systematic review of the frog family hylidae,with special
DESCRIPTION
Systematic Review of the Frog Family Hylidae,With SpecialTRANSCRIPT
SYSTEMATIC REVIEW OF THE FROG
FAMILY HYLIDAE, WITH SPECIAL
REFERENCE TO HYLINAE:
PHYLOGENETIC ANALYSIS AND
TAXONOMIC REVISION
JULIAN FAIVOVICHDivision of Vertebrate Zoology (Herpetology), American Museum of Natural History
Department of Ecology, Evolution, and Environmental Biology (E3B)Columbia University, New York, NY
CELIO F.B. HADDADDepartamento de Zoologia, Instituto de Biociencias, Unversidade Estadual Paulista, C.P. 199
13506-900 Rio Claro, Sao Paulo, Brazil([email protected])
PAULO C.A. GARCIAUniversidade de Mogi das Cruzes, Area de Ciencias da Saude
Curso de Biologia, Rua Candido Xavier de Almeida e Souza 20008780-911 Mogi das Cruzes, Sao Paulo, Brazil
and Museu de Zoologia, Universidade de Sao Paulo, Sao Paulo, Brazil([email protected])
DARREL R. FROSTDivision of Vertebrate Zoology (Herpetology), American Museum of Natural History
JONATHAN A. CAMPBELLDepartment of Biology, The University of Texas at Arlington
Arlington, Texas 76019([email protected])
WARD C. WHEELERDivision of Invertebrate Zoology, American Museum of Natural History
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 294, 240 pp., 16 figures, 2 tables, 5 appendices
Issued June 24, 2005
Copyright q American Museum of Natural History 2005 ISSN 0003-0090
2
CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Resumo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Taxon Sampling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Outgroup Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Basal Neobatrachians . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Ranoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Hyloidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
The Ingroup: Hylidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Hemiphractinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Pelodryadinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Phyllomedusinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19Hylinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Gladiator Frogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21Andean Stream-Breeding Hyla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26The 30-Chromosome Hyla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27Middle American/Holarctic Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32Casque-Headed Frogs and Related Genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38Species and Species Groups of Hyla Not Associated with Other Clades . . . . . 40Other Genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Character Sampling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43Gene Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43DNA Isolation and Sequencing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Da Silva’s (1998) Dissertation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45Phylogenetic Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Major Patterns of Relationships of Hylidae and Outgroups . . . . . . . . . . . . . . . . . . . . . . . 49Pelodryadinae and Phyllomedusinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Pelodryadinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53Phyllomedusinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Major Patterns of Relationships Within Hylinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54South American I Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Andean Stream-Breeding Hyla and the Tepuian Clade . . . . . . . . . . . . . . . . . . . . . . . 57Gladiator Frogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59Atlantic/Cerrado Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59Green Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60True Gladiator Frog Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
South American II Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61Relationships of Scinax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61Lysapsus, Pseudis, and Scarthyla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62Sphaenorhynchus, Xenohyla, and the 30-Chromosome Hyla . . . . . . . . . . . . . . . . . . 63
Middle American/Holarctic Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66Lower Central American Lineage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70Real Hyla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
South American/West Indian Casque-Headed Frogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72Taxonomic Conclusions: A New Taxonomy of Hylinae and Phyllomedusinae . . . . . . . . . 74
Hylinae Rafinesque, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
2005 3FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Cophomantini Hoffmann, 1878 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75Aplastodiscus A. Lutz in B. Lutz, 1950 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Aplastodiscus albofrenatus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82Aplastodiscus albosignatus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82Aplastodiscus perviridis Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Bokermannohyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82Bokermannohyla circumdata Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82Bokermannohyla claresignata Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83Bokermannohyla martinsi Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83Bokermannohyla pseudopseudis Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Hyloscirtus Peters, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84Hyloscirtus armatus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84Hyloscirtus bogotensis Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84Hyloscirtus larinopygion Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Hypsiboas Wagler, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85Hypsiboas albopunctatus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86Hypsiboas benitezi Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86Hypsiboas faber Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87Hypsiboas pellucens Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87Hypsiboas pulchellus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87Hypsiboas punctatus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88Hypsiboas semilineatus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88Species of Hypsiboas Unassigned to Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Myersiohyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89Dendropsophini Fitzinger, 1843 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Dendropsophus Fitzinger, 1843 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90Dendropsophus columbianus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90Dendropsophus garagoensis Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90Dendropsophus labialis Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91Dendropsophus leucophyllatus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91Dendropsophus marmoratus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91Dendropsophus microcephalus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91Dendropsophus minimus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92Dendropsophus minutus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92Dendropsophus parviceps Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Species of Dendropsophus Unassigned to Group . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Lysapsus Cope, 1862 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Pseudis Wagler, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Scarthyla Duellman and de Sa, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94Scinax Wagler, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Scinax catharinae Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95Scinax ruber Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Sphaenorhynchus Tschudi, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97Xenohyla Izecksohn, 1996 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Hylini Rafinesque, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98Acris Dumeril and Bibron, 1841 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99Anotheca Smith, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99Bromeliohyla new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99Charadrahyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99Duellmanohyla Campbell and Smith, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100Ecnomiohyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100Exerodonta Brocchi, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
Exerodonta sumichrasti Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
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Species of Exerodonta Unassigned to Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101Hyla Laurenti, 1768 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Hyla arborea Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101Hyla cinerea Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102Hyla eximia Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102Hyla versicolor Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102Species of Hyla Unassigned to Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Isthmohyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102Isthmohyla pictipes Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103Isthmohyla pseudopuma Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Megastomatohyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103Plectrohyla Brocchi, 1877 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Plectrohyla bistincta Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104Plectrohyla guatemalensis Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Pseudacris Fitzinger, 1843 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105Pseudacris crucifer Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105Pseudacris ornata Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106Pseudacris nigrita Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106Pseudacris regilla Clade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Ptychohyla Taylor, 1944 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106Smilisca Cope, 1865 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106Tlalocohyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107Triprion Cope, 1866 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Lophiohylini Miranda-Ribeiro, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107Aparasphenodon Miranda-Ribeiro, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108Argenteohyla Trueb, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108Corythomantis Boulenger, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108Itapotihyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109Nyctimantis Boulenger, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109Osteocephalus Steindachner, 1862 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109Osteopilus Fitzinger, 1843 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109Phyllodytes Wagler, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Phyllodytes auratus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110Phyllodytes luteolus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110Phyllodytes tuberculosus Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110Species of Phyllodytes Unassigned to Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
Tepuihyla Ayarzaguena and Senaris, ‘‘1992’’ [1993] . . . . . . . . . . . . . . . . . . . . . . . . 110Trachycephalus Tschudi, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Incertae Sedis and Nomina Dubia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111Phyllomedusinae Gunther, 1858 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Agalychnis Cope, 1864 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113Cruziohyla, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113Hylomantis Peters, ‘‘1872’’ [1873] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Hylomantis aspera Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115Hylomantis buckleyi Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
Pachymedusa Duellman, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115Phasmahyla Cruz, 1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115Phrynomedusa Miranda-Ribeiro, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116Phyllomedusa Wagler, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Phyllomedusa burmeisteri Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117Phyllomedusa hypochondrialis Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117Phyllomedusa perinesos Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117Phyllomedusa tarsius Group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
2005 5FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Species of Phyllomedusa Unassigned to Group . . . . . . . . . . . . . . . . . . . . . . . . . . 118Biogeographic Commentary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Relationships Between Australia and South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118Relationships Between South and Middle America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Guayana Highlands–Andes–Atlantic Forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119Guayana Highlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119Impact of the Andes in Hyline Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119Atlantic Forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
Origin of West Indian Hylids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123Middle America and the Holarctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124A Rough Temporal Framework: Clues from the Hylid Fossil Record . . . . . . . . . . . . . 125Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130Appendix 1: Summary of Taxonomic Changes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152Appendix 2: Locality Data and GenBank Accessions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174Appendix 3: Morphological Characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200Appendix 4: Additional Comments on Some Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202Appendix 5: List of Molecular Synapomorphies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205Note added in proof . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
6
ABSTRACT
Hylidae is a large family of American, Australopapuan, and temperate Eurasian treefrogsof approximately 870 known species, divided among four subfamilies. Although some groupsof Hylidae have been addressed phylogenetically, a comprehensive phylogenetic analysis hasnever been presented.
The first goal of this paper is to review the current state of hylid systematics. We focus onthe very large subfamily Hylinae (590 species), evaluate the monophyly of named taxa, andexamine the evidential basis of the existing taxonomy. The second objective is to perform aphylogenetic analysis using mostly DNA sequence data in order to (1) test the monophyly ofthe Hylidae; (2) determine its constituent taxa, with special attention to the genera and speciesgroups which form the subfamily Hylinae, and c) propose a new, monophyletic taxonomyconsistent with the hypothesized relationships.
We present a phylogenetic analysis of hylid frogs based on 276 terminals, including 228hylids 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 thespecies-rich genus Hyla. The included taxa allowed us to test the monophyly of 24 of the 35nonmonotypic genera and 25 species groups of Hyla. The phylogenetic analysis includesapproximately 5100 base pairs from four mitochondrial (12S, tRNA valine, 16S, and cyto-chrome b) and five nuclear genes (rhodopsin, tyrosinase, RAG-1, seventh in absentia, and28S), and a small data set from foot musculature.
Concurring with previous studies, the present analysis indicates that Hemiphractinae are notrelated to the other three hylid subfamilies. It is therefore removed from the family and ten-tatively considered a subfamily of the paraphyletic Leptodactylidae. Hylidae is now restrictedto Hylinae, Pelodryadinae, and Phyllomedusinae. Our results support a sister-group relation-ship between Pelodryadinae and Phyllomedusinae, which together form the sister taxon ofHylinae. Agalychnis, Phyllomedusa, Litoria, Hyla, Osteocephalus, Phrynohyas, Ptychohyla,Scinax, Smilisca, and Trachycephalus are not monophyletic. Within Hyla, the H. albomargin-ata, H. albopunctata, H. arborea, H. boans, H. cinerea, H. eximia, H. geographica, H. gra-nosa, H. microcephala, H. miotympanum, H. tuberculosa, and H. versicolor groups are alsodemonstrably nonmonophyletic. Hylinae is composed of four major clades. The first of theseincludes the Andean stream-breeding Hyla, Aplastodiscus, all Gladiator Frogs, and a Tepuianclade. The second clade is composed of the 30-chromosome Hyla, Lysapsus, Pseudis, Scar-thyla, Scinax (including the H. uruguaya group), Sphaenorhynchus, and Xenohyla. The thirdmajor clade is composed of Nyctimantis, Phrynohyas, Phyllodytes, and all South American/West Indian casque-headed frogs: Aparasphenodon, Argenteohyla, Corythomantis, Osteoce-phalus, Osteopilus, Tepuihyla, and Trachycephalus. The fourth major clade is composed ofmost of the Middle American/Holarctic species groups of Hyla and the genera Acris, Anotheca,Duellmanohyla, Plectrohyla, Pseudacris, Ptychohyla, Pternohyla, Smilisca, and Triprion. Anew monophyletic taxonomy mirroring these results is presented where Hylinae is dividedinto four tribes. Of the species currently in ‘‘Hyla’’, 297 of the 353 species are placed in 15genera; of these, 4 are currently recognized, 4 are resurrected names, and 7 are new. Hyla isrestricted 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, andPternohyla is placed in the synonymy of Smilisca. The genus Dendropsophus is resurrectedto include all former species of Hyla known or suspected to have 30 chromosomes. Exerodontais resurrected to include the former Hyla sumichrasti group and some members of the formerH. miotympanum group. Hyloscirtus is resurrected for the former Hyla armata, H. bogotensis,and H. larinopygion groups. Hypsiboas is resurrected to include several species groups—manyof them redefined here—of Gladiator Frogs. The former Hyla albofrenata and H. albosignatacomplexes of the H. albomarginata group are included in Aplastodiscus.
New generic names are erected for (1) Agalychnis calcarifer and A. craspedopus; (2) Os-teocephalus 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 ofthe H. pictipes and H. pseudopuma groups; and (10) a clade composed of the H. circumdata,H. claresignata, H. martinsi, and H. pseudopseudis groups.
2005 7FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
RESUMO
A famılia Hylidae e constituıda por aproximadamente 870 especies, agrupadas em quatrosubfamılias e com distribuicao nas Americas, Australia/Papua-Nova Guine e Eurasia. Apesarde alguns grupos de Hylidae terem sido estudados separadamente, uma hipotese filogeneticacompreensiva para a famılia nunca foi proposta.
O objetivo inicial desse estudo e rever o atual estado sistematico da famılia Hylidae. Atencaoespecial e dada a subfamılia Hylinae (590 especies), para a qual nos avaliamos o monofiletismodos taxa atualmente reconhecidos e examinamos as bases do arranjo taxonomico aceito nopresente. O segundo objetivo e realizar uma analise filogenetica usando caracteres obtidos, emsua maioria, a partir de sequencias de ADN, a fim de (1) testar o monofiletismo da famıliaHylidae;(2) determinar sua constituicao taxonomica, dando enfase aos generos e grupos deespecies incluıdos na subfamılia Hylinae; e (3) propor uma nova taxonomia baseada em gruposmonofileticos e consistente com a hipotese filogenetica aqui apresentada.
Neste trabalho apresentamos, juntamente com uma revisao sistematica dos hilıdeos, umaanalise filogenetica baseada em 275 terminais, sendo 227 de hilıdeos, mais 48 taxas represen-tando grupos externos. Na analise filogenetica estao representados 40 dos 41 generos de Hy-lidae das quatro subfamılias reconhecidas e 39 dos 41 grupos atualmente reconhecidos para ogrande genero Hyla. Os taxons incluıdos permitem testar a monofilia de 24 dos generos naomonotıpicos e 25 grupos de especies de Hyla. A analise filogenetica inclui aproximadamente5100 pares de bases de quatro genes mitocondriais (12S, tval, 16S, citocromo b) e cinco genesnucleares (rodopsina, tirosinase, RAG-1, seventh in absentia e 28s), alem de um pequenoconjunto de dados sobre a musculatura do pe.
De forma similar ao que tem sido observado em outros estudos, a presente analise indicaque os Hemiphractinae nao sao relacionados as outras tres subfamılias de hilıdeos, portanto,sendo removidos desta famılia e tentativamente considerados como uma subfamılia dos par-afileticos Leptodactylidae. Hylidae e agora restrita a Hylinae, Pelodryadinae e Phyllomedusi-nae. Nossos resultados corroboram uma relacao de grupos irmaos entre estas duas ultimassubfamılias, que juntas correspondem ao taxon irmao de Hylinae. Phyllomedusa, Agalychnis,Litoria, Hyla, Osteocephalus, Phrynohyas, Pseudis, Ptychohyla, Scinax, Smilisca e Trachy-cephalus nao sao monofileticos. Dentro do genero Hyla, os grupos de H. albomarginata, H.albopunctata, H. arborea, H. boans, H. cinerea, H. eximia, H. geographica, H. granosa, H.microcephala, H. miotympanum, H. tuberculosa e H. versicolor nao sao monofileticos.
Em nossa analise, Hylinae aparece composta por quatro grandes clados. O primeiro delesincluindo todas as ras-gladiadoras, as especies andinas de Hyla que se reproduzem em ria-chos e um clado dos Tepuis. O segundo grande clado e composto por Scinax, Sphaenorhyn-chus, ‘‘pseudıdeos’’, Scarthyla e as especies de Hyla com 30 cromossomos. O terceiro gran-de clado e composto por Phyllodytes, Phrynohyas, Nyctimantis e todas as seguintes perer-ecas-de-capacete da America do Sul e Indias Ocidentais: Argenteohyla, Aparasphenodon,Corythomantis, Osteocephalus, Osteopilus, Tepuihyla e Trachycephalus. O quarto e ultimogrande clado e composto pela maioria dos grupos de especies de Hyla centro-americanos/holarticos e pelos generos Acris, Anotheca, Duellmanohyla, Plectrohyla, Pseudacris, Pty-chohyla, Pternohyla, Smilisca e Triprion. E apresentada uma nova taxonomia monofiletica,espelhando estes resultados, onde Hylinae e dividida em quatro tribos. Das especies corren-temente incluıdas em Hyla, 297 de 353 sao alocadas em 15 generos, dos quais quatro saocorrentemente reconhecidos, quatro sao nomes revalidados e seis sao novas descricoes. Ogenero Hyla fica restrito aos grupos de H. arborea, H. cinerea, H. eximia, H. femoralis eH. versicolor, sendo o conteudo de alguns destes grupos redefinido. Phrynohyas e sinon-imizada a Trachycephalus, Pternohyla e sinonimizada a Smilisca e Duellmanohyla e sinon-imizada a Ptychohyla. O genero Dendropsophus e revalidado para as especies de Hyla com,ou presumivelmente tendo, 30 cromossomos. Exerodonta e revalidado, passando a incluiros grupos de Hyla sumichrasti e H. pinorum. Hyloscirtus e revalidado para acomodar osgrupos de Hyla armata, H. bogotensis e H. larinopygion. Hypsiboas e revalidada para acom-odar diversos grupos de especies—muitos deles aqui redefinidos—de ras-gladiadoras. Oscomplexos de Hyla albofrenata e H. albosignata, do grupo de H. albomarginata, sao in-cluıdos no genero Aplastodiscus.
Nomes genericos novos sao apresentados para (1) Agalychnis calcarifer e A. craspedopus,(2) Osteocephalus langsdorffii e para os grupos de especies de (3) Hyla aromatica, (4) H.
8 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
bromeliacia, (5) H. godmani, (6) H. mixomaculata, (7) H. taeniopus, (8) H. tuberculosa, epara o clado compostos pelos grupos de especies de (9) H. pictipes e H. pseudopuma e parao clado composto pelos grupos de especies de (10) H. circumdata, H. claresignata, H. martinsie H. pseudopseudis.
