Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2009-05-11
Page range: 1–97
Abstract views: 60
PDF downloaded: 20

Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni

Museo de Zoología, Escuela de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Apartado 17-01-2184, Quito, Ecuador Natural History Museum & Biodiversity Research Center, Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-7561, USA
Evolutionary Biology Centre, Department of Evolution, Genomics and Systematics, Uppsala University, SE-752 36 Uppsala, Sweden
Natural History Museum & Biodiversity Research Center, Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, Kansas 66045-7561, USA
Asociación Amigos de Doñana, Panama 6, 41012 Sevilla, Spain. Museo de Historia Natural, Fundación La Salle de Ciencias Naturales, Apartado 1930 Caracas 1010-A, Venezuela
Anfibios y Reptiles, Conservación Internacional-Colombia & Laboratorio de Herpetología, Pontificia Universidad Javeriana. Bogotá D.C., Colombia
Evolutionary Biology Centre, Department of Evolution, Genomics and Systematics, Uppsala University, SE-752 36 Uppsala, Sweden Estación Biológica de Doñana-CSIC, Pabellón del Perú, Avd. María Luisa s/n, 41013 Sevilla, Spain
Amphibia Allophrynidae Amazon Andes Anura Centrolenid frogs Classification Cordillera de la Costa Central America Diversity Guiana Shield Hyloidea Neobatrachia Neotropics Phylogeny South America Systematics

Abstract

Based on a molecular phylogeny, a new phylogenetic taxonomy that is compatible with both the International Code of Zoological Nomenclature (ICZN) and the PhyloCode is proposed for Glassfrogs and their sister taxon, Allophryne ruthveni. The arrangement presented herein emphasizes the recognition of clades having (i) significant statistical support and congruence among phylogenetic estimation methods (i.e., parsimony, maximum likelihood, and Bayesian inference criteria), (ii) congruence among genetic markers, and (iii) morphological and/or behavioral distinctiveness. Also, when previously recognized groups are recovered as monophyletic or nearly monophyletic, we propose taxa that minimize the number of name changes required to make these groups monophyletic, preserving the names and contents of previous classifications (i.e., nomenclatural stability). The evolutionary proximity of Centrolenidae and Allophrynidae is recognized by combining these families into an unraked taxon, Allocentroleniae—a proposal that maintains the traditional names and species contents of Centrolenidae and Allophrynidae. We arrange centrolenid diversity in two subfamilies: Centroleninae and Hyalinobatrachinae. Within Centroleninae, the diagnosis and species content of the genera Centrolene, Cochranella, and Nymphargus are modified; Teratohyla is resurrected and modified, and Chimerella, Espadarana, Rulyrana, Sachatamia, and Vitreorana are proposed as new genera. The other subfamily, Hyalinobatrachinae, contains the new genus Celsiella and a modified Hyalinobatrachium that fully corresponds to the former fleischmanni Group. Additionally, the genus Ikakogi is described. Ikakogi could not be assigned with confidence to either subfamily and it is placed as incertae sedis in Centrolenidae. The data at hand suggest that Ikakogi tayrona is a lineage as old as the subfamilies Hyalinobatrachinae and Centroleninae. The revised taxonomy differs markedly from previous arrangements, which were based on phenetics and few morphological characters. Most of the genera defined herein are confined to distinct biogeographic regions, highlighting the importance of geography in the speciation of Glassfrogs. The principal limitation of this proposal is that it is based on an incomplete sampling of taxa (54% of the recognized Glassfrogs). Although diagnoses are based on phenotypic traits, there are several cases (16% of all species) in which the allocation of species is ambiguous because of morphological homoplasy and the lack of molecular data. Finally, in an attempt to facilitate species identification, comparison, and generic placement, we provide photographs for most (~ 96%) of the recognized centrolenid species.

References

  1. Aguayo, R. & Harvey, M.B. (2006) A new glassfrog of the Cochranella granulosa group (Anura: Centrolenidae) from a Bolivian cloud forest. Herpetologica, 62, 323–330.

    Alfaro, M.E., Zoller, S. & Lutzoni, F. (2003) Bayes or bootstrap? A simulation study comparing the performance of Bayesian Markov chain Monte Carlo sampling and bootstrapping in assessing phylogenetic confidence. Molecular Biology and Evolution, 20, 255–266.

    AmphibiaWeb. (2009) Information on amphibian biology and conservation. Berkeley, California. Available from http://amphibiaweb.org/ (accessed 19 April 2009).

    Austin, J.D., Lougheed, S.C., Tanner, K., Chek, A.A., Bogart, J.P. & Boag, P.T. (2002) A molecular perspective on the evolutionary affinities of an enigmatic neotropical frog, Allophryne ruthveni. Zoological Journal of the Linnean Society, 134, 335–346.

