treefrogs: Genetic diversification in the Brazilian Atlantic Forest and the origin of a polyploid species

Molecular Phylogenetics and Evolution

Volumn 57, Issue 3, 2010, Pages 1120-1133

  Brunes, Tuliana O ^a,b   Sequeira, Fernando ^a   Haddad, Célio F B ^b   Alexandrino, João ^b  

^a CENTRO DE INVESTIGAÇÃO EM BIODIVERSIDADE E RECURSOS GENÉTICOS   (Portugal)

^b SÃO PAULO STATE UNIVERSITY UNESP   (Brazil)

Author keywords

Amphibian; Brazilian Atlantic Forest; Hylidae; Multilocus species tree; NDNA; Neotropics mtDNA; Phyllomedusa; Phylogeny; Polyploidy

Indexed keywords

MITOCHONDRIAL DNA;

ANIMAL; ARTICLE; BAYES THEOREM; CELL NUCLEUS; CLASSIFICATION; DNA SEQUENCE; FROGS AND TOADS; GENETICS; GEOGRAPHY; MOLECULAR EVOLUTION; PHYLOGENY; PHYLOGEOGRAPHY; SEQUENCE ALIGNMENT; STATISTICAL MODEL; TETRAPLOIDY;

ANIMALS; ANURA; BAYES THEOREM; CELL NUCLEUS; DNA, MITOCHONDRIAL; EVOLUTION, MOLECULAR; GEOGRAPHY; LIKELIHOOD FUNCTIONS; PHYLOGENY; PHYLOGEOGRAPHY; SEQUENCE ALIGNMENT; SEQUENCE ANALYSIS, DNA; TETRAPLOIDY;

AMPHIBIA; HYLA; HYLIDAE; PHYLLOMEDUSA; PHYLLOMEDUSA BURMEISTERI; PHYLLOMEDUSA DISTINCTA; PHYLLOMEDUSA IHERINGII; PHYLLOMEDUSA TETRAPLOIDEA;

EID: 78649651735     PISSN: 10557903     EISSN: 10959513     Source Type: Journal    
DOI: 10.1016/j.ympev.2010.08.026     Document Type: Article

References (114)

