Abstract
A phylogenetic analysis of sequences from 503 species of hylid frogs and four outgroup taxa resulted in 16,128 aligned sites of 19 genes. The molecular data were subjected to a maximum likelihood analysis that resulted in a new phylogenetic tree of treefrogs. A conservative new classification based on the tree has (1) three families composing an unranked taxon, Arboranae, (2) nine subfamilies (five resurrected, one new), and (3) six resurrected generic names and five new generic names. Using the results of a maximum likelihood timetree, times of divergence were determined. For the most part these times of divergence correlated well with historical geologic events. The arboranan frogs originated in South America in the Late Mesozoic or Early Cenozoic. The family Pelodryadidae diverged from its South American relative, Phyllomedusidae, in the Eocene and invaded Australia via Antarctica. There were two dispersals from South America to North America in the Paleogene. One lineage was the ancestral stock of Acris and its relatives, whereas the other lineage, subfamily Hylinae, differentiated into a myriad of genera in Middle America.
References
Agnolin, F. (2012) A new Calyptocephalidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Studia Geologica Salmanticensia, 48, 129–178.
Aitchison, J.C., Ali, J.R. & Davis, A.M. (2007) When and where did India and Asia collide? Journal of Geophysical Research, 112, B05423.
http://dx.doi.org/10.1029/2006JB004706
Ali, J.R. (2012) Colonizing the Caribbean: is the GAARlandia land-bridge hypothesis gaining a foothold? Journal of Biogeography, 39, 431–433.
http://dx.doi.org/10.1111/j.1365-2699.2011.02674.x
Almeida, C.J. & Cardoso, A.J. (1985) Variabilidade em medidas dos espermatozóides de Hyla fuscovaria (Amphibia, Anura) e seu significado taxonômico. Revista Brasileira de Biologia, 35, 387–391.
Almendáriz, A., Brito, J., Batallas, D. & Ron, S. (2014) Una especie nueva de rana arbórea del género Hyloscirtus (Amphibia: Anura: Hylidae) de la Cordillera del Cóndor. Papéis Avulsos de Zoologia, 54 (4), 33–49.
http://dx.doi.org/10.1590/0031-1049.2014.54.04
Alonso, R., Crawford, A.J. & Bermingham, E. (2011) Molecular phylogeny of an endemic radiation of Cuban toads (Bufonidae: Peltophryne) based on mitochondrial and nuclear genes. Journal of Biogeography, 39, 434–451.
http://dx.doi.org/10.1111/j.1365-2699.2011.02594.x
AmphibiaWeb (2014) AmphibiaWeb: Information on Amphibian Biology and Conservation. University of California, Berkeley, California. Available from: http://amphibiaweb.org/ (accessed 23 April 2015)
Ananjeva, N.B., Orlov, N.L., Khalikov, R.G., Darevsky, I.S., Ryabov, S.A. & Barabanov, A.V. (2006) The Reptiles of Northern Eurasia. Pensoft, Sofia, Bulgaria, 250 pp.
Anstis, M. (2013) Tadpoles and Frogs of Australia. New Holland Press, London, 829 pp.
Axelrod, D.I. (1958) Evolution of the Madro-Tertiary geoflora. Botanical Review, 24, 433–509.
http://dx.doi.org/10.1007/BF02872570
Axelrod, D.I. (1972) Edaphic aridity as a factor in angiosperm evolution. American Naturalist, 106, 311–320.
http://dx.doi.org/10.1086/282773
Bacon, C.D., Silvestro, D., Jaramillo, C., Smith, B.T., Chakrabarty, P. & Antonelli, A. (2015) Biological evidence supports an early and complex emergence of the Isthmus of Panama. Proceedings of the National Academy of Science USA, 112, 6110–6115.
http://dx.doi.org/10.1073/pnas.1423853112
Báez, A.M. (1981) Redescription and relationships of Saltenia ibanezi, a Late Cretaceous pipid frog from northwestern Argentina. Ameghiniana, 18, 127–154.
Báez, A. & Gómez, R.O. (2014) Is hyperossification concealing the phylogenetic signal in osteological traits in anurans? [abstract]. In: 74th Annual Meeting of Vertebrate Paleontology, November 5–8, 2014. Berlin, Germany.
Báez, A.M., Gómez, R.O., Ribeiro, L.C.B., Martinelli, A.G., Teixeira, V.P.A. & Ferraz, M.L.F. (2012) The diverse Cretaceous neobatrachian fauna of South America: Uberobatrachus carvalhoi, a new frog from the Maastrichtian Marília Formation, Minas Gerais, Brazil. Gondwana Research, 22, 1141–1150.
http://dx.doi.org/10.1016/j.gr.2012.02.021
Báez, A.M., Moura, G.J.B. & Gómez, R.O. (2009) Anurans from the Lower Cretaceous Crato Formation of northeastern Brazil; implications for the early divergence of neobatrachians. Cretaceous Research, 30, 829–846.
http://dx.doi.org/10.1016/j.cretres.2009.01.002
Báez, A.M. & Perí, S. (1989) Baurubatrachus pricei, nov. gen. et sp., un anuro del Cretácico Superior de Minas Gerais, Brasil. Anals da Academia Brasileira de Ciências, 61, 447–458.
Báez, A.M. & Rage, J.-C. (1998) Pipid frogs from the Upper Cretaceous of In Beceten, Niger. Paleontology, 41, 669–691.
Báez, A.M. & Trueb, L. (1997) Redescription of the Paleogene Shelania pascuali from Patagonia and its bearing on the relationships of fossil and Recent pipoid frogs. Scientific Papers, Natural History Museum. The University of Kansas, 4, 1–41.
Báez, A.M., Trueb, L. & Calvo, J.O. (2000) The earliest known pipoid frog from South America: a new genus from the Middle Cretaceous of Argentina. Journal of Vertebrate Paleontology, 20, 490–500.
http://dx.doi.org/10.1671/0272-4634(2000)020[0490:TEKPFF]2.0.CO;2
Barker, P.F. & Burrell, J. (1977) The opening of the Drake Passage. Marine Geology, 25, 15–34.
http://dx.doi.org/10.1016/0025-3227(77)90045-7
Batista, A., Hertz, A., Mebert, K., Köhler, G., Lotzkat, S., Ponce, M. & Vesely, M. (2014) Two new fringe-limbed frogs of the genus Ecnomiohyla (Anura: Hylidae) from Panama. Zootaxa, 3826 (3), 449–474.
http://dx.doi.org/10.11646/zootaxa.3826.3.2
Beck, R.M.D., Godthelp, H., Weisbecker, V., Archer, M. & Hand, S.J. (2008) Australia's oldest marsupial fossils and their biogeographical implications. PLoS ONE, 3 (3), e1858.
http://dx.doi.org/10.1371/journal.pone.0001858
Beurlen, K. (1970) Geologie von Brasilien. Beiträge zur Regionalen Geologie der Erde. Band 9. Borntraeger, Berlin, 444 pp.