    Avise, J. C. & Johns, G. C. (1999) Proposal for a standardized temporal scheme of biological classification for extant species. Proceedings of the National Academy of Sciences of the United States of America, 96, 7358–7363.

    Ayarzagüena, J. (1992) Los Centrolenidos de la Guayana Venezolana. Publicaciones de la Asociación de Amigos de Doñana, 1, 1–48.

    Ayarzagüena, J. & Señaris, J.C. (1996) Dos nuevas especies de Cochranella (Anura; Centrolenidae) para Venezuela. Publicaciones de la Asociación de Amigos de Doñana, 8, 1–16.

    Barrera-Rodríguez, M. (2000 “1999”) Estudio anatómico de cuatro especies de ranitas de cristal del género Hyalinobatrachium Ruiz and Lynch 1991 grupo fleischmanni (Amphibia: Anura: Centrolenidae). Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 23 (suplemento especial), 245–260.

    Barrera-Rodríguez, M. & Ruiz-Carranza, P.M. (1989) Una nueva especie del género Centrolenella Noble 1920 (Amphibia: Anura: Centrolenidae) de la Cordillera Occidental de Colombia. Trianea, 3, 77–84.

    Barrio-Amoros, C.L. & Brewer-Carias, C. (2008) Herpetological results of the 2002 expedition to Sarisarinama, a tepui in Venezuelan Guayana, with the description of five new species. Zootaxa, 1942, 3–68.

    Biju, S.D. & Bossuyt, F. (2003) New frog family from India reveals an ancient biogeographical link with the Seychelles. Nature, 425, 711–714.

    Boettger, O. (1892) Katalog der Batrachier-Sammlung im Museum der Naturforschenden Gesellschaft in Frankfurt am Main. Knauer, Frankfurt am Main.

    Boettger, O. (1893) Ein neuer Laubfrosch aus Costa Rica. Berichte über die Senckenbergische naturforschende Gesellschaft in Frankfurt am Main, 1893, 251–252.

    Bolívar-G., W., Grant, T. & Osorio, L.A. (1999) Combat behavior in Centrolene buckleyi and other centrolenid frogs. Alytes, 16, 77–83.

    Boore, J.L. (2006) The use of genome-level characters for phylogenetic reconstruction. Trends in Ecology & Evolution, 21, 439–446.

    Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. Trustees, British Museum (Natural History), London, 495 pp.

    Boulenger, G.A. (1895) Third report on additions to the batrachian collection in the Natural-History Museum. Proceedings of the Zoological Society of London, 1894, 640–646.

    Boulenger, G.A. (1918) Descriptions of new South-American Batrachians. Annual Magazine of Natural History, 9, 427–433.

    Burton, T.C. (1998) Pointing the way: The distribution and evolution of some characters of the finger muscles of frogs. American Museum Novitates, 3229, 1–13.

    Burton, T.C. (2004) Muscles of the pes of hylid frogs. Journal of Morphology, 260, 209–233.

    Cadle, J.E. & McDiarmid, R.W. (1990) Two New Species of Centrolenella (Anura, Centrolenidae) from Northwestern Peru. Proceedings of the Biological Society of Washington, 103, 746–768.

    Caldwell, J.P. & Hoogmoed, M.S. (1998) Allophrynidae, Allophryne, A. ruthveni. Catalogue of American Amphibians and Reptiles, 666, 1–3.

    Cannatella, D.C. (1980) Two new species of Centrolenella from Bolivia (Anura: Centrolenidae). Proceedings of the Biological Society of Washington, 93, 714–724.

    Cannatella D.C. & Duellman W.E. (1982) Two new species of Centrolenella, with a brief review of the genus in Perú and Bolivia. Herpetologica, 38, 380–388.

    Cantino, P.D. & de Queiroz, K. (2007) PhyloCode: International Code of Phylogenetic Nomenclature. Version 4b. Available at: http://www.ohiou.edu/phyloCode3a.pdf.

    Castroviejo-Fisher, S., De la Riva, I. & Vilà, C. (2007a) Transparent frogs show potential of natural world. Nature, 449, 972.

    Castroviejo-Fisher, S., Ayarzagüena, J. & Vilà, C. (2007b) A new species of Hyalinobatrachium (Centrolenidae: Anura) from Serrania de Perijá, Venezuela. Zootaxa, 1441, 51–62.

    Castroviejo-Fisher, S., Señaris, J.C., Ayarzagüena, J. & Vilà, C. (2008) Resurrection of Hyalinobatrachium orocostale and notes on the Hyalinobatrachium orientale species complex (Anura: Centrolenidae). Herpetologica, 64, 472–484.

    Castroviejo-Fisher, S., Guayasamin, J.M. & Kok, P.J.R. (2009) Species status of Centrolene lema Duellman and Señaris, 2003 (Amphibia: Centrolenidae) revealed by Integrative Taxonomy. Zootaxa, 1980, 16–28.