1. Akaike H. A new look at the statistical model identification. IEEE Transactions on Automatic Control 1974, 19:716-723.
2. Arbogast B.S., Edwards S.V., Wakeley J., Beerli P., Slowinski J.B. Estimating divergence times from molecular data on phylogenetic and population genetic timescales. Annual Review of Ecology of Systematic 2002, 33:707-740.
3. Bachtrog D., Thornton K., Clark A., Andolfatto P. Extensive introgression of mitochondrial DNA relative to nuclear gene flow in the Drosophila yakuba species group. Evolution: International Journal of Organic Evolution 2006, 60:292-302.
4. Behling H. South and southeast Brazilian grasslands during Late Quaternary times: a synthesis. Palaeogeography, Palaeoclimatology, Palaeoecology 2002, 177:19-27.
5. Behling H., Lichte M. Evidence of dry and cold climatic conditions at glacial times in tropical southeastern Brazil. Quaternary Research 1997, 48:348-358.
6. Belfiore N.M., Liu L., Moritz C. Multilocus phylogenetics of a rapid radiation in the genus Thomomys. Systematic Biology 2008, 57:294-310.
7. Bofkin L., Goldman N. Variation in evolutionary processes at different codon positions. Molecular Biology and Evolution 2007, 24:513-521.
8. Borge T., Webster M.T., Andersson G., Saetre G.P. Contrasting patterns of polymorphism and divergence on the Z chromosome and autosomes in two Ficedula flycatcher species. Genetics 2005, 171:1861-1873.
9. Bossu C.M., Near T.J. Gene trees reveal repeated instances of mitochondrial DNA introgression in orangethroat darters (Percidae: Etheostoma). Systematic Biology 2009, 58:114-129.
10. Brandley M.C., Schmitz A., Reeder T.W. Partitioned Bayesian analyses, partition choice, and the phy-logenetic relationships of scincid lizards. Systematic Biology 2005, 54:373-390.
11. Broughton R.E., Harrison R.G. Nuclear gene genealogies reveal historical, demographic and selective factors associated with speciation in field crickets. Genetics 2003, 163:1389-1401.
12. Bruen T.C., Philippe H., Bryant D. A simple and robust statistical test for detecting the presence of recombination. Genetics 2006, 172:2665-2681.
13. Bull V., Beltrán M., Jiggins C.D., McMillan W.O., Bermingham E., Mallet J. Polyphyly and gene flow between non-sibling Heliconius species. BMC Biology 2006, 4:11.
14. Cabanne G.S., Santos F.R., Miyaki C.Y. Phylogeography of Xiphorhynchus fuscus (Passeriformes, Dendrocolaptidae): vicariance and recent demographic expansion in southern Atlantic forest. Biological Journal of the Linnean Society 2007, 91:73-84.
15. Cabanne G.S., d'Horta F.M., Sari H.E.R., Santos F.R., Miyaki C.Y. Nuclear and mitochondrial phylogeography of the Atlantic forest endemic Xiphorhynchus fuscus (Aves: Dendrocolaptidae): biogeography and systematics implications. Molecular Phylogenetics and Evolution 2008, 49:760-773.
16. Carnaval A.C., Bates J.M. Amphibian DNA shows marked genetic structure and tracks Pleistocene climate change in northeastern Brazil. Evolution 2007, 61:2942-2957.
17. Carnaval A.C., Moritz C. Historical climate modeling predicts patterns of current biodiversity in the Brazilian Atlantic forest. Journal of Biogeography 2008, 35:1187-1201.
18. Carnaval A.C., Hickerson M.J., Haddad C.F.B., Rodrigues M.T., Moritz C. Stability predicts genetic diversity in the Brazilian Atlantic Forest hotspot. Science 2009, 323:785-789.
19. Charlesworth B., Bartolomé C., Noel V. The detection of shared and ancestral polymorphisms. Genetical Research 2005, 86:149-157.
20. Chenuil A., Galtier N., Berrebi P. A test of the hypothesis of an autopolyploid vs allopolyploid origin for a tetraploid lineage: application to the genus Barbus (Cyprinidae). Heredity 1999, 82:373-380.
21. Costa L.P., Leite Y.L.R., da Fonseca G.A.B., da Fonseca M.T. Biogeography of South American forest mammals: endemism and diversity in the Atlantic forest. Biotropica 2000, 32:872-881.
22. Costa L.P. The historical bridge between the Amazon and the Atlantic forest of Brazil: a study of molecular phylogeography with small mammals. Journal of Biogeography 2003, 30:71-86.