RESUMEN
Hylidae es una familia muy grande (aproximadamente 870 especies conocidas) de ranasarboricolas de America, Australia y Papua, y Eurasia, y esta dividida en cuatro subfamilias.Aunque existen algunos estudios que analizan las relaciones filogeneticas de algunos gruposaislados de Hylidae, no existe ningun analisis filogenetico de toda la familia.
Los objetivos de este trabajo son, primero, revisar el estado actual de la sistematica de lafamilia, haciendo enfasis en la subfamilia Hylinae, que es la mas grande (590 especies), yevaluando la evidencia existente para la monofilia de los distintos agrupamientos taxonomicos.El segundo objetivo es realizar un analisis filogenetico basado principalmente en secuenciasde ADN, con el proposito de a) testear la monofilia de la familia Hylidae, b) determinar quetaxa la constituyen, con especial atencion en los generos y grupos de especies de la subfamiliaHylinae, c) proponer una nueva taxonomia monofiletica, consistente con nuestra hipotesisfilogenetica.
Se presenta una revision completa del estado de conocimiento de la sistematica de la familiaHylidae, junto con un analisis filogenetico de 276 terminales, incluyendo 228 Hylidae y 48grupos externos. Se incluyen representantes de 40 de los 41 generos de de las cuatro subfam-ilias de Hylidae, y de 39 de los 41 grupos de especies del genero Hyla. Asimismo, los taxaincluidos permitieron testear la monofilia de 24 de los 35 generos no monotipicos, y 25 gruposde especies de Hyla. El analisis filogenetico incluye aproximadamente 5100 pares de bases decuatro genes mitocondriales (12S, tRNA valina, 16S, cytocromo b) y cinco genes nucleares(rhodopsina, RAG-1, seventh in absentia, tyrosinasa y 28S), y una matriz de 38 characteresde musculatura del pie.
En coincidencia con estudios anteriores, los resultados del analisis indican que los Hemi-phractinae no pertenecen a Hylidae, y por lo tanto se los excluye de la familia, que hora esrestringida a Hylinae, Pelodryadinae, y Phyllomedusinae. Nuestros resultados soportan la mo-nofilia de estas dos ultimas subfamilias, que a su vez son el taxon hermano de Hylinae.Phyllomedusa, Agalychnis, Litoria, Hyla, Osteocephalus, Phrynohyas, Trachycephalus, Smi-lisca, Ptychohyla, y Scinax no son monofleticos. Ademas, dentro de Hyla, los grupos de H.albomarginata, H. albopunctata, H. arborea, H. boans, H. cinerea, H. eximia, H. geographica,H. granosa, H. microcephala, H. miotympanum, H. tuberculosa, y H. versicolor tampoco sonmonofileticos. Hylinae resulta estar compuesto por cuatro clados principales. El primero deestos incluye a Aplastodiscus y todos los grupos de especies de Hyla incluidos en las ranasgladiadoras, las Hyla que se reproducen en arroyos de los Andes, y un clado de los TepuiesGuayaneses. El segundo clado principal esta compuesto por Scarthyla, Scinax, Sphaenorhyn-chus, ‘‘pseudidos’’, y las Hyla de 30 cromosomas. El tercer clado principal esta compuestopor Nyctimantis, Phyllodytes, Phrynohyas, y todas las ranas ‘‘cabeza de casco’’ sudamericanasy de las Indias Occidentales: Argenteohyla, Aparasphenodon, Corythomantis, Osteopilus, Os-teocephalus, Trachycephalus, y Tepuihyla. El cuarto clado principal esta compuesto por lamayoria de los grupos de especies de Hyla Centro Americanos y holarticos y los generosAcris, Anotheca, Duellmanohyla, Plectrohyla, Pseudacris, Ptychohyla, Pternohyla, Smilisca,y Triprion.
Con base en estos resultados, se presenta una nueva taxonomia monofiletica, adonde Hylinaees dividida en cuatro tribus. Ademas, 297 de las 353 especies hasta ahora incluidas en Hylason divididas en 15 generos, cuatro de los cuales son generos que ya estaban en uso, cuatroson nombres resucitados de la sinonimia de Hyla, y siete son nuevos. Hyla es restringido aH. femoralis y los grupos de H. arborea, H. cinera, H. eximia, e H. versicolor, cuyos conten-idos son en algunos casos redefinidos. Asi mismo, Phrynohyas es incluido en la sinonimia deTrachycephalus, y Pternohyla en la sinonimia de Smilisca. Dendropsophus es revalidado paraincluir todas las especies previamente incluidas en los grupos de especies de 30 cromosomas,o sospechadas de tener 30 cromosomas. Exerodonta es revalidado para incluir el grupo deHyla sumichrasti, y un fragmento de especies incluidas en el grupo de H. miotympanum.
2005 9FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Hyloscirtus es revalidado para incluir los grupos de H. armata, H. bogotensis, e H. larino-pygion. Hypsiboas es revalidado para incluir muchos grupos de especies—varios de ellos aquiredefinidos—de ranas gladiadoras. Los complejos de Hyla albofrenata e H. albosignata delgrupo de H. albomarginata son incluidos en Aplastodiscus.
Nuevos nombres genericos son propuestos para (1) Agalychnis calcarifer y A. craspedopus,(2) Osteocephalus langsdorffii, los grupos de (3) Hyla aromatica, (4) H. bromeliacia, (5) H.godmani, (6) H. mixomaculata,(7) H. taeniopus, (8) H. tuberculosa, y para los clados com-puestos por los grupos de (9) H. pictipes y H. pseudopuma, y por los grupos de (10) H.circumdata, H. claresignata, H. martinsi, e H. pseudopseudis.
INTRODUCTIONHylidae is a large family of American,
Australopapuan, and temperate Eurasiantreefrogs of approximately 870 known spe-cies, composed of four subfamilies (Duell-man, 2001; Darst and Cannatella, 2004;Frost, 2004). Although some groups of Hy-lidae have been addressed phylogenetically(e.g., Campbell and Smith, 1992; Duellmanand Campbell, 1992; Mendelson et al., 2000;Faivovich, 2002; Haas, 2003; Moriarty andCannatella, 2004; Faivovich et al., 2004), acomprehensive phylogenetic analysis hasnever been presented.
The first goal of this paper is to review thestate of hylid systematics. We focus on thevery large subfamily Hylinae, evaluate themonophyly of named taxa, and examine theevidential basis of the existing taxonomy.The second objective is to perform a phylo-genetic analysis using four mitochondrial andfive nuclear genes in order to (1) test themonophyly of the family Hylidae; (2) deter-mine its constituent taxa, with special atten-tion to the genera and species groups whichform the subfamily Hylinae; and (3) proposea new, monophyletic taxonomy consistentwith the hypothesized relationships.
MATERIALS AND METHODSIn most revisionary studies involving ma-
jor taxonomic rearrangements and phyloge-netic analyses it is normal to have a sectionon the history of taxonomic changes. Thescale of this particular study makes that goalimpractical. A discussion of the state of thetaxonomy of hylid frogs is dealt with simul-taneously with discussions of the taxa chosenfor the purpose of phylogenetic analysis.
TAXON SAMPLING
Any phylogenetic analysis has an impor-tant component of experimental design as-
sociated with the selection of the terminaltaxa. In an ideal phylogenetic study, all ter-minal descendants of a given hypotheticalancestor should to be included in order toavoid ‘‘problems’’ due to taxon sampling.This ideal condition is unattainable, and allnotions of relationships among organisms areaffected to an unknown degree by incom-plete taxon sampling. Because there is noway of ever knowing all the hylid speciesthat have become extinct, we concentrate onthe diversity that we do know. Furthermore,due to the unavailability of samples we can-not include sequences of all of the nearly 860currently described species of Hylidae. What,therefore, is the best taxon sampling for thisstudy? Because our primary goal is to testthe monophyly of all available genera andspecies groups of Hylidae, the most appro-priate terminals to include are those that pro-vide the strongest test of previously hypoth-esized relations. By considering morpholog-ical divergence as a rough guide to DNA se-quence diversity, maximally diverse taxawithin a given group are likely to pose astronger test of its monophyly than do mor-phologically similar taxa (Prendini, 2001).Groups for which no apparent synapomor-phies are known are a priori more likely tobe nonmonophyletic, and therefore good rep-resentations of the morphological diversity ofthese groups are especially appropriate. Oursuccess varied in securing multiple represen-tatives of these groups.
The following discussion deals in partwith the state of knowledge of frog phylo-genetics. Included within the discussion is alist of terminals used in this analysis alongwith a justification and explanation for ourchoices. A summary of the species includedis presented in table 1. To conserve space,species authorships are not mentioned in the
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TABLE 1Species Included in this Analysis and Species Groups or Genera They Represent
2005 11FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
TABLE 1(Continued)
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TABLE 1(Continued)
2005 13FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
text but are given in the section ‘‘TaxonomicConclusions: A New Taxonomy of Hylinaeand Phyllomedusinae’’ and in appendix 1.For museum abbreviations used throughoutthis paper see appendix 2.
OUTGROUP SELECTION
Recent studies (Haas, 2003; Darst andCannatella, 2004) have suggested that theHylidae as traditionally understood is notmonophyletic, with the Hemiphractinae dis-placed phylogenetically from the Hylinae,Phyllomedusinae, and Pelodryadinae. Theaforementioned studies did not provide ex-tensive outgroup comparisons. In order toavail ourselves of a strong test of hylidmonophyly, we included 48 nonhylid out-group taxa representing 14 neobatrachianfamilies.
Basal Neobatrachians
Heleophrynidae, Sooglossidae, Limnodyn-astinae and Myobatrachinae1 have been re-lated to each other by several authors (Lynch,1973; Duellman and Trueb, 1986; Ford andCannatella, 1993; Hay et al., 1995; Ruvinskyand Maxson, 1996; Biju and Bossuyt, 2003;Darst and Cannatella, 2004). While in thepast they were considered part of Hyloidea,they were recently excluded by Darst andCannatella (2004). The evidence indicatesthat they are basal neobatrachians distantlyrelated to the apparently monophyletic Hy-loidea; however, their exact positions and in-terrelationships are still unclear (Darst andCannatella, 2004; Haas, 2003). We includeone heleophrynid (Heleophryne purcelli),one myobatrachine (Pseudophryne bibroni),and two limnodynastines (Limnodynastessalmini and Neobatrachus sudelli) in ourstudy. Furthermore, because some membersof the Australopapuan hylids have been pos-ited to be related to the Myobatrachidae(Lynch, 1971; Savage, 1973), their inclusionprovides a strong test of hylid monophyly.
1 We are referring to these two myobatrachid subfam-ilies separately; we are not aware of any putative syna-pomorphy supporting the monophyly of Myobatrachi-dae.
Ranoidea
Recent papers dealing with ranoid groups(Emerson et al., 2000; Vences et al., 2003a)or at least ranoid exemplars (Biju and Bos-suyt, 2003; Darst and Cannatella, 2004) havesuggested the existence of three majorclades, although this remains to be elucidat-ed. The three major clades are composed of(1) Arthroleptidae, Astylosternidae, Hemi-sotidae, and Hyperoliidae; (2) Microhylidae;and (3) the remaining ranoids (including Pe-tropedetidae, Mantellidae, Rhacophoridae,and the paraphyletic Ranidae). We includeexemplars of Astylosternidae, Hyperoliidae,Hemisotidae, Microhylidae, Ranidae, Man-tellidae, and Rhacophoridae.
Hyloidea
The Hylidae has long been considered tobe embedded within a poorly defined majorgroup of neobatrachians (Nicholls, 1916; No-ble, 1922; Lynch, 1971, 1973; Ford and Can-natella, 1993) for which no morphologicalevidence of monophyly exists, although mo-lecular evidence (Hay et al., 1995; Ruvinskyand Maxson, 1996; Darst and Cannatella,2004) does support its monophyly. As rede-fined by Darst and Cannatella (2004) Hylo-idea includes the nonmonophyletic Lepto-dactylidae (Ford and Cannatella, 1993; Haas,2003), Dendrobatidae, Hylidae, Bufonidae,Brachycephalidae, Centrolenidae, Rhinoder-matidae, and the monotypic Allophrynidae.In order to have a strong test of the mono-phyly of hylids, we include representativesof most of the nonhylid hyloid families.
The monophyly of Dendrobatidae is notcontroversial (see Grant et al., 1997), al-though recent phylogenetic analyses using ri-bosomal mitochondrial sequences (Vences etal., 2000; Santos et al., 2003; Vences et al.,2003b) show that there is a serious need toredefine most of the currently recognizedgenera. According to the results of Vences etal. (2003b), there are two major clades ofdendrobatids: (1) one composed of Dendro-bates, Phyllobates, Cryptophyllobates, Epi-pedobates (paraphyletic), Minyobates, andseveral groups of the rampantly polyphyleticColostethus; and (2) another clade composedof Mannophryne, Nephelobates, Allobates,and two separate clades of Colostethus (none
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of the aforementioned analyses included theapparently primitive genus Aromobates; seeMyers et al. 1991). We include as exemplarsof the first major clade Dendrobates auratusand Phyllobates bicolor, and as exemplar ofthe second clade, Colostethus talamancae.
Bufonidae is a monophyletic group (for alist of morphological synapomorphies, seeFord and Cannatella, 1993; Haas, 2003) forwhich no study addressing comprehensivelyits internal relationships has been published.Partial studies (Graybeal, 1997; Darst andCannatella, 2004; Haas, 2003) support theidea that Melanophryniscus is one of its mostbasal clades in the family. As a rough sampleof bufonid diversity we include representa-tives of Melanophryniscus, Dendropryniscus,Atelopus, Didynamipus, Schismaderma,Bufo, Pedostibes, and Osornophryne.
The notion that the presumably monophy-letic Centrolenidae (Ruiz-Carranza andLynch, 1991) is closely related to hylids(Lynch, 1973; Duellman and Trueb, 1986;Ford and Cannatella, 1993) was recentlychallenged by the phylogenetic analyses ofHaas (2003), who used larval morphology,and Darst and Cannatella (2004), who usedmitochondrial ribosomal genes. Austin et al.(2002) recently provided molecular evidencesupporting a relationship between Centrolen-idae and the monotypic Allophrynidae. Theinternal relationships of Centrolenidae re-main virtually unstudied. As a rough repre-sentation of centrolenid diversity, in thisstudy we include Centrolene prosoblepon,Cochranella bejaranoi, and Hyalinobatrach-ium eurygnathum. We also include Allophry-ne ruthveni.
Leptodactylid nonmonophyly has been ac-cepted for some time (Lynch, 1971) and wasconfirmed in an explicit cladistic frameworkby analyses using morphology (Haas, 2003)and DNA sequences (Ruvinsky and Maxson,1996; Darst and Cannatella, 2004; Vences etal., 2003b). In the analysis by Darst and Can-natella (2004), Leptodactylidae is rampantlyparaphyletic because all the other hyloid ex-emplars are nested within it.
From the five currently recognized sub-families of leptodactylids (Laurent, 1986)there is more or less convincing evidence ofmonophyly for two of them: Eleutherodac-tylinae (direct development, eggs relatively
large, few in number; Lynch, 19712) andLeptodactylinae (presence of a bony elementin the sternum; Lynch, 1971). The analysisof Darst and Cannatella (2004) corroboratedthe monophyly of these two groups, albeitwith limited taxon sampling. In the analysisby Haas (2003), the exemplars of Leptodac-tylinae were not monophyletic. No demon-strable synapomorphies are known for Cer-atophryinae, Cycloramphinae or Telmatobi-inae. Considering this situation, we includeseveral leptodactylid exemplars (see table 1);our poorest sampling is within Cycloramphi-nae, where we only have representation forone genus, Crossodactylus.
The single representative of Brachyce-phalidae included by Darst and Cannatella(2004) in their analysis was nested within theexemplars of the leptodactylid subfamilyEleutherodactylinae, as suggested earlier byIzecksohn (1988). We include Brachycephal-us ephippium in our analysis.
Of the hyloid families, only Rhinoderma-tidae is not represented in our analysis. Thisgroup was suggested to be nested in the sub-family Telmatobiinae by Barrio and Rinaldide Chieri (1971) based on a similar karyo-type and by Manzano and Lavilla (1995a)based on the presence of the m. pelvocuta-neus in Rhinoderma and Eupsophus. In theDNA sequences analyses by Ruvinsky andMaxson (1996) and Biju and Bossuyt (2003)Rhinoderma appears in different positionswithin Hyloidea.
THE INGROUP: HYLIDAE
Inasmuch as the hylids are the primary fo-cus of this study, our sampling is most densefor this taxon and requires substantially moredetailed discussion than does our outgroupselection.
Duellman (1970) arranged the family infour subfamilies: Amphignathodontinae,Hemiphractinae, Hylinae (including both theAustralian and American groups), and Phyl-lomedusinae. Trueb (1974) subsequently syn-onymized the Amphignathodontinae andHemiphractinae. On the basis of evidence
2 Note that Lynch (1971) presented an extensive def-inition of the group; it is unclear if any of the othercharacter states he mentioned could be considered syn-apomorphic.
2005 15FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
presented by Tyler (1971), Duellman (1977)placed the Australian hylids in their own sub-family, Pelodryadinae, although Savage(1973) had previously regarded it as a dif-ferent family and suggested that it was de-rived from Myobatrachidae.
