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.

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. (2013). Systematics of spiny-backed treefrogs (Hylidae: Osteocephalus): an Amazonian puzzle. -Zoologica Scripta, 42, 351-380. 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. Corresponding author: Karl-Heinz Jungfer, Institute of Integrated Sciences, Department of Biology, University of Koblenz-Landau, Universit€ atsstr. 1, 56070 Koblenz, Germany. E-mail: khjungfer@aol.com Juli an Faivovich, Divisi on Herpetolog ıa, Museo Argentino de Ciencias Naturales 'Bernardino Rivadavia'-CONICET, Angel Gallardo 470, C1405DJR, Buenos Aires, Argentina and Departamento de Biodiversidad y Biologia Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Buenos Aires, Argentina. E-mail: julian@macn.gov.ar Jos e M. Padial, Section of Amphibians and Reptiles, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA, 15213-4080 Systematics of spiny-backed treefrogs K.-H. Jungfer et al. IntroductionTreefrogs of the genus Osteocephalus constitute an important component of the amphibian fauna of the Amazonian and Guianan regions of South America. Their distributions...

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Aim We used frogs of the clade Allocentroleniae (Centrolenidae + Allophrynidae; c. 170 species endemic to Neotropical rain forests) as a model system to address the historical biogeography and diversification of Neotropical rain forest biotas.Location Neotropical rain forests.Methods We used an extensive taxon (109 species) and gene (seven nuclear and three mitochondrial genes) sampling to estimate phylogenetic relationships, divergence times, ancestral area distributions, dispersal-vicariance events, and the temporal pattern of diversification rate. ResultsThe Allocentroleniae started to diversify in the Eocene in South America and by the early Miocene were present in all major Neotropical rain forests except in Central America, which was colonized through 11 late range expansions. The initial uplifts of the Andes during the Oligocene and early Miocene, as well as marine incursions in the lowlands, are coincidental with our estimates of the divergence times of most clades of Allocentroleniae. Clades with broad elevational distributions occupy more biogeographical areas. Most dispersals involve the Andes as a source area but the majority were between the Central and the Northern Andes, suggesting that the Andes did not play a major role as a species pump for the lowlands. The diversification of glassfrogs does not follow a south-to-north pattern of speciation for Andean clades, and the establishment of a transcontinental Amazon drainage system is coincidental in time with the isolation of the Atlantic Forest glassfrogs. Diversification analyses indicated that a model of constantly increasing diversity best fits the data, compatible with the 'evolutionary museum' hypothesis or 'ancient cradle' hypothesis.Main conclusions Our work illustrates how the different geological and climatic historical events of the Neotropics shaped, at different levels of the phylogeny, the diversity of a species-rich clade, highlighting the importance of studying large evolutionary radiations at a continental scale.

Deciphering the products of evolution at the species level: the need for an integrative taxonomy. -Zoologica Scripta , 38 , 431-447. Progress in molecular techniques together with the incorporation of phylogenetic analyses of DNA into taxonomy have caused an increase in the number of species' discoveries in groups with morphological characters that are difficult to study or in those containing polytypic species. But some emerged criticisms plead for a taxonomic conservatism grounded either on the requirement of providing evidences of morphological distinctiveness or reproductive barriers to erect new species names. In a case study of taxonomic research on Neotropical frogs, we combine several lines of evidence (morphological characters, prezygotic reproductive isolation and phylogenetic analyses of mitochondrial DNA) to test the status of 15 nominal species and to assess the degree of agreement of the different lines of evidence. Our study reveals that morphology alone is not sufficient to uncover all species, as there is no other single line of evidence independently. Full congruence between lines of evidence is restricted to only four out of the 15 species. Five species show congruence of two lines of evidence, whereas the remaining six are supported by only one. The use of divergence in morphological characters seems to be the most conservative approach to delineate species boundaries because it does not allow the identification of some sibling reciprocally monophyletic species differing in their advertisement calls. The separate analysis of differences in advertisement calls (evidence of reproductive isolation) or of phylogenetic data alone also shows limitations, because they do not support some morphological species. Our study shows that only an integrative approach combining all sources of evidence provides the necessary feedback to evaluate the taxonomic status of existing species and to detect putative new ones. Furthermore, the application of integrative taxonomy enables the identification of hypotheses about the existence of species that will probably be rejected or changed, and those that can be expected to persist.