23. Crawford A.J. Huge populations and old species of Costa Rican and Panamanian dirt frogs inferred from mitochondrial and nuclear gene sequences. Molecular Ecology 2003, 12:2525-2540.
24. Crawford A.J. Relative rates of nucleotide substitution in frogs. Journal of Molecular Evolution 2003, 57:636-641.
25. Dewet J.M.J. Origins of polyploids. Polyploidy, Biological Relevance 1980, 3-16. Plenum Press, New York. W.H. Lewis (Ed.).
26. Dolman G., Moritz C. Multilocus perspective on refugial isolation and divergence in rainforest skinks (Carlia). Evolution 2006, 60:573-582.
27. Drummond A.J., Ho S.Y.W., Phillips M.J., Rambaut A. Relaxed phylogenetics and dating with confidence. PLoS Biology 2006, 4:e88.
28. Drummond A.J., Rambaut A. BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology 2007, 7:214.
29. Edwards S.V., Beerli P. Perspective: gene divergence, population divergence, and the variance in coalescence time in phylogeography studies. Evolution 2000, 54:1839-1854.
30. Edwards S.V., Liu L., Pearl D.K. High-resolution species trees without concatenation. Proceedings of the National Academy of Sciences of the United States of America 2007, 104:5936-5941.
31. Evans B.J., Kelley D.B., Tinsley R.C., Melnick D.J., Cannatella D.C. A mitochondrial DNA phylogeny of African clawed frogs: phylogeography and implications for polyploid evolution. Molecular Phylogenetics and Evolution 2004, 33:197-213.
32. Evans B.J., Kelley D.B., Melnick D.J., Cannatella D.C. Evolution of RAG-1 in polyploid clawed frogs. Molecular Biology and Evolution 2005, 22:1193-1207.
33. Faivovich J., Haddad C.F.B., Baêta D., Jungferd K., Álvares G.F.R., Brandão R.A., Sheilf C., Barrientos L.S., Barrio-Amorós C.L., Cruz C.A.G., Wheeler W.C. The phylogenetic relationships of the charismatic poster frogs, Phyllomedusinae (Anura, Hylidae). Cladistics 2009, 25:1-35.
34. Fitzpatrick S.W., Brasileiro C.A., Haddad C.F.B., Zamudio K.R. Geographical variation in genetic structure of an Atlantic Coastal Forest frog reveals regional differences in habitat stability. Molecular Ecology 2009, 18:2877-2896.
35. Flot J.F., Tillier A., Samadi S., Tillier S. Phase determination from direct sequencing of length-variable DNA regions. Molecular Ecology Notes 2006, 6:627-630.
36. Funk D.J., Omland K.E. Species-level paraphyly and polyphyly: frequency, causes, and consequences, with insights from animal mitochondrial DNA. Annual Review of Ecology and Systematics 2003, 34:397-423.
37. Godinho R., Crespo E.G., Ferrand N., Harris D.J. Phylogeny and evolution of the green lizards, Lacerta spp. (Squamata: Lacertidae) based on mitochondrial and nuclear DNA sequences. Amphibia-Reptilia 2005, 26:271-286.
38. Gonçalves H., Martínez-Solano I., Ferrand N., García-París M. Conflicting phylogenetic signal of nuclear vs mitochondrial DNA markers in midwife toads (Anura, Discoglossidae, Alytes): Deep coalescence or ancestral hybridization?. Molecular Phylogenetics and Evolution 2007, 44:494-500.
39. Gouveia S.E.M., Pessenda L.C.R., Aravena R., Boulet R., Scheel-Ybert R., Bendassoli J.A., Ribeiro A.S., Freitas H.A. Carbon isotopes in charcoal and soils in studies of paleovegetation and climate changes during the late Pleistocene and the Holocene in the southeast and centerwest regions of Brazil. Global and Planetary Change 2002, 33:95-106.
40. Grazziotin F.G., Monzel M., Echeverrigaray S., Bonatto S.L. Phylogeography of the Bothrops jararaca complex (Serpentes: Viperidae): past fragmentation and island colonization in the Brazilian Atlantic Forest. Molecular Ecology 2006, 15:3969-3982.
41. Gregory T.R., Mable B.K. Polyploidy in animals. In The evolution of the genome 2005, 427-517. Elsevier, San Diego. T.R. Gregory (Ed.).
42. Haddad C.F.B., Pompal J.P., Batistic R.F. Natural hybridization between diploid and tetraploid species of leaf-frogs, genus Phyllomedusa (Amphibia). Journal of Herpetology 1994, 28:425-430.
43. Haffer J. Speciation in Amazonian forest birds. Science 1969, 165:131-137.
44. Hall, T.A., 1999. BioEdit: a User-Friendly Biological Sequence Alignment Editor and Analysis Program for Windows 95/98/NT. Nucleic Acids Symposium Series 41, pp. 95-98.
