Revealing cryptic diversity using molecular phylogenetics and phylogeography in frogs of the Scinax ruber and Rhinella margaritifera species groups

Molecular Phylogenetics and Evolution

Volumn 43, Issue 2, 2007, Pages 567-582

  Fouquet, Antoine ^a,f   Vences, Miguel ^b   Salducci, Marie Dominique ^a   Meyer, Axel ^c   Marty, Christian ^d   Blanc, Michel ^e   Gilles, André ^a  

^a AIX MARSEILLE UNIVERSITY   (France)

^b TECHNICAL UNIVERSITY OF BRAUNSCHWEIG   (Germany)

^c UNIVERSITY OF KONSTANZ   (Germany)

^d Impasse Jean Galot   (French Guiana)

^e NONE   (French Guiana)

^f UNIVERSITY OF CANTERBURY   (New Zealand)

Author keywords

18S rDNA; Amphibia; Bufonidae; Hylidae; Mitochondrial genes; Neotropics; Rhinella; Scinax; Tyrosinase

Indexed keywords

MITOCHONDRIAL DNA; MONOPHENOL MONOOXYGENASE; RIBOSOME DNA;

ANIMAL; ARTICLE; CLASSIFICATION; FRENCH GUIANA; FROGS AND TOADS; GENETICS; MOLECULAR EVOLUTION; PHYLOGENY;

ANIMALS; ANURA; DNA, MITOCHONDRIAL; DNA, RIBOSOMAL; EVOLUTION, MOLECULAR; FRENCH GUIANA; MONOPHENOL MONOOXYGENASE; PHYLOGENY;

AMPHIBIA; ANURA; BUFONIDAE; HYLIDAE; LEPTODACTYLIDAE; MARGARITIFERA; SCINAX; SCINAX RUBRA;

EID: 34248139068     PISSN: 10557903     EISSN: 10959513     Source Type: Journal    
DOI: 10.1016/j.ympev.2006.12.006     Document Type: Article

References (118)