    Cisneros-Heredia, D.F. (2007) A new species of glassfrog of the genus Centrolene from the foothills of Cordillera Oriental of Ecuador (Anura: Centrolenidae). Herpetozoa, 20, 27–34.

    Cisneros-Heredia, D.F. & Guayasamin, J.M. (2006) Amphibia, Anura, Centrolenidae, Centrolene mariaelenae: Distribution extension, Ecuador. Check List, 2, 93–95.

    Cisneros-Heredia, D.F. & McDiarmid, R.W. (2006a) A new species of the genus Centrolene (Amphibia : Anura : Centrolenidae) from Ecuador with comments on the taxonomy and biogeography of Glassfrogs. Zootaxa, 1344, 1–32.

    Cisneros-Heredia, D.F. & McDiarmid, R.W. (2006b) Review of the taxonomy and conservation status of the Ecuadorian glassfrog Centrolenella puyoensis Flores & McDiarmid (Amphibia: Anura: Centrolenidae). Zootaxa,1361, 21–31.

    Cisneros-Heredia, D.F. & McDiarmid, R.W. (2007) Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1–82.

    Cisneros-Heredia, D.F. & Meza-Ramos, P. (2007) An enigmatic new species of glassfrog (Amphibia: Anura: Centrolenidae) from the Amazonian Andean slopes of Ecuador. Zootaxa, 1485, 33–41.

    Cisneros-Heredia, D.F. & Morales-Mite, M. (2008) A new species of glassfrog from elfin forests of the Cordillera del Cóndor, southeastern Ecuador. Herpetozoa, 21 (1/2), 49–56.

    Cisneros-Heredia, D.F. & Yánez-Muñoz, M. (2007) A new species of Glassfrog (Centrolenidae) from the southern Andean foothills on the west Ecuadorian region. South American Journal of Herpetology, 2, 1–10.

    Cochran, D.M. & Goin, C.J. (1970) Frogs of Colombia. United States National Museum Bulletin, Smithsonian Institution Press, Washington, D.C., 655 pp.

    da Silva, H. (1998). Phylogenetic relationships of the family Hylidae with emphasis on the relationships within the subfamily Hylinae (Amphibia: Anura). Ph.D. Dissertation, University of Kansas, Lawrence.

    Darst, C.R. & Cannatella, D.C. (2004) Novel relationships among hyloid frogs inferred from 12S and 16S mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 31, 462–475.

    Darwin, C.R. (1859) On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life, ed 1. John Murray, London, x + 502 pp.

    de Pinna, M.C. (1991) Concepts and tests of homology in the cladistic paradigm. Cladistics, 7, 367–394.

    de Queiroz, K. (2005) Ernst Mayr and the modern concept of species. Proceedings of the National Academy of Sciences of the United States of America, 102, 6600–6607.

    Dubois A. (2005) Amphibia Mundi. 1.1. An ergotaxonomy of recent amphibians. Alytes, 23, 1–24.

    Duellman, W.E. (1976) Centrolenids frogs from Perú. The University of Kansas Museum of Natural History Occasional Papers, 52, 1–11.

    Duellman, W.E. (1980) The identity of Centrolenella grandisonae Cochran and Goin (Anura, Centrolenidae). Transactions of the Kansas Academy of Sciences, 83, 26–32.

    Duellman, W.E. (1981) Three new species of centrolenid frogs from the Pacific versant of Ecuador and Colombia. The University of Kansas Museum of Natural History Occasional Papers, 88, 1–9.

    Duellman, W.E. (2001) Hylid Frogs of Middle America. Society for the Study of Amphibians and Reptiles, Ithaca, New York, + 92 plates, 1159 pp.

    Duellman, W.E. & Burrowes, P.A. (1989) New species of frogs, Centrolenella, from the Pacific Versant of Ecuador and Southern Colombia. The University of Kansas Museum of Natural History Occasional Papers, 132, 1–14.

    Duellman, W.E. & Savitzky, A.H. (1976) Aggressive behavior in a centrolenid frog, with comments on territoriality in anurans. Herpetologica, 32, 401–404.

    Duellman W.E. & Schulte R. (1993) New species of centrolenid frogs from northern Peru. The University of Kansas Museum of Natural History Occasional Papers, 155, 1–33.

    Duellman, W.E. & Señaris, J.C. (2003) A new species of glass frog (Anura: Centrolenidae) from the Venezuelan Guayana. Herpetologica, 59, 247–252.

    Duellman, W.E. & Toft, C.A. (1979) Anurans from Serrania de Sira, Amazonian Peru: taxonomy and biogeography. Herpetologica, 35, 60–70.