45. Heads M. Dating nodes on molecular phylogenies: a critique of molecular biogeography. Cladistics 2005, 21:62-78.
46. Heled J., Drummond A.J. Bayesian inference of species trees from multilocus data. Molecular Biology and Evolution 2010, 27:570-580.
47. Hudson R.R., Kaplan N.L. Statistical properties of the numberof recombination events in the history of a sample of DNA sequences. Genetics 1985, 111:147-164.
48. Hudson R.R., Coyne J.A. Mathematical consequences of the genealogical species concept. Evolution 2002, 56:1557-1565.
49. Huelsenbeck J.P., Ronquist F. MRBAYES v.3.0b4: Bayesian inference of phylogenetic trees. Bioinformatics 2001, 17:754-755.
50. Johnson K.P., Clayton D.H. Nuclear and mitochondrial genes contain similar phylogenetic signal for pigeons and doves (Aves: Columbiformes). Molecular Phylogenetics and Evolution 2000, 14:141-151.
51. Kass R.E., Raftery A.E. Bayes factors. Journal of the American Statistical Association 1995, 90:773-795.
52. Keller M.J., Gerhardt H.C. Polyploidy alters advertisement call structure in grey treefrogs. Proceedings of the Royal Society of London B 2001, 268:341-345.
53. King M. Animal Cytogenetics 4: Chordata 2, Amphibia 1990, Gebriider Borntraeger, Berlin.
54. Kotaki M., Kurabayashi A., Matsui M., Khonsue W., Djong T.H., Tandon M., Sumida M. Genetic divergences and phylogenetic relationships among the Fejervarya limnocharis complex in Thailand and neighboring countries revealed by mitochondrial and nuclear genes. Zoological Science 2008, 25:381-390.
55. Lara M.C., Patton J.L. Evolutionary diversification of spiny rats (genus Trinomys, Rodentia: Echimyidae) in the Atlantic Forest of Brazil. Zoological Journal of the Linnean Society 2000, 130:661-686.
56. Ledru M.P. Late Quaternary and climatic changes in central Brazil. Quaternary Research 1993, 39:90-98.
57. Ledru M.P., Braga P.I.S., Soubiès F., Fournier M., Martin L., Suguio K., Turq B. The last 50000 years in the Neotropics (Southern Brazil): evolution of vegetation and climate. Palaeogeography, Palaeoclimatology, Palaeoecology 1996, 123:239-257.
58. Ledru M.P., Mourguiart P., Riccomini C. Related changes in biodiversity, insolation and climate in the Atlantic rainforest since the last interglacial. Palaeogeography, Palaeoclimatology, Palaeoecology 2009, 271:140-152.
59. Liu L., Edwards S.V. Phylogenetic analysis in the anomaly zone. Systematic Biology 2009, 58:452-460.
60. Lynch J.D. The amphibians of the lowland tropical rainforests. South American Herpetofauna Its Origin, Evolution, and Dispersal The Museum of Natural History 1979, The University of Kansas, Kansas. W.E. Duellman (Ed.).
61. Mable B.K. 'Why polyploidy is rarer in animals than plants': myths and mechanisms. Biological Journal of Linnean Society 2004, 82:453-466.
62. Mable B.K., Roberts J.D. Mitochondrial DNA evolution of tetraploids in the genus Neobatrachus (Anura:Myobatrachidae). Copeia 1997, 1997:680-689.
63. Macey J., Larson A., Ananjeva N., Fang Z., Papenfuss T. Two novel gene orders and the role of light-strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular Biology and Evolution 1997, 14:91-104.
64. Machado C.A., Hey J. The causes of phylogenetic conflict in a classic Drosophila species group. Proceedings of the Royal Society of London B 2003, 270:1193-1202.
65. Maddison W.P. Gene trees in species trees. Systematic Biology 1997, 46:523-536.
66. Mallet J. Hybridization, ecological races and the nature of species: empirical evidence for the ease of speciation. Philosophical Transactions of the Royal Society B 2008, 363:2971-2986.
67. Martino A.L., Sinsch U. Speciation by polyploidy in Odontophrynus americanus. Journal of Zoology 2002, 257:67-81.
68. Morellato L.P.C., Haddad C.F.B. Introduction: the Brazilian Atlantic forest. Biotropica 2000, 32:786-792.
69. Moriyama E.N., Powell J.R. Synonymous substitution rates in Drosophila: mitochondrial versus nuclear genes. Journal of Molecular Evolution 1997, 45:378-391.