1. Avise J.C. Phylogeography: The History and Formation of Species (2000), Harvard University Press
2. Ballard J.W.O., and Whitlock M.C. The incomplete natural history of mitochondria. Mol. Ecol. 13 (2004) 729-744
3. Biju S.D., and Bossuyt F. New frog family from India reveals an ancient biogeographical link with the Seychelles. Nature 425 (2003) 711-714
4. Blaustein A.R., Wake D.B., and Sousa W.P. Amphibian declines-judging stability, persistence, and susceptibility of populations to local and global extinctions. Conservation Biol. 8 (1994) 60-71
5. Borkin L.Y., Litvinchuk S.N., Rozanov Y.M., and Skorinov D.V. On cryptic species (from the example of amphibians). Zoologichesky Zh. 83 (2004) 936-960
6. Bossuyt F., Meegaskumbura M., Beenaerts N., Gower D.J., Pethiyagoda R., Roelants K., Mannaert A., Wilkinson M., Bahir M.M., Manamendra-Arachchi K., Ng P.K.L., Schneider C.J., Oommen O.V., and Milinkovitch M.C. Local endemism within the western Ghats-Sri Lanka biodiversity hotspot. Science 306 (2004) 479-481
7. Bossuyt F., and Milinkovitch M.C. Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proc. Natl. Acad. Sci. USA 97 (2000) 6585-6590
8. Boujard, T., Pascal, M., Meunier, J.F., Le Bail, P.Y., 1997. Poissons de Guyane. Guide ecologique de l'Approuague et de la réserve des Nouragues. INRA.
9. Bourne G.R. Lekking behavior in the Neotropical frog Ololygon rubra. Behav. Ecol. Sociobiol. 31 (1992) 173-180
10. Bourne G.R., and York H. Vocal behaviors are related to nonrandom structure of anuran breeding assemblages in Guyana. Ethol. Ecol. Evol. 13 (2001) 313-329
11. Caron H., Dumas S., Marque G., Messier C., Bandou E., Petit R.J., and Kremer A. Spatial and temporal distribution of chloroplast DNA polymorphism in a tropical tree species. Mol. Ecol. 9 (2000) 1089-1098
12. Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol. Biol. Evol. 17 (2000) 540-552
13. Charles-Dominique P., Blanc P., Larpin D., Ledru M.P., Riera B., Sarthou C., Servant M., and Tardy C. Forest perturbations and biodiversity during the last ten thousand years in French Guiana. Acta Oecol. Int. J. Ecol. 19 (1998) 295-302
14. Chek A.A., Austin J.D., and Lougheed S.C. Why is there a tropical-temperate disparity in the genetic diversity and taxonomy of species?. Evol. Ecol. Res. 5 (2003) 69-77
15. Chek A.A., Lougheed S.C., Bogart J.P., and Boag P.T. Perception and history: molecular phylogeny of a diverse group of neotropical frogs, the 30-chromosome Hyla Anura: Hylidae. Mol. Phylogenet. Evol. 18 (2001) 370-385
16. Cherry L.M., Case S.M., and Wilson A.C. Relative rates of morphological evolution in primates, carnivores and frogs. Am. Zool. 17 (1977) 910
17. Chiari Y., Vences M., Vieites D.R., Rabemananjara F., Bora P., Ravoahangimalala O.R., and Meyer A. New evidence for parallel evolution of colour patterns in Malagasy poison frogs (Mantella). Mol. Ecol. 13 (2004) 3763-3774
18. Clement M., Posada D., and Crandall K.A. TCS: a computer program to estimate gene genealogies. Mol. Ecol. 9 (2000) 1657-1659
19. Crawford A.J. Huge populations and old species of Costa Rican and Panamanian dirt frogs inferred from mitochondrial and nuclear gene sequences. Mol. Ecol. 12 (2003) 2525-2540
20. Darst C.R., and Cannatella D.C. Novel relationships among hyloid frogs inferred from 12S and 16S mitochondrial DNA sequences. Mol. Phylogenet. Evol. 31 (2004) 462-475
21. De Granville J.-J. Phytogeographical characteristics of the Guianan forests. Taxon 37 (1988) 578-594
22. De Granville J.-J. Rain forest and xeric flora refuges in french Guiana. In: Prance G.T. (Ed). Biological Diversification in the Tropics (1982), Columbia Univ. Press 159-181
23. De la Riva I., Köhler J., Lötters S., and Reichle S. Ten years of research on Bolivian amphibians: updated checklist, distribution, taxonomic problems, literature and iconography. Rev. Esp. Herpetol. (2000) 14
24. de Queiroz K. The general lineage concept of species, species criteria, and the process of speciation. In: Howard D.J., and Berlocher S.H. (Eds). Endless Forms: Species and Speciation (1998), Oxford University Press 57-75