Bokermann, W.C.A. & Sazima, I. (1973) Anfibios da Serra Serra di Cipó, Minas Gerais, Brasil. II. Duas espécies novas de Hyla (Anura: Hylidae). Revista Brasileira de Biologia, 33, 521–528.
Bossuyt, F. & Roelants, K. (2009) Anura. In: Hedges, S.B. & Kumar, S. (Eds.), Timetree of Life. Oxford University Press., New York, pp. 357–364.
Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. 2nd Ed. Taylor and Francis, London, xvi + 503 pp.
Brocchi, P. (1877) Note sur quelques batraciens hylaeformes recuilles au Mexique et au Guatemala. Bulletin de la Société Philomathique de Paris, Series 7, 1, 122–132.
Brochu, C.A. (1999) Phylogenetics, taxonomy, and historical biogeography of Alligatoroidea. Society of Vertebrate Paleontology Memoir, 6, 9–100.
http://dx.doi.org/10.2307/3889340
Brundin, L. (1965) On the real nature of transantarctic relationships. Evolution, 19, 496–505.
http://dx.doi.org/10.2307/2406246
Brunetti, A.E., Hermida, G.N., Luna, M.C., Barsotti, A.M.G., Jared, C., Antoniazzi, M.M., Rivera-Correa, M., Berneck, B.V.M. & Faivovich, J. (2015) Diversity and evolution of sexually dimorphic mental and lateral glands in Cophomantini treefrogs (Anura: Hylidae: Hylinae). Biological Journal of the Linnean Society, 114, 12–34.
http://dx.doi.org/10.1111/bij.12406
Caminer, M.A. & Ron, S.R. (2014) Systematics of treefrogs of the Hypsiboas calcaratus and Hypsiboas fasciatus species complex (Anura: Hylidae) with the description of four new species. ZooKeys, 370, 1–68.
http://dx.doi.org/10.3897/zookeys.370.6291
Campbell, J.A. (1999) Distribution patterns of amphibians in Middle America. In: Duellman, W.E. (Ed.), Patterns of Distribution of Amphibians, A Global Perspective. Johns Hopkins University Press, Baltimore, Maryland, pp. 111–210.
Campbell, J.A & Smith, E.N. (1992) A new frog of the genus Ptychohyla (Hylidae) from the Sierra de Santa Cruz, Guatemala, and description of a new genus of Middle American stream-breeding treefrogs. Herpetologica, 48, 153–167.
Cannatella, D.C. (1980) A review of the Phyllomedusa buckleyi group (Anura: Hylidae). Occasional Papers, Museum of Natural History University of Kansas, 87, 1–40.
Cannatella, D.C. (1982) Leaf-frogs of the Phyllomedusa perinesos Group (Anura: Hylidae). Copeia, 1982, 501–513.
http://dx.doi.org/10.2307/1444649
Castroviejo-Fisher, S., Guayasamin, J.M., Gonzales-Voyer, A. & Vilà, C. (2014) Neotropical diversification seen through glassfrogs. Journal of Biogeography, 41, 66–80.
http://dx.doi.org/10.1111/jbi.12208
Castroviejo-Fisher, S., Padial, J.M., Da Silva, H.R., Rojas-Runjaic, F.J.M., Medina-Méndez, E. & Frost, D.R. (2015) Phylogenetic systematics of egg-brooding frogs (Anura: Hemiphractidae) and the evolution of direct development. Zootaxa, 4004 (1), 1–75.
http://dx.doi.org/10.11646/zootaxa.4004.1.1
Coates, A.G. & Obando, J.A. (1996) The geological evolution of the Central American isthmus. In: Jackson, J.B.C., Budd, A.F. & Coates, A.G. (Eds.), Evolution & Environment in Tropical America. University of Chicago Press, Chicago, pp 21–56.
Cocroft, R.B. (1994) A cladistic analysis of chorus frog phylogeny (Hylidae: Pseudacris). Herpetologica 50, 420–437.
Coloma, L.A., Carvajal-Endara, S., Dueñas, J.F., Paredes-Recalde, A., Morales-Mite, M.A., Almeida-Reinoso, D., Tapia, E.E., Hutter, C.R., Toral-Contreras, E. & Guayasamin, J.M. (2012) Molecular phylogenetics of stream treefrogs of the Hyloscirtus larinopygion group (Anura: Hylidae), and description of two new species from Ecuador. Zootaxa, 3364 (4), 1–78.
http://dx.doi.org/10.11646/zootaxa.3686.4.3
Cook, L.G. & Crisp, M.D. (2005) Not so ancient: the extant crown group of Nothofagus represents a post-Gondwanan radiation. Proceedings of the Biological Society, 272, 2535–2544.
http://dx.doi.org/10.1098/rspb.2005.3219
Crawford, A.J. & Smith, E.N. (2005) Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution, 35, 536–555.
http://dx.doi.org/10.1016/j.ympev.2005.03.006
Crowhurst, P.V., Hill, K.C., Foster, D.A. & Bennett, A.P. (1996) Thermochronological and geochemical constraints on the tectonic evolution of northern Papua New Guinea. In: Hall, R. & Blundell, B.J. (Eds.), Tectonic Evolution of South-east Asia. Geological Society of London Special Publication, 106, 525–537.
http://dx.doi.org/10.1144/gsl.sp.1996.106.01.33
Darlington, P.J. (1965) Bio-geography of the southern end of the world. Harvard University Press, Cambridge, Massachusetts, 236 pp.
http://dx.doi.org/10.4159/harvard.9780674492073
Daudin, F.M. (1802) Histoire Naturelle des Rainettes, des Grenouilles, et des crapauds. Levrault, Paris, 71 pp.
Davies, H.L. (2012) The geology of New Guinea – the cordilleran margin of the Australian continent. Episodes, 35, 87–102.
de la Fuente, M.S., Sterli, J. & Maniel, I. (2014) Origin, Evolution and Biogeographic History of South American Turtles. Springer, London, 170 pp.
http://dx.doi.org/10.1007/978-3-319-00518-8
De la Riva, I., Köhler, J., Lötters, S. & Reiche, S. (2000) Ten years of research on Bolivian amphibians: updated checklist, distribution, taxonomic problems, literature, and iconography. Revista Española de Herpetología, 14, 18–164.