    Duellman, W.E. & Trueb, L. (1994) Biology of Amphibians. The Johns Hopkins University Press, Baltimore and London, i–xxi + 670 pp.

    Dunn, E.R. (1931) New frogs from Panama and Costa Rica. Occasional Papers of the Boston Society of Natural History, 5, 385–401.

    Erixon, P., Svennblad, B., Britton, T. & Oxelman, B. (2003) Reliability of Bayesian posterior probabilities and bootstrap frequencies in phylogenetics. Systematic Biology, 52, 665–673.

    Fabrezi, M. (1992) El carpo de los anuros. Alytes, 10, 1–29.

    Fabrezi, M. (1993) The anuran tarsus. Alytes, 11, 47–63.

    Fabrezi, M. & Alberch, P. (1996) The carpal elements of anurans. Herpetologica, 52, 188–204.

    Fabrezi, M. & Langone, J.A. (2000) Los caracteres morfológicos del controvertido Neobatrachia arborícola Allophryne ruthveni Gaige, 1926. Cuadernos de herpetología 14, 1, 47–59.

    Faivovich, J., Haddad, C.F.B., Garcia, P.C.A., Frost, D.R., Campbell, J.A. & Wheeler, W.C. (2005) Systematic review of the frog family hylidae, with special reference to hylinae: phylogenetic analysis and taxonomic revision. Bulletin of the American Museum of Natural History, 294, 1–240.

    Flores, G. (1985) A new Centrolenella (Anura) from Ecuador, with comments on nuptial pads and prepollical spines in Centrolenella. Journal of Herpetology, 19, 313–320.

    Flores, G. (1987) A new Centrolenella from the Amazonian Lowlands of Colombia. Journal of Herpetology, 21, 185–190.

    Flores, G. & McDiarmid, R.W. (1989) Two new species of South American Centrolenella (Anura: Centrolenidae) related to C. mariae. Herpetologica, 45, 401–411.

    Ford, L.S. & Cannatella, D.C. (1993) The major clades of frogs. Herpetological Monographs, 7, 94–117.

    Frost, D.R. (2009) Amphibian Species of the World: an Online Reference. Version 5.3 (12 February, 2009). American Museum of Natural History, New York, USA. Available from http://research.amnh.org/herpetology/amphibia/index.php (accessed 06 February 2009).

    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. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370.

    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., & Wheeler, W.C. (2007) Is The Amphibian Tree of Life really fatally flawed? Cladistics, 23, 1–11.

    Gaige, H.T. (1926) A new frog from British Guiana. University of Michigan, Ocassional Papers of the Museum of Natural History, 176, 1–3.

    Gatesy, J., DeSalle, R. & Wahlberg, N. (2007) How many genes should a systematist sample? Conflicting insights from a phylogenomic matrix characterized by replicated incongruence. Systematic Biology, 56, 355–363.

    Goin, C.J. (1961) Three new centrolenid frogs from Ecuador. Zoologischer Anzeiger, 166, 95–104.

    Goin, C.J. (1963) A new Centrolenid frog from Venezuela. Acta Biologica Venezuelica, 3, 283–286.

    Goin, C.J. (1964) Distribution and synonymy of Centrolenella fleischmanni in northern South America. Herpetologica, 20, 1–8.

    Goin, C.J. (1966) A new frog of the genus Centrolenella from Suriname. Studies on the fauna of Suriname and other Guyanas, 8, 77–80.

    Goin, C.J. (1968) A New Centrolenid Frog from Guyana. Quarterly Journal of the Florida Academy of Sciences, 30, 115–118.

    Goin, C., Goin, O. & Zug, G. (1978) Introduction To Herpetology, ed 3. W. H. Freeman and Company, San Francisco, California.

    Grant, T., Bolívar-G., W. & Castro, F. (1998) The advertisement call of Centrolene geckoideum. Journal of Herpetology, 32, 452–455.

    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. & Wheeler, W.C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1–262.

    Guayasamin, J.M. & Barrio-Amorós, C. (2005) Combat behavior in Centrolene andinum (Rivero, 1968) (Anura: Centrolenidae). Salamandra, 41,153–155.

    Guayasamin, J.M. & Bonnacorso, E. (2004). A new species of glass frog (Centrolenidae: Cochranella) from the lowlands of northwestern Ecuador, with comments on the Cochranella granulosa group. Herpetologica, 60, 485–494.

    Guayasamin, J.M., Castroviejo-Fisher, S., Ayarzagüena, J., Trueb, L. & Vilà, C. (2008a) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574–595.

    Guayasamin, J.M., Cisneros-Heredia, D.F., Castroviejo-Fisher, S. (2008b) Taxonomic identity of Cochranella petersi Goin, 1961 and Centrolenella ametarsia Flores, 1987. Zootaxa, 1815, 25–34..