25. Duellman W.E. Quaternary climatic-ecological fluctuations in the lowland tropics: frogs and forests. In: Prance G.T. (Ed). Biological Diversification in the Tropics (1982), Columbia Univ. Press 389-402
26. Duellman W.E. Patterns of Distribution of Amphibians: A Global Perspective (1999), Johns Hopkins University Press
27. Duellman, W.E., Wiens, J.J., 1993. Hylid frogs of the genus Scinax Wagler, 1830, in Amazonian Ecuador and Peru. Museum of Natural History, the University of Kansas.
28. Dutech C., Joly H.I., and Jarne P. Gene flow, historical population dynamics and genetic diversity within French Guianan populations of a rainforest tree species, Vouacapoua americana. Heredity 92 (2004) 69-77
29. Dutech C., Maggia L., and Joly H.I. Chloroplast diversity in Vouacapoua americana (Caesalpiniaceae), a neotropical forest tree. Mol. Ecol. 9 (2000) 1427-1432
30. Dutech C., Maggia L., Tardy C., Joly H.I., and Jarne P. Tracking a genetic signal of extinction-recolonization events in a neotropical tree species: Vouacapoua americana Aublet in French Guiana. Evolution 57 (2003) 2753-2764
31. Efron B. 1977 Rietz Lecture-bootstrap methods-another look at the jackknife. Ann. Stat. 7 (1979) 1-26
32. Faivovich J., García P.C.A., Ananias F., Lanari L., Basso N.G., and Wheeler W.C. A molecular perspective on the phylogeny of the Hyla pulchella species group (Anura, Hylidae). Mol. Phylogenet. Evol. 32 (2004) 938-950
33. Faivovich J., Haddad C.F.B., García P.C.A., Frost D.R., Campbell J.A., and Wheeler W.C. Systematic review of the frog family hylidae, with special reference to Hylinae: phylogenetic analysis and taxonomic revision. Bull. Am. Museum Nat. Hist. 294 (2005) 1-240
34. Farris J.S., Kallersjo M., Kluge A.G., and Bult C. Testing Significance of Incongruence. Cladistics-The International Journal of the Willi Hennig Society 10 (1994) 315-319
35. Felsenstein J. Confidence limits on phylogenies-an approach using the bootstrap. Evolution 39 (1985) 783-791
36. Frailey C.D., Lavina E.L., Rancy A., and Filho J.P.S. A proposed Pleistocene/Holocene lake in the Amazona basin and its significance to Amazonian geology and biogeography. Acta Amazonica 18 (1989) 119-143
37. Frost, D.R., 2004. Amphibian Species of the World: an Online Reference. Version 3.0. New York: American Museum of Natural History. [Electronic Database accessible at ].
38. Frost D.R., and Hillis D.M. Species in concept and practice-herpetological applications. Herpetologica 46 (1990) 87-104
39. Frost D.R., Grant T., Faivovich J., Bain R.H., Haas A., Haddad C.F.B., De Sa R.O., Channing A., Wilkinson M., Donnellan S.C., Raxworthy C.J., Campbell J.A., Blotto B.L., Moler P., Drewes R.C., Nussbaum R.A., Lynch J.D., Green D.M., and Wheeler W.C. The Amphibian tree of life. Bull. Am. Museum Nat. Hist. 297 (2006) 1-370
40. Funk D.J., and Omland K.E. Species-level paraphyly and polyphyly: frequency, causes, and consequences, with insights from animal mitochondrial DNA. Ann. Rev. Ecol. Evol. Syst. 34 (2003) 397-423
41. Garcia-Paris M., Alcobendas M., Buclkey D., and Wake D.B. Dispersal of viviparity across contact zones in Iberian populations of fire salamanders (Salamandra) inferred from discordance of genetic and morphological traits. Evolution 57 (2003) 129-143
42. Gaston K.J., and Williams P.H. Spatial patterns in taxonomic diversity. In: Gaston K.J. (Ed). Biodiversity: A Biology of Numbers and Difference (1996), Blackwell Science 202-229
43. Glaw F., and Köhler J. Amphibian species diversity exceeds that of mammals. Herpetol. Rev. 29 (1998) 11-12
44. Goldstein P., Wyner K., Doukakis P., Egan M.G., Amato G., Rosenbaum H., and DeSalle R. Theory and methods for diagnosing species and populations in conservation. Ann. Mo. Bot. Garden 92 (2005) 12-27
45. Goldstein P.Z., DeSalle R., Amato G., and Vogler A.P. Conservation genetics at the species boundary. Conservation Biol. 14 (2000) 120-131
46. Gorzula, S., Señaris, C., 1998. Contribution to the herpetofauna of the Venezuelan Guayana I.A data base. xviii+270+32pp.