Donnelly, T.W. (1985) Mesozoic and Cenozoic plate evolution of the Caribbean Region. In: Stehli, F.G. & Webb, S.D. (Eds.), The Great American Biotic Interchange. Plenum Press, New York, pp. 89–121.
http://dx.doi.org/10.1007/978-1-4684-9181-4_4
Donovan, S.K. (2002) A karst of thousands: Jamaica’s limestone scenery. Geology Today, 18, 143–151.
http://dx.doi.org/10.1046/j.0266-6979.2003.00356.x
dos Santos, S.P., Ibáñez, R. & Ron, S.R. (2015) Systematics of the Rhinella margaritifera complex (Anura: Bufonidae) from western Ecuador and Panama with insights in the biogeography of Rhinella alata. ZooKeys, 501, 109–145.
http://dx.doi.org/10.3897/zookeys.501.8604
Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7, 214.
http://dx.doi.org/10.1186/1471-2148-7-214
Duellman, W.E. (1960) A distributional study of the amphibians of the Isthmus of Tehuantepec, Mexico. University of Kansas Publications Museum of Natural History, 13, 19–72.
Duellman, W.E. (1972a) South American frogs of the Hyla rostrata Group (Amphibia, Anura, Hylidae). Zoologische Mededelingen, Rijksmuseum van Natuurlijke Historie Leiden, 47, 177–192, pls. 1–3.
Duellman, W.E. (1972b) A review of the neotropical frogs of the Hyla bogotensis Group. Occasional Papers, Museum of Natural History University of Kansas, 11, 1–31.
Duellman, W.E. (1977) Liste der rezenten Amphibien und Reptilien Hylidae, Centrolenidae, Pseudidae. Das Tierreich, 95, xix + 225 pp.
Duellman, W.E. (1999) Distribution patterns of amphibians in South America. In: Duellman, W.E. (Ed.), Patterns of Distribution of Amphibians, A Global Perspective. Johns Hopkins University Press, Baltimore, Maryland, pp. 255–328.
Duellman, W.E. (2001) Hylid frogs of Middle America. Society for the Study of Amphibians and Reptiles, Ithaca, New York, xvi + 1159 pp.
Duellman, W.E. (2005) Cusco Amazónico. The Lives of Amphibians and Reptiles in an Amazonian Rainforest. Cornell University Press, Ithaca, New York, xv + 433 pp.
Duellman, W.E. (2015) Marsupial Frogs, Gastrotheca and Allied Genera. Johns Hopkins University Press, Baltimore, Maryland, xv + 407 pp.
Duellman, W.E., Cadle, J.E. & Cannatella, D.C. (1988) A new species of terrestrial Phyllomedusa (Anura: Hylidae) from southern Peru. Herpetologica, 44, 91–95.
Duellman, W.E., De la Riva, I. & Wild, E.R. (1997) Frogs of the Hyla armata and Hyla pulchella groups in the Andes of South America, with definitions and analyses of phylogenetic relationships of Andean groups of Hyla. Scientific Papers, Natural History Museum University of Kansas, 3, 1–41.
http://dx.doi.org/10.5962/bhl.title.48689
Duellman, W.E. & Hillis, D.M. (1990) Systematics of frogs of the Hyla larinopygion Group. Occasional Papers, Museum of Natural History University of Kansas, 134, 1–23.
Duellman, W.E. & Lehr, E. (2009) Terrestrial Breeding Frogs (Strabomantidae) in Peru. Natur und Tier-Verlag, Münster, Germany, 382 pp.
Duellman, W.E. & Mendelson, J.R. (1995) Amphibians and reptiles from northern Departamento Loreto, Peru: taxonomy and biogeography. University of Kansas Science Bulletin, 55, 329–376.
Duellman, W.E. & Sweet, S.S. (1999) Distribution patterns of amphibians in the Nearctic Region of North America. In: Duellman, W.E. (Ed.), Patterns of Distribution of Amphibians, A Global Perspective. Johns Hopkins University Press, Baltimore, Maryland, pp. 31–109.
Duellman, W.E. & Trueb, L. (1983) Frogs of the Hyla columbiana group: taxonomy and phylogenetic relationships. In: Rhodin, A.G.J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary Biology. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, pp. 33–51.
Duellman, W.E. & Wiens, J.J. (1992) The status of the hylid frog genus Ololygon and the recognition of Scinax Wagler, 1830. Occasional Papers, Museum of Natural History University of Kansas, 151, 1–23.
Duméril, A.M.C. & Bibron, G. (1841) Erpétologie Générale ou Histoire Naturelle Compléte des Reptiles. Vol. 8. Roret, Paris, 792 pp.
Dunbar, C.O. (1961) Historical Geology, 2nd Ed. John Wiley and Sons, New York, 500 pp.
Duque-Caro, H. (1990) Neogene stratigraphy, paleoceanography and paleobiogeography in northwest South America and the evolution of the Panama seaway. Palaeogeography, Palaeoclimatology, Palaeoecology, 77, 203–234.
http://dx.doi.org/10.1016/0031-0182(90)90178-A
Durham, J.W., Arellano, A.R.V. & Peck Jr., J.H. (1955) Evidence for no Cenozoic Isthmus of Tehuantepec seaways. Bulletin of the Geological Society of America, 66, 977–992.
http://dx.doi.org/10.1130/0016-7606(1955)66[977:EFNCIO]2.0.CO;2
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.
http://dx.doi.org/10.1093/nar/gkh340
Estes, R. (1977) Relationships of the South African fossil frog Eoxenopoides reuningi (Anura, Pipidae). Annals of the South African Museum, 73, 49–80.
Estes, R. & Báez, A.M. (1985) Herpetofaunas of North and South America during the late Cretaceous and Cenozoic: Evidence for Interchange? In: Stehli, F.G. & Webb, S.D. (Eds.), The Great American Biotic Interchange. Plenum Press, New York, pp. 139–197.
http://dx.doi.org/10.1007/978-1-4684-9181-4_6
Estes, R. & Reig, O.A. (1973) The early fossil record of frogs. A review of the evidence. In: Vial, J.L. (Ed.), Evolutionary Biology of the Anurans. University of Missouri Press, Columbia, Missouri, pp. 11–63.