Egg-brooding frogs (Hemiphractidae) are a group of 105 currently recognized Neotropical species, with a remarkable diversity of developmental modes, from direct development to free-living and exotrophic tadpoles. Females carry their eggs on the back and embryos have unique bell-shaped gills. We inferred the evolutionary relationships of these frogs and used the resulting phylogeny to review their taxonomy and test hypotheses on the evolution of developmental modes and bell-shaped gills. Our inferences relied on a total evidence parsimony analysis of DNA sequences of up to 20 mitochondrial and nuclear genes (analyzed under tree-alignment), and 51 phenotypic characters sampled for 83% of currently valid hemiphractid species. Our analyses rendered a well-resolved phylogeny, with both Hemiphractidae (sister of Athesphatanura) and its six recognized genera being monophyletic. We also inferred novel intergeneric relationships [((Cryptobatrachus, Flectonotus), (Stefania, (Fritziana, (Hemiphractus, Gastrotheca))))], the non-monophyly of all species groups previously proposed within Gastrotheca and Stefania, and the existence of several putative new species within Fritziana and Hemiphractus. Contrary to previous hypotheses, our results support the most recent common ancestor of hemiphractids as a direct-developer. Free-living aquatic tadpoles apparently evolved from direct-developing ancestors three to eight times. Embryos of the sister taxa Cryptobatrachus and Flectonotus share a pair of single gills derived from branchial arch I, while embryos of the clade including the other four genera have two pairs of gills derived from branchial arches I and II respectively. Furthermore, in Gastrotheca the fusion of the two pairs of gills is a putative synapomorphy. We propose a revised taxonomy concordant with our optimal topologies.

Despite recent efforts to accelerate exploration and species description, the diversity of high Andean frogs remains highly underestimated. We report high levels of species diversity in direct-developing frogs or terraranas inhabiting the wet puna and adjacent cloud forests of the Amazonian versant of the Andes in Bolivia and Peru. Descriptive evidence of external morphology, distribution patterns and molecular phylogenetic analyses support the existence of nine unnamed species in two clades, which represents a 30% increase in species diversity for those clades. The relationships of these species and their relatives in Holoadeninae are tested using nuclear and mitochondrial genes for 159 terminals representing the 11 genera in this subfamily and 25 species of previously unknown relationships. Our results corroborate species monophyly in all but three cases and support the monophyly of all Holoadeninae genera, albeit the position of some differs between analyses. We propose a new genus (Microkayla gen. nov.) for the clade containing all Bolivian species formerly in Psychrophrynella plus five species from southern Peru. The new genus is monophyletic and supported by anatomical synapomorphies. Psychrophrynella is re-diagnosed and redefined to include three species from the Andes of southern Peru. We discuss the taxonomic instability associated with Noblella and Psychrophrynella due to the fact that the type species of both genera share a number of traits that support a close relationship. We also name and describe three new species of Bryophryne and two of Microkayla from Peru, provide baseline data for the future description of four Bolivian species of Microkayla, and describe the unknown mating calls of two species. Our results support that the grasslands of the Amazonian versant of the Andes harbour a large diversity of species with small altitudinal and horizontal distributions that replace each other along a latitudinal axis. These species belong to different lineages whose closest relatives are forest species, often from distant parts of the continent. These patterns suggest that high Andean environments were colonized several times independently by species with forest ancestors and which radiated into a multitude of species with remarkably similar ecomorphologies. The extent of these radiations remains obscured by a still rudimentary knowledge of species diversity due to insufficient fieldwork and taxonomic research.