    Guayasamin, J.M., Lehr, E., Rodriguez, D. & Aguilar, C. (2006a) A new species of glass frog (Centrolenidae: Cochranella ocellata group) from central Peru. Herpetologica, 62, 163–172.

    Guayasamin, J.M., Bustamante, M.R., Almeida-Reinoso, D. & Funk, W.C. (2006b) Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics. Zoological Journal of the Linnean Society, 147, 489–513.

    Guayasamin, J.M. & Trueb, L. (2007) A new species of Glassfrog (Anura: Centrolenidae) from the lowlands of northwestern Ecuador, with comments on centrolenid osteology. Zootaxa, 27–45.

    Haas, A. (2003) Phylogeny of frogs as inferred from primarily larval characters (Amphibia : Anura). Cladistics, 19, 23–89.

    Harvey, M.B. (1996) A new species of glass frog (Anura: Centrolenidae: Cochranella) from Bolivia and the taxonomic status of Cochranella flavidigitata. Herpetologica, 52, 427–435.

    Harvey, M.B. & Noonan, B.P. (2005) Bolivian glass frogs (Anura : Centrolenidae) with a description of a new species from Amazonia. Proceedings of the Biological Society of Washington, 118, 428–441.

    Hayes, M.P. & Starrett, P.H. (1980) Notes on a collection of Centrolenid Frogs from the Colombian Chocó. Bulletin of the Southern California Academy of Science, 79, 89–96.

    Hedges, S.B., Duellman, W.E. & Heinicke, M.P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1–182.

    Heinicke, M.P., Duellman, W.E. & Hedges, S.B. (2007) Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. Proceedings of the National Academy of Sciences of the United States of America, 104, 10092–10097.

    Hennig, W. (1966) Phylogenetic systematics. University of Illinois Press, Urbana, Chicago & London, 284 pp.

    Hillis, D.M. & Bull, J.J. (1993) An Empirical-Test of Bootstrapping as a Method for Assessing Confidence in Phylogenetic Analysis. Systematic Biology, 42, 182–192.

    Hillis, D.M. & Wilcox, T.P. (2005) Phylogeny of New World true frogs. Molecular Phylogenetics and Evolution, 34, 299–314.

    Hooghiemstra, H., Wijninga, V.M., & Cleef, A.M. (2006) The paleobotanical record of Colombia: implications for biogeography and biodiversity. Annals of the Missouri Botanical Garden, 93, 297–325.

    Hoorn, C., Guerrero, J., Sarmiento, G.A. & Lorente, M.A. (1995) Andean tectonics as a cause for changing drainage patterns in Miocene northern South America. Geology, 23, 237–240.

    ICZN (1999) International Code of Zoological Nomenclature, ed 4. International Trust for Zoological Nomenclature, London, xxix + 306 pp.

    Jiménez de la Espada, M. (1872) Nuevos batracios americanos. Anales de la Sociedad Española de Historia Natural, 1, 85–88.

    Kok, P.J.R., & Castroviejo-Fisher, S. (2008) Glassfrogs (Anura: Centrolenidae) of Kaieteur National Park, Guyana, with notes on the distribution and taxonomy of some species of the family in the Guiana Shield. Zootaxa, 1680, 25–53.

    Kubicki, B. (2007) Ranas de vidrio de Costa Rica / Glass frogs of Costa Rica. Editorial INBio, Santo Domingo de Heredia, 304 pp.

    Langone, J.A. & Segalla, M.V. (1997) Ampliación de la distribución del género Allophryne (Amphibia, Anura, Allophrynidae). Cuadernos de Herpetología, 11, 87–88.

    Lescure, J. (1975) Contribution a l'étude des amphibiens de Guyane Francaise. V. Les centrolenidae. Bulletin de la Société Zoologique de France, 100, 385–394.

    Lescure, J. & Marty, C. (2001 "2000") Atlas des Amphibiens de Guyane. SPN/MNHN, Paris, Patrimoines Naturels 45, 388 pp.

    Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, ed 10, reformata. Tomus I. Laurentii Salvii, Holmiae, 828 pp.

    Lutz, A. (1925) Batraciens du Brésil. Comptes Rendus des Séances de la Société de Biologie et de Ses Filiales, Paris, 93, 137–139.

    Lutz, B. (1947). Trends towards non-aquatic and direct development in frogs. Copeia, 4, 242–252.

    Lutz, B. & Kloss, G.R. (1952) Anfíbios anuros do alto Solimões e Rio Negro, Apontamentos sôbre algunas formas e suas vicariantes. Memorias do Instituto Oswaldo Cruz, 50, 625–678.

    Lynch, J.D. (1973) The transition from archaic to advanced frogs. In: Vial, J.L. (Ed.), Evolutionary biology of anurans: Contemporary research on major problems. University of Missouri Press, Columbia, Missouri, pp. 133–182.