Evans, S.E., Groenke, J.R., Jones, M.E.H., Turner, A.H. & Krause, D.W. (2014) New material of Beelzebufo, a hyperossified frog (Amphibia: Anura) from the Late Cretaceous of Madagascar. PLoS ONE, 9 (1), e87236.
http://dx.doi.org/10.1371/journal.pone.0087236
Faivovich, J. (2002) A cladistic analysis of Scinax (Anura: Hylidae). Cladistics, 18, 367–393.
http://dx.doi.org/10.1111/j.1096-0031.2002.tb00157.x
Faivovich, J., Haddad, C.F.B., Baêta, D., Jungfer, K.-H., Álvares, G.F.R., Brandão, R.A., Sheil, C., Barrientos, L.S., Barrio-Amorós, C.L., Cruz, C.A.G. & Wheeler, W.C. (2010) The phylogenetic relationships of the charismatic poster frogs, Phyllomedusinae (Anura, Hylidae). Cladistics, 26, 227–261.
http://dx.doi.org/10.1111/j.1096-0031.2009.00287.x
Faivovich, J., Haddad, C.F.B., García, 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.
http://dx.doi.org/10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2
Faivovich, J., McDiarmid, R.W. & Myers, C.W. (2013) Two new species of Myersiohyla (Anura: Hylidae) from Cerro de la Neblina, Venezuela, with comments on other species of the genus. American Museum Novitates, 3792, 1–63.
http://dx.doi.org/10.1206/3792.1
Farris, D.W., Jaramillo, C., Bayona, G., Restrepo-Moreno, S.A., Montes, C., Cardona, A., Mora, A., Speakman, R.J., Glascock, M.D. & Valencia, V. (2011) Fracturing of the Panamanian Isthmus during initial collision with South America. Geology, 39, 1007–1010.
http://dx.doi.org/10.1130/G32237.1
Fouquette Jr., M.J. & Delahoussaye, A.J. (1977) Sperm morphology in the Hyla rubra group (Amphibia, Anura, Hylidae) and its bearing on generic status. Journal of Herpetology, 11, 387–396.
http://dx.doi.org/10.2307/1562720
Fouquette Jr., M.J. & Dubois, A. (2014) A Checklist of North American Amphibians and Reptiles. Seventh Edition. Volume 1—Amphibians. Xlibris LLC, Bloomington, Indiana, 586 pp.
Freitas, R.O. de. (1951) Ensaio sobre de a tectonica moderno do Brasil. Universidade São Paulo Facultad de Filosofia, Ciencias, Letras Boletim de Geologia, 130, 1–120.
Frost, D.R. (2015) Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History, New York, USA. Available from: http://research.amnh.org/herpetology/amphibia/index.html (accessed 15 September 2015)
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, 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, 1–291.
http://dx.doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2
Funkhouser, A. (1957) A review of the neotropical tree-frogs of the genus Phyllomedusa. Occasional Papers of the Natural History Museum Stanford University, 5, 2–90.
Galbreath, K.E. & Hoberg, E.P. (2012) Return to Beringia: parasites reveal cryptic biogeographic history of North American pikas. Proceedings of the Royal Society B, 279, 371–378.
http://dx.doi.org/10.1098/rspb.2011.0482
Galloway, R.W. & Kemp, E.M. (1981) Late Cenozoic environments in Australia. In: Keast, A. (Ed.), Ecological Biogeography of Australia. W. Junk, The Hague, pp. 51–80.
http://dx.doi.org/10.1007/978-94-009-8629-9_4
Galvez, V.J. & Morca, J. (1994) Geologia. In: Leyva, P. (Ed.), Colombia Pacífico. Vol. 1. Financiera Energética Nacional, Bogotá, pp. 80–95.
Gamble, T., Bauer, A.M., Greenbaum, E. & Jackman, T.R. (2008) Evidence for Gondwanan vicariance in an ancient clade of gecko lizards. Journal of Biogeography, 35, 88–104.
http://dx.doi.org/10.1111/j.1365-2699.2007.01770.x
Gansser, R. (1954) The Guiana Shield (S. America). Eclogae Geologicae Helvetiae, 44, 77–112.
Gehara, M., Crawford, A.J., Orrico, V.G.D., Rodríguez, A., Lötters, S., Fouquet, A., Barrientos, L.S., Brusquetti, F., De la Riva, I., Ernst, R., Urrutia, G.G., Glaw, F., Guayasamin, J.M., Hölting, M., Jansen, M., Kok, P.J.R., Kwet, A., Lingnau, R., Lyra, M., Moravec, J., Pombal, J.P., Jr., Rojas-Runjaic, F.J.M., Schulze, A., Señaris, J.C., Solé, M., Rodrigues, M.T., Twomey, E., Haddad, C.F.B., Vences, M. & Köhler, J. (2014) High levels of diversity uncovered in a widespread nominal taxon: continental phylogeography of the neotropical tree frog Dendropsophus minutus. PLoS One, 9 (9), e103958.
http://dx.doi.org/10.1371/journal.pone.0103958
Gregory-Wodzicki, K.M. (2000) Uplift history of the central and northern Andes: a review. Bulletin of the Geological Society of America, 112, 1091–1105.
http://dx.doi.org/10.1130/0016-7606(2000)112<1091:UHOTCA>2.0.CO;2
Guayasamin, J.M., Rivera-Correa, M., Arteaga, A., Culebras, J., Bustamante, L., Pyron, R.A., Peñafiel, N., Morochz, C. & Hutter, C.R. (2015) Molecular phylogeny of stream treefrogs (Hylidae: Hyloscirtus bogotensis Group), with a new species from the Andes of Ecuador. Neotropical Biodiversity, 1, 2–21.
http://dx.doi.org/10.1080/23766808.2015.1074407
Günther, A.C.L.G. (“1858” 1859) Catalogue of the Batrachia Salientia in the Collection of the British Museum. Taylor and Francis, London, xvi + 160 pp.
Günther, R. & Richards, S.J. (2005) Three new mountain stream dwelling Litoria (Amphibia: Anura: Hylidae) from western New Guinea. Russian Journal of Herpetology, 12, 195–212.
Haddad, C.F.B., Pombal Jr., J.P. & Batistic, R.F. (1994) Natural hybridization between diploid and tetraploid species of leaf-frogs, genus Phyllomedusa (Amphibia). Journal of Herpetology, 28, 425–430.
http://dx.doi.org/10.2307/1564953
Haffer, J. (1974) Avian speciation in tropical South America. Publications of the Nuttall Ornithological Club, 14, 1–390.
