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      Systematics of Ecnomiohyla tuberculosa with the description of a new species and comments on the taxonomy of Trachycephalus typhonius (Anura, Hylidae)

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          Abstract

          Abstract

          Ecnomiohyla tuberculosa is an Amazonian hylid of uncertain phylogenetic position. Herein DNA sequences of mitochondrial and nuclear genes are used to determine its phylogenetic relationships. New sequences and external morphology of Trachycephalus typhonius are also analyzed to assess the status of Ecuadorian and Peruvian populations. The phylogeny shows unequivocally that Ecnomiohyla tuberculosa is nested within the genus Tepuihyla , tribe Lophiohylini . This position was unexpected because the remaining species of Ecnomiohyla belong to the tribe Hylini . To solve the paraphyly of the genus Ecnomiohyla , Ecnomiohyla tuberculosa is transferred to the genus Tepuihyla . Comparisons of DNA sequences, external morphology, and advertisement calls between populations of Ecnomiohyla tuberculosa from Ecuador and Peru indicate that the Peruvian population represents an undescribed species. The new species is described and a species account is provided for Ecnomiohyla tuberculosa . Trachycephalus typhonius is paraphyletic relative to Trachycephalus cunauaru , Trachycephalus hadroceps , and Trachycephalus resinifictrix . The phylogenetic position of populations from western Ecuador indicates that they represent a species separate from Trachycephalus typhonius sensu stricto. We resurrect the name Hyla quadrangulum ( Trachycephalus quadrangulum comb. n.) for those populations. Amazonian populations of “ Trachycephalus typhonius ” from Ecuador and Peru are genetically and morphologically distinct from Trachycephalus typhonius sensu stricto and are conspecific with the holotype of Hyla macrotis . Therefore, we also resurrect Hyla macrotis , a decision that results in Trachycephalus macrotis comb. n.

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          Evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs: treefrog trees unearth the roots of high tropical diversity.

          Why are there more species in the tropics than in temperate regions? In recent years, this long-standing question has been addressed primarily by seeking environmental correlates of diversity. But to understand the ultimate causes of diversity patterns, we must also examine the evolutionary and biogeographic processes that directly change species numbers (i.e., speciation, extinction, and dispersal). With this perspective, we dissect the latitudinal diversity gradient in hylid frogs. We reconstruct a phylogeny for 124 hylid species, estimate divergence times and diversification rates for major clades, reconstruct biogeographic changes, and use ecological niche modeling to identify climatic variables that potentially limit dispersal. We find that hylids originated in tropical South America and spread to temperate regions only recently (leaving limited time for speciation). There is a strong relationship between the species richness of each region and when that region was colonized but not between the latitudinal positions of clades and their rates of diversification. Temperature seasonality seemingly limits dispersal of many tropical clades into temperate regions and shows significant phylogenetic conservatism. Overall, our study illustrates how two general principles (niche conservatism and the time-for-speciation effect) may help explain the latitudinal diversity gradient as well as many other diversity patterns across taxa and regions.
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            Biogeographic analysis reveals ancient continental vicariance and recent oceanic dispersal in amphibians.

            R Pyron (2014)
            Amphibia comprises over 7000 extant species distributed in almost every ecosystem on every continent except Antarctica. Most species also show high specificity for particular habitats, biomes, or climatic niches, seemingly rendering long-distance dispersal unlikely. Indeed, many lineages still seem to show the signature of their Pangaean origin, approximately 300 Ma later. To date, no study has attempted a large-scale historical-biogeographic analysis of the group to understand the distribution of extant lineages. Here, I use an updated chronogram containing 3309 species (∼ 45% of extant diversity) to reconstruct their movement between 12 global ecoregions. I find that Pangaean origin and subsequent Laurasian and Gondwanan fragmentation explain a large proportion of patterns in the distribution of extant species. However, dispersal during the Cenozoic, likely across land bridges or short distances across oceans, has also exerted a strong influence. Finally, there are at least three strongly supported instances of long-distance oceanic dispersal between former Gondwanan landmasses during the Cenozoic. Extinction from intervening areas seems to be a strong factor in shaping present-day distributions. Dispersal and extinction from and between ecoregions are apparently tied to the evolution of extraordinarily adaptive expansion-oriented phenotypes that allow lineages to easily colonize new areas and diversify, or conversely, to extremely specialized phenotypes or heavily relictual climatic niches that result in strong geographic localization and limited diversification.
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              Hylid frog phylogeny and sampling strategies for speciose clades.