    Lynch, J.D. (1981) The identity of Hylopsis platycephala Werner, a centrolenid frog from northern Colombia. Journal of Herpetology, 15, 283–291.

    Lynch, J.D. (1990) A new ocellated frog (Centrolenidae) from western Colombia. Proceedings of the Biological Society of Washington, 103, 35–38.

    Lynch, J.D. (1993) A new centrolenid frog from the Andes of Western Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 18, 567–570.

    Lynch, J.D. & Duellman, W.E. (1973) A review of the Centrolenid frogs of Ecuador, with descriptions of new species. The University of Kansas Museum of Natural History Occasional Papers, 16, 1–66.

    Lynch, J.D. & Freeman, H.L. (1966) Systematic status of a South American frog, Allophryne ruthveni Gaige. The University of Kansas Museum of Natural History Miscellaneous Publications, 17, 493–502.

    Lynch, J.D., Ruiz-Carranza, P.M. & Rueda, J.V. (1983) Notes on the distribution and reproductive biology of Centrolene geckoideum Jimenez de la Espada in Colombia and Ecuador (Amphibia: Centrolenidae). Studies on Neotropical Fauna and Environment, 18, 239–243.

    Lynch, J.D. & Ruiz-Carranza, P.M. (1996) A remarkable new centrolenid frog from Colombia with a review of nuptial excrescences in the family. Herpetologica, 52, 525–535.

    Manzano, A.S., Fabrezi, M. & Vences, M. (2007) Intercalary elements, treefrogs, and the early differentiation of a complex system in the Neobatrachia. The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology, 290, 1551–1567.

    Mayr, E. (2004) What Makes Biology Unique? Considerations on the Autonomy of a Scientific Discipline. Cambridge University Press, Cambridge, U.K., 256 pp.

    McCranie, J.R. & Wilson, L.D. (2002) The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles, Ithaca, New York, 625 pp.

    McDiarmid, R.W. (1978) Evolution of parental care in frogs. In: Burghardt, G.M. & Bekoff, M. (Eds.), The Development of Behavior: Comparative and Evolutionary Aspects. Garland STPM Press, New York, pp. 127–147.

    McDiarmid, R.W. & Savage, J.M. (1984) Taxonomic status of the frog genus Centrolenella Noble (Anura: Centrolenidae). Journal of Herpetology, 18, 213–215.

    Moore, W.S. (1995) Inferring phylogenies from mtDNA variation: mitochondrial-gene trees versus nuclear-gene trees. Evolution, 49, 718–726.

    Müller, L. (1924) Neue LaubfrOsche aus dem Staate Santa Catherina, S.O. Brazilien. Zoolgischen Anzeiger, 59, 233–238.

    Myers, C.W. & Donnelly, M.A. (1997) A tepui herpetofauna on a granitic mountain (Tamacuari) in the borderland between Venezuela and Brazil: report from the Phipps Tapirapecó Expedition. American Museum Novitates, 3213, 1–71.

    Myers, C.W. & Donnelly, M.A. (2001) Herpetofauna of the Yutajé-Corocoro Massif, Venezuela: Second report from the Robert G. Goelet American Museum-Terramar Expedition to northwestern Tepuis. Bulletin of the American Museum of Natural History, 261, 1–85.

    Noble, G.K. (1920) Two new batrachians from Colombia. Bulletin of the American Museum of Natural History, 42, 441–446.

    Noble, G.K. (1922) The phylogeny of the salientia: I. the osteology and the thigh musculature; their bearing on the classification and phylogeny. Bulletin of the American Museum of Natural History, 46, 1–87.

    Noble, G.K. (1924) Some Neotropical batrachians preserved in the United States National Museum with a note on the secondary sexual characters of these and other amphibians. Proceedings of the Biological Society of Washington, 37, 65–72.

    Noble, G. K. (1931) The Biology of the Amphibia. McGraw-Hill, New York, 577 pp.

    Noonan, B.P. & Bonett, R.M. (2003) A new species of Hyalinobatrachium (Anura: Centrolenidae) from the highlands of Guyana. Journal of Herpetology, 37, 91–97.

    Padial, J.M. & De la Riva, I. (2007) Taxonomy, the Cinderella of science, hidden by its evolutionary stepsister. Zootaxa 1577, 1–2.

    Peters, W.C.H. (1873) Über eine neue Schildkrötenart, Cinosternon Effeldtii und einige andere neue oder weniger bekannte Amphibien. Monatsberichte der Königlich preussischen Akademic der Wissenschaften zu Berlin, 1873, 603–618.

    Peterson, A.T., Soberón, J. & Sanchez-Cordero V. (1999) Conservatism of ecological niches in evolutionary time. Science, 285, 1265–1267.