Duellman (2001), based mostly on da Sil-va’s results (1998), presented a phylogeneticanalysis where hylids, including the subfam-ily Pseudinae, were considered to be mono-phyletic. Synapomorphies suggested byDuellman (2001) as being common to histhree most parsimonious trees are the pos-session of claw-shaped terminal phalangesand the three articular surfaces on metacarpalIII. The possibility of hylid polyphyly hasnot been seriously considered by most frogsystematists, even after Ruvinsky and Max-son’s (1996) results (which showed their sin-gle Hemiphractinae exemplar not closely re-lated with the other hylid exemplars), untilthe idea was suggested on morphologicalgrounds by Haas (2003), followed by Darstand Cannatella (2004) on the basis of molec-ular evidence. These authors found no evi-dence of a relationship between Hemiphrac-tinae and the other hylid subfamilies, whichwere thought to form a monophyletic group.Beyond this result, Haas (2003) presentedevidence from larval morphology that sug-gested that Pelodryadinae is paraphyleticwith respect to Hylinae (Hylinae not beingdemonstrably monophyletic), with Pseudinaeand Phyllomedusinae possibly being imbed-ded within it.3
3 Burton (2004) presented a study of hylid foot my-ology, a valuable collection of observations on manyspecies, including the definition of numerous characters,and a phylogenetic analysis of the hylid subfamiliescombining his characters with those employed by Duell-man (2001). The list of synapomorphies provided(Burton 2004: 228) is the result of optimizing the dataset on his strict consensus tree. When only unambiguoussynapomorphies common to all the most parsimonioustrees are considered, the list is reduced considerably, andthere are no unambiguous transformations from footmusculature that support relationships among the sub-families (these are supported solely by the charactersfrom Duellman, 2001). Instead, some foot muscle char-acter states are autapomorphic for Allophrynidae, Hem-iphractinae, Phyllomedusinae, and Pseudinae (Burton’spaper was submitted before Darst and Cannatella’s paperwas published). Throughout the present paper, all thesynapomorphies reported from Burton’s (2004) study areonly those that occur in all equally parsimonious trees.
Hemiphractinae
Mendelson et al. (2000) and Duellman(2001) presented brief taxonomic histories ofthis taxon. The monophyly of Hemiphracti-nae is supported by the presence of bell-shaped gills in larvae and by female transportof eggs in a specialized depression or sac inthe dorsum (Noble, 1927). Burton (2004)added the broad m. abductor brevis plantaehallucis. In Duellman’s (2001) cladogram,Hemiphractinae is considered to be the sistergroup of Phyllomedusinae, with the evidenceof this relationship being the proximal headof metacarpal II not between prepollex anddistal prepollex, and the larval spiracle sinis-tral and ventrolateral.
Haas’s (2003) exemplar4 of the Hemi-phractinae was Gastrotheca riobambae. Hisresults suggested that Hemiphractinae are un-related to other hylids, although the positionof Hemiphractinae within Neobatrachia isstill unresolved. A similar result regardingHemiphractinae as being unrelated to hylidswas advanced by Ruvinsky and Maxson(1996) and corroborated by Darst and Can-natella (2004). These authors also did not re-cover the exemplars of Hemiphractinae thatthey used (Gastrotheca pseustes and Cryp-tobatrachus sp.) as forming a monophyleticgroup.5
Hemiphractinae includes five genera:Cryptobatrachus, Flectonotus, Gastrotheca,Hemiphractus, and Stefania. Mendelson etal. (2000) studied the relationships amongthese genera, performing a phylogeneticanalysis using morphological and life-historycharacters, arriving at the topology (Crypto-batrachus Flectonotus (Stefania (‘‘Gastro-theca’’ 1 Hemiphractus))). This analysis in-cluded five outgroups, all of which were rep-resentatives of the other hylid subfamilies.The results suggested that Cryptobatrachusand Flectonotus are each monophyletic, andthat Gastrotheca is paraphyletic with Hemi-phractus nested within it. As Hass (2003)
4 Haas (2003) noted that the larval morphology ofseveral species of Gastrotheca is quite similar, so pre-sumably his selection of G. riobambae as an exemplarwould have little effect on the analysis.
5 However, a reanalysis of their data using parsimonyand considering insertions/deletions as a fifth state didrecover a monophyletic Hemiphractinae (Faivovich, per-sonal obs.).
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noted, however, Mendelson et al.’s (2000)outgroup structure could not test the propo-sition of hylid diphyly.
Cryptobatrachus: In the analysis per-formed by Mendelson et al. (2000), themonophyly of the two representatives ofCryptobatrachus is supported by several os-teological characters, among them the pres-ence of an anteromedial process in the neo-palatine.6 In their analysis, the relationshipbetween Cryptobatrachus and the otherHemiphractinae is unresolved. This genuscomprises three described species; in thisstudy we include sequences of an unidenti-fied species available from GenBank.
Gastrotheca: Mendelson et al. (2000) sug-gested that Hemiphractus is nested withinGastrotheca, a result that contrasts with theopinions of previous workers (Noble, 1927;Trueb, 1974; Duellman and Hoogmoed,1984) who considered Hemiphractus basalamong marsupial frogs because they lack abrooding pouch. However, Mendelson et al.(2000) continued to recognize Hemiphractus(the older of both names) and Gastrothecapending a more complete phylogenetic study.Synapomorphies of the clade composed ofthese two genera are: cultriform process be-coming distinctly narrow anteriorly; anteriorprocess of vomer articulating only with max-illa; pre- and postchoanal process bifurcatingat the level of the dentigerous process; natureof occipital artery pathway (a groove);brooding pouch with posterior opening; andbell-shaped gills fused distally.
Most of the 49 currently recognized spe-cies of Gastrotheca are placed in four speciesgroups, the G. marsupiata, G. nicefori, G.plumbea, and G. ovifera groups (Duellmanet al., 1988a). These groups are generally de-fined on the basis of overall similarity. Theonly character-based test of their monophyly,the analysis of Mendelson et al. (2000), in-cluded 17 exemplars and suggested that noneof the three nonmonotypic groups is mono-phyletic.
Dubois (1987) placed the species withinthree subgenera: Gastrotheca, Duellmania
6 For the synapomorphy list, we copied the data setfrom the. pdf file of Mendelson et al. (2000) and eval-uated character distribution in TNT (Goloboff et al.,2000).
(part of the G. plumbea group), and Opistho-delphys (G. ovifera as well as parts of the G.marsupiata and G. plumbea groups). Duell-mania and Opisthodelphys were shown to beparaphyletic by Mendelson et al. (2000), al-though they did not test the monophyly ofthe subgenus Gastrotheca. In this study weinclude two species of the Gastrotheca mar-supiata group (G. cf. marsupiata and G.pseustes) and two of the G. ovifera group (G.cornuta and G. fissipes).
Hemiphractus: Relationships of this genuswere recently reviewed by Sheil et al. (2001),who provided a number of unambiguous syn-apomorphies to support its monophyly (suchas the cultriform process of the parasphenoidthat becomes distinctly narrow anteriorly, thepresence of a zygomatic ridge, and the pres-ence of a supraorbital ridge). Hemiphractushas six described species; we include Hemi-phractus helioi in our study.
Stefania: Duellman and Hoogmoed(1984), Senaris et al. (‘‘1996’’ [1997]), andMacCulloch and Lathrop (2002) reviewedthis genus. Senaris et al. (‘‘1996’’ [1997])suggested that the zygomatic ramus of thesquamosal being close to or in contact withthe maxilla was a diagnostic character stateof Stefania, ‘‘at least for the Venezuelan spe-cies’’ (translated from the Spanish). Mendel-son et al. (2000) did not test the monophylyof Stefania since they included only one ex-emplar (S. evansi). It is unclear if any of theautapomorphies of S. evansi in that study areactually synapomorphies of Stefania.
Stefania was divided by Rivero (1970)into two species groups based on head shape(‘‘as broad as long or longer than broad’’ inthe S. evansi group; ‘‘much broader thanlong’’ in the S. ginesi group). The only testof the monophyly of these two groups is thephylogenetic analysis of the seven speciesthen known, performed by Duellman andHoogmoed (1984). In that analysis, the S. gi-nesi group was monophyletic and nestedwithin the paraphyletic S. evansi group. Al-though Senaris et al. (‘‘1996’’ [1997]) sug-gested the origin of the S. ginesi group fromthe S. evansi group, they continued to rec-ognize of both groups.
Since the revision of Duellman and Hoog-moed (1984), another 11 species assigned toboth species groups of Stefania have been
2005 17FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
named (see Barrio Amoros and Fuentes,2003; MacCullogh and Lathrop, 2002). Inour analysis we include one exemplar of theStefania evansi group (S. evansi) and one ofthe S. ginesi group (S. schuberti).
Flectonotus: Duellman and Gray (1983)reviewed these frogs as two genera, Flecto-notus and Fritziana, even though their phy-logenetic analysis indicated that Flectonotuswas paraphyletic with respect to Fritziana.Subsequently, Weygoldt and Carvalho e Sil-va (1991) placed Fritziana in the synonymyof Flectonotus to render a monophyletic tax-onomy. Flectonotus is composed of five de-scribed species.
In the analysis by Mendelson et al. (2000),the position of Flectonotus remained unre-solved with respect to Cryptobatrachus andthe clade composed of Stefania plus Gastro-theca. Synapomorphies of Flectonotus in thatanalysis are: quadratojugal that does not ar-ticulate with maxilla; brooding pouch formedby dorsolateral folds of skin; overlap be-tween m. intermandibularis and m. submen-talis; and absence of supplementary elementsof m. intermandibularis. Because of verylimited availability of species of Flectonotus,in this study we include only Flectonotus sp.,an unidentified species from southeasternBrazil, whose female has a brooding pouchwith a middorsal slit.
Pelodryadinae
The monophyly of this Australopapuangroup is supported by a single possible syn-apomorphy: presence of supplementary api-cal elements of m. intermandibularis (Tyler,1971). Although relationships between Aus-tralian and New World hylids were recog-nized very early (most species of Litoriawere named as Hyla), hypotheses regardingthe relationships of Pelodryadinae with othergroups have been rarely advanced. Trewavas(1933), Duellman (1970), and Bagnara andFerris (1975) suggested a relationship be-tween Pelodryadinae with Phyllomedusinae.Trewavas (1933) observed similarities in la-ryngeal structures in the limited data set ather disposal (only three pelodryadines andtwo phyllomedusines). Duellman (1970) re-ferred to ‘‘similarities in vertebral charac-ters’’ without further details and to identical
number of chromosomes. Bagnara and Ferris(1975) noticed the presence in some speciesof Litoria of large melanosomes containingthe red pigment rhodomelanochrome (lateridentified as pterorhodin; Misuraca et al.,1977), a character state previously knownonly for some species of Phyllomedusinae.Specifically, Bagnara and Ferris (1975) sug-gested that some species of Litoria could berelated to the Phyllomedusinae, an implicitsuggestion of Pelodryadinae paraphyly. Thisidea was discussed by Tyler and Davies(1978a), who rejected the possibility of pe-lodryadine paraphyly but did not address apossible sister-taxon relationship of Pelo-dryadinae and Phyllomedusinae. This alter-native was suggested again, based on chro-mosome morphology, by Kuramoto and Al-lison (1991). In the phylogenetic analysespresented by Duellman (2001), Pelodryadi-nae was placed as the sister of a clade com-posed of the remaining subfamilies, whichare united in having a distally bifid tendo su-perficialis. In this same cladogram, the onlysynapomorphy of Pelodryadinae is the ante-rior differentiation of the m. intermandibu-laris, although the monophyly of Pelodry-adinae was assumed and not tested in thatanalysis.
The phylogenetic analysis performed byHaas (2003) presents the most extensive testof Pelodryadinae monophyly so far pub-lished; his Pelodryadinae exemplars form aparaphyletic series with respect to his Neo-tropical hylid exemplars. More recently,Darst and Cannatella (2004) and Hoegg et al.(2004) presented evidence from the ribosom-al mitochondrial and nuclear genes support-ing the monophyly of their Pelodryadinaesample and the monophyly of Pelodryadinae1 Phyllomedusinae.
Litoria: The diagnosis and contents of Li-toria were reviewed by Tyler and Davies(1978b). It is unclear whether any of thecharacter states included in their extensivediagnosis are synapomorphic. However, con-sidering subsequent comments by several au-thors (King et al., 1979; Tyler, 1979; Tylerand Davies, 1979; Maxson et al., 1985; Haasand Richards, 1998), the available evidencesuggests that Litoria is paraphyletic with re-spect to the other genera of Pelodryadinae,
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Nyctimystes and Cyclorana, with the latterbeing the sister taxon of the L. aurea group.
The 132 currently recognized species (up-dated from Frost, 2004) placed in 37 speciesgroups (Tyler and Davies, 1978b) make anexhaustive sampling of the group a goal be-yond the present analysis.7 Relationshipsamong some species groups of Litoria wereaddressed by means of albumin immunolog-ical distances as determined through micro-complement fixation (Maxson et al., 1982;Hutchinson and Maxson, 1986, 1987). Froma character-based (as opposed to a distance-based) perspective, relationships among thespecies groups of Litoria remain unknown,and the monophyly of most of those groupswith more than single species remains un-tested.
Tyler and Davies (1978b) tentatively di-vided the 37 species groups into three ‘‘Cat-egories’’, A (8 species groups), B (14 speciesgroups), and C (7 species groups). Tyler(1982) added a fourth category (D), wherehe included the Litoria nannotis group.These groupings were criticized by King(1980) on karyotypic grounds. Besides, themonophyly of these four categories remainlargely untested, with the only possible ex-ception being the work by Cunningham(2002) on the L. nannotis group; however,his lack of sufficient outgroup sampling pre-cluded a rigorous test.
In our analyses we include representativesof two species groups of category A, Litoriaaurea (the L. aurea group) and L. freycineti(the L. freycineti group); three representa-tives of category B, L. caerulea (the L. ca-erulea group), L. infrafrenata (the L. infraf-renata group), and L. meiriana (the L. mei-riana group); and one representative of cat-egory C, L. arfakiana (the L. arfakianagroup).
Nyctimystes: This genus was rediagnosedby Tyler and Davies (1979). Among the listof characters provided by them, the synapo-morphies of Nyctimystes seem to be the ver-tical pupil and the presence of palpebral ve-nation. Tyler and Davies (1979) suggested
7 A detailed analysis of Pelodryadinae is currently be-ing carried out by S. Donnellan. Taxon sampling here isprovided to optimize characters effectively to the baseof the Pelodryadinae and not to reevaluate the taxonomyof Litoria and its generic satellites.
that Nyctimystes was most closely related tosome species groups of Litoria from NewGuinea, implying that Nyctimystes is nestedwithin Litoria. Specifically, they referred tothe L. angiana, L. arfakiana, L. becki, L. dor-sivena, L. eucnemis, and L. infrafrenatagroups as the most likely to be related toNyctimystes, because they share with Nycti-mystes similarities in cranial structure (the L.infrafrenata and L. eucnemis groups) or thepresence of large unpigmented ova and lotictadpoles bearing large, ventral, suctorialmouths (the other groups). Nyctimystes cur-rently comprises 24 described species, 5 ofwhich (N. disruptus, N. oktediensis, N. trach-ydermis, N. tyleri, and N. papua) were in-cluded in the N. papua species group byZweifel (1983) and Richards and Johnston(1993). We could not locate tissues of anymembers of the N. papua group, so we can-not test its monophyly. Nevertheless, we in-clude the available species N. kubori, N. na-rinosus, and N. pulcher.
Cyclorana: This genus was thought to berelated to the Australian leptodactylids (nowMyobatrachidae) by Parker (1940), and wasconsidered as such by Lynch (1971). Tyler(1972) first proposed its relationship to Aus-tralian hylids on the basis of the presence ofa differentiated apical element of the m. in-termandibularis. Subsequently, Tyler (1978)transferred Cyclorana to Hylidae. Tyler(1979), King et al. (1979), and Tyler et al.(1981) considered it to be related to the Li-toria aurea group, a result that was coinci-dent with the analyses of albumin immuno-logical distances generated by microcomple-ment fixation (Hutchinson and Maxson,1987). Burton (1996) suggested that havingthe m. palmaris longus divided into two seg-ments (as opposed to three) is a synapomor-phy supporting the monophyly of Cyclorana,L. dahlii, and L. alboguttata (two species ofthe L. aurea group). Based on sperm mor-phology, Meyer et al. (1997) transferred L.alboguttata to Cyclorana. The only morpho-logical synapomorphy suggested for Cyclor-ana is the anterior ossification of the sphe-nethmoid to incorporate a portion of the tec-tum nasi (Tyler and Davies, 1993).
The 13 species of Cyclorana have beenseparated into different groups based on kar-yotypes, sperm morphology, and immuno-
2005 19FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
logical distances (King et al., 1979; Maxsonet al., 1982; Maxson et al., 1985; Meyer etal., 1997). These are the C. brevipes, C. aus-tralis, and C. platicephala ‘‘lineages’’. In thepresent study, we include only C. australis.
Phyllomedusinae
Cruz (1990) and Duellman (2001) provid-ed taxonomic histories of this group, mostlyat the generic level. The monophyly of Phyl-lomedusinae has not been controversial; sev-eral character states have been advanced tosupport it. Some of the muscular characterstates include the supplementary posterolat-eral elements of the m. intermandibularis(Tyler, 1971); tendo superficialis pro digiti II(pes) arising from a deep, triangular muscle,which originates on the distal tarsal 2–3; ten-do superficialis pro digiti III arising entirelyfrom the aponeurosis plantaris; and m. exten-sor brevis superficialis digiti IV with a singleslip (Burton, 2004). There are also severallarval character states that support the mono-phyly of this group; for example, the arcussubocularis of larval chondrocranium withdistinct lateral processes (Fabrezi and Lavil-la; 1992, Haas, 2003); ultralow suspensorium(Haas, 2003); secondary fenestrae parietales(Haas, 2003); and absence of a passage be-tween ceratohyal and ceratobranchial I(Haas, 2003).