              How should characters and taxa be sampled to resolve efficiently the phylogeny of ancient and highly speciose groups? We addressed this question empirically in the treefrog family Hylidae, which contains > 800 species and may be nonmonophyletic with respect to other anuran families. We sampled 81 species (54 hylids and 27 outgroups) for two mitochondrial genes (12S, ND1), two nuclear genes (POMC, c-myc), and morphology (144 characters) in an attempt to resolve higher-level relationships. We then added 117 taxa to the combined data set, many of which were sampled for only one gene (12S). Despite the relative incompleteness of the majority of taxa, the resulting trees placed all taxa in the expected higher-level clades with strong support, despite some taxa being > 90% incomplete. Furthermore, we found no relationship between the completeness of a taxon and the support (parsimony bootstrap or Bayesian posterior probabilities) for its localized placement on the tree. Separate analysis of the data set with the most taxa (12S) gives a somewhat problematic estimate of higher-level relationships, suggesting that data sets scored only for some taxa (ND1, nuclear genes, morphology) are important in determining the outcome of the combined analysis. The results show that hemiphractine hylids are not closely related to other hylids and should be recognized as a distinct family. They also show that the speciose genus Hyla is polyphyletic, but that its species can be arranged into three monophyletic genera. A new classification of hylid frogs is proposed. Several potentially misleading signals in the morphological data are discussed.
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                Author and article information

                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2016
                9 November 2016
                : 630
                : 115-154
                Affiliations
                [1 ]Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Av. 12 de Octubre y Roca, Aptdo. 17-01-2184, Quito, Ecuador
                [2 ]División de Herpetología-Centro de Ornitología y Biodiversidad (CORBIDI), Santa Rita N˚105 Of. 202, Urb. Huertos de San Antonio, Surco, Lima, Perú
                [3 ]Laboratorio de Biogeografía, Red de Biología Evolutiva, Instituto de Ecología A.C., Carretera antigua a Coatepec 351, El Haya, CP 91070, Xalapa, Veracruz, México
                [4 ]Programa de Investigación en Biodiversidad Amazónica, Instituto de Investigaciones de la Amazonia Peruana (IIAP), Av. Quiñones Km 2.5, Iquitos, Perú
                [5 ]Current Address: Laboratório de Sistemática de Vertebrados, Pontifícia Universidade Católica do Rio Grande do Sul - PUCRS. Av. Ipiranga, 6681, Porto Alegre, RS 90619-900, Brazil
                [6 ]Department of Biology, Colorado State University, 1878 Campus Delivery, Fort Collins, CO 80523, USA
                [7 ]Current Address: IKIAM, Universidad Regional Amazónica, km 7 vía Muyuna, Tena, Ecuador
                [8 ]Museo Ecuatoriano de Ciencias Naturales, Sección de Vertebrados, División de Herpetología, calle Rumipamba 341 y Av. de los Shyris, Quito, Ecuador
                Author notes
                Corresponding author: Santiago R. Ron ( santiago.r.ron@ 123456gmail.com )

                Academic editor: A. Crottini

                Article
                10.3897/zookeys.630.9298
                5126525
                f984b2b5-0be6-4bcb-8c8c-875fd826c357
                Santiago R. Ron, Pablo J. Venegas, H. Mauricio Ortega-Andrade, Giussepe Gagliardi-Urrutia, Patricia E. Salerno

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 24 May 2016
                : 4 October 2016
                Categories
                Research Article

                Animal science & Zoology
                advertisement call,amazon basin,biodiversity,ecuador,lophiohylini,peru,phylogeny,tepuihyla,animalia,anura,hylidae

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