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Systematic Review of the Frog Family Hylidae, with Special Reference to Hylinae: Phylogenetic Analysis and Taxonomic Revision

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Systematic Review of the Frog Family Hylidae, with Special Reference to Hylinae: Phylogenetic Analysis and Taxonomic Revision SYSTEMATIC REVIEW OF THE FROG FAMILY HYLIDAE, WITH SPECIAL REFERENCE TO HYLINAE: PHYLOGENETIC ANALYSIS AND TAXONOMIC REVISION JULIAÂ N FAIVOVICH Division of Vertebrate Zoology (Herpetology), American Museum of Natural History Department of Ecology, Evolution, and Environmental Biology (E3B) Columbia University, New York, NY ([email protected]) CEÂ LIO F.B. HADDAD Departamento de Zoologia, Instituto de BiocieÃncias, Unversidade Estadual Paulista, C.P. 199 13506-900 Rio Claro, SaÄo Paulo, Brazil ([email protected]) PAULO C.A. GARCIA Universidade de Mogi das Cruzes, AÂ rea de CieÃncias da SauÂde Curso de Biologia, Rua CaÃndido Xavier de Almeida e Souza 200 08780-911 Mogi das Cruzes, SaÄo Paulo, Brazil and Museu de Zoologia, Universidade de SaÄo Paulo, SaÄo Paulo, Brazil ([email protected]) DARREL R. FROST Division of Vertebrate Zoology (Herpetology), American Museum of Natural History ([email protected]) JONATHAN A. CAMPBELL Department of Biology, The University of Texas at Arlington Arlington, Texas 76019 ([email protected]) WARD C. WHEELER Division of Invertebrate Zoology, American Museum of Natural History ([email protected]) BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 294, 240 pp., 16 ®gures, 2 tables, 5 appendices Issued June 24, 2005 Copyright q American Museum of Natural History 2005 ISSN 0003-0090 CONTENTS Abstract ....................................................................... 6 Resumo ........................................................................ 7 Resumen ....................................................................... 8 Introduction .................................................................... 9 Materials and Methods .......................................................... 9 Taxon Sampling .............................................................. 9 Outgroup Selection ........................................................ 13 Basal Neobatrachians .................................................... 13 Ranoidea ............................................................... 13 Hyloidea ............................................................... 13 The Ingroup: Hylidae ...................................................... 14 Hemiphractinae ......................................................... 15 Pelodryadinae ........................................................... 17 Phyllomedusinae ........................................................ 19 Hylinae ................................................................ 20 Gladiator Frogs ....................................................... 21 Andean Stream-Breeding Hyla .......................................... 26 The 30-Chromosome Hyla ............................................. 27 Middle American/Holarctic Clade ....................................... 32 Casque-Headed Frogs and Related Genera ............................... 38 Species and Species Groups of Hyla Not Associated with Other Clades ..... 40 Other Genera ......................................................... 41 Character Sampling .......................................................... 43 Gene Selection ............................................................ 43 DNA Isolation and Sequencing .............................................. 43 Morphology .............................................................. 44 Da Silva's (1998) Dissertation .......................................... 45 Phylogenetic Analysis ........................................................ 45 Results ....................................................................... 48 Discussion .................................................................... 49 Major Patterns of Relationships of Hylidae and Outgroups ....................... 49 Pelodryadinae and Phyllomedusinae ........................................... 50 Pelodryadinae ............................................................. 53 Phyllomedusinae .......................................................... 54 Major Patterns of Relationships Within Hylinae ................................. 54 South American I Clade .................................................... 54 Andean Stream-Breeding Hyla and the Tepuian Clade ....................... 57 Gladiator Frogs ......................................................... 59 Atlantic/Cerrado Clade ................................................... 59 Green Clade ............................................................ 60 True Gladiator Frog Clade ................................................ 60 South American II Clade ................................................... 61 Relationships of Scinax .................................................. 61 Lysapsus, Pseudis, and Scarthyla .......................................... 62 Sphaenorhynchus, Xenohyla, and the 30-Chromosome Hyla .................. 63 Middle American/Holarctic Clade ........................................... 66 Lower Central American Lineage ......................................... 70 Real Hyla .............................................................. 72 South American/West Indian Casque-Headed Frogs ............................ 72 Taxonomic Conclusions: A New Taxonomy of Hylinae and Phyllomedusinae ......... 74 Hylinae Ra®nesque, 1815 ..................................................... 75 2 2005 FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE 3 Cophomantini Hoffmann, 1878 .............................................. 75 Aplastodiscus A. Lutz in B. Lutz, 1950 .................................... 76 Aplastodiscus albofrenatus Group ....................................... 82 Aplastodiscus albosignatus Group ....................................... 82 Aplastodiscus perviridis Group ......................................... 82 Bokermannohyla, new genus .............................................. 82 Bokermannohyla circumdata Group ..................................... 82 Bokermannohyla claresignata Group .................................... 83 Bokermannohyla martinsi Group ........................................ 83 Bokermannohyla pseudopseudis Group .................................. 83 Hyloscirtus Peters, 1882 ................................................. 84 Hyloscirtus armatus Group ............................................. 84 Hyloscirtus bogotensis Group ........................................... 84 Hyloscirtus larinopygion Group ......................................... 85 Hypsiboas Wagler, 1830 .................................................. 85 Hypsiboas albopunctatus Group ........................................ 86 Hypsiboas benitezi Group .............................................. 86 Hypsiboas faber Group ................................................ 87 Hypsiboas pellucens Group ............................................. 87 Hypsiboas pulchellus Group ............................................ 87 Hypsiboas punctatus Group ............................................ 88 Hypsiboas semilineatus Group .......................................... 88 Species of Hypsiboas Unassigned to Group .............................. 89 Myersiohyla, new genus .................................................. 89 Dendropsophini Fitzinger, 1843 ............................................. 90 Dendropsophus Fitzinger, 1843 ........................................... 90 Dendropsophus columbianus Group ..................................... 90 Dendropsophus garagoensis Group ...................................... 90 Dendropsophus labialis Group .......................................... 91 Dendropsophus leucophyllatus Group .................................... 91 Dendropsophus marmoratus Group ...................................... 91 Dendropsophus microcephalus Group ................................... 91 Dendropsophus minimus Group ......................................... 92 Dendropsophus minutus Group ......................................... 92 Dendropsophus parviceps Group ........................................ 93 Species of Dendropsophus Unassigned to Group .......................... 93 Lysapsus Cope, 1862 .................................................... 93 Pseudis Wagler, 1830 .................................................... 93 Scarthyla Duellman and de SaÂ, 1988 ...................................... 94 Scinax Wagler, 1830 ..................................................... 94 Scinax catharinae Clade ............................................... 95 Scinax ruber Clade .................................................... 96 Sphaenorhynchus Tschudi, 1838 .......................................... 97 Xenohyla Izecksohn, 1996 ................................................ 98 Hylini Ra®nesque, 1815 .................................................... 98 Acris DumeÂril and Bibron, 1841 .......................................... 99 Anotheca Smith, 1939 ................................................... 99 Bromeliohyla new genus ................................................. 99 Charadrahyla, new genus ................................................ 99 Duellmanohyla Campbell and Smith, 1992 ................................ 100 Ecnomiohyla, new genus ................................................ 100 Exerodonta Brocchi, 1879 ............................................... 100 Exerodonta sumichrasti Group ......................................... 101 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 294 Species of Exerodonta Unassigned to Group ............................ 101 Hyla Laurenti, 1768
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