The subfamily is comprised of six nominalgenera: Agalychnis (8 species), Hylomantis(2 species), Pachymedusa (1 species), Phas-mahyla (4 species), Phrynomedusa (5 spe-cies), and Phyllomedusa (32 species). Cruz(1990) discussed the taxonomic distributionof several character states shared by subsetsof these genera. A cladistic analysis testingthe monophyly of each of these and their in-terrelationships remains to be completed.
Agalychnis: Duellman (2001) presented aphylogenetic analysis of Agalychnis and Pa-chymedusa, using a vector of character statespresent in the Phyllomedusa buckleyi groupas an outgroup (data taken from Cannatella,1980). His analysis suggested no synapo-morphies for Agalychnis. In our analysis weinclude all species available to us: A. calcar-ifer, A. callidryas, A. litodryas, and A. sal-tator (species not included are A. annae, A.craspedopus, A. moreletii, and A. spurrelli).
Hylomantis: This genus was resurrected byCruz (1990) for the species formerly placedin the Phyllomedusa aspera group (Cruz,‘‘1988’’ [1989]). From the extensive diag-nosis presented by Cruz (1990), the only ap-parent synapomorphy of Hylomantis seemsto be the lanceolate discs of fingers and toes.Cruz (1990: 725), however, considered likelythat Hylomantis was paraphyletic with re-spect to Phasmahyla. Hylomantis has twodescribed species, H. aspera and H. granu-losa. Only H. granulosa was available forthis study.
Pachymedusa: This monotypic genus wasrecognized by Duellman (1968a) to reflecthis view that a remnant of the ancestral stockgave rise to the other Phyllomedusinae.However, Duellman (2001) found no evi-dence supporting the monophyly of Agaly-chnis independent of Pachymedusa. The sin-gle species Pachymedusa dacnicolor is in-cluded in our analysis.
Phasmahyla: This genus was erected byCruz (1990) for the species formerly con-tained in the Phyllomedusa guttata group(Bokermann and Sazima, 1978; Cruz, 1982).Cruz (1990) provided an extensive definitionof the genus based on adult and larval mor-phology. Probable synapomorphies of Phas-mahyla are the lack of a vocal sac in adultmales and larval modifications presumablyassociated with surface film feeding, such asthe anterodorsal position of the oral disc, re-duction in number and size of labial toothrows, distribution and shape of submarginalpapillae, and the upper jaw sheath with a me-dial projection (see Cruz, 1982, 1990). Thegenus is composed of four species, P. coch-ranae, P. exilis, P. guttata, and P. jandaia.We include Phasmahyla cochranae and P.guttata in our study.
Phrynomedusa: This genus was resurrect-ed by Cruz (1990) for the species formerlyplaced in the Phyllomedusa fimbriata group(Izecksohn and Cruz, 1976; Cruz, 1982).From the extensive definition provided byCruz (1990), possible synapomorphies ofPhrynomedusa appear to be the presence ofa bicolored iris and the complete marginalpapillae in the oral disc of the larva. Phry-nomedusa contains five described species;unfortunately, we could not secure any rep-
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resentatives of this taxon for the presentstudy.
Phyllomedusa: No synapomorphies areknown to support the monophyly of the 32species of Phyllomedusa. This genus in-cludes simply those species that are not in-cluded in the other five genera of Phyllo-medusinae. There are currently five speciesgroups recognized within Phyllomedusa: theP. buckleyi group (Cannatella, 1980), P. bur-meisteri group (Lutz, 1950; Pombal andHaddad, 1992), P. hypochondrialis group(Bokermann, 1965a), P. perinesos group(Cannatella, 1982), and P. tarsius group (Dela Riva, 1999). The monophyly of thesegroups had not been tested, and relationshipsamong them remain unstudied. Furthermore,several species (e.g., P. bicolor, P. palliata,P. tomopterna, and P. vaillanti) have notbeen assigned to any species group. Someauthors (Funkhouser, 1957; Duellman,1968a, 1969; Cannatella, 1980; Jungfer andWeygoldt, 1994) have suggested that thePhyllomedusa buckleyi group deserves ge-neric recognition.
Duellman et al. (1988b) posited the exis-tence of a clade composed of most speciesof Phyllomedusa (excluding the P. buckleyigroup and the P. guttata group, now Phas-mahyla, but see below), implicitly includingthe species now placed in Hylomantis. Ap-parent synapomorphies of this clade are thewell-developed parotoid glands; the presenceof the slip of the m. depressor mandibulaethat originates from the dorsal fascia at thelevel of the m. dorsalis scapulae; first toe lon-ger than second; and the eggs wrapped inleaves. Duellman et al. (1988b) explicitly ex-cluded the species then included in the P.guttata group (now Phasmahyla) from thisapparent clade. However, Lutz (1954), Bok-ermann and Sazima (1978), Weygoldt(1984), and Haddad (personal obs.) reportedP. guttata, P. jandaia, P. exilis, and P. coch-ranae, respectively, to oviposit in foldedleaves, and Cruz (1990) reported in Hylo-mantis the presence of the slip of the m. de-pressor mandibulae that originates from thedorsal fascia at the level of the m. dorsalisscapulae. In this study we include represen-tatives of the Phyllomedusa buckleyi group(P. lemur), P. burmeisteri group (P. tetraplo-idea), P. hypochondrialis group (P. hypo-
chondrialis), and P. tarsius group (P. tar-sius). We include also P. bicolor, P. palliata,P. tomopterna, and P. vaillanti, four speciesunassigned to groups.
Hylinae
This taxon is the primary focus of thisstudy. Hyline monophyly is supported bytwo synapomorphies: the tendo superficialisdigiti V with an additional tendon that arisesventrally from m. palmaris longus (da Silva,1998, as cited by Duellman, 2001), and kar-yotype with 24 (or more) chromosomes(Duellman, 2001). The published molecularevidence is ambiguous regarding the issue.8
No comprehensive study of hyline system-atics has been published, although the resultsof one unpublished dissertation (da Silva,1998) have been widely circulated (e.g.,Duellman, 2001). Although the problems ofhylid systematics have been recognized forsome time, almost all work has been done atthe level of satellite genera (e.g., Duellma-nohyla, Plectrohyla, Ptychohyla, Scinax) orspecies groups of Hyla.
For the purpose of our analysis we includ-ed representatives of all 27 genera of Hyli-nae. Within the genus Hyla, we included ex-emplars of 39 of the 41 species groups thathad been recognized (we lack exemplars forthe H. claresignata and H. garagoensisgroups). We are not recognizing monotypicspecies group because (1) they do not rep-resent testable hypothesis, and (2) they are
8 In the analysis of Darst and Cannatella (2004), Hy-linae (including pseudids) is monophyletic only in theirmaximum likelihood analysis, not in their parsimonyanalysis. If, unlike Darst and Cannatella (2004), inser-tion/deletion events are considered as informative vari-ations, their results still show a paraphyletic Hylinae,having Leptodactylus pentadactylus 1 Lithodytes linea-tus nested within Hylinae. In the same analysis Hemi-phractinae is monophyletic. We do not think that thesedifferences in results reflect relative merits of the differ-ent approaches but instead represent problems in the tax-on sampling of the analysis of Darst and Cannatella(2003) (which was not designed to test the monophylyof Hylinae). Salducci et al. (2002) presented a molecularphylogenetic analysis using a fragment of 16S of a sam-pling restricted to sequences available in GenBank andHylidae of French Guyana. Rana palmipes and Hyali-nobatrachium taylori were the only outgroups. Consid-ering the restricted taxon sampling and the minimalnumber of outgroups, their results are difficult to inter-pret or to compare with other studies.
2005 21FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
not a rank in the Linnean taxonomy andtherefore are not required for consistency andstability purposes.
Hyla is the most species-rich genus of hy-lid frogs. It is currently placed in 41 speciesgroups, plus other species that had not beenassociated with any group. In turn, majorclades composed of several of these speciesgroups have been suggested. Because nostudy has ever suggested Hyla to be mono-phyletic and several have suggested that it isparaphyletic with respect to other hyline gen-era (Duellman and Campbell, 1992; Cocroft,1994; Faivovich, 2002; Haas, 2003; Darstand Cannatella, 2004; Faivovich et al.,2004), we refer further discussion to theheadings for the various genera and speciesgroups. Comments about apparent majorclades and proposed relationships amongspecies groups are mostly reserved for thediscussion section of this paper.
Gladiator Frogs
Kluge (1979: 1) referred to the frogs thenplaced in the Hyla boans group as ‘‘gladiatorfrogs’’ ‘‘in view of their extremely pugna-cious behavior and the well developed pre-pollical spines that they use when fighting.’’Following Duellman (1970, 1977), Kluge(1979) included in the group H. boans, H.circumdata, H. crepitans, H. faber, H. par-dalis, and H. pugnax. Nevertheless, a pre-pollical spine has been reported for severalspecies groups. Furthermore, territorial fight-ing has been reported or suspected to occurin several of these species.9 Because of this,and due to the lack of a better term, we preferto use the term ‘‘Gladiator Frogs’’ to refercollectively to all the mostly South Americanspecies having a prepollical spine, as was
9 It remains to be studied if these territorial behaviorsare homologous. Species of this putative clade, otherthan some members of the H. boans group, where com-bat was observed or are suspected to occur, are Hylacircumdata (Haddad, personal obs.), H. cordobae (Fai-vovich, personal obs.), H. goiana (Menin et al., 2004),H. joaquini (Garcia et al., 2003), H. marginata (Garciaet al., 2001b), H. marianitae (Duellman et al., 1997), H.prasina (Haddad, personal obs.), H. melanopleura (Lehrand May, 2004), H. pulchella (Gallardo, 1970; Langone,‘‘1994’’ [1995]), H. punctata (Sehinkman and Faivov-ich, personal obs.), H. raniceps (Guimaraes et al., 2001),H. riojana (Blotto and Baldo, personal commun.), andH. semilineata (Haddad, personal obs.).
done by Faivovich et al. (2004), instead ofrestricting its use to the H. boans group. Thespecies groups that are currently referred tothe Gladiator Frog clade are the H. albom-arginata, H. albopunctata, H. boans, H. cir-cumdata, H. claresignata, H. geographica,H. granosa, H. martinsi, H. pseudopseudis,H. pulchella, and H. punctata10 groups (Bok-ermann, 1972, Hoogmoed, 1979, Duellmanet al., 1997, Eterovick and Brandao, 2001,Duellman, 2001, Faivovich et al., 2004).
Hyla albomarginata Group: The H. al-bomarginata group was first recognized for-mally by Cochran (1955) for a group of spe-cies (H. albomarginata, H. albosignata, H.albofrenata, H. musica) that Lutz (‘‘1948’’[1949]) referred to as the green species ofHyla of southeastern Brazil. Cochran (1955)included H. prasina (latter included in the H.pulchella group by Lutz, 1973) in this group.Duellman (1970) presented a definition ofthe group and included, besides the speciesconsidered by Cochran (1955), H. rufitela, H.pellucens, H. albopunctulata (now consid-ered a member of the H. bogotensis group;see below), and H. albolineata (now consid-ered a species of Gastrotheca; see Sachsse etal., 1999).
Cruz and Peixoto (‘‘1985’’ [1987]) dividedthe Hyla albomarginata group into three‘‘complexes’’: the H. albomarginata com-plex, containing H. albomarginata and H.rufitela; the H. albofrenata complex, con-taining H. albofrenata, H. arildae, H. ehr-hardti (as H. arianae; see Faivovich et al.,2002), H. musica, and H. weygoldti; and theH. albosignata complex, containing H. al-bosignata, H. callipygia, H. cavicola, H. flu-minea, and H. leucopygia. Hyla pellucensshould also be included in the albomarginatacomplex, because this species was includedby Duellman (1970) but was overlooked byCruz and Peixoto (‘‘1985’’ [1987]); verylikely the same applies for H. rubracyla, aspecies that was resurrected from the syn-onymy of H. pellucens by Duellman (1974)and included in the H. albomarginata group
10 Faivovich et al. (2004) mentioned that Duellman etal. (1997) did not include the Hyla punctata group with-in a putative clade of gladiator frogs, but overlooked thefact that Duellman (2001: 776) stated that ‘‘members ofthe . . . H. punctata group might be included’’ in thisclade.
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by Duellman in Frost (1985). The only pos-sible synapomorphies that were proposed forthese complexes were the red coloration ofthe webbing in the two species of the H. al-bomarginata complex studied by the authors,and presence of cloacal ornamentation in theH. albosignata complex (several instances ofhomoplasy within hylids). Haddad and Sa-waya (2000) and Hartmann et al. (2004) fur-ther suggested that the H. albofrenata and H.albosignata complexes share a reproductivemode where the male constructs a subterra-nean nest in the muddy side of streams andpools that is completely concealed afterspawning, a nest where larvae spend earlystages of development; subsequent to flood-ing, the exotrophic larvae live in ponds orstreams.
Cruz et al. (2003) added Hyla ibirapitangaand H. sibilata to the H. albosignata com-plex. Note that species included in both theH. albofrenata and H. albosignata complex-es do not posses a prepollical spine, as dospecies in the H. albomarginata complex. Onrecent occasions, some authors (Haddad andSawaya, 2000; Garcia et al., 2001a) referreddirectly to the H. albofrenata and H. albo-signata groups without further comment. Weinclude in the present analysis representa-tives of the three complexes: H. albosignata,H. callipygia, H. cavicola, and H. leucopygiaas representatives of the H. albosignata com-plex; H. arildae, H. weygoldti, and a Hylasp. 1, a new species similar to H. ehrhardti,as representatives of the H. albofrenata com-plex; and H. albomarginata, H. pellucens,and H. rufitela as exemplars of the H. albo-marginata complex.
Hyla albopunctata Group: Cochran (1955)recognized the H. albopunctata group on thebasis of the ‘‘more streamlined body shape,by lacking an outer metatarsal tubercle, andby having the fingers webbed only at thebase . . . ’’. She included in the group H. al-bopunctata, H. raniceps, and several speciesfrom southeastern Brazil that are now in theH. claresignata and H. pulchella groups.Cochran and Goin (1970) recognized a H.lanciformis group (on the basis of large size,a white margin on the upper lip, pointedheads, and reduced webbing between the fin-gers) in which they included H. lanciformis,H. multifasciata, and H. boans (name applied
incorrectly to H. albopunctata; see Duell-man, 1971a). Duellman (1971a) implicitlyunited these two groups and considered thelarger H. albopunctata group to be composedof H. albopunctata, H. lanciformis, H. mul-tifasciata, and H. raniceps. De Sa (1995,1996) stated that there was no evidence sup-porting the monophyly of the group. Cara-maschi and Niemeyer (2003) added H. leu-cocheila to the group and suggested that itwas monophyletic, but they presented no ev-idence to this effect. We include all speciesexcept H. leucocheila in our analysis.
Hyla boans Group: The constitution of theH. boans group as well as the definition ofthe group present a rather confusing history.Affinities between species of what is cur-rently called the H. boans group were firstrecognized by Cochran (1955), who includedin what she called the H. faber group thespecies H. crepitans, H. faber, H. langsdorffii(now a species of Osteocephalus, see Duell-man, 1974), and H. pardalis. Some of thediagnostic characters of this group were largesize and the presence of what she called aprominent spurlike prepollex in males. Coch-ran and Goin (1970) included H. faber, H.pardalis, H. rosenbergi, and H. maxima(now a junior synonym of H. boans; seeDuellman, 1971a) in the H. maxima group,and they excluded H. crepitans, placing it inits own group. Duellman (1970) presented aformal definition of the H. boans group, inwhich he included H. boans, H. circumdata(now in the H. circumdata group), H. cre-pitans, H. faber, H. langsdorffii, H. pardalis,and H. rosenbergi. Lutz (1973) included thespecies in three different groups,11 in one ofwhich she also included several species nowincluded in the H. circumdata, H. pseudop-seudis, and H. martinsi groups (the ‘‘specieswith long, sharp pollex rudiment’’). Kluge(1979) resurrected H. pugnax from the syn-onymy of H. crepitans, including it also inthe H. boans group. Martins and Haddad(1988) included in the group H. lundii (usingthe name H. biobeba, a junior synonym, seeCaramaschi and Napoli, 2004), based on ob-
11 The ‘‘species with long, sharp pollex rudiment’’, the‘‘species with undulated glandular outline’’, and the‘‘species with pattern on the transparent part of the lowereyelid’’.
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servations of nest construction done by Jim(1980). Implicitly, they also removed H. cir-cumdata from the group. Hoogmoed (1990)resurrected H. wavrini from the synonymy ofH. boans and retained it in the group. Duell-man (2001) omitted H. lundii from the groupwithout comment. Caramaschi and Ro-drigues (2003) added H. exastis, suggestingthat it was related to H. lundii and H. par-dalis on the basis of its lichenous color pat-tern and the rugose skin texture. Caramaschiand Napoli (2004) presented a formal defi-nition of the group. In summary, and follow-ing Caramaschi and Napoli (2004), we re-gard the H. boans group to be composed ofH. boans, H. crepitans, H. exastis, H. faber,H. lundii, H. pardalis, H. pugnax, H. rosen-bergi, and H. wavrini. The only synapomor-phy that has ever been proposed for thisgroup is the nest-building behavior of males,which has been observed in most species(see Martins and Moreira, 1991 for a re-view). Early reports of H. crepitans indicatedthat males do not construct nests; this wasshown to be facultative by Caldwell (1992).This behavior is still unknown in H. pugnax.From this group we include in our analysisH. boans, H. crepitans, H. faber, H. lundii,and H. pardalis.