70. Myers N., Mittermeier R.A., Mittermeier C.G., da Fonseca G.A.B., Kent J. Biodiversity hotspots for conservation priorities. Nature 2000, 403:853-858.
71. Nei M. Molecular Evolutionary Genetics 1987, Colombia University Press, New York.
72. Nichols R. Gene trees and species trees are not the same. Trends in Ecology and Evolution 2001, 16:358-364.
73. Nylander J.A.A., Wilgenbusch J.C., Warren D.L., Swofford D.L. AWTY (are we there yet?): a system 803 for graphical exploration of MCMC convergence in Bayesian phylogenetics. Bioinformatics 2008, 24:581-583.
74. Oliveira-Filho A.T., Fontes M.A. Patterns of floristic differentiation among Atlantic Forests in Southeastern Brazil and the influence of climate. Biotropica 2000, 32:793-810.
75. Osborn T.C., Pires J.C., Birchler J.A., Auger D.L., Chen Z.J., Lee H.S., Comai L., Madlung A., Doerge R.W., Colot V., Martienssen R.A. Understanding mechanisms of novel gene expression in polyploids. Trends in Genetics 2003, 19:141-147.
76. Otto S.P. The evolutionary consequences of polyploidy. Cell 2007, 131:452-462.
77. Otto S.P., Whitton J. Polyploid incidence and evolution. Annual Review of Genetics 2000, 34:401-437.
78. Pamilo P., Nei M. Relationships between gene trees and species trees. Molecular Biology and Evolution 1988, 5:568-583.
79. Pellegrino K.C.M., Rodrigues M.T., Waite A.N., Morando M., Yonenaga-Yassuda Y., Sites J.W. Phylogeography and species limits in the Gymnodactylus darwinii complex (Gekkonidae: Squamata): genetic structure coincides with river systems in the Brazilian Atlantic Forest. Biological Journal of the Linnean Society 2005, 85:13-26.
80. Pinho C., Harris D.J., Ferrand N. Non-equilibrium estimates of gene flow inferred from nuclear genealogies suggest that Iberian and North African wall lizards (Podarcis spp.) are an assemblage of incipient species. BMC Evolutionary Biology 2008, 8:63.
81. Pombal J.P., Haddad C.F.B. Espécies de Phyllomedusa do grupo burmeisteri do Brasil Oriental, com descrição de uma espécie nova (Amphibia, Hylidae). Revista Brasileira de Biologia 1992, 52:217-229.
82. Posada D. JModelTest: phylogenetic model averaging. Molecular Biology and Evolution 2008, 25:1253-1256.
83. Posada D., Buckley T.R. Model selection and model averaging in phylogenetics: advantages of the AIC and Bayesian approaches over likelihood ratio tests. Systematic Biology 2004, 53:793-808.
84. Prado, C.P.A., Haddad, C.F.B., Zamudio, K., submitted for publication. High genetic structure among populations of a widespread hylid frog of the Brazilian Cerrado. Molecular Ecology.
85. Ptacek M.G., Gerhardt H.C., Sage R.D. Speciation by polyploidy in treefrogs: multiple origins of the tetraploid Hyla versicolor. Evolution 1994, 48:898-908.
86. Prychitko T.M., Moore W.S. The utility of DNA sequences of an intron from the β-fibrinogen gene in phylogenetic analysis of woodpeckers (Aves: Picidae). Molecular Phylogenetics and Evolution 1997, 8:193-204.
87. Rambaut, A., 2008. FigTree v.1.1.2. Available from: http://tree.bio.ed.ac.uk/software/figtree/.
88. Rambaut, A., Drummond, A.J., 2007. Tracer v1.4.1. Available from: http://beast.bio.ed.ac.uk/Tracer.
89. Ribas C.C., Miyaki C. Molecular systematics in Aratinga parakeets: species limits and historical biogeography in the 'solstitialis' group, and the systematic position of Nandayus nenday. Molecular Phylogenetics and Evolution 2004, 30:663-675.
90. Ribeiro A.C. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotropical Ichthyologists 2006, 4:225-246.
91. Ribeiro M.C., Metzger J.P., Martensen A.C., Ponzoni F.J., Hirota M.M. The Brazilian Atlantic Forest: how much is left, and how is the remaining forest distributed? Implications for conservation. Biological Conservation 2009, 142:1141-1153.
92. Ronquist F., Heulsenbeck J.P. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 2003, 19:1572-1574.
93. Rozas J., Sanchez-DelBarrio J.C., Messeguer X., Rozas R. DnaSP, DNA polymorphism analyses by the coalescent and other methods. Bioinformatics 2003, 19:2496-2497.
94. Rull V. Blackwell Publishing Ltd Speciation timing and Neotropical biodiversity: the Tertiary-Quaternary debate in the light of molecular phylogenetic evidence. Molecular Ecology 2008, 17:2722-2729.
