Systematic review of the frog family Hylidae, with special reference to Hylinae: Phylogenetic analysis and taxonomic revision

dc.contributor.authorFaivovich, Julián
dc.contributor.authorHaddad, Célio F. B.
dc.contributor.authorGarcia, Paulo C. A.
dc.contributor.authorFrost, Darrel R.
dc.contributor.authorCampbell, Jonathan A.
dc.contributor.authorWheeler, Ward C.
dc.contributor.institutionColumbia University
dc.contributor.institutionUniversidade Estadual Paulista (Unesp)
dc.contributor.institutionUniversidade de Mogi das Cruzes (UMC)
dc.contributor.institutionUniversidade de São Paulo (USP)
dc.contributor.institutionAmerican Museum of Natural History
dc.contributor.institutionUniversity of Texas at Arlington
dc.date.accessioned2014-05-27T11:21:46Z
dc.date.available2014-05-27T11:21:46Z
dc.date.issued2005-12-27
dc.description.abstractHylidae is a large family of American, Australopapuan, and temperate Eurasian treefrogs of approximately 870 known species, divided among four subfamilies. Although some groups of Hylidae have been addressed phylogenetically, a comprehensive phylogenetic analysis has never been presented. The first goal of this paper is to review the current state of hylid systematics. We focus on the very large subfamily Hylinae (590 species), evaluate the monophyly of named taxa, and examine the evidential basis of the existing taxonomy. The second objective is to perform a phylogenetic analysis using mostly DNA sequence data in order to (1) test the monophyly of the Hylidae; (2) determine its constituent taxa, with special attention to the genera and species groups which form the subfamily Hylinae, and c) propose a new, monophyletic taxonomy consistent with the hypothesized relationships. We present a phylogenetic analysis of hylid frogs based on 276 terminals, including 228 hylids and 48 outgroup taxa. Included are exemplars of all but 1 of the 41 genera of Hylidae (of all four nominal subfamilies) and 39 of the 41 currently recognized species groups of the species-rich genus Hyla. The included taxa allowed us to test the monophyly of 24 of the 35 nonmonotypic genera and 25 species groups of Hyla. The phylogenetic analysis includes approximately 5100 base pairs from four mitochondrial (12S, tRNA valine, 16S, and cytochrome b) and five nuclear genes (rhodopsin, tyrosinase, RAG-1, seventh in absentia, and 28S), and a small data set from foot musculature. Concurring with previous studies, the present analysis indicates that Hemiphractinae are not related to the other three hylid subfamilies. It is therefore removed from the family and tentatively considered a subfamily of the paraphyletic Leptodactylidae. Hylidae is now restricted to Hylinae, Pelodryadinae, and Phyllomedusinae. Our results support a sister-group relationship between Pelodryadinae and Phyllomedusinae, which together form the sister taxon of Hylinae. Agalychnis, Phyllomedusa, Litoria, Hyla, Osteocephalus, Phrynohyas, Ptychohyla, Scinax, Smilisca, and Trachycephalus are not monophyletic. Within Hyla, the H. albomarginata, H. albopunctata, H. arborea, H. boons, H. cinerea, H. eximia, H. geographica, H. granosa, H. microcephala, H. miotympanum, H. tuberculosa, and H. versicolor groups are also demonstrably nonmonophyletic. Hylinae is composed of four major clades. The first of these includes the Andean stream-breeding Hyla, Aplastodiscus, all Gladiator Frogs, and a Tepuian clade. The second clade is composed of the 30-chromosome Hyla, Lysapsus, Pseudis, Scarthyla, Scinax (including the H. uruguaya group), Sphaenorhynchus, and Xenohyla. The third major clade is composed of Nyctimantis, Phrynohyas, Phyllodytes, and all South American/West Indian casque-headed frogs: Aparasphenodon, Argenteohyla, Corythomantis, Osteocephalus, Osteopilus, Tepuihyla, and Trachycephalus. The fourth major clade is composed of most of the Middle American/Holarctic species groups of Hyla and the genera Acris, Anotheca, Duellmanohyla, Plectrohyla, Pseudacris, Ptychohyla, Pternohyla, Smilisca, and Triprion. A new monophyletic taxonomy mirroring these results is presented where Hylinae is divided into four tribes. Of the species currently in Hyla, 297 of the 353 species are placed in 15 genera; of these, 4 are currently recognized, 4 are resurrected names, and 7 are new. Hyla is restricted to H. femoralis and the H. arborea, H. cinerea, H. eximia, and H. versicolor groups, whose contents are