Hyla circumdata Group: This group wasfirst mentioned by Bokermann (1967a,1972), without providing any diagnosis.Heyer (1985) provided the first formal defi-nition, diagnosing the group by the combi-nation of a well-developed prepollex and theposterior face of the thigh having dark ver-tical stripes. The group was further discussedand expanded by Caramaschi and Feio(1990), Pombal and Haddad (1993), Napoli(2000), Caramaschi et al. (2001), and Napoliand Pimenta (2003). Three other speciesgroups, the H. claresignata, H. martinsi, andH. pseudopseudis groups, as well as H. al-varengai, historically had been satellites ofthe H. circumdata group, with these speciesbeing alternatively included or excludedfrom the group. These groups and H. alvar-engai are treated separately. With the rec-ognition of these three groups being separatefrom the H. circumdata group, it is unclearwhich synapomorphies support its monophy-ly as currently defined.
Duellman et al. (1997) suggested that all
species of the Hyla circumdata group shouldbe transferred to the H. pulchella group. Fai-vovich et al. (2004) demonstrated by usingDNA sequences from four mitochondrialgenes that the two groups are not closely re-lated. In the analysis of Faivovich et al.(2004), the three exemplars of the H. circum-data group then available (H. astartea, H.circumdata, and H. hylax) formed a mono-phyletic group that is the sister taxon of theremaining Gladiator Frogs they included intheir analysis. Napoli and Pimenta (2003),Napoli and Caramaschi (2004), and Napoli(2005) recognized 15 species in the group:H. ahenea, H. astartea, H. caramaschii, H.carvalhoi, H. circumdata, H. feioi, H. gou-veai, H. hylax, H. ibitipoca, H. izecksohni,H. lucianae, H. luctuosa, H. nanuzae, H. rav-ida, and H. sazimai. We include five speciesin our analysis: H. astartea, H. circumdata,H. hylax, as well as Hyla sp. 3 and Hyla sp.4, two undescribed species from littoral areasof northern Sao Paulo (state) and southrn Riode Janeiro (state), Brazil, respectively.
Hyla claresignata Group: A close relation-ship between H. clepsydra and H. claresig-nata was suggested by Bokermann (1972),who noticed striking similarities in larval andadult morphology. Bokermann (1972) sug-gested a possible relationship of these specieswith the H. circumdata group; followinghim, Jim and Caramaschi (1979) included H.clepsydra and H. claresignata in the H. cir-cumdata group. However, subsequent work-ers (Caramaschi and Feio, 1990; Pombal andHaddad, 1993) who referred to the H. cir-cumdata group did not include them in thegroup. The H. claresignata group was rec-ognized in the restricted form by Duellmanet al. (1997). Possible synapomorphies of theH. claresignata group are character states as-sociated with the torrent-dwelling larvae ofthese species: oral disc completely surround-ed by marginal papillae, and 7/12–8/13 labialtooth rows. We were not able to secure sam-ples of either of the two species of thisgroup.
Hyla geographica Group: This group wasdelimited by Cochran (1955: 180) as beingcharacterized by its ‘‘extremely attenuatelimbs’’. Cochran and Goin (1970) character-ized the species of this group as ‘‘moderate-sized tree frogs with elongate dermal ap-
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pendages on the heels and reduced webbingbetween the fingers.’’ Duellman (1973a) pre-sented an extensive characterization of thegroup (including vomers large with angulardentigerous processes, each bearing as manyas 20 teeth; nuptial excrescences present inbreeding males; projecting prepollices absentin both sexes; calcars present; palpebralmembrane clear or reticulated). However, itis unclear from his account if any of thesecharacter states could be considered syna-pomorphic of the group.
Duellman (1973a) included in this groupHyla calcarata, H. fasciata, and H. geogra-phica. Later, Pyburn (1977, 1984) added H.microderma and H. hutchinsi. Goin andWoodley (1969) considered H. kanaima re-lated to the H. geographica group, and Py-burn (1984) included H. kanaima in thegroup. Lutz (1963, 1973) and Bokermann(1966a) stressed similarities between H. se-cedens and H. semilineata (as H. geographi-ca), but Caramaschi et al. (2004a) suggestedthat actually this species is closer to H. bis-choffi (of the H. pulchella group). Duellman(in Frost, 1985) and Duellman and Hoog-moed (1992), respectively, included H. pic-turata and H. roraima in the H. geographicagroup. D’Heursel and de Sa (1999) arguedfor the recognition of H. semilineata, a spe-cies that had previously been placed in thesynonymy of H. geographica. Lescure andMarty (2000) included H. dentei, a speciesthat Bokermann (1967b) considered to havecharacter states of both H. raniceps (H. al-bopunctata group) and the H. geographicagroup. Caramaschi et al. (2004a) added H.pombali to the group. In summary, the H.geographica group is currently composed of11 species: H. calcarata, H. dentei, H. fas-ciata, H. geographica, H. hutchinsi, H. kan-aima, H. microderma, H. picturata, H. pom-bali, H. roraima, and H. semilineata. We in-clude H. fasciata, H. calcarata, H. kanaima,H. microderma, H. picturata, H. roraima,and H. semilineata in our study.
Hyla granosa Group: This group was firstdefined by Cochran and Goin (1970) asgreen frogs that share the vomerine teeth be-ing in rather heavy, arched series, and withmales having a ‘‘protruding spine in the pre-pollex’’. These authors included H. granosa,H. rubracyla (now in the H. albomarginata
complex of the H. albomarginata group, seeabove), and H. guibei (now a junior synonymof H. pellucens; see Duellman, 1974). Pre-viously, Rivero (1964) stated that H. alemaniwas allied with H. granosa. Rivero (‘‘1971’’[1972]) considered H. sibleszi to be relatedto H. granosa. Hoogmoed (1979) mentioned,without any diagnosis, the H. granosa groupin the Guayanas, in which he included H.ornatissima. Mijares-Urrutia (1992a) consid-ered H. alemani, H. granosa, H. ornatissima,and H. sibleszi to be the members of thisgroup, and he provided a characterizationbased on larval features. We are not awareof any synapomorphies for this group. In thisanalysis we include H. granosa and H. sib-leszi.
Hyla martinsi Group: This group was rec-ognized by Bokermann (1965b) for two spe-cies, H. langei and H. martinsi, characterizedby the presence of an extensive hooklike hu-meral crest and by a bifid prepollex. Boker-mann (1964a) noticed ‘‘superficial similari-ties’’ of H. martinsi with H. circumdata. Car-amaschi and Feio (1990) and Cardoso (1983)included H. martinsi in the H. circumdatagroup for having the diagnostic charactersestablished by Heyer (1985). However, basedon the presence of bifid prepollex and a hu-meral spine, Caramaschi et al. (2001) pre-ferred to keep it as a separate species group.As a representative of this group we includeH. martinsi in the analysis.
Hyla pseudopseudis Group: This groupwas recognized by Pombal and Caramaschi(1995) as closely related to the H. circum-data group, from which it was differentiatedmostly by its color pattern. Eterovick andBrandao (2001) further differentiated bothgroups based on the presence of short, lateralirregular tooth rows and for having addition-al posterior tooth rows (6–8 rows) in the oraldiscs of the larvae of the H. pseudopseudisgroup (a maximum of 5 posterior rows in theH. circumdata group). Caramaschi et al.(2001) transferred H. ibitiguara from the H.circumdata group to the H. pseudopseudisgroup on the basis of its similar externalmorphology, color pattern, and habits. Thegroup currently comprises three species, H.ibitiguara, H. pseudopseudis and H. saxico-la, plus Hyla sp. 6 (aff. H. pseudopseudis), anew species from Bahia, Brazil, that is being
2005 25FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
described by Lugli and Haddad (in prep.).Only tissues of this new species were avail-able for this study.
Hyla pulchella Group: The history of thisgroup was recently reviewed by Faivovich etal. (2004). These authors also presented aphylogenetic analysis based on mitochondri-al DNA sequences of four genes, and includ-ed 10 of the then 14 species included in thegroup, plus exemplars of the former H. po-lytaenia group and several outgroups. Theirresults indicate that the H. polytaenia group,as defined by Cruz and Caramaschi (1998),is nested within the H. pulchella group. Con-sequently, species included in the H. poly-taenia group were transferred to the H. pul-chella group, where they are recognized asthe polytaenia clade. Considering this actionand the species status given to H. cordobaeand H. riojana, Faivovich et al. (2004) raisedthe number of species included in the H. pul-chella group to 25. Carnaval and Peixoto(2004) recently added H. freicanecae to thegroup. Caramaschi et al. (2004a) suggestedthat H. secedens is related to H. bischoffi,therefore adding implicitly the species to theH. pulchella group. Caramaschi and Cruz(2004) added H. beckeri and H. latistriata tothe H. polytaenia clade, adding two morespecies to the H. pulchella group. Faivovichet al. (2004) had doubts regarding the rec-ognition of H. callipleura. Duellman et al.(1997) included this name as a junior syno-nym of H. balzani, but Kohler (2000) res-urrected it using the combination H. cf. cal-lipleura for some populations in Bolivia. Wetentatively recognize H. callipleura as valid,but stress the necessity of further studies toclarify its status.
While the monophyly of this redefinedHyla pulchella group is supported by molec-ular evidence, no morphological synapomor-phies have been proposed so far (see alsocomments for the H. circumdata and H. lar-inopygion groups). In summary, there are 30species included in this group: H. albonigra;H. andina; H. balzani; H. beckeri; H. bis-choffi; H. buriti; H. caingua; H. callipleura;H. cipoensis; H. cordobae; H. cymbalum; H.ericae; H. freicanecae; H. goiana; H. guen-theri; H. joaquini; H. latistriata; H. leptoli-neata; H. marginata; H. marianitae; H. me-lanopleura; H. palaestes; H. phaeopleura; H.
polytaenia; H. prasina; H. pulchella; H. rio-jana; H. secedens; H. semiguttata; and H.stenocephala. In this analysis we include thesame species that were available to Faivovichet al. (2004) (H. andina; H. balzani; H. bis-choffi; H. caingua; H. cordobae; H. ericae;H. guentheri; H. joaquini; H. leptolineata; H.marginata; H. marianitae; H. prasina; H.pulchella; H. riojana; H. semiguttata, and anundescribed species), plus H. polytaenia. Thespecies that Faivovich et al. (2004) calledHyla sp. 2 corresponds to what Caramaschiand Cruz (2004) recently described as H. la-tistriata, and so is included under that name.
Hyla punctata Group: This group was firstrecognized by Cochran and Goin (1970),who included H. punctata, H. rhodoporus,and H. rubeola (these last two were subse-quently considered to be synonyms of H.punctata by Duellman, 1974). They charac-terized the group as ‘‘moderately small greentree frogs with small vomerine tooth patches,reduced webbing between the fingers, with-out spines on the pollex, and without ulnaror tarsal ridges.’’ Hoogmoed (1979) men-tioned this group without defining it. No syn-apomorphies have been proposed for thisgroup. Besides H. punctata, two other spe-cies could be included on this poorly definedgroup: H. hobbsi, a species resurrected fromthe synonymy of H. punctata by Pyburn(1978), and H. atlantica, a name recently ap-plied by Caramaschi and Velosa (1996) forthe populations on eastern Brazil previouslyconsidered as H. punctata. In this analysiswe include H. punctata.
Species of Probable Gladiator Frogs Un-assigned to Species Group: Hyla alvarengai:This bizarre species was said by Bokermann(1964a) to share some character states withH. martinsi and H. saxicola (now placed inthe H. martinsi and H. pseudopseudisgroups, respectively), such as the notable de-velopment of the prepollex and the shape ofthe sacral diapophyses. Lutz (1973) includedit in the group of the ‘‘species with long,sharp pollex rudiment’’, together with H.crepitans, H. faber, and species now includ-ed in the H. circumdata and H. martinsigroups. She referred to it as Hyla (Plectro-hyla?) alvarengai and stated that it was ‘‘de-void of affinities with the very large speciesof Hyla’’, suggesting instead a possible re-
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lationship with Plectrohyla. A similar opin-ion was presented by Sazima and Bokermann(1977), who noticed ‘‘superficial similari-ties’’ with Plectrohyla, but they argued thatthey differed in larval morphology, spawn,and vocalizations. Duellman et al. (1997) in-cluded H. alvarengai in the H. circumdatagroup, presumably because it shares the di-agnostic characters of the group. Eterovickand Brandao (2001) and Caramaschi et al.(2001) did not consider it as a member of theH. circumdata group. Unfortunately, wecould not secure this species for our analysis,although we include a new species similar toH. alvarengai that is in the process of beingdescribed by Lugli and Haddad (in prep.).
Hyla fuentei: This species was describedby Goin and Goin (1968) based on a singleadult female from Surinam. Hoogmoed(1979) mentioned the existence of two ad-ditional specimens collected close to the typelocality. Since then, no author has referred tothis species. From the original description,there are few characters that allow the asso-ciation of this species with any other groupof Hyla. The angulate dentigerous process ofthe vomer suggests that this species could beassociated with certain Gladiator Frogs, assome species currently placed in the H. al-bopunctata, H. boans, and H. geographicagroups have this character state. A study ofthe holotype and discovery of male speci-mens should clarify the matter.
Hyla heilprini: This West Indian hylid wasassociated with the H. albomarginata groupby Duellman (1970) based on the presenceof a ‘‘green’’ peritoneum (actually it is whiteparietal peritoneum, like in species of the H.albomarginata, H. bogotensis, H. granosa,and H. punctata groups; Lynch and Ruiz-Carranza [1991: 4]; Faivovich, personal obs.)and external pigmentation. This was fol-lowed by Trueb and Tyler (1974), who no-ticed that its morphology was ‘‘highly rem-iniscent’’ of those from that species group.While it seems clear that H. heilprini is aGladiator Frog, we are not aware of any syn-apomorphy relating it to the H. albomargin-ata group. This species was included in theanalysis.
Three species of Hyla from the Venezue-lan Tepuis: There are three species of Hylafrom the Venezuelan Tepuis that have not
been posited to be related to any other spe-cies or group of species: H. benitezi, H. le-mai, and H. rhythmicus. The presence of aprepollical spine (Rivero, ‘‘1971’’ [1972];Donnelly and Myers, 1991; Senaris and Ay-arzaguena, 2002) associates these specieswith Gladiator Frogs. Hyla benitezi and H.lemai were included in the analysis.
Hyla varelae: Carrizo (1992) describedthis species based on a single adult male. Itwas suggested to be related to H. raniceps inthe description. No additional specimenshave been collected since the description,and it was not included in this analysis.
Andean Stream-Breeding Hyla
Duellman et al. (1997) presented a phy-logenetic analysis restricted to wholly or par-tially Andean species groups of Hyla. Ontheir most parsimonious tree, the H. armata,H. bogotensis, and H. larinopygion groupstogether form a monophyletic group sup-ported by three transformations in larvalmorphology: the enlarged, ventrally orientedoral disc; the complete marginal papillae;and labial tooth rows formula 4/6 or more.
Hyla armata Group: The H. armata groupwas first recognized by Duellman et al.(1997) for its single species, H. armata. Koh-ler (2000) and De la Riva et al. (2000) sub-sequently reported that H. charazani was asecond member of the H. armata group.Duellman et al. (1997) described four syna-pomorphies for the H. armata group: thepresence in males of keratin-covered bonyspines on the proximal ventral surface of thehumerus, on the expanded distal element ofthe prepollex, and on the first metacarpal;tadpole tail long with low fins and bluntlyrounded tip; forearms hypertrophied; and thepresence of a ‘‘shelf’’ on the larval upper jawsheath. We include both species in our anal-ysis.
Hyla bogotensis Group: This group wasreviewed by Duellman (1970, 1972b, 1989)and Duellman et al. (1997). The only syna-pomorphy that has been suggested for thisgroup is the presence in males of a mentalgland.12 Hyla albopunctulata was redescribed
12 See Duellman (2001) and La Marca (1985) for com-ments on taxonomic distribution and morphological var-iation of this gland, and Romero de Perez and Ruiz-Carranza (1996) for its histological structure.
2005 27FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
by Duellman and Mendelson (1995), who re-jected a possible relationship with the H. bo-gotensis group, as suggested by Goin in Riv-ero (1969) and Duellman (in Frost, 1985),and they simply stated that its relationshipswere unclear. Faivovich et al. (in prep.) stud-ied two male syntypes (BMNH 1880.12.5159 and 1880.12.5160), which posses a no-ticeable mental gland. For this reason, we as-sociate this species with the H. bogotensisgroup. The Hyla bogotensis group is thencomposed of 15 species: H. albopunctulata,H. alytolylax, H. bogotensis, H. callipeza, H.colymba, H. denticulenta, H. jahni, H. las-cinia, H. lynchi, H. palmeri, H. phyllognata,H. piceigularis, H. platydactyla, H. simmon-si, and H. torrenticola. In this analysis wewere able to include only H. colymba and H.palmeri.
Hyla larinopygion Group: Duellman andHillis (1990) and Duellman and Coloma(1993) reviewed this group, and Duellmanand Hillis (1990) and Duellman et al. (1997)provided a formal definition, although it isunclear whether any of the morphologicalcharacter states employed in these character-izations is synapomorphic for the group.Duellman and Hillis (1990) performed a phy-logenetic analysis using isozymes of fivespecies of the group. In the phylogeneticanalysis of Duellman et al. (1997), the au-thors did not identify any synapomorphy forthe H. larinopygion group; it merely lacksthe apparent synapomorphies of the H. ar-mata and H. bogotensis groups. Because ofproblems with the limits of the H. larinopy-gion group, Kizirian et al. (2003) were un-certain about the placement of H. tapichal-aca, a species that they considered most sim-ilar to the H. larinopygion, H. armata, andH. pulchella groups.13 Faivovich et al. (2004)showed that H. tapichalaca and H. armata(the only exemplars of Andean stream-breed-ing Hyla they included) were sister taxa, andonly very distantly related to the H. pulchellagroup. The H. larinopygion group currentlycomprises nine species: H. caucana, H. lar-inopygion, H. lindae, H. pacha, H. pantos-
13 The only character state that led Kizirian et al.(2003) to consider Hyla tapichalaca similar to the H.pulchella group is the presence of an enlarged, pointed,recurved prepollex.
ticta, H. psarolaima, H. ptychodactyla, H.sarampiona, and H. staufferorum. In thisanalysis, we include H. pacha, H. pantostic-ta, and H. tapichalaca.