Hanken, J. & Wake, D.B. (1982) Genetic differentiation among plethodontid salamanders (genus Bolitoglossa) in Central and South America: implications for the South American invasion. Herpetologica, 38, 272–287.
Haq, B.U., Hardenbol, J. & Vail, P.R. (1987) Chronology of fluctuating sea levels since the Triassic. Science, 235, 1156–1167.
http://dx.doi.org/10.1126/science.235.4793.1156
Hardy Jr., J.D. & Burrows, R.J. (1986) Systematic status of the Spring Peeper, Hyla crucifer (Amphibia: Hylidae). Bulletin of the Maryland Herpetological Society, 22, 68–89.
Hedges, S.B. (1986) An electrophoretic analysis of Holarctic hylid frog evolution. Systematic Zoology, 35, 1–21.
http://dx.doi.org/10.2307/2413287
Hedges, S.B. (1992) The number of replications needed for accurate estimation of the bootstrap p-value in phylogenetic studies. Molecular Biology and Evolution, 9, 366–369.
Hedges, S.B. (1996) Historical biogeography of West Indian vertebrates. Annual Review of Ecology and Systematics, 27, 163–196.
http://dx.doi.org/10.1146/annurev.ecolsys.27.1.163
Hedges, S.B. (2001) Caribbean biogeography: an outline. In: Woods, C.A. & Sergile, F.E. (Eds.), Biogeography of the West Indies: Patterns and Perspectives. CRC Press, Boca Raton, Florida, pp. 15–33.
Hedges, S.B. (2006) Paleogeography of the Antilles and origin of West Indian terrestrial vertebrates. Annals of the Missouri Botanical Garden, 93, 231–244.
http://dx.doi.org/10.3417/0026-6493(2006)93[231:POTAAO]2.0.CO;2
Hedges, S.B. (2013) Revision shock in taxonomy. Zootaxa, 3681 (3), 297–298.
http://dx.doi.org/10.11646/zootaxa.3681.3.11
Hedges, S.B. (2014) The high-level classification of skinks (Reptilia, Squamata, Scincomorpha). Zootaxa, 3765 (4), 317–338.
http://dx.doi.org/10.11646/zootaxa.3765.4.2
Hedges, S.B. & Conn, C.E. (2012) A new skink fauna from Caribbean islands (Squamata, Mabuyidae, Mabuyinae). Zootaxa, 3288, 1–244.
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, 104, 10092–97.
http://dx.doi.org/10.1073/pnas.0611051104
Heinicke, M.P., Duellman, W.E., Trueb, L., Means, D.B., MacCulloch, R.D. & Hedges, S.B. (2009) A new frog family (Anura: Terrarana) from South America and an expanded direct-developing clade revealed by molecular phylogeny. Zootaxa, 2211, 1–35.
Hillis, D.M. & de Sa, R. (1988) Phylogeny and taxonomy of the Rana palmipes Group (Salientia: Ranidae). Herpetological Monographs, 2, 1–26.
http://dx.doi.org/10.2307/1467024
Hillis, D.M. & Wilcox, T.P. (2005) Phylogeny of the New World true frogs (Rana). Molecular Phylogenetics and Evolution, 34, 299–314.
http://dx.doi.org/10.1016/j.ympev.2004.10.007
Hoffmann, C.K. (1878) Klassen und Ordnungen der Amphibien wissenschaftlich dargestelldt in Wort un Bild. In: Bronn, H.G. (Ed.), Die Klassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelldt in Wort und Bild. Vol. 6. Pt 2. C.F. Winter, Leipzig and Heidelberg, pp. 1–726.
Holman, J.A. (1961) A new hylid genus from the Lower Miocene of Florida. Copeia, 1961, 354–355.
http://dx.doi.org/10.2307/1439818
Hoorn, C. (2006) The birth of the mighty Amazon. Scientific American, 294, 52–59.
http://dx.doi.org/10.1038/scientificamerican0506-52
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.
http://dx.doi.org/10.1130/0091-7613(1995)023<0237:ATAACF>2.3.CO;2
Hua, X., Fu, C., Li, J., Nieto Montes de Oca, A. & Wiens, J.J. (2009) A revised phylogeny of holarctic treefrogs (genus Hyla) based on nuclear and mitochondrial DNA sequences. Herpetologica, 65, 246–259.
http://dx.doi.org/10.1655/08-058R1.1
Ickert-Bond, S.M., Murray, D.F. & DeChaime, E. (2009) Contrasting patterns of plant distribution in Beringia. Alaska Park Science, 8, 26–32.
Iturralde-Vinent, M.A. & MacPhee, R.D.E. (1999) Paleogeography of the Caribbean region: implications for Cenozoic biogeography. Bulletin of the American Museum of Natural History, 238, 1–95.
IUCN (2014) The IUCN Red List of Threatened Species. International Union for the Conservation of Nature, Cambridge, UK. Available from: http://www.iucnredlist.org (accessed 1 January 2014)
Jaillard, E., Hérail, G., Monfret, T., Diaz-Martinez, E., Baby, P., Lavenu, A. & Dumont, J.F. (2000) Tectonic evolution of the Andes of Ecuador, Peru, Bolivia, and northernmost Chile. In: Cordani, U.G., Milani, E.J., Thomaz-Filho, A. & Campos, D.A. (Eds.), Tectonic Evolution of South America. 31st International Geological Congress, Rio de Janeiro, pp. 481–559.
Jeannel, R. (1967) Biogeographie de l’Amerique austral. In: Delamere Dehoutteville, C. & Rapaport, E. (Eds.), Biologie de l’Amerique Austral. Vol. 3. C. N. R. S. Groupe Francais Argiles C. R. Reun Etud, Paris, pp. 401–460.
Jungfer, K.-H., Faivovich, J., Padial, J.M., Castroviejo-Fisher, S., Lyra, M.M., Berneck, B.V.M., Iglesias, P.P., Kok, P.J.R., MacCulloch, R.D., Rodrigues, M.T., Verdade, V.K., Torres Gastello, C.P., Chaparro, J.C., Valdujo, P.H., Reichle, S., Moravek, J., Gvoždík, V., Gagliardi-Urrutia, G., Ernst, R., De la Riva, I., Means, D.B., Lima, A.P., Señaris, J.C., Wheeler, W.C. & Haddad, C.F.B. (2013) Systematics of spiny-backed treefrogs (Hylidae: Osteocephalus): an Amazonian puzzle. Zoologica Scripta, 42, 351–380.
http://dx.doi.org/10.1111/zsc.12015
King, P.B. (1958) Evolution of modern surface features in western North America. In: Hubbs, C.L. (Ed.), Zoogeography. American Association for the Advancement of Science, Washington, DC, pp. 3–60.