95. Saadi, A., Machette, M.N., Haller, K. M., Dart, R.L, Bradley, L., Souza, A.M.P.D., 2002. Map and database of Quaternary faults and lineaments in Brazil. U.S. Geological Survey, Open-File Report 02-230, version 1.0. Available from: http://pubs.usgs.gov/of/2002/ofr-02-230/.
96. Saadi A., Bezerra F.H.R.B., Costa R.D., Igreja H.L.S., Franzinelli E. Neotectônica da plataforma brasileira. Quaternário do Brasil 2005, 211-230. Holos editora, Ribeirão Preto. C.R.G. Souza, K. Suguio, A.M.S. Oliveira, P.E. Oliveira (Eds.).
97. Saia S.E.M.G., Pessenda L.C.R., Gouveia S.E.M., Aravena R., Bendassolli J.A.B. Last glacial vegetation changes in the Atlantic Forest, southeastern Brazil. Quaternary International 2008, 184:195-201.
98. Seehausen O. Hybridization and adaptive radiation. Trends in Ecology & Evolution 2004, 19:198-207.
99. Sequeira F., Ferrand N., Harris D.J. Assessing the phylogenetic signal of the nuclear ß-Fibrinogen intron 7 in salamandrids (Amphibia: Salamandridae). Amphibia-Reptilia 2006, 27:409-418.
100. Sigrist M.S., Carvalho C.J.B. Detection of areas of endemism on two spatial scales using Parsimony Analysis of Endemicity (PAE): the Neotropical region and the Atlantic Forest. Biota Neotrop. 2008, 8:33-42.
101. Silva-Filho I.S., Juncá F.A. Evidence of full species status of the neotropical leaffrog Phyllomedusa burmeisteri bahiana (A. Lutz, 1925) (Amphibia, Anura, Hylidae). Zootaxa 2006, 1113:51-64.
102. Silvestro, D., Michalak, I., 2010. RAxML GUI: a graphical front-end for RAML. Available at: http://sourceforge.net/projects/raxmlgui/.
103. Slade R.W., Moritz C., Heideman A., Hale P.T. Multiple nuclear gene phylogenies: Application to pinnipeds and comparation with a mitochondrial DNA gene phylogeny. Molecular Ecology 1994, 11:341-356.
104. Stamatakis A. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 2006, 22:2688-2690.
105. Stephens M., Smith N.J., Donnelly P. A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics 2001, 68:978-989.
106. Stock M. Mating call differences between diploid and tetraploid green toads (Bufo viridis complex) in Middle Asia. Amphibia-Reptilia 1998, 19:29-42.
107. Stöck M., Moritz C., Hickerson M., Frynta D., Dujsebayeva T., Eremchenko V., Macey J.R., Papenfuss T.J., Wake D.B. Evolution of mitochondrial relationships and biogeography of Palearctic green toads (Bufo viridis subgroup) with insights in their genomic plasticity. Molecular Phylogenetics and Evolution 2006, 41:663-689.
108. Stöck M., Ustinova J., Lamatsch D.K., Schartl M., Perrin N., Moritz C. A vertebrate reproductive system involving three ploidy levels: hybrid origin of triploids in a contact zone of diploid and tetraploid palearctic green toads (Bufo viridis subgroup). Evolution 2010, 64:944-959.
109. Tamura K., Dudley J., Nei M., Kumar S. MEGA4: molecular evolutionary genetics analysis (MEGA) software version 4.0. Molecular Biology and Evolution 2007, 24:1596-1599.
110. Thomé M.T.C., Zamudio K.R., Giovanelli J.G.R., Haddad C.F.B., Baldissera Jr.F.A., Alexandrino J. Phylogeography of endemic toads and post-Pliocene persistence of the Brazilian Atlantic Forest. Molecular Phylogenetic and Evolution 2010, 55:1018-1031.
111. Tymowska J. Polyploidy and cytogenetic variation in frogs of the genus Xenopus. Amphibian Cytogenetics, Evolution 1991, Academic Press, San Diego. D.M. Green, S.K. Sessions (Eds.).
112. Watterson G.A. On the number of segregation sites. Theoretical Population Biology 1975, 7:256-276.
113. Wiens J.J., Kuczynski C.A., Arif S., Reeder T.W. Molecular Phylogenetics and Evolution Phylogenetic relationships of phrynosomatid lizards based on nuclear and mitochondrial data, and a revised phylogeny for Sceloporus. Molecular Phylogenetics and Evolution 2010, 54:150-161.
114. Wüster W., Ferguson J.E., Quijada-Mascareñas A., Pook C.E., Salomão M.G., Thorpe R.S. Tracing an invasion: landbridges, refugia, and the phylogeography of the Neotropical rattlesnake (Serpentes: Viperidae: Crotalus durissus). Molecular Ecology 2005, 14:1095-1108.