The 30-Chromosome Hyla
Only the species of the Hyla microcephalagroup were initially reported to have 30 chro-mosomes (Duellman and Cole, 1965; Duell-man, 1967). However, as species of othergroups were reported to have 30 chromo-somes (Duellman, 1970; Bogart, 1973), it be-came evident that this was a characteristic ofseveral species groups. Currently, the H. co-lumbiana, H. decipiens, H. garagoensis, H.labialis, H. leucophyllata, H. marmorata, H.microcephala, H. minima, H. minuta, H.parviceps, and H. rubicundula groups, plusseveral species unassigned to any group arebelieved to conform to a monophyletic groupsupported by this character state (Duellman,1970; Duellman and Trueb; 1983; Duellmanet al., 1997; Napoli and Caramaschi, 1998;Carvalho e Silva et al., 2003).
Hyla columbiana Group: This group wasfirst proposed by Duellman and Trueb (1983)for three species: H. carnifex, H. columbi-ana, and H. praestans. Kaplan (1991, 1999)found no evidence of monophyly for thegroup. Kaplan (1997) resurrected H. bogertifrom the synonymy of H. carnifex, adding afourth species to the group. Kaplan (1999)suggested that ‘‘the presence of two close,triangular lateral spaces between the cricoidand arytenoids at the posterior part of thelarynx’’ is a synapomorphy of the H. colum-biana group excluding H. praestans, whichhe considered to be closely related to the H.garagoensis group. The group therefore iscomposed of H. bogerti, H. carnifex, and H.columbiana. In the present analysis, we in-clude H. carnifex.
Hyla decipiens Group: While describingthe tadpoles of H. oliveirai and H. decipiens,Pugliese et al. (2000) noticed that they havemarginal papillae (unlike other known larvaeof the H. microcephala group), and theypointed out that they may not be members ofthe H. microcephala group as considered byBastos and Pombal (1996). Pugliese et al.(2000) noticed similarities in tadpole mor-phology with H. berthalutzae, with which
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these two species also share oviposition onleaves outside the water. Pugliese et al.(2000) also associated H. haddadi with thesethree species based on external similarity.
Carvalho e Silva et al. (2003) suggestedthe recognition of the Hyla decipiens groupfor these species. The group was defined bylarval features that include one row of mar-ginal papillae, an ovoid body in lateral view,eyes in the anterior third of the body, dorsalfin arising at the end of the body, tail withtransverse dark stripes on a light background,and pointed tip without a flagellum. It is un-clear which of these character states could beconsidered to be possible synapomorphies.Perhaps one apparent synapomorphy of thegroup, not mentioned by the authors, couldbe the oviposition on leaves outside the wa-ter. The group is composed of H. berthalut-zae, H. decipiens, H. haddadi, and H. oliv-eirai. In our analysis, we include H. ber-thalutzae.
Hyla garagoensis Group: This speciesgroup was first recognized by Kaplan andRuiz-Carranza (1997), who diagnosed it bythe presence of alternated pigmented and un-pigmented longitudinal stripes on the hind-limbs of larvae. The H. garagoensis group iscurrently composed of three species, H. gar-agoensis, H. padreluna, and H. virolinensis.Unfortunately, no species of this group wasavailable for our analysis.
Hyla labialis Group: This group was firstrecognized by Cochran and Goin (1970) forH. labialis (including what is now H. platy-dactyla, a species of the H. bogotensisgroup). They characterized the group by thevomerine teeth being in two rounded patchesand by the presence of a well-developed ax-illary membrane (referred to as a patagium)that is bright blue in life. Duellman (1989)presented a more extensive definition of thegroup, noting that the axillary membrane wasabsent, a point with which we agree. A sim-ilar definition was presented by Duellman etal. (1997). No synapomorphies were sug-gested for this species group. The group cur-rently comprises three species, H. labialis, H.meridensis, and H. pelidna. In this study weinclude H. labialis.
Hyla leucophyllata Group: This group wasdefined and later partly reviewed by Duell-man (1970, 1974). From Duellman’s (1970)
extensive definition, the only possible syna-pomorphy seems to be the presence of twoglandular patches in the pectoral region (thischaracter state, however, was ignored in ev-ery subsequent paper dealing with phyloge-netic relationships of 30-chromosome Hyla).In the phylogenetic analysis presented byDuellman and Trueb (1983), the only syna-pomorphy they proposed for the group wasviolin larval body shape. This character stateseems problematic in that it is present in lar-vae of some species associated with the H.microcephala and H. rubicundula groups(see Lavilla, 1990; Pugliese et al., 2001), andtherefore the level of inclusiveness of theclade that is supported by this transformationis unclear. From the perspective of adult mor-phology, we suggest that glandular patchesin the pectoral region is a synapomorphy ofthe H. leucophyllata group; we observed itclearly on specimens of both sexes in all spe-cies of the group. Another character state thatis either a likely synapomorphy of the H. leu-cophyllata group or of a more inclusive cladeis the oviposition on leaves hanging over wa-ter (this oviposition mode occurs also in oth-er 30-chromosome species; see comments inthe H. microcephala and H. parvicepsgroups).
The Hyla leucophyllata group is com-posed of seven species: H. bifurca, H. ebrac-cata, H. elegans, H. leucophyllata, H. trian-gulum, H. rossalleni, and H. sarayacuensis.In our analysis, we include H. ebraccata, H.sarayacuensis, and H. triangulum.
Hyla marmorata Group: This group wasrecognized by Cochran (1955) for H. mar-morata, H. microps, and H. giesleri based onthe presence of an axillary membrane, wartyskin around the margin of the lower lip, shortsnout, crenulated margin of limbs, shorthindlimbs, developed finger and toe web-bing, dorsal marbled pattern, and orange col-oration in thighs and webbings. Bokermann(1964b) further diagnosed the group by itspossession of a very large vocal sac. Severalof these character states are possible syna-pomorphies for the group (such as the wartyskin around the margin of the lower lip, thecrenulated margin of limbs, the dorsal mar-bled pattern). Duellman and Trueb (1983)suggested that this group could be diagnosed
2005 29FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
by the presence of a row of small marginalpapillae in the larval oral disc.
Bokermann (1964b), like Cochran (1955),included in the Hyla marmorata group otherspecies as well: H. parviceps, H. microps, H.schubarti, and H. moraviensis (now consid-ered a synonym of H. lancasteri; see Duell-man, 1966). Subsequent authors transferredthese species to other groups. This group isnow composed of eight species: H. acreana,H. dutrai, H. marmorata, H. melanargyrea,H. nahdereri, H. novaisi, H. senicula, and H.soaresi. In our analysis we include H. mar-morata and H. senicula.
Hyla microcephala Group: This group wasdefined by Duellman and Fouquette (1968)and Duellman (1970). Despite their extensivecharacterization, the only character statementioned by these authors that subsequentlyhas been considered a possible synapomor-phy of this group is the lack of marginal pa-pillae in the oral disc. Later, Duellman andTrueb (1983) added the depressed bodyshape of the larvae, another likely synapo-morphy.
While the study of Duellman and Fou-quette (1968) was focused on Middle Amer-ican species, Duellman (1970) referred anumber of South American species to theHyla microcephala group. Overall, he in-cluded in the group H. elongata (a juniorsynonym of H. rubicundula; see Napoli andCaramaschi, 1999), H. microcephala, H.minuta, H. nana, H. phlebodes, H. robert-mertensi, H. sartori, and H. werneri. Duell-man (1972a) also included H. mathiassoniand H. rhodopepla in the group, and he ex-cluded H. minuta. Cochran and Goin (1970)also had excluded H. minuta by placing it intheir H. minuta group. Duellman (1973b) in-cluded H. gryllata, and Langone and Basso(1987) added H. minuscula, H. sanborni, andH. walfordi.
Cruz and Dias (1991) placed Hyla bipunc-tata in the group on the basis of larval char-acters.14 Marquez et al. (1993) included H.leali in the H. microcephala group, withoutmentioning that it was included by Duellman
14 Duellman (2001) stated that the relationships ofHyla bipunctata were uncertain. Presumably he was notaware of the description of its tadpole by Cruz and Dias(1991).
(1982) in the H. minima group. Furthermore,Marquez et al. (1993) noticed that H. lealiand H. rhodopepla share vocalizations withshort, pulsatile notes with extremely lowdominant frequencies. Because of these sim-ilarities in vocalizations of H. leali with aspecies of the H. microcephala group, in theabsence of other evidence we prefer to keepH. leali in this group.
Bastos and Pombal (1996) suggested thatHyla branneri, H. decipiens, H. haddadi, andH. oliveirai were closely related species thatcould be tentatively associated with the H.microcephala group based on overall simi-larity of adult morphology, but this wasquestioned by Pugliese et al. (2000) and Car-valho e Silva et al. (2003), who excludedthese species from the group (see commentsfor the H. decipiens group). Pombal and Bas-tos (1998) also added H. berthalutzae, H.cruzi, and H. meridiana. Cruz et al. (2000)added H. pseudomeridiana. Kohler and Lot-ters (2001a) tentatively included H. joannae,based on its similarities in vocalization andadult morphology with H. leali (but see com-ments regarding H. leali above). Carvalho eSilva et al. (2003) added H. studerae and ex-cluded H. berthalutzae (see comments for theH. decipiens group).
Of the 20 species currently included in theHyla microcephala group, tadpoles are onlyknown for 9 species: H. bipunctata, H. mer-idiana, H. microcephala, H. nana, H. phle-bodes, H. pseudomeridiana, H. rhodopepla,H. sanborni, and H. studerae (Bokermann,1963, Duellman, 1970, 1972a, Lavilla, 1990,Cruz and Dias, 1991, Cruz et al., 2000, Pug-liese et al., 2000, Carvalho e Silva et al.,2003). All of these species have the two ap-parent synapomorphies of the H. microce-phala group (see comments for the H. rubi-cundula group).
The 20 species currently included in theHyla microcephala group are H. bipunctata,H. branneri, H. cruzi, H. gryllata, H. joan-nae, H. leali, H. mathiassoni, H. meridiana,H. microcephala, H. minuscula, H. nana, H.phlebodes, H. pseudomeridiana, H. rhodo-pepla, H. robertmertensi, H. sanborni, H.sartori, H. studerae, H. walfordi, and H. wer-neri. In this analysis we include H. bipunc-tata, H. microcephala, H. nana, H. rhodo-pepla, H. sanborni, and H. walfordi.
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Hyla minima Group: Duellman (1982) ten-tatively grouped together five species fromthe Upper and Middle Amazon Basin andeastern Andes for which data on larval mor-phology, vocalizations, and osteology weremostly absent. He based the grouping ofthese species on small body size and distri-bution. The species he included were H.aperomea, H. leali, H. minima, H. riveroi,and H. rossalleni. The group as such was notnamed until Duellman in Frost (1985) re-ferred to it as the H. minima group. Vigleand Goberdhan-Vigle (1990) added H. mi-yatai to this group; Marquez et al. (1993)implicitly transferred H. leali to the H. mi-crocephala group (see comments for the H.microcephala group above); De la Riva andDuellman (1997) redescribed H. rossalleniand placed it in the H. leucophyllata group.
Small size is a difficult criterion to applyfor the Hyla minima group, considering thatits constituent species are not smaller thanseveral species of the H. microcephalagroup. Duellman (2001) suggested that theH. minima group should be associated withthe H. parviceps group. Unfortunately, thisdoes not improve the systematics of thesefrogs, because no synapomorphies are knownfor either of these two groups (see the H.parviceps group for more comments). In thepresent analysis, we could include only H.miyatai.
Hyla minuta Group: This group was firstdefined by Cochran (1955); most of the spe-cies then included subsequently were trans-ferred by several authors to the H. leuco-phyllata, H. microcephala, or H. rubicundulagroups. The character state Cochran (1955)used to distinguish the H. minuta group wasthe immaculate anterior and posterior surfac-es of the thigh, a character state that is sharedby several 30-chromosome Hyla.
Martins and Cardoso (1987) describedHyla xapuriensis and referred it to the H.minuta group without providing any defini-tion. Kohler and Lotters (2001b) describedH. delarivai and tentatively suggested that itis close to H. minuta. Duellman and Trueb(1983) stated that the H. minuta group con-tained two species, but they did not statewhich one was the second species, and theyprovided no evidence for its monophyly. Thegroup is currently composed of H. minuta
and H. xapuriensis, and, following Kohlerand Lotters (2001b), we tentatively includeH. delarivai. For the purpose of our analysis,only H. minuta was available.
Hyla parviceps Group: This group wasfirst defined by Duellman (1970), and byDuellman and Crump (1974), who also re-viewed it. According to Duellman andCrump (1974), the species in the H. parvi-ceps group differ from other 30-chromosomeHyla species by having: (1) a pronouncedsexual dimorphism in size; (2) shorter snout;(3) tympanic ring indistinct or absent; (5)small (or reduced) axillary membrane; (9)sexual dimorphism in width of dorsolateralstripes; (10) suborbital bars; (11) thighsmarked with spots; (13) iris pale gray withred ring around pupil; (15) more perichon-dral ossification in the tectum nasi and solumnasi; and (17) squamosal articulating withprootics. Duellman and Crump (1974) char-acterized the tadpoles as having ovoid bodiesand xiphicercal tails with moderately deepfins not extending into body; anteroventraloral disc, large marginal papillae, robust ser-rated jaw sheaths, and no more than one rowof labial teeth. It is unclear which of thesecharacter states could be synapomorphies ofthe group. Duellman and Trueb (1983) didnot provide any synapomorphy for the group.
Duellman and Crump (1974) included inthe Hyla parviceps group H. bokermanni, H.brevifrons, H. luteoocellata, H. microps, H.parviceps, and H. subocularis. Heyer (1977)added H. pauiniensis. Heyer (1980) resur-rected H. giesleri from the synonymy of H.microps. Weygoldt and Peixoto (1987) ten-tatively included H. ruschii in the group.Martins and Cardoso (1987) added H. tim-beba. Duellman and Trueb (1989) addedH. allenorum and H. koechlini. Duellman(2001) added H. grandisonae and H. schu-barti. Lescure and Marty (2000) referred H.luteoocellata to a separate group, the H. lu-teoocellata group, which they characterizedby the presence of a cream-colored suborbit-al stripe. Because they did not discuss dif-ferences with the H. parviceps group, we stillconsider H. luteoocellata a member of the H.parviceps group. However, the species theydescribed, H. gaucheri, does not seem tohave this stripe.
The Hyla parviceps group is currently
2005 31FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
composed of 15 species: H. allenorum, H.bokermanni, H. brevifrons, H. gaucheri, H.giesleri, H. grandisonae, H. koechlini, H. lu-teoocellata, H. microps, H. parviceps, H.pauiniensis, H. ruschii, H. schubarti, H. su-bocularis, and H. timbeba. In our analysis weinclude H. brevifrons, H. giesleri, and H.parviceps.
Hyla rubicundula Group: This group wasrecently defined by Napoli and Caramaschi(1998) and was diagnosed by small body size,patternless thighs, and green dorsum in lifethat changes to pinkish or violet when pre-served. Bogart (1970), Rabello (1970) andGruber (2002) reported a 30-chromosomekaryotype in H. rubicundula and H. elianeae.Historically, species of this group were asso-ciated with species now placed in the H. mi-crocephala group, such as H. nana and H.sanborni (Lutz, 1973). The group is currentlycomposed of H. anataliasiasi, H. araguaya,H. cachimbo, H. cerradensis, H. elianeae, H.jimi, H. rhea, H. rubicundula, and H. tritaen-iata. In our analysis, we include H. rubicun-dula.
Species Known or Presumed to Have 30Chromosomes Not Assigned to Any Group:Hyla anceps: Lutz (1948, 1973) associatedH. anceps with H. leucophyllata based on ithaving an axillary membrane, tadpoles withxiphicercal tail, and vivid flash colors. Coch-ran (1955) considered this species to have noknown close relatives and placed it on itsown species group. Bogart (1973) reported a30-chromosome karyotype for this species,and he tentatively associated it with H. mi-crocephala, H. bipunctata, H. elongata (a ju-nior synonym of H. rubicundula), and H.rhodopepla for sharing a single pair of telo-centric chromosomes. Regardless of havingbeen reported to have 30 chromosomes, Hylaanceps has been ignored in all subsequentliterature dealing with phylogenetic relationsof 30-chromosome species of Hyla. Wogel etal. (2000) redescribed the tadpole of H. an-ceps and stated that its morphology support-ed the idea of this species belonging to itsown species group, without giving furtherdetails. We include this species in the anal-ysis.
Hyla amicorum: This species was consid-ered to be similar to H. battersbyi and H.minuta (Mijares-Urrutia, 1998). On this basis
we consider it a 30-chromosome species. Wecould not include this species in the analysis.
Hyla battersbyi: Since its original descrip-tion (Rivero, 1961) this species has beenscarcely mentioned in the literature. We con-sider it tentatively as a 30-chromosome spe-cies based on the association made by Mi-jares-Urrutia (1998) of this species with H.minuta based on overall similarity. We couldnot include this species in the analysis.