Kluge, A.G. (1979) The gladiator frogs of Middle America and Colombia—a reevaluation of their systematics (Anura: Hylidae). Occasional Papers of the Museum of Zoology, University of Michigan, 688, 1–24.
Köhler, J., Koscinski, D., Padial, J.M., Chaparro, J.C., Handford, P., Lougheed, S.C. & De la Riva, I. (2010) Systematics of Andean gladiator frogs of the Hypsiboas pulchellus species group (Anura, Hylidae). Zoologica Scripta, 39, 572–590.
http://dx.doi.org/10.1111/j.1463-6409.2010.00448.x
Kolenc, F., Borteiro, C. & Tedros, M. (2003) La larva de Hyla uruguaya Schmidt, 1944 (Anura: Hyodae), con comentarios sobre su biología en Uruguay y su status taxonómico. Cuadernos de Herpetología, 17, 87–100.
Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29, 1695–1701.
http://dx.doi.org/10.1093/molbev/mss020
Lanfear, R., Calcott, B., Kainer, D., Mayer, C. & Stamatakis, A. (2014) Selecting optimal partitioning schemes for phylogenomic datasets. BMC Evolutionary Biology, 14, 82.
http://dx.doi.org/10.1186/1471-2148-14-82
Latrubesse, E.M., Cozzuol, M., da Silva-Caminha, S.A.F., Rigsby, C.A., Absy, M.L. & Jaramillo, C. (2010) The Late Miocene paleogeography of the Amazon Basin and the evolution of the Amazon River system. Earth-Science Reviews, 99, 99–124.
http://dx.doi.org/10.1016/j.earscirev.2010.02.005
Lawyer, L.A. & Gahagan, L.M. (1998) Opening of the Drake Passage and its impact on Cenozoic ocean circulation. In: Crowley, T.J. & Burke, K.C. (Eds.), Tectonic Boundary Conditions for Climate Reconstruction. Oxford Monographs on Geology and Geophysics, 39, Oxford, UK, pp. 212–223.
Lemmon, E.M., Lemmon, A.R. & Cannatella, D.C. (2007) Geological and climatic forces driving speciation in the continentally distributed trilling chorus frogs (Pseudacris). Evolution, 61, 2086–2103.
http://dx.doi.org/10.1111/j.1558-5646.2007.00181.x
Li, Z.X. & Powell, C.McA. (2001) An outline of the palaeogeographic evolution of the Australasian region since the beginning of the Neoproterozoic. Earth-Science Reviews, 53, 237–277.
http://dx.doi.org/10.1016/S0012-8252(00)00021-0
Lötters, S., Reichle, S., Faivovich, J. & Bain, R.H. (2005) The stream-dwelling tadpole of Hyloscirtus charazani (Anura: Hylidae) from Andean Bolivia. Studies on Neotropical Fauna and Environment, 40, 181–185.
http://dx.doi.org/10.1080/01650520500309735
Lutz, B. (1966) Pithecopus ayeaye, a new Brazilian hylid with vertical pupils and grasping feet. Copeia, 1966, 236–240.
http://dx.doi.org/10.2307/1441130
Lynch, J.D. & Duellman, W.E. (1997) Frogs of the genus Eleutherodactylus in western Ecuador. Special Publications, Natural History Museum, University of Kansas, 23, 1–236.
Macqueen, P., Seddon, J.M., Austin, J.J., Hamilton, S. & Goldizen, A.W. (2010) Phylogenetics of the pademelons (Macropodidae: Thylogale) and historical biogeography of the Australo-Papuan region. Molecular Phylogenetics and Evolution, 57, 1134–1148.
http://dx.doi.org/10.1016/j.ympev.2010.08.010
McCranie, J.R. & Wilson, L.D. (2002) The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles, Ithaca, New York, x + 625 pp.
McKenna, M.C. (1975) Fossil mammals and Early Eocene North Atlantic land continuity. Annals of the Missouri Botanical Garden, 62, 335–353.
http://dx.doi.org/10.2307/2395200
McLoughlin, S. (2001) The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany, 49, 271–300.
http://dx.doi.org/10.1071/BT00023
Mendelson, J.R., III, Savage, J.M., Griffith, E., Ross, H., Kubicki, B. & Gagliardo, R. (2008) Spectacular new gliding species of Ecnomiohyla (Anura: Hylidae) from central Panama. Journal of Herpetology, 42, 750–759.
http://dx.doi.org/10.1670/08-025R1.1
Menzies, J. (2006) The Frogs of New Guinea and the Solomon Islands. Pensoft Publishers, Sofia, Bulgaria, x + 345 pp.
Menzies, J.I. (2014) Notes on Nyctimystes (Anura: Hylidae), tree frogs of New Guinea, with descriptions of four new species. Alytes, 30, 42–68.
Menzies, J.I. & Tippet, J. (1976) Chromosome numbers of Papuan hylid frogs and the karyotype of Litoria infrafrenata (Amphibia, Anura, Hylidae). Journal of Herpetology, 10, 167–173.
http://dx.doi.org/10.2307/1562977
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, Louisiana, pp. 1–8.
http://dx.doi.org/10.1109/GCE.2010.5676129
Min, M.S., Yang, S.Y., Bonett, R.M., Vieites, D.R., Brandon, R.A. & Wake, D.B. (2005) Discovery of the first Asian plethodontid salamander. Nature, 435, 87–90.
http://dx.doi.org/10.1038/nature03474
Mittleman, M.B. & List, J.C. (1953) The generic differentiation of the swamp treefrogs. Copeia, 1953, 80–83.
http://dx.doi.org/10.2307/1440129
Montes, C., Cardona, A., Jaramillo, C., Pardo, A., Silva, J.C., Valencia, V., Ayala, C., Pérez-Angel, L.C., Rodriguez-Parra, L.A., Ramirez, V. & Niño, H. (2015) Middle Miocene closure of the Central American Seaway. Science, 348 (6231), 226–229.
http://dx.doi.org/10.1126/science.aaa2815
Mora, A., Baby, P., Roddaz, M., Parra, M., Brusset, S., Hermoza, W. & Espurt, N. (2010) Tectonic history of the Andes and sub-Andean zones: implications for the development of the Amazon drainage basin. In: Hoorn, C. & Wesselingh, F.P. (Eds.), Amazonia, Landscape and Species Evolution: A Look into the Past. Wiley–Blackwell Publishing, Ltd., Hoboken, New Jersey, pp. 38–60.