47. Grant T., Frost D.R., Caldwell J.P., Gagliardo R., Haddad C.F.B., Kok P.J.R., Means D.B., Noonan B.P., Schargel W.E., and Wheeler W.C. Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bull. Am. Museum Nat. Hist. 299 (2006) 1-262
48. Grant T., and Kluge A.G. Data exploration in phylogenetic inference: scientific, heuristic, or neither. Cladistics 19 (2003) 379-418
49. Graybeal A. Phylogenetic relationships of bufonid frogs and tests of alternate macroevolutionary hypotheses characterizing their radiation. Zool. J. Linnean Soc. 119 (1997) 297-338
50. Haffer J. Speciation in amazonian forest birds. Science 165 (1969) 131-137
51. Haffer, J., 1990. Geoscientific aspects of allopatric speciation. In: Peters, G., Hutterer, R. (Eds.), Vertebrates in the Tropics, Museum A. Koenig, pp. 45-60.
52. Haffer J. Alternative models of vertebrate speciation in Amazonia: an overview. Biodiversity Conservation 6 (1997) 451-476
53. Hanken J. Why are there so many new amphibian species when amphibians are declining?. Trends Ecol. Evol. 14 (1999) 7-8
54. Harrison R.G. Hybrid Zones and the Evolutionary Process (1993), Oxford University Press
55. Hass C.A., Dunski J.F., Maxson L.R., and Hoogmoed M.S. Divergent lineages within the Bufo margaritifer complex (Amphibia, Anura, Bufonidae) revealed by albumin immunology. Biotropica 27 (1995) 238-249
56. Haas W. Beitrag zur Biologie und Verbreitung dreier Vertreter des neotropischen Bufo typhonius-Komplexes (Anura: Bufonidae). Salamandra 40 3/4 (2004) 207-216
57. Hay J.M., Ruvinsky I., Hedges S.B., and Maxson L.R. Phylogenetic relationships of amphibian families inferred from DNA sequences of mitochondrial 12s and 16s ribosomal RNA genes. Mol. Biol. Evol. 12 (1995) 928-937
58. Hoegg S., Brinkmann H., Taylor J.S., and Meyer A. Phylogenetic timing of the fish-specific genome duplication correlates with the diversification of teleost fish. J. Mol. Evol. 59 (2004) 190-203
59. Hoogmoed, M.S., 1979. The herpetofauna of the Guianan region. In: Duellman, W.E. (Ed.), The South American Herpetofauna: Its Origin, Evolution and Dispersal., Museum of Natural History, University of Kansas, pp. 241-279.
60. Hoogmoed, M.S., 1990. Biosystematics of South American Bufonidae, with special reference to the Bufo "typhonius group. In: Peters, G., Hutterer, R. (Eds.), Vertebrates in the tropics, Museum Alexander Koenig, pp. 113-123.
61. Houlahan J.E., Findlay C.S., Schmidt B.R., Meyer A.H., and Kuzmin S.L. Quantitative evidence for global amphibian population declines. Nature 404 (2000) 752-755
62. Jarman S.N., and Elliott N.G. DNA evidence for morphological and cryptic Cenozoic speciations in the Anaspididae, 'living fossils' from the Triassic. J. Evol. Biol. 13 (2000) 624-633
63. Jegu M., and Keith P. Lower Oyapock River as northern limit for the Western Amazon fish fauna or only a stage in its northward progression. C. R. Acad. Sci. III 322 (1999) 1133-1143
64. Joyce D.A., Lunt D.H., Bills R., Turner G.F., Katongo C., Duftner N., Sturmbauer C., and Seehausen O. An extant cichlid fish radiation emerged in an extinct Pleistocene lake. Nature 435 7038 (2005) 90-95
65. Köhler J., Vieites D.R., Bonett R.M., Garcia F.H., Glaw F., Steinke D., and Vences M. New amphibians and global conservation: a boost in species discoveries in a highly endangered vertebrate group. Bioscience 55 (2005) 693-696
66. Kuchta S.R., and Tan A.M. Isolation by distance and post-glacial range expansion in the rough-skinned newt, Taricha granulosa. Mol. Ecol. 14 (2005) 225-244
67. Kumar S., Tamura K., Jakobsen I.B., and Nei M. MEGA2: Molecular Evolutionary Genetics Analysis Software. Bioinformatics 17 (2001) 1244-1245
68. Ledru M.P., Blanc P., Charles-Dominique P., Fournier M., Martin L., Riera B., and Tardy C. Palynological reconstruction of the rain forest in French Guiana during the past 3000 years. C. R. Acad. Sci. II Fascicule a-Sciences De La Terre Et Des Planetes 324 (1997) 469-476
69. Lescure, J., Marty, C., 2000. Atlas des Amphibiens de Guyane. Muséum national d'Histoire naturelle.