Hyla haraldschultzi: Bokermann (1962)stated that the relationships of this specieswere uncertain. Lutz (1973), while treatingthe ‘‘small to minute forms’’, where she in-cluded most of the 30-chromosome Hyla,considered H. haraldschultzi to be insuffi-ciently known. We could not include thisspecies in the analysis.
Hyla limai: In the original description,Bokermann (1962) associated this specieswith H. minuta and H. werneri. Apart froma brief comment by Lutz (1973), it has notbeen referred to again in the literature. Had-dad (unpubl. data), based on morphologicalvariation in populations of H. minuta, findsthat H. limai could be a junior synonym ofthis species. We could not include this nom-inal species in the analysis.
Hyla praestans: Duellman and Trueb(1983) originally placed this species in theH. columbiana group. Kaplan (1999), basedon the presence of a small medial depressionin the internal surface of each arytenoid, sug-gested that H. praestans was related to theH. garagoensis group and possibly its sistertaxon. We could not secure samples for theanalysis.
Hyla stingi: This species, externally simi-lar to H. minuta, has been suggested to bethe sister group of a clade composed of theH. minuta, H. marmorata, H. parviceps, H.leucophyllata, and H. microcephala groups(this being supported by the reduction in thelabial tooth row formula from 1/2 to 0/1;Kaplan, 1994). The apparent synapomorphyuniting H. stingi with this clade is the ante-rior position of the oral disc. This speciescould not be included in the analysis.
Hyla tintinnabulum: This species was as-sociated with H. branneri and H. rubicun-dula by Lutz (1973). This fact and our studyof one of the syntypes (NHMG 473; adultmale) suggest that it is a 30-chromosome
32 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Hyla. We could not include this species inthe analysis.
Hyla yaracuyana: This species was asso-ciated with the 30-chromosome Hyla by Mi-jares-Urrutia and Rivero (2000), but it wasnot included in any species group. We couldnot include this species in the analysis.
Middle American/Holarctic Clade
The monophyly of most Middle Americanand Holarctic Hylinae was suggested byDuellman (1970, 2001) based on biogeo-graphic grounds. This clade would includemost Middle American and Holarctic speciesgroups of Hyla plus the genera Acris, Anoth-eca, Duellmanohyla, Plectrohyla, Pseuda-cris, Pternohyla, Ptychohyla, Smilisca, andTriprion.
Acris: This very distinctive taxon was re-viewed by Duellman (1970) and was includ-ed in several studies of Holarctic hylids(Gaudin, 1974; Hedges, 1986; Cocroft, 1994;da Silva, 1997). In the strict consensus of theanalysis by Cocroft (1994), the position ofAcris was unresolved with respect of the oth-er Holarctic hylids. In the reanalysis done byda Silva (1997), it is sister to a clade com-posed of the species of the Hyla cinerea andH. versicolor groups. The two species A. cre-pitans and A. gryllus are included in ouranalysis.
Anotheca: This monotypic genus was re-viewed by Duellman (1970, 2001). Autapo-morphies of this taxon include the uniqueskull ornamentation composed of sharp, dor-sally pointed spines in the margins of fron-toparietal, maxilla, nasal (including canthalridge), and squamosal, and character statesthat result in its reproductive mode, includingthe female feeding tadpoles with trophic eggs(see Jungfer, 1996). We include the singlespecies Anotheca spinosa in this analysis.
Duellmanohyla: This genus was reviewedby Duellman (2001). Duellmanohyla wasproposed by Campbell and Smith (1992),based on four suggested synapomorphies in-volving larval morphology: a greatly en-larged, pendant oral disc (referred to as fun-nel-shaped mouth by Duellman, 2001); longand pointed serrations on the jaw sheaths;upper jaw sheath lacking lateral processes;and greatly shortened tooth rows. Duellman
(2001) added the bright red iris and the labialstripe expanded below the orbit. Duellma-nohyla contains the following eight species:D. chamulae, D. ignicolor, D. lythrodes, D.rufioculis, D. salvavida, D. schmidtorum, D.soralia, and D. uranochroa. In our analysis,we include D. rufioculis and D. soralia.
Hyla arborea Group: All of the species ofHyla of Europe, North Africa, and Asia havelong been recognized as composing a singlespecies group (Stejneger, 1907; Pope, 1931).Probably because species of this group havegenerally been discussed in isolation of otherhylids, we are not aware of any author hav-ing provided a diagnosis of this group thatcould differentiate them, at least phenotypi-cally, from the Nearctic species placed in theH. cinerea and H. eximia groups. No syna-pomorphy has ever been suggested; themonophyly of the group has apparently beenassumed based on geography and the exter-nal similarity of most species (9 of the 16currently recognized species have been con-sidered at some point of it taxonomic historyto be subspecies or varieties of H. arborea).
Various authors considered the Hyla ar-borea group to be closely related with someNorth American species. Based on similari-ties in advertisement calls, Kuramoto (1980)suggested that the H. arborea group is close-ly related to the H. eximia group of temperateMexico and southwestern United States.
The assumed monophyly of the Hyla ar-borea group was challenged by Anderson(1991) on karyotypic grounds, and this wassupported by Borkin (1999). These authorssuggested the presence of at least two line-ages resulting from two independent inva-sions to Eurasia. According to Anderson(1991), at least H. japonica and H. suweo-nensis (she did not include other easternAsian species) share the presence of a NORin chromosome 6 with the representativesshe studied of the H. eximia group (H. ar-enicolor, H. euphorbiacea, H. eximia), someof the H. versicolor group (H. avivoca, H.chrysoscelis, H. versicolor; H. femoralis),and with H. andersonii. Hyla arborea, H.chinensis, H. meridionalis, and H. savignyishare with most Nearctic species of Hyla, aswell as several of the outgroups that Ander-son (1991) included, the NOR located inchromosomes 10/11 (she considered the
2005 33FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
chromosome pairs to be homologous in thesespecies, with the shift in NOR position pre-sumably corresponding to modulation of het-erochromatin material and not to a translo-cation event; Anderson 1991: 323).
The Hyla arborea group as currently un-derstood is composed of the following 16species: H. annectans, H. arborea, H. chi-nensis, H. hallowellii, H. immaculata, H. in-termedia, H. japonica, H. meridionalis, H.sanchiangensis, H. sarda, H. savignyi, H.simplex, H. suweonensis, H. tsinlingensis, H.ussuriensis, and H. zhaopingensis. In thisanalysis we include H. arborea, H. annec-tans, H. japonica, and H. savignyi.
Hyla bistincta Group: This group was re-viewed by Duellman (2001). The lack of ev-idence supporting the monophyly of the H.bistincta group, together with the possibilitythat Plectrohyla is nested within it, has beenrepeatedly noted (Duellman and Campbell:1992; Toal, 1994; Wilson et al., 1994a; Men-delson and Toal, 1995; Ustach et al., 2000;Canseco-Marquez et al., 2002).
Duellman (2001) presented a cladisticanalysis of the Hyla bistincta group that in-cluded the 17 species known at that time,using a vector of character states of Plectro-hyla and H. miotympanum as the root. Hereported that the analysis resulted in 89 mostparsimonious trees, from which he chose onethat is presented in his figure 400 (Duellman2001: 952). In this tree, the H. bistinctagroup is monophyletic, being supported by asingle transformation, the loss of vocal slits.A reanalysis of his data set15 indicates thatPlectrohyla is nested within the H. bistinctagroup on several of the most parsimonioustrees, and that the strict consensus tree is al-most entirely collapsed. Therefore, Duell-man’s analysis provides no evidence for themonophyly of the H. bistincta group.
The Hyla bistincta group currently com-prises the following 18 described species: H.ameibothalame, H. bistincta, H, calthula, H.calvicollina, H. celata, H. cembra, H. cha-
15 This reanalysis, like that attempted by Canseco-Marquez et al. (2002), could not reproduce the resultspresented by Duellman (2001). The present discussionis based on the results that were obtained using the ad-ditivities specified by Duellman. It resulted in 45 mostparsimonious trees of 56 steps, with CI 5 0.57 and RI5 0.52.
radricola, H. chryses, H. crassa, H. cyanom-ma, H. labedactyla, H. mykter, H. pachyder-ma, H. pentheter, H. psarosema, H. robert-sorum, H. sabrina, and H. siopela. In thisanalysis we include H. bistincta and H. cal-thula.
Hyla bromeliacia Group: The taxonomy,composition, and history of this group werereviewed by Duellman (1970, 2001). Possi-ble synapomorphies suggested by Duellman(2001) are those modifications of the phyto-telmic larvae, mostly the depressed body andthe elongated tail. The other characters heused to separate this group from the brome-liad-dwelling species of the H. pictipes groupare likely plesiomorphic, such as the lack ofmassive temporal musculature (present onlyin H. zeteki and H. picadoi), the presence ofmore than one tooth row (only one in H. pi-cadoi and H. zeteki), and the oral disc notentirely bordered by a single row of papillae(as in H. zeteki and H. picadoi). The groupcomprises two species, H. bromeliacia andH. dendroscarta. In this study we includeonly H. bromeliacia.
Hyla cinerea Group: This group was firstrecognized by Blair (1958) based on adver-tisement call structure. For a review of itshistory see Anderson (1991). The group iscurrently composed of four species, H. ci-nerea, H. femoralis, H. gratiosa, and H. squi-rella, and the group is monophyletic in theanalyses of Hedges (1986), Cocroft (1994),and da Silva (1998), being supported by al-lozyme data (taken in the last two studiesfrom Hedges, 1986). We included all fourspecies of the group in the present analysis.
Hyla eximia Group: The taxonomy, com-position, and history of this group were thor-oughly reviewed by Duellman (1970, 2001).Duellman (2001) presented an extensive def-inition; however, it is unclear which of thecharacter states could be taken as evidenceof monophyly of the group. Hedges (1986)and Cocroft (1994) included two species, H.arenicolor and H. eximia, in their analyses.While in Hedges’s (1986) results these twospecies form a monophyletic group, on thestrict consensus of Cocroft’s (1994) mostparsimonious trees, both species are notmonophyletic and form a basal polytomywith several other species of Hyla. da Silva’s(1997) reanalysis of Cocroft’s (1994) data set
34 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
yielded on its strict consensus H. arenicolorand H. eximia as a monophyletic group,which appears to be supported by the allo-zyme data of Hedges (1986). Besides theseven species included by Duellman (2001)in the group, Eliosa Leon (2002) resurrectedH. arboricola from the synonymy of H. ex-imia, adding an eigth species to the group.See the H. arborea group for additional com-ments. The H. eximia group is currently com-posed of eight species: Hyla arboricola, H.arenicolor, H. bocourti, H. euphorbiacea, H.eximia, H. plicata, H. walkeri, and H. wrigh-torum. In this analysis we include H. areni-color, H. euphorbiacea, H. eximia, and H.walkeri.
Hyla godmani Group: The taxonomy,composition, and history of this group werereviewed by Duellman (1970, 2001). Duell-man (2001) included in this group the onlylowland pond-breeders from Middle Americathat do not have 30 chromosomes. He sug-gested as tentative evidence of monophylythe weakly ossified skulls and the presenceof an axillary membrane. This group cur-rently comprises four species: H. godmani,H. loquax, H. picta, and H. smithii. We in-clude H. picta and H. smithii in our analysis.
Hyla miotympanum Group: The taxonomy,composition, and history of this group werethoroughly reviewed by Duellman (1970,2001). Duellman (2001), using a hypotheti-cal outgroup, suggested eight synapomor-phies for the group16: abbreviated axillarymembrane; indistinct fold on wrist; fingersless than one-half webbed; tarsal fold presentthrough the entire length of the tarsus; cloa-cal opening directed posteroventrally; nuptialexcrescences present; medial ramus of pter-ygoid short, not in contact with prootic; andlarval oral disc in ventral position. It seemsunlikely that any of these character transfor-mations will hold as synapomorphies of thegroup in the context of a more inclusive anal-ysis. The group currently contains 11 spe-cies: H. abdivita, H. arborescandens, H. biv-ocata, H. catracha, H. cyclada, H. hazelae,H. juanitae, H. melanomma, H. miotympan-
16 Note that in ‘‘the preferred’’ cladogram, there is acharacter 27 as one of the synapomorpies of the Hylamiotympanum group; presumably this is an error for 17,since the data set has 22 characters, and character 17 isinclusive of the ingroup.
um, H. perkinsi, and H. pinorum. Our anal-ysis includes H. arborescandens, H. cyclada,H. melanomma, H. miotympanum, and H.perkinsi.
Hyla mixomaculata Group: Duellman(1970) reviewed the group and listed severalcharacter states that define it. Some of theseare probably synapomorphies, such as(known) larvae with an enlarged oral discwith 7/10 or 11 labial tooth rows (the largestnumber of posterior tooth rows known for aMiddle American hylid group); the taxonom-ic distribution of other character states indi-cate that they are likely synapomorphies ofa more inclusive clade, such as the largefrontoparietal fontanelle and the absence (orreduction, unclear on fig. 211 of Duellman,1970) of the quadratojugal. The group is cur-rently composed of four species, H. mixe, H.mixomaculata, H. nubicola, and H. pellita.Only H. mixe was available for this analysis.
Hyla pictipes Group: The taxonomy, com-position, and history of this group were re-viewed by Duellman (1970, 2001). Duellman(2001) provided a phylogenetic analysis17 ofthe montane species of Hyla of lower CentralAmerica, where he included the H. pseudop-uma and H. pictipes groups.
Duellman (2001) suggested six synapo-morphies for the group: slender nasals inadults; tadpoles with stream-dwelling habits;oral disc ventral; complete marginal papillae;only one row of marginal papillae; and pres-ence of submarginal papillae. Note that withthe possible exception of slender nasals inadults, the other three character states as de-fined by Duellman (2001) are also present inseveral other groups of Middle Americanstream-breeding frogs (i.e., the Hyla bistinctagroup, H. mixomaculata group, H. sumi-chrasti group, and Plectrohyla).
The Hyla pictipes group includes 11 spe-cies: H. calypsa, H. debilis, H. insolita, H.lancasteri, H. picadoi, H. pictipes, H. rivu-
17 We could not reproduce the results of Duellman(2001) using his data matrix under the same additivities.This is most probably due to an editorial problem withthe data matrix; the scores for Hyla debilis are all 0.Evidently the data set as printed is different from thatused to choose the trees shown, where H. debilis is thesister species of H. rivularis. Because of this situation,we discuss the synapomorphies shown in the book, notthose from our reanalysis.
2005 35FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
laris, H. thorectes, H. tica, H. xanthosticta,and H. zeteki. Considering the uncertaintiesregarding the monophyly of this group, anappropriate taxon sampling should ideally in-clude representatives of the four former spe-cies groups currently combined into the H.pictipes group. Unfortunately, the only rep-resentatives available are H. rivularis andHyla sp. 5 (aff. H. thorectes), an undescribedspecies from Mexico similar to H. thorectes.
Hyla pseudopuma Group: The taxonomy,composition, and history of this group werereviewed by Duellman (1970, 2001), whocould not provide a synapomorphy for it. Inhis phylogenetic analysis of the H. pseudop-uma and H. pictipes groups, the H. pseudop-uma group appears as a basal unresolvedgrade, although this is a consequence of con-straining the nonmonophyly of the H. pseu-dopuma group by using H. angustilineata asthe root. The H. pseudopuma group includesfour species: H. angustilineata, H. graceae,H. infucata, and H. pseudopuma. In thisstudy we include only H. pseudopuma.
Hyla sumichrasti Group: The taxonomy,composition, and history of this group werereviewed by Duellman (1970, 2001). Possi-ble synapomorphies for the group (Duell-man, 2001) are the presence of massive na-sals, and tadpoles with immense oral discs,with 3/6 to 3/7 labial tooth rows instead ofthe 2/3 to 2/6 of the H. miotympanum group.The group currently includes four species: H.chimalapa, H. smaragdina, H. sumichrasti,and H. xera. In this study we include H. chi-malapa and H. xera.
Hyla taeniopus Group: This group was re-viewed by Duellman (1970, 2001), who de-fined it as having well-ossified quadratoju-gals in contact with maxillaries, and tadpolesthat have ventral mouths with two or threeanterior rows of teeth and three or four pos-terior rows. While the presence of enlargedtestes is a possible synapomorphy of a sub-group composed of H. altipotens, H. trux,and H. taeniopus, there is no evidence for themonophyly of the entire group (Mendelsonand Campbell, 1999; Duellman, 2001). Fivespecies are currently included in this group:H. altipotens, H. chaneque, H. nephila, H.taeniopus, and H. trux. In this analysis, weinclude H. nephila and H. taeniopus.
Hyla tuberculosa Group: Affinities among
species of fringe-limbed treefrogs were firstsuggested by Dunn (1943) and by Taylor(1948, 1952) based on their overall appear-ance. Firschein and Smith (1956) suggestedthat the presence of a ‘‘prepollex’’ (presum-ably referring to an enlarged prepollex) andsimilarity of external habitus, size, skin tex-ture, and fringed limbs were indicative of acommon origin. Duellman (1970, 2001) pre-sented a formal definition of the group, as-serting that these frogs be placed in the samegroup based on their large size, presence ofdermal fringes on the limbs (although absentin H. dendrophasma), extensive webbing onhand and feet, and modified prepollices. (Im-portantly, note that apparently the modifiedprepollices involve three different morphol-ogies: modification into a projecting spine ora spadelike blade or a clump of spines;Duellman, 1970.) Duellman (2001) addedthat there is no compelling evidence that thegroup is monophyletic, an opinion that weshare. Furthermore, he tentatively suggestedthat this group could be related with theGladiator Frogs rather than with the MiddleAmerican/ Holarctic clade. The Hyla tuber-culosa group has been referred to as the H.miliaria group (Campbell et al., 2000; Duell-man, 1970, 2001; Savage and Heyer, ‘‘1968’’[1969]) and is composed of H. dendrophas-ma, H. echinata, H. fimbrimembra, H. mili-aria, H. minera, H. phantasmagoria, H. sal-vaje, H. thysanota, H. tuberculosa, and H.valancifer. In this analysis, we include H.dendrophasma and H. miliaria.