Muzzoppapa, P. & Báez, A.M. (2009) Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus. Ameghiniana, 46, 113–125.
Nascimento, F.A.C.d. & Skuk, G.O. (2007) Description of the tadpole of Hylomantis granulosa (Anura: Hylidae). Zootaxa, 1663, 59-65.
Nieden, F. (1923) Anura I. Subordo Aglossa und phaneroglossa, Sectio 1, Arcifera. Das Tierreich, 46, xxxii + 584.
Ortega-Andrade, H.M., Bermingham, J., Aulestia, C. & Paucar, C. (2010) Herpetofauna of the Bilsa Biological Station, Province of Esmeraldas, Ecuador. Checklist, 6, 119–154.
Padial, J.M., Grant, T. & Frost D.R. (2014) Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa, 3825 (1), 1–132.
http://dx.doi.org/10.11646/zootaxa.3825.1.1
Pauly, G.B., Hillis, D.M. & Cannatella, D.C. (2004) The history of a Nearctic colonization: molecular phylogenetics and biogeography of the Nearctic toads (Bufo). Evolution, 58, 2517–2535.
http://dx.doi.org/10.1111/j.0014-3820.2004.tb00881.x
Peixoto, O.L. (1987) Characterização do grupo “perpusillus” e revalidação da posição taxonómica de Ololygon perpusilla perpusilla e Ololygon perpusilla v-signata (Amphibia, Anura, Hyidae). Arquivos da Universidade Federal Rural do Rio de Janeiro, 10, 37–49.
Pimenta, B.V.S., Faivovich, J. & Pombal, J. (2007a) On the identity of Hyla strigilata Spix, 1824 (Anura: Hylidae): redescription and neotype designation for a “ghost” taxon. Zootaxa, 1441, 35–49.
Pimenta, B.V.S., Nunes, I. & Cruz, C.A.G. (2007b) Notes on the poorly known phyllomedusine frog Hylomantis aspera Peters 1872 (Anura, Hylidae). South American Journal of Herpetology, 2, 206–214.
http://dx.doi.org/10.2994/1808-9798(2007)2[206:NOTPKP]2.0.CO;2
Pinto-Sánchez, N.R., Ibáñez, R., Madriñán, S., Sanjur, O.I., Bermingham, E. & Crawford, A.J. (2012) The Great American biotic interchange in frogs: multiple and early colonization of Central America by the South American genus Pristimantis (Anura: Craugastoridae). Molecular Phylogenetics and Evolution, 62, 954–972.
http://dx.doi.org/10.1016/j.ympev.2011.11.022
Pittman, W.C., III, Cande, S., LaBrecque, J. & Pindell, J. (1993) Fragmentation of Gondwana: the separation of Africa from South America. In: Goldblatt, P. (Ed.), Biological Relationships between Africa and South America. Yale University Press, New Haven, Connecticut, pp. 15–34.
Poinar, G., Jr. & Wake, D.B. (2015) Paleoplethodon hispaniolae gen. n., sp. n. (Amphibia: Caudata), a fossil salamander from the Caribbean. Palaeodiversity, 8, 21–28.
Pombal Jr., J.P. & Gordo, M. (1991) Duas novas espécies de Hyla da floresta atlântica no Estado do São Paulo (Amphibia, Anura). Memorias do Instituto Butantan, 53, 135–144.
Pombal Jr., J.P. & Haddad, C.F.B. (1992) Espécies de Phyllomedusa do grupo burmeisteri do Brasil oriental, com descrição de uma espécie nova (Amphibia, Hylidae). Revista Brasileira de Biologia, 52, 217–229.
Poole, I. & Cantrill, D.J. (2006) Cretaceous and Cenozoic vegetation of Antarctica integrating the fossil wood record. In: Francis, J.E. Pirrie, D. & Crame, J.A. (Eds.), Cretaceous-Tertiary High-Latitude Palaeoenvironments, James Ross Basin, Antarctica. Geological Society, London, Special Publications 258, London, pp. 63–81.
Pyron, R.A. (2014) Biogeographic analysis reveals ancient continental vicariance and recent oceanic dispersal in amphibians. Systematic Biology, 63, 779–797.
http://dx.doi.org/10.1093/sysbio/syu042
Pyron, R.A., Burbrink, F.T., Colli, G.R., Montes de Oca, A.N., Vitt, L.J., Kuczynski, C.A. & Wiens, J.J. (2011) The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees. Molecular Phylogenetics and Evolution, 58, 329–342.
http://dx.doi.org/10.1016/j.ympev.2010.11.006
Pyron, R.A., Burbrink, F.T. & Wiens, J.J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13, 93.
http://dx.doi.org/10.1186/1471-2148-13-93
Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2,800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543–583.
http://dx.doi.org/10.1016/j.ympev.2011.06.012
Pyron, R.A. & Wiens, J.J. (2013) Large-scale phylogenetic analyses reveal the causes of high tropical amphibian diversity. Proceedings of the Royal Society B, 280, 1–10.
http://dx.doi.org/10.1098/rspb.2013.1622
Richards, S.J. & Oliver, P.M. (2006) Two new species of large green canopy-dwelling frogs (Anura: Hylidae: Litoria) from Papua New Guinea. Zootaxa, 1295, 41–60.
Rivera-Correa, M. & Faivovich, J. (2013) A new species of Hyloscirtus (Anura: Hylidae) from Colombia, with a rediagnosis of Hyloscirtus larinopygion (Duellman, 1973). Herpetologica, 69, 298–313.
http://dx.doi.org/10.1655/HERPETOLOGICA-D-12-00059
Roca, A.L., Bar-Gal, G.K., Eizirik, E., Helgen, K.M., Maria, R., Springer, M.S., O’Brien, S.J. & Murphy, W.J. (2004) Mesozoic origin for West Indian insectivores. Nature, 429, 649–651.
http://dx.doi.org/10.1038/nature02597
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 USA, 104, 887–892.
http://dx.doi.org/10.1073/pnas.0608378104
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–42.
http://dx.doi.org/10.1093/sysbio/sys029
Rosen, D.E. (1975) A vicariance model of Caribbean biogeography. Systematic Zoology, 24, 431–464.
http://dx.doi.org/10.2307/2412905
Ruane, S., Pyron, R.A. & Burbrink, F.T. (2011) Phylogenetic relationships of the Cretaceous frog Beelzebufo from Madagascar and the placement of fossil constraints based on temporal and phylogenetic evidence. Journal of Evolutionary Biology, 24, 274–285.
http://dx.doi.org/10.1111/j.1420-9101.2010.02164.x
Ruibal, R. & Shoemaker, V. (1984) Osteoderms in anurans. Journal of Herpetology, 18, 313–328.
http://dx.doi.org/10.2307/1564085
Sánchez, D.A. (2010) Larval development and synapomorphies for species groups of Hyloscirtus Peters, 1882 (Anura: Hylidae: Cophomantini). Copeia, 2010, 351–363.
http://dx.doi.org/10.1643/CH-10-010
Sanchiz, B. (1998a) Salientia. Encyclopedia of Paleoherpetology. Verlag Dr. Friedrich Pfeil, München, xii + 275 pp.
