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Systematics of spiny‐backed treefrogs (Hylidae: Osteocephalus): an Amazonian puzzle
Authors:Karl‐Heinz Jungfer  Julián Faivovich  José M Padial  Santiago Castroviejo‐Fisher  Mariana M Lyra  Bianca V M Berneck  Patricia P Iglesias  Philippe J R Kok  Ross D MacCulloch  Miguel T Rodrigues  Vanessa K Verdade  Claudia P Torres Gastello  Juan Carlos Chaparro  Paula H Valdujo  Steffen Reichle  Ji?í Moravec  Václav Gvo?dík  Giussepe Gagliardi‐Urrutia  Raffael Ernst  Ignacio De la Riva  Donald Bruce Means  Albertina P Lima  J Celsa Señaris  Ward C Wheeler  Célio F B Haddad
Institution:1. División Herpetología, Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’‐CONICET, , C1405DJR Buenos Aires, Argentina;2. Departamento de Biodiversidad y Biologia Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, , Buenos Aires, Argentina;3. Section of Amphibians and Reptiles, Carnegie Museum of Natural History, , Pittsburgh, PA, 15213‐4080 USA;4. Department of Herpetology, American Museum of Natural History, , New York, NY, 10024‐5192 USA;5. Laboratorio de Sistemática de Vertebrados, Pontifícia Universidade Católica do Rio Grande do Sul, , Porto Alegre, RS, Brazil;6. Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista, , CEP 13506‐900 Rio Claro, S?o Paulo, Brazil;7. Departamento de Ecología, Genética y Evolución, Facultad de Ciencias Exactas y Naturales, Instituto de Ecología, Genética y Evolución de Buenos Aires, IEGEBA‐CONICET, Universidad de Buenos Aires, , C1428EHA Buenos Aires, Argentina;8. Amphibian Evolution Lab, Department of Biology, Vrije Universiteit Brussel, , B‐1050 Brussels, Belgium;9. Department of Vertebrates, Royal Belgian Institute of Natural Sciences, , B‐1000 Brussels, Belgium;10. Department of Natural History, Royal Ontario Museum, , Toronto, ON, M5S 2C6 Canada;11. Departamento de Zoologia, Instituto de Biociências, Universidade de S?o Paulo, , CEP 05508‐090 S?o Paulo, Brazil;12. Centro de Ciências Naturais e Humanas, Universidade Federal do ABC, , 5001 CEP 09210‐971 Santo André, S?o Paulo, Brazil;13. Departamento de Herpetología, Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, , Apto. 14‐0434 Lima 14, Peru;14. Museo de Historia Natural, Universidad Nacional de San Antonio Abad del Cusco, , Cusco, Peru;15. Departamento de Ecologia, Instituto de Biociências, Universidade de S?o Paulo, , 05508‐900 S?o Paulo, SP, Brazil;16. Fundación para la Conservación del Bosque Chiquitano, , Santa Cruz de la Sierra, Bolivia;17. Department of Zoology, National Museum, , Prague 9, Czech Republic;18. Programa de Investigación en Biodiversidad Amazónica del Instituto de Investigaciones de la Amazonia Peruana, , Iquitos, Perú;19. Museum für Tierkunde, Senckenberg Natural History Collections Dresden, , D‐01109 Dresden, Germany;20. Department of Biodiversity and Evolutionary Biology, Museo Nacional de Ciencias Naturales‐CSIC, , 28006 Madrid, Spain;21. Coastal Plains Institute and Land Conservancy, , Tallahassee, FL, 32303 USA;22. Instituto Nacional de Pesquisas da Amaz?nia, , CEP 69011‐970 Manaus, Amazonas, Brazil;23. Laboratorio de Ecología y Genética de Poblaciones, Centro de Ecología, Instituto Venezolano de Investigaciones Científicas, , Caracas, 1020‐A Venezuela;24. Division of Invertebrate Zoology, American Museum of Natural History, , New York, NY, 10024‐5192 USA
Abstract:Spiny‐backed tree frogs of the genus Osteocephalus are conspicuous components of the tropical wet forests of the Amazon and the Guiana Shield. Here, we revise the phylogenetic relationships of Osteocephalus and its sister group Tepuihyla, using up to 6134 bp of DNA sequences of nine mitochondrial and one nuclear gene for 338 specimens from eight countries and 218 localities, representing 89% of the 28 currently recognized nominal species. Our phylogenetic analyses reveal (i) the paraphyly of Osteocephalus with respect to Tepuihyla, (ii) the placement of ‘Hyla’ warreni as sister to Tepuihyla, (iii) the non‐monophyly of several currently recognized species within Osteocephalus and (iv) the presence of low (<1%) and overlapping genetic distances among phenotypically well‐characterized nominal species (e.g. O. taurinus and O. oophagus) for the 16S gene fragment used in amphibian DNA barcoding. We propose a new taxonomy, securing the monophyly of Osteocephalus and Tepuihyla by rearranging and redefining the content of both genera and also erect a new genus for the sister group of Osteocephalus. The colouration of newly metamorphosed individuals is proposed as a morphological synapomorphy for Osteocephalus. We recognize and define five monophyletic species groups within Osteocephalus, synonymize three species of Osteocephalus (O. germani, O. phasmatus and O. vilmae) and three species of Tepuihyla (T. celsae, T. galani and T. talbergae) and reallocate three species (Hyla helenae to Osteocephalus, O. exophthalmus to Tepuihyla and O. pearsoni to Dryaderces gen. n.). Furthermore, we flag nine putative new species (an increase to 138% of the current diversity). We conclude that species numbers are largely underestimated, with most hidden diversity centred on widespread and polymorphic nominal species. The evolutionary origin of breeding strategies within Osteocephalus is discussed in the light of this new phylogenetic hypothesis, and a novel type of amplexus (gular amplexus) is described.
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