70. Lougheed S.C., Gascon C., Jones D.A., Bogart J.P., and Boag P.T. Ridges and rivers: a test of competing hypotheses of Amazonian diversification using a dart-poison frog (Epipedobates femoralis). Proc. R. Soc. Lond. B 266 (1999) 1829-1835
71. Maxson L.R. Molecular probes of phylogeny and biogeography in toads of the widespread genus Bufo. Mol. Biol. Evol. 1 (1984) 345-356
72. Mayr E. Systematics and the Origin of Species (1942), Columbia University Press, New York
73. Meegaskumbura M., Bossuyt F., Pethiyagoda R., Manamendra-Arachchi K., Bahir M., Milinkovitch M.C., and Schneider C.J. Sri Lanka: an amphibian hot spot. Science 298 (2002) 379
74. Meyer A., Salzburger W., and Schartl M. Hybrid origin of a swordtail species (Teleostei: Xiphophorus clemenciae) driven by sexual selection. Mol. Ecol. 15 3 (2006) 721
75. Meyer C.P., and Paulay G. DNA barcoding: Error rates based on comprehensive sampling. PLoS Biol. 3 12 (2005) 1-10
76. Miquelis A., Martin J.-F., Carson E.W., Brun G., and Gilles A. Performance of 18S rDNA helix E23 for phylogenetic relationships within and between the Rotifera-Acanthocephala clades. C. R. Acad. Sci. III 323 (2000) 925-941
77. Monsen K.J., and Blouin M.S. Genetic structure in a montane ranid frog: restricted gene flow and nuclear-mitochondrial discordance. Mol. Ecol. 12 (2003) 3275-3286
78. Moritz C. Applications of mitochondrial DNA analysis in conservation-a critical-review. Mol. Ecol. 3 (1994) 401-411
79. Moritz C. Defining evolutionarily significant units for conservation. Trends Ecol. Evol. 9 (1994) 373-375
80. Moritz C., Schneider C.J., and Wake D.B. Evolutionary relationships within the Ensatina eschscholtzii complex confirm the ring species interpretation. Syst. Biol. 41 (1992) 273-291
81. Myers N., Mittermeier R.A., Mittermeier C.G., Da Fonseca G.A.B., and Kent J. Biodiversity hotspots for conservation priorities. Nature 403 (2000) 853-858
82. Noonan B.P., and Gaucher P. Phylogeography and demography of Guianan harlequin toads (Atelopus): diversification within a refuge. Mol. Ecol. 14 (2005) 3017-3031
83. Nores M. An alternative hypothesis for the origin of Amazonian bird diversity. J. Biogeogr. 26 (1999) 475-485
84. Patton J.L., and Smith M.F. Paraphyly, polyphyly, and the nature of species boundaries in pocket gophers (genus Thomomys). Syst. Biol. 43 (1994) 11-26
85. Posada D., and Crandall K.A. MODELTEST: testing the model of DNA substitution. Bioinformatics 14 (1998) 817-818
86. Pramuk J.B. Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence. Zool. J. Linnean Soc. 146 (2006) 407-452
87. Räsänen M.E., Salo J.S., and Jungner H. Holocene floodplain lake sediments in the Amazon-C-14 dating and paleoecological use. Quat. Sci. Rev. 10 (1991) 363-372
88. Räsänen M.E., Salo J.S., Jungner H., and Pittman L.R. Evolution of the western Amazon lowland relief: impact of andean foreland dynamics. Terra Nova 2 (1990) 320-332
89. Richards C.M., and Moore W.S. A phylogeny for the African treefrog family Hyperoliidae based on mitochondrial rDNA. Mol. Phylogenet. Evol. 5 (1996) 522-532
90. Ronquist F., and Huelsenbeck J.P. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19 (2003) 1572-1574
91. Ruvinsky I., and Maxson L.R. Phylogenetic relationships among bufonoid frogs (Anura: Neobatrachia) inferred from mitochondrial DNA sequences. Mol. Phylogenet. Evol. 5 (1996) 533-547
92. Salducci M.-D., Marty C., Chappaz R., and Gilles A. Molecular phylogeny of French Guiana Hylinae: implications for the systematic and biodiversity of the neotropical frogs. C. R. Biol. 325 (2002) 141-153
93. Salducci M.-D., Marty C., Fouquet A., and Gilles A. Phylogenetic relationships and biodiversity in Hylids (Anura: Hylidae) from French Guiana. C. R. Biol. 328 (2005) 1009-1024
94. Salzburger W., and Meyer A. The species flocks of East African cichlid fishes: recent advances in molecular phylogenetics and population genetics. Naturwissenschaften 91 (2004) 277-290
95. Sequeira F., Alexandrino J., Rocha S., Arntzen J.W., and Ferrand N. Genetic exchange across a hybrid zone within the Iberian endemic golden-striped salamander, Chioglossa lusitanica. Mol. Ecol. 14 (2005) 245-254