Sanchiz, B. (1998b) Vertebrates from the Early Miocene lignite deposits of the opencast mine Oberdorf (western Styrian Basin, Austria). 2. Amphibia. Annalen des Naturhistorischen Museums in Wien, 1999, 13–29.
Sanmartín, I. & Ronquist, F. (2004) Southern Hemisphere biogeography inferred by event-based models: Plant versus animal patterns. Systematic Biology, 53, 216–243.
http://dx.doi.org/10.1080/10635150490423430
Santos, J.C., Coloma, L.A., Summers, K., Caldwell, J.P., Ree, R. & Cannatella, D.C. (2009) Amazonian amphibian diversity is primarily derived from late Miocene Andean lineages. PLoS Biology, 7 (3), e1000056.
http://dx.doi.org/10.1371/journal.pbio.1000056
Savage, J.M. & Kubicki, B. (2010) A new species of fringe-limbed frog, genus Ecnomiohyla (Anura: Hylidae) from the Atlantic slope of Costa Rica, Central America. Zootaxa, 2719, 21–34.
Schmidt, K.P. (1953) A Checklist of North American Amphibians and Reptiles, 6th Ed. University of Chicago Press, Chicago, 280 pp.
Schuchert, C. (1935) Historical Geology of the Antillean-Caribbean Region. John Wiley and Sons, New York, xxvi + 811 pp.
Shackleton, N.J. & Kennett, J.P. (1975) Palaeotemperature history of the Cenozoic, and the initiation of Antarctic glaciation: oxygen and carbon isotope analysis in DSDP sites 277, 279, 281. Initial Report of the Deep Sea Drilling Project, 29, 743–755.
Sindaco, R. & Jeremcenko, V.K. (2008) The Reptiles and Amphibians of the Western Palearctic. Edizioni Belevedere, Latina, Italy, 580 pp.
Sioli, H. (Ed.) (1984) The Amazon: Limnology and Landscape Ecology of a Mighty Tropical River and its Basin. W. Junk, Dordrecht, The Netherlands, 762 pp.
http://dx.doi.org/10.1007/978-94-009-6542-3
Springer, M.S., Westerman, M., Kavanagh, J.R., Burk, A., Woodburne, M.O., Kao, D.J. & Krajewski, C. (1998) The origin of the Australasian marsupial fauna and the phylogenetic affinities of the enigmatic monito del monte and marsupial mole. Proceedings of the Royal Society of London Series B–Biological Sciences, 265, 2381–2386.
http://dx.doi.org/10.1098/rspb.1998.0587
Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
http://dx.doi.org/10.1093/bioinformatics/btu033
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML Web Servers. Systematic Biology, 57, 758–771.
http://dx.doi.org/10.1080/10635150802429642
Stejneger, L. (1907) Herpetology of Japan and adjacent territory. Bulletin of the United States National Museum, 58, xx + 577 pp.
Stöck, M., Dubey, S., Klütsh, C., Litvinchuk, S.N., Scheidt, U. & Perrin, N. (2008) Mitochondrial and nuclear phylogeny of circum-Mediterranean treefrogs from the Hyla arborea group. Molecular Phylogenetics and Evolution, 49, 1019–1024.
http://dx.doi.org/10.1016/j.ympev.2008.08.029
Suárez, P., Cardozo, D., Baldo, D., Pereyra, M.O., Faivovich, J., Orrico, V.G.D., Catroli, G.F., Grabiele, M., Bernarde, P.S., Nagamachi, C.Y., Haddad, C.F.B. & Pieczarka, J.C. (2013) Chromosome evolution in Dendropsophini (Amphibia, Anura, Hylinae). Cytogenetic and Genome Research, 141, 295–308.
http://dx.doi.org/10.1159/000354997
Szczerbak, N.N. (2003) Guide to the Reptiles of the Eastern Palearctic. Krieger Publishing Company, Malabar, Florida, 350 pp.
Tamura, K., Battistuzzi, F.U., Billing-Ross, P., Murillo, O., Filipski, A. & Kumar, S. (2012) Estimating divergence times in large molecular phylogenies. Proceedings of the National Academy of Sciences USA, 109, 19333–19338.
http://dx.doi.org/10.1073/pnas.1213199109
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular Evolutionary Genetics and Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods. Molecular Biology and Evolution, 28, 2731-2739.
http://dx.doi.org/10.1093/molbev/msr121
Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
http://dx.doi.org/10.1093/molbev/mst197
Townsend, T.M., Mulcahy, D.G., Noonan, B.P., Sites, J.W., Jr., Kuczynski, C.A., Wiens, J.J. & Reeder, T.W. (2011) Phylogeny of iguanian lizards inferred from 29 nuclear loci, and a comparison of concatenated and species-tree approaches for an ancient, rapid radiation. Molecular Phylogenetics and Evolution, 61, 363–380.
http://dx.doi.org/10.1016/j.ympev.2011.07.008
Trueb, L. (1970) Evolutionary relationships of casque-headed tree frogs with co-ossified skulls (family Hylidae). University of Kansas Publications Museum of Natural History, 18, 547–716.
Trueb, L. (1999) The Early Cretaceous pipoid anuran, Thoraciliacus: redescription, revaluation, and taxonomic status. Herpetologica, 55, 139–157.
Trueb, L., Ross, C.F. & Smith, R. (2005) A new pipoid anuran from the Late Cretaceous of South Africa. Journal of Vertebrate Paleontology, 25, 533–547.
http://dx.doi.org/10.1671/0272-4634(2005)025[0533:ANPAFT]2.0.CO;2
Tschudi,