    Philippe, H. & Telford, M.J. (2006) Large-scale sequencing and the new animal phylogeny. Trends in Ecology & Evolution, 21, 614–620.

    Rafinesque, C.S. (1815) Analyse de nature, ou tableau de l’universe et des corps organisés. Jean Barravecchia, Palermo, 224 pp.

    Rivero, J.A. (1968) Los Centrolénidos de Venezuela. Memoria de la Sociedad de Ciencias Naturales La Salle, 28, 301–334.

    Rivero, J.A. (1985) Nuevos Centrolenidos de Colombia y Venezuela. Brenesia, 23, 335–373.

    Roelants, K., Gower, D.J., Wilkinson, M., Loader, S.P., Biju, S.D., Guillaume, K., Moriau, L. & Bossuyt, F. (2007) Global patterns of diversification in the history of modern amphibians. Proceedings of the National Academy of Sciences of the United States of America, 104, 887–892.

    Rokas, A., Williams, B.L., King, N. & Carroll, S.B. (2003) Genome-scale approaches to resolving incongruence in molecular phylogenies. Nature, 425, 798–804.

    Rueda-Almonacid, J.V. (1994) Estudio anatómico y relaciones sistemáticas de Centrolene geckoideum (Salientia: Anura: Centrolenidae). Trianea, 5, 133–187.

    Ruiz-Carranza, P.M., Hernández-Camacho, J. & Ardila-Robayo, M.C. (1986) Una nueva especie colombiana del género Centrolene Jiménez de la Espada 1872 (Amphibia: Anura) y redefinición del género. Caldasia, 15, 431–444.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1989) Una nueva especie de Centrolenella Noble, 1920 (Amphibia: Anura: Centrolenidae) de la Cordillera Oriental de Colombia. Trianea, 3, 67–75.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1991a) Ranas Centrolenidae de Colombia I. Propuesta de una nueva clasificación genérica. Lozania, 57, 1–30.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1991b) Ranas Centrolenidae de Colombia II. Nuevas especies de Centrolene de laCordillera Oriental y Sierra Nevada de Santa Marta. Lozania, 58, 1–26.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1991c) Ranas Centrolenidae de Colombia III. Nuevas especies de Cochranella del Grupo granulosa. Lozania, 59, 1–18.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1991d) Ranas Centrolenidae de Colombia IV. Nuevas especies de Cochranella del Grupo ocellata de la Cordillera Oriental. Lozania, 60, 1–13.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1995a) Ranas Centrolenidae de Colombia V. Cuatro nuevas especies de Cochranella de la Cordillera Central. Lozania, 62, 1–24.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1995b) Ranas Centrolenidae de Colombia VI. Cuatro nuevas especies de Cochranella de la Cordillera Occidental. Lozania, 63, 1–15.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1995c) Ranas Centrolenidae de Colombia VIII. Cuatro nuevas especies de Centrolene de la Cordillera Central. Lozania, 65, 1–16.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1996) Ranas Centrolenidae de Colombia IX. Dos nuevas especies del suroeste de Colombia. Lozania, 68, 1–11.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1997) Ranas Centrolenidae de Colombia X. Los centrolénidos de un perfil del flanco oriental de la Cordillera Central en el Departamento de Caldas. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 21, 541–553.

    Ruiz-Carranza, P.M. & Lynch, J.D. (1998) Ranas Centrolenidae de Colombia XI. Nuevas especies de ranas de cristal del genero Hyalinobatrachium. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 22, 571–586.

    Sanchiz, B. & De la Riva, I. (1993) Remarks on the tarsus of Centrolenid frogs (Amphibia: Anura). Graellsia, 49, 115–117.

    San Mauro, D., Gower, D.J., Oommen, O.V., Wilkinson, M. & Zardoya, R. (2004) Phylogeny of caecilian amphibians (Gymnophiona) based on complete mitochondrial genomes and nuclear RAG1. Molecular Phylogenetics and Evolution, 33, 413–427.

    Savage, J.M. (1967) A New Tree-Frog (Centrolenidae) from Costa Rica. Copeia, 1967, 325–331.

    Savage, J.M. (1973) The geographic distribution of frogs: patterns and predictions. In: Vial, J.L. (Ed.), Evolutionary biology of the anurans: contemporary research on major problems. University of Missouri Press, Columbia, Missouri, pp. 351–445.

    Savage, J. M. (2002) The amphibians and reptiles of Costa Rica: a herpetofauna between two continents, between two seas. The University of Chicago Press, Chicago and London, 934 pp.

    Savage, J.M. & Starrett, P.H. (1967) A new fringe-limbed tree-frog (family Centrolenidae) from lower Central America. Copeia, 1967, 604–609.