96. Shimodaira H., and Hasegawa M. Multiple comparison of log-likelihoods with applications to phylogenetic inference. Mol. Biol. Evol. 16 (1999) 1114-1116
97. Sites J.W., and Marshall J.C. Delimiting species: a Renaissance issue in systematic biology. Trends Ecol. Evol. 18 (2003) 462-470
98. Sites J.W., and Marshall J.C. Operational criteria for delimiting species. Ann. Rev. Ecol. Evol. Syst. 35 (2004) 199-227
99. Sota T., and Vogler A.P. Incongruence of mitochondrial and nuclear gene trees in the carabid beetles Ohomopterus. Syst. Biol. 50 (2001) 39-59
100. Steiner C., and Catzeflis F.M. Mitochondrial diversity and morphological variation of Marmosa murina (Didelphidae) in French Guiana. J. Mammal. 84 (2003) 822-831
101. Stuart B.L., Inger R.F., and Voris H.K. High level of cryptic species diversity revealed by sympatric lineages of Southeast Asian forest frogs. Biol. Lett. 2 (2006) 470-474
102. Stuart S.N., Chanson J.S., Cox N.A., Young B.E., Rodrigues A.S.L., Fischman D.L., and Waller R.W. Status and trends of amphibian declines and extinctions worldwide. Science 306 (2004) 1783-1786
103. Swofford, D.L., 2002. PAUP*: Phylogenetic Analysis Using Parsimony (*and Other Methods), version 4.0b10 [computer program]. Sunderland (Massachusetts): Sinauer.
104. Takahata N., and Slatkin M. Mitochondrial gene flow. Proc. Natl. Acad. Sci. USA 81 (1984) 1764-1767
105. Thompson J.D., Gibson T.J., Plewniak F., Jeanmougin F., and Higgins D.G. The ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res. 24 (1997) 4876-4882
106. Van de Peer Y., Caers A., De Rijk P., and De Wachter R. Database on the structure of small ribosomal subunit RNA. Nucleic Acids Res. 26 (1998) 179-182
107. Van Vuuren B.J., Kinet S., Chopelet J., and Catzeflis F. Geographic patterns of genetic variation in four Neotropical rodents: conservation implications for small game mammals in French Guiana. Biol. J. Linnean Soc. 81 (2004) 203-218
108. Vélez-Rodríguez, C.M., 2004. Sistemática de los sapos neotropicales pertenecientes al grupo Bufo typhonius (Bufonidae). Tesis Doctoral. Universidad austral de Chile. Valdivia Chile. 159 pp.
109. Vences M., Kosuch J., Boistel R., Haddad C.F.B., La Marca E., Lötters S., and Veith M. Convergent evolution of aposematic coloration in Neotropical poison frogs: a molecular phylogenetic perspective. Organ. Divers. Evol. 3 (2003) 215-226
110. Vences M., Kosuch J., Lötters S., Widmer A., Jungfer K.H., Köhler J., and Veith M. Phylogeny and classification of poison frogs (Amphibia: Dendrobatidae), based on mitochondrial 16S and 12S ribosomal RNA gene sequences. Mol. Phylogenet. Evol. 15 (2000) 34-40
111. Vences M., Thomas M., Bonett R.M., and Vieites D.R. Deciphering amphibian diversity through DNA barcoding: chances and challenges. Philos. Trans. R. Soc. Lond. B 360 (2005) 1859-1868
112. Vences, M., Thomas, M., Van der Meijden, A.Y., Chiari, Y., Vieites, D.R., 2005b. Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers Zool. 2.
113. Wake D.B., and Jockusch E.L. Detecting species borders using diverse data sets. In: Bruce R.C., Jaeger R.G., and Houck L.D. (Eds). The Biology of Plethodontid Salamanders (2000), Kluwer Academic/Plenum Publishers 95-119
114. Wiley E.O. Evolutionary species concept reconsidered. Syst. Zool. 27 (1978) 17-26
115. Wilson E.O. The Diversity of Life (1992), Belknap Press of Harvard University Press
116. Wynn A., and Heyer W.R. Do geographically widespread species of tropical amphibians exist? An estimate of genetic relatedness within the neotropical frog Leptodactylus fuscus (Schneider 1799) (Anura Leptodactylidae). Trop. Zool. 14 (2001) 255-285
117. Young, B.E., Stuart, S.N., Chanson, J.S., Cox, N.A., Boucher, T.M., 2004. Disappearing Jewels: The Status of NewWorld Amphibians. NatureServe.
118. Zangari F., Cimmaruta R., and Nascetti G. Genetic relationships of the western Mediterranean painted frogs based on allozymes and mitochondrial markers: evolutionary and taxonomic inferences (Amphibia, Anura, Discoglossidae). Biol. J. Linnean Soc. 87 (2006) 515-536