redefined. Phrynohyas is placed in the synonymy of Trachycephalus, and Pternohyla is placed in the synonymy of Smilisca. The genus Dendropsophus is resurrected to include all former species of Hyla known or suspected to have 30 chromosomes. Exerodonta is resurrected to include the former Hyla sumichrasti group and some members of the former H. miotympanum group. Hyloscirtus is resurrected for the former Hyla armata, H. bogotensis, and H. larinopygion groups. Hypsiboas is resurrected to include several species groups - many of them redefined here - of Gladiator Frogs. The former Hyla albofrenata and H. albosignata complexes of the H. albomarginata group are included in Aplastodiscus. New generic names are erected for (1) Agalychnis calcarifer and A. craspedopus; (2) Osteocephalus langsdorffii; the (3) Hyla aromatica, (4) H. bromeliacia, (5) H. godmani, (6) H. mixomaculata, (7) H. taeniopus, (8) and H. tuberculosa groups; (9) the clade composed of the H. pictipes and H. pseudopuma groups; and (10) a clade composed of the H. circumdata, H. claresignata, H. martinsi, and H. pseudopseudis groups. Copyright © American Museum of Natural History 2005.en
dc.description.affiliationDivision of Vertebrate Zoology (Herpetology) Department of Ecology, Evolution, and Environmental Biology (E3B) Columbia University, New York, NY
dc.description.affiliationDepartamento de Zoologia Instituto de Biociências Unversidade Estadual Paulista, C.P. 199, 13506-900 Rio Claro, São Paulo
dc.description.affiliationUniversidade de Mogi Das Cruzes Área de Ciências da Saúde Curso de Biologia, R. C. Xavier de Almeida e Souza 200, 08780-911 Mogi das Cruzes, Sao Paulo
dc.description.affiliationMuseu de Zoologia Universidade de São Paulo, São Paulo
dc.description.affiliationDivision of Vertebrate Zoology (Herpetology) American Museum of Natural History
dc.description.affiliationDepartment of Biology University of Texas at Arlington, Arlington, TX 76019
dc.description.affiliationDivision of Invertebrate Zoology American Museum of Natural History
dc.format.extent1-227
dc.identifierhttp://dx.doi.org/10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2
dc.identifierhttp://hdl.handle.net/11449/68699
dc.identifier.citationBulletin of the American Museum of Natural History, n. 294, p. 1-227, 2005.
dc.identifier.doi10.1206/0003-0090(2005)294[0001:SROTFF]2.0.CO;2
dc.identifier.doihttp://hdl.handle.net/11449/68699
dc.identifier.issn0003-0090
dc.identifier.scopus2-s2.0-27944485147
dc.identifier.urihttp://hdl.handle.net/11449/68699
dc.language.isoeng
dc.relation.ispartofBulletin of the American Museum of Natural History
dc.relation.ispartofjcr2.938
dc.relation.ispartofsjr1,430
dc.rights.accessRightsAcesso restrito
dc.sourceScopus
dc.subjectAcris
dc.subjectAgalychnis
dc.subjectAgalychnis calcarifer
dc.subjectAgalychnis craspedopus
dc.subjectAnotheca
dc.subjectAnura
dc.subjectAparasphenodon
dc.subjectCorythomantis
dc.subjectDuellmanohyla
dc.subjectHemiphractinae
dc.subjectHyla
dc.subjectHyla arborea
dc.subjectHyla armata
dc.subjectHyla bogotensis
dc.subjectHyla cinerea
dc.subjectHyla circumdata
dc.subjectHyla eximia
dc.subjectHyla femoralis
dc.subjectHyla geographica
dc.subjectHyla godmani
dc.subjectHyla larinopygion
dc.subjectHyla microcephala
dc.subjectHyla miotympanum
dc.subjectHyla mixomaculata
dc.subjectHyla pseudopuma
dc.subjectHyla sumichrasti
dc.subjectHyla taeniopus
dc.subjectHyla versicolor
dc.subjectHylidae
dc.subjectHylinae
dc.subjectLeptodactylidae
dc.subjectLitoria
dc.subjectLysapsus
dc.subjectNyctimantis
dc.subjectOsteocephalus
dc.subjectOsteocephalus langsdorffii
dc.subjectOsteopilus
dc.subjectPelodryadinae
dc.subjectPhrynohyas
dc.subjectPhyllodytes
dc.subjectPhyllomedusa
dc.subjectPhyllomedusinae
dc.subjectPlectrohyla
dc.subjectPseudacris
dc.subjectPseudis
dc.subjectPternohyla
dc.subjectPtychohyla
dc.subjectScarthyla
dc.subjectScinax
dc.subjectSmilisca
dc.subjectSphaenorhynchus
dc.subjectTepuia
dc.subjectTrachycephalus
dc.subjectTriprion
dc.subjectUruguaya
dc.titleSystematic review of the frog family Hylidae, with special reference to Hylinae: Phylogenetic analysis and taxonomic revisionen
dc.typeArtigo
dcterms.licensehttp://www.bioone.org/page/terms_of_use
unesp.author.lattes0458077399058762[2]
unesp.author.orcid0000-0002-7044-5764[2]
unesp.campusUniversidade Estadual Paulista (Unesp), Instituto de Biociências, Rio Claropt

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