    Señaris, J.C. & Ayarzagüena, J. (2001) Una nueva especie de rana de cristal del género Hyalinobatrachium (Anura: Centrolenidae) del Delta del Río Orinoco, Venezuela. Revista de Biología Tropical, 49, 1083–1093.

    Señaris, J.C. (2001 “1999”) Una nueva especie de Hyalinobatrachium (Anura: Centrolenidae) de la Cordillera de la Costa, Venezuela. Memoria de la Fundación La Salle de Ciencias Naturales, 59, 133–147.

    Señaris, J.C. & Ayarzegüena, J. (2005) Revisión taxonómica de la Familia Centrolenidae (Amphibia; Anura) de Venezuela. Publicaciones del Comité Español del Programa Hombre y Biosfera – Red IberoMaB de la UNESCO, Sevilla, 337 pp.

    Schwalm, P.A. & McNulty, J.A. (1980) The morphology of dermal chromatophores in the infrared-reflecting glass-frog Centrolenella fleischmanni. Journal of Morphology, 163, 37–44.

    Shubin, N. & Alberch, P. (1986) A morphogenetic approach on the origin and basic organization of the tetrapod limb. In: Hecht, M., Wallace, B., & Prance, G. (Eds.), Evolutionary Biology. Plenum Press, New York, pp. 319– 387.

    Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 21, 2688–2690.

    Starrett, P.H. (1960) Description of tadpoles of Middle American frogs. Miscellaneous Publications of the Museum of Zoology University of Michigan, 110, 1–37.

    Starrett, P. & Savage, J.M. (1973) The systematic status and distribution of Costa Rican glass-frogs, genus Centrolenella (Family Centrolenidae), with description of a new species. Bulletin of the Southern California Academy of Sciences, 72, 57–78.

    Taylor, E.H. (1949) Costa Rican frogs of the genera Centrolene and Centrolenella. University of Kansas Science Bulletin, 33, 257–270.

    Taylor, E.H. (1951) Two new genera and a new family of tropical American frogs. Proceedings of the Biological Society of Washington, 64, 33–40.

    Taylor, E.H. (1952) A review of the frogs and toads of Costa Rica. University of Kansas Science Bulletin, 35, 577–942.

    Taylor, E.H. (1958) Notes on Costa Rican Centrolenidae with descriptions of new forms. University of Kansas Science Bulletin, 39, 41–68.

    Torres-Gastello, C.P., Suárez-Segovia, J. & Cisneros-Heredia, D.F. (2008) Cochranella erminea, a new species of Centrolenidae (Amphibia: Anura: Athesphatanura) from Amazonian Peru. Journal of the National Museum (Prague), Natural History Series, 176, 1–12.

    Trueb, L. (1973) Bones, frogs, and evolution. In: Vial, J.L. (Ed.), Evolutionary biology of the anurans: contemporary research on major problems. University of Missouri Press, Columbia, Missouri, pp. 65–132.

    Tschanz, C.M., Marvin, R.F., Cruz, J., Mehnert, H., & Cebulla, G. (1974) Geologic Evolution of the Sierra Nevada de Santa Marta area, Colombia. Geological Society of America Bulletin, 85, 273–284.

    Velásquez-Álvarez, A.A., Rada, M., Sánchez-Pacheco, S.J. & Acosta, A.R. (2007) A new species of glassfrog (Anura: Centrolenidae) from the western slope of the Cordillera Oriental, Colombia. South American Journal of Herpetology, 2, 191–197.

    Werner, F. (1894) Uber einige Novitaten der Herpetologischen Sammlung des Wiener zoolog. vergl. anatom. Instituts. Zoologischer Anzeiger, 17, 155–157.

    Whitfield, J.B. & Lockhart, P.J. (2007) Deciphering ancient rapid radiations. Trends in Ecology & Evolution, 22, 258–265.

    Wiens, J.J. (2007) Review of "The amphibian tree of life" by Frost et al. Quarterly Review of Biology, 82, 55–56.

    Wiens, J.J. (2008) Systematics and herpetology in the age of genomics. Bioscience, 58, 297–307.

    Wiens, J.J., Fetzner, J.W., Parkinson, C.L. & Reeder, T.W. (2005) Hylid frog phylogeny and sampling strategies for speciose clades. Systematic Biology, 54, 719–748.

    Wild, E.R. (1994) Two new species of centrolenid frogs from the Amazonian slope of the Cordillera-Oriental, Ecuador. Journal of Herpetology, 28, 299–310.

    Wilson, E.O. (2005) Systematics and the future of biology. Proceedings of the National Academy of Sciences of the United States of America, 102, 6520–6521.

    Zink, R.M., Barrowclough, G. F. (2008) Mitochondrial DNA under siege in avian phylogeography. Molecular Ecology, 17, 2107–2121.