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Taxonomy and Systematics of Spiny-Backed Treefrogs, Genus Osteocephalus (Amphibia: Anura: Hylidae)
(2015)
The pan-Amazonian treefrog genus Ostecephalus is poorly understood both on a taxonomic and phylogenetic level. The status of several frogs already or not yet referred to the genus is unclear and the relationships among the genus and with respect to related genera is not understood. In this work O. cabrerai (Cochran and Goin, 1970) from Colombia and Peru is redescribed and O. festae (Peracca, 1904) from the foothills of the Andes in Ecuador is revalidated. Hyla inframaculata Boulenger, 1882, from the lower Amazon in Brazil, is reallocated to Osteocephalus and O. elkejungingerae (Henle, 1981) from the Andean foothills in Peru is shown to be a synonym of Hyla mimetica (Melin, 1941), the valid name being O. mimeticus. Hyla vilarsi Melin, 1941 is considered a valid species in the genus Osteocephalus and revalidated from the synonymies of several other frogs. Three new species, O. castaneicola from northern Bolivia and southern Peru, O. duellmani from a sub-Andean mountain range in southern Ecuador, and O. camufatus from central Amazonian Brazil, are described. A phylogenetic analysis based on up to nine mitochondrial genes and one nuclear one reveals the paraphyly of the genus as previously understood with respect to the genus Tepuihyla. A new taxonomy is proposed, securing the monophyly of Osteocephalus and Tepuihyla by rearranging and redefining the content of both genera. A new genus, Dryaderces, is erected for the sister group of Osteocephalus. The colouration of newly metamorphosed individuals is proposed as a morphological synapomorphy for Osteocephalus. Five monophyletic species groups within Osteocephalus are recognized, three species of Osteocephalus (O. germani, O.rnphasmatus, O. vilmae) and three species of Tepuihyla (T. celsae, T. galani, T. talbergae) synonymized and three species (Hyla helenae to Osteocephalus, O.rnexophthalmus to Tepuihyla and O. pearsoni to Dryaderces gen. n.) reallocated. Furthermore, nine putative new species are flagged (an increase to 138% of the current diversity), an indication 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 phylogeny and a novel type of amplexus (gular amplexus) is described.