Hyla versicolor Group: This group wasfirst recognized by Blair (1958) on the basisof advertisement call structure. See Duellman(1970) and Anderson (1991) for a brief tax-onomic history. Both Blair (1958) and Duell-man (1970) included H. arenicolor in thegroup until Hedges (1986), Cocroft (1994),and da Silva (1997) showed on successiveanalyses that this species was more closelyrelated to H. eximia (see the H. eximia groupfor further comments), always on the basisof the allozyme data collected by Hedges(1986).
Although placed originally in the Hyla ci-nerea group by Blair (1958), H. andersoniiwas transferred to the H. versicolor group byWiley (1982), with this being supported byHedges (1986), Cocroft (1994), and da Silva
36 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
(1997) on the basis of the allozyme data col-lected by Hedges (1986). Similarly, H. fe-moralis was considered a member of the H.versicolor group until Hedges (1986) trans-ferred it to the H. cinerea group, an assign-ment also corroborated by da Silva (1997).The group currently comprises four species:H. andersonii, Hyla avivoca, H. chrysoscelis,and H. versicolor. In this study we includeall but H. chrysoscelis.
Pseudacris: The phylogenetic relation-ships of this genus were reviewed by Hedges(1986), Cocroft (1994), da Silva (1997), andMoriarty and Cannatella (2004). Hedges(1986) presented an electrophoretic analysisof 33 presumed genetic loci, where all spe-cies of Pseudacris were monophyletic, andwhich provided evidence to include the for-mer Hyla cadaverina, H. crucifer, and H. re-gilla in Pseudacris. Cocroft (1994) per-formed a phylogenetic analysis of Pseuda-cris, including several Holarctic hylids asoutgroups, with characters from varioussources (osteology, vocalizations, karyo-types, allozymes, sperm morphology) andprevious analyses (Hedges, 1986). The strictconsensus of his most parsimonious treesshows P. crucifer as the sister taxon of theremaining species of Pseudacris; the twoclassically recognized species groups (the P.ornata and the P. nigrita groups) each beingmonophyletic; and P. ocularis being the sis-ter taxon of the P. nigrita group. However,Hedges (1986) found no evidence supportingthe inclusion of P. cadaverina and P. regillain Pseudacris, and for this reason he treatedthem as Hyla. Furthermore, the relationshipsof the remaining, monophyletic species ofPseudacris with the other Holarctic hylineare unresolved. In the modified reanalysisperformed by da Silva (1997), the strict con-sensus shows that the clade composed of H.regilla and H. cadaverina is sister to Pseu-dacris and is supported apparently by allo-zyme data. Because of this, these two speciesare considered again to be within Pseudacris(da Silva, 1997).
Moriarty and Cannatella (2004) presenteda phylogenetic analysis of the mitochondrialribosomal genes 12S, tRNA valine, and 16Sthat included all species of Pseudacris andthree outgroups, Hyla andersonii, H. chry-soscelis, and H. eximia. The analysis identi-
fied four major clades: (1) the P. regillaclade (P. cadaverina and P. regilla; Moriartyand Cannatella referred to it as the WestCoast clade); (2) the Pseudacris ornata clade(P. ornata, P. streckeri, and P. illinoiensis;Moriarty and Cannatella called it fat frogsclade); (3) the P. crucifer clade (P. cruciferand P. ocularis); and (4) the P. nigrita clade(including P. brimleyi, P. brachyphona, P.clarkii, P. feriarum, P. maculata, and P. tris-eriata; Moriarty and Cannatella called itTrilling Frogs clade). In our study we includePseudacris cadaverina, P. crucifer, P. ocu-laris, P. regilla, and P. triseriata.
Pternohyla: This Mexican casque-headedfrog genus was reviewed by Trueb (1969)and Duellman (1970, 2001). In Duellman’s(2001) phylogenetic analysis of Pternohyla,Smilisca, and Triprion, the monophyly ofPternohyla is supported by four synapomor-phies: small discs on fingers; supernumerarytubercles diffuse or absent; large inner meta-tarsal tubercle18; and large marginal papillaein the larval oral disc. Unfortunately, smalldiscs on fingers are a synapomorphy only un-der an accelerated optimization (ACCT-RAN), and therefore they have no evidentialvalue for the clade.
In Duellman’s (2001) analysis, Triprionplus Pternohyla forms a monophyletic groupnested within Smilisca. The synapomorphiessupporting Triprion plus Pternohyla are19:nasals with broad medial contact; median ra-mus of pterygoid not in contact with prootic;maxilla moderately expanded laterally; cra-nial-integumentary co-ossification present;webbing between fingers absent; and innermetatarsal tubercle small. Pternohyla is com-posed of two similar species, P. dentata, and
18 For the character ‘‘inner metatarsal tubercle’’,Duellman (2001) defined four character states: moderate(0), small (1), large (2), and spadelike (3). He stated thathe considered this character as additive in the order0→1→2→3; unless there is an editorial mistake, thisseems a rather peculiar and unjustified ordering.
19 Duellman maps on his preferred tree character states4.1, 7.2, 8.1, 9.1, 13.1, 16.1, and 19.1. However, an ex-amination of the character list and matrix shows thatthere is no state 2 defined for character 7, and that char-acter 13 is actually an autapomorphy of Triprion spa-tulatus; it is very likely that 13 is a typographical errorfor character 14, which according to the data set is asynapomorphy for the group; in the synapomorphy listgiven above, we assume that these problems were fixed,and so we ignore character state 7.2.
2005 37FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
P. fodiens. In this analysis we include P. fod-iens.
Plectrohyla: This genus was reviewed byDuellman (2001) and discussed by McCranieand Wilson (2002). Its phylogenetic relation-ships were addressed by Duellman andCampbell (1992), Wilson et al. (1994a), andDuellman (2001). The monophyly of Plec-trohyla does not appear to be controversial.Duellman and Campbell (1992) listed sixsynapomorphies: bifurcated alary process ofpremaxilla; sphenethmoid ossified anteriorly,incorporating the septum nasi and projectingforward to the leading margins of the nasals;frontoparietals abutting broadly anteriorlyand posteriorly, exposing a small area of thefrontoparietal fontanelle; hypertrophied fore-arms; and absence of lateral folds in the oraldisc. Wilson et al. (1994a) added ‘‘prepollexenlarged, elongated, ossified, flat, terminallyblunt.’’ Duellman (2001) interpreted that thisdefinition corresponded to more than onecharacter, and so he divided it into two char-acters, the derived state of the first one being‘‘enlarged and ossified prepollex in both sex-es’’, and the derived state of the second onebeing ‘‘enlarged and truncate prepollex.’’ Seecomments under the Hyla bistincta group.
Plectrohyla currently contains 18 species:P. acanthodes, P. avia, P. chrysopleura, P.dasypus, P. exquisitia, P. glandulosa, P. gua-temalensis, P. hartwegi, P. ixil, P. lacertosa,P. matudai, P. pokomchi, P. psiloderma, P.pycnochila, P. quecchi, P. sagorum, P. te-cunumani, and P. teuchestes. In this analysiswe include P. guatemalensis, P. glandulosa,and P. matudai.
Ptychohyla: This group was reviewed byDuellman (2001). Campbell and Smith(1992) suggested three synapomorphies forPtychohyla: the presence of two rows of mar-ginal papillae, an increased number of toothrows in larvae (from 3/5 to 6/9), and astrongly developed lingual flange of the parspalatina of the premaxilla. Duellman (2001)also suggested as synapomorphies the pres-ence of ventrolateral glands in breedingmales, and the coalescence of tubercles toform a distinct ridge on the ventrolateraledge of the forearm. The increase in thenumber of tooth rows could actually be asynapomorphy not of Ptychohyla but for amore inclusive clade containing Ptychohyla
plus other species of stream-breeding hylidsthat also have a tooth row formula largerthan 2/3. Similarly, ventrolateral glands arepresent also in some species of Duellmano-hyla (Campbell and Smith, 1992; Duellman,2001).
For an unstated reason, Savage (2002a)excluded Ptychohyla legleri and P. salvador-ensis from Ptychohyla, placing them back inHyla. These two species were originally inHyla (former H. salvadorensis group; seeDuellman, 1970) until Campbell and Smith(1992) transferred them to Ptychohyla. Be-cause we are not aware of any evidence sup-porting Savage’s action, we consider them tobe members of Ptychohyla.
Ptychohyla is composed of 12 species: P.acrochorda, P. erythromma, P. euthysanota,P. hypomykter, P. legleri, P. leonhard-schultzei, P. macrotympanum, P. panchoi, P.salvadorensis, P. sanctaecrucis, P. spinipol-lex, and P. zophodes. In this analysis weinclude P. euthysanota, P. hypomykter, P.leonhardschultzei, P. spinipollex, P. zopho-des, and Ptychohyla sp., an undescribed spe-cies from Oaxaca, Mexico.
Smilisca: This genus was reviewed byDuellman and Trueb (1966) and Duellman(1970, 2001). Duellman (2001) could ad-vance no evidence for the monophyly ofSmilisca. He presented a phylogenetic anal-ysis rooted with a hypothetical ancestor,whose strict consensus showed Pternohylaplus Triprion nested within Smilisca, beingmore closely related to S. baudinii and S.phaeota. The synapomorphies supportingPternohyla 1 Triprion 1 ‘‘Smilisca’’ are thepresence of lateral flanges on the frontopari-etals, and the unexposed frontoparietal fon-tanelle. The species of Smilisca have beendivided (Duellman and Trueb, 1966) into theS. sordida group (S. puma and S. sordida),the S. baudinii group (S. baudinii, S. cy-anosticta, and S. phaeota), and S. sila, a formconsidered intermediate between these twogroups. In Duellman’s (2001) phylogeneticanalysis, S. sila plus the S. sordida group ismonophyletic, with its synapomorphy beingthe short maxillary process of the nasal. Smi-lisca contains six species: S. baudinii, S. cy-anosticta, S. phaeota, S. puma, S. sila, andS. sordida. In this analysis we include thethree species in the S. baudinii group, S. bau-
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dinii, S. cyanosticta, S. phaeota, and one spe-cies of the S. sordida group, S. puma.
Triprion: This genus was reviewed byTrueb (1969) and Duellman (1970, 2001). InDuellman’s (2001) phylogenetic analysis ofPternohyla, Smilisca, and Triprion, themonophyly of Triprion is supported by threesynapomorphies20: maxilla greatly expandedlaterally, prenasal bone present, and presenceof parasphenoid odontoids. See commentsfor Smilisca and Pternohyla. Triprion iscomposed of two species, T. petasatus andT. spatulatus. In the analysis we include T.petasatus.
Casque-Headed Frogs and Related Genera
Duellman’s (2001) suggestion of MiddleAmerican/Holarctic frogs being monophylet-ic clearly separates the Middle Americancasque-headed frogs (Triprion, Pternohyla)from the South American and West Indiancasque-headed frogs. This is not surprisingconsidering that traditionally the groupknown as the casque-headed frogs was con-sidered to be nonmonophyletic (Trueb,1970a, 1970b). However, the position of theSouth American and West Indian casque-headed frogs remains controversial, and noauthor has presented evidence indicatingwhether they form a monophyletic group.
Aparasphenodon: This genus of casque-headed frogs was reviewed and characterizedby Trueb (1970a) and Pombal (1993). Thepresence of a prenasal bone is a likely syn-apomorphy of Aparasphenodon (with aknown homoplastic occurrence in Triprion,as reported by Trueb, 1970a). This genus cur-rently comprises three species, A. bokerman-ni, A. brunoi, and A. venezolanus. We includeA. brunoi in the analysis.
Argenteohyla: This monotypic genus wasdescribed and reviewed by Trueb (1970b),who segregated it from Trachycephalus,where it had been placed by Klappenbach(1961). Motives for this segregation were theabsence in Argenteohyla of several characterstates of Trachycephalus as redefined byTrueb (1970a), such as the dermal spheneth-
20 On his preferred tree (his fig. 410), one of the char-acter transformations is numbered 18; this is a typo-graphical error for 12, the only other character that sup-ports this clade but not shown in the tree.
moid, the poorer development of ossificationand cranial sculpturing, and vocal sacs thatwhen inflated protrude posteroventrally tothe angles of the jaw. Possible autapomor-phies of this taxon include the fusion of thezygomatic ramus of the squamosal with thepars facialis of the maxilla. The genus com-prises a single species, A. siemersi, for whicha northern subspecies, A. s. pederseni, wasdescribed by Williams and Bosso (1994). Inthis analysis we included a specimen thatcorresponds to the northern form.
Corythomantis: This monotypic genus wasreviewed by Trueb (1970a). Autapomorphiesof this genus include the absence of pala-tines, and nasals that conceal the allaryprocesses of premaxillaries (Trueb, 1970a).We include the single species Corythomantisgreeningi in this analysis.
Osteocephalus: This genus was diagnosedby Goin (1961) and Trueb (1970a) and stud-ied in detail by Trueb and Duellman (1971).These authors recognized five species: Osteo-cephalus verruciger, O. taurinus, O. buckleyi,O. leprieurii, and O. pearsoni. In the last 20years, several new species were described,adding to a total of 18 currently recognizedspecies (see Jungfer and Hodl, 2002; Lynch,2002). Trueb and Duellman (1971) employed20 character states to characterize Osteoce-phalus. Jungfer and Hodl (2002) modifiedsome of these characters to take into accountsubsequently discovered species. As stated byRon and Pramuk (1999), referring to the di-agnostic states employed earlier by Trueb andDuellman (1971), it is unclear which, if any,of the character states are synapomorphic forthe genus. Trueb (1970a) and Trueb andDuellman (1971) suggested, based on thepresence of paired lateral vocal sacs in thefive species then recognized, that Osteoce-phalus was related to a group composed ofArgenteohyla, Trachycephalus, and Phryno-hyas.
Martins and Cardoso (1987) described Os-tecephalus subtilis that, unlike the other spe-cies known at that time, is characterized bya single, subgular vocal sac that expands lat-erally; a similar morphology was describedby Smith and Noonan (2001) in O. exoph-thalmus. Jungfer and Schiesari (1995) de-scribed O. oophagus, a species with a single,median vocal sac, a reproductive mode in-
2005 39FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
volving oviposition in bromeliads, and phy-totelmous oophagous larvae. Jungfer et al.(2000), Jungfer and Lehr (2001), and Lynch(2002) described four species, O. deridens,O. fuscifacies, O. leoniae, and O. heyeri,which also have a single, median vocal sac.According to Lynch (2002), O. cabrerai alsoshares this characteristic. Reproductivemodes are unknown for O. cabrerai, O. ex-ophthalmus, O. heyeri, O. leoniae, and O.subtilis; spawning in bromeliads is suspectedfor O. deridens and O. fuscifacies (Jungfer etal., 2000). Note that Lynch (2002) doubted apossible relationship between O. heyeri andwhat he called the ‘‘presumed clade of oop-hagous species’’ (where he included O. der-idens, O. fuscifacies, and O. oophagus), sug-gesting instead that it could be related towhat he called O. rodriguezi (at that time al-ready transferred to the new genus Tepuihylaby Ayarzaguena et al., ‘‘1992’’ [1993b]).While the species known or suspected tospawn in bromeliads could be monophyletic,we are not aware of any synapomorphy sup-porting the monophyly of all remaining spe-cies of Osteocephalus.
The species currently included in Osteo-cephalus are O. buckleyi, O. cabrerai, O.deridens, O. elkejungingerae, O. exophthal-mus, O. fuscifacies, O. heyeri, O. langsdorf-fii, O. leoniae, O. leprieurii, O. mutabor, O.oophagus, O. pearsoni, O. planiceps, O. sub-tilis, O. taurinus, O. verruciger, and O. ya-suni. Considering the uncertainties regardingOsteocephalus, we attempted to include rep-resentatives of the morphological and repro-ductive diversity within the genus: O. ca-brerai, O. langsdorffii, O. leprieurii, O. oop-hagus, and O. taurinus.
Osteopilus: The genus Osteopilus was res-urrected by Trueb and Tyler (1974) for threeapparently related species that were often re-ferred to collectively as the Hyla septentrion-alis group (see Dunn, 1926; Trueb, 1970a).Trueb and Tyler (1974) provided a diagnosticdefinition of the genus; a possible synapo-morphy is the differentiation of the m. inter-mandibularis to form supplementary apicalelements. Trueb and Tyler (1974) also main-tained, due to the impressive morphologicaldivergence, that Osteopilus, the other Antil-lean groups then considered to be in Hyla (H.heilprini, H. marianae, H. pulchrilineata, H.
vasta, H. wilderi), and the new genus theyerected, Calyptahyla, represented several in-dependent invasions from the mainland.
Maxson (1992) and Hass et al. (2001), us-ing albumin immunological distances, sug-gested that Osteopilus is paraphyletic withrespect to most other West Indian hylids(with the exception of Hyla heilprini, a Glad-iator Frog). Hedges (1996) mentioned thatunpublished DNA sequence data confirmedthese findings. Anderson (1996) presented akaryological study of the three species of Os-teopilus, indicating that her data were com-patible with a monophyletic Osteopilus.Based on the comments by Hedges (1996),and immunological results of Hass et al.(2001), Franz (2003), Powell and Henderson(2003a, 2003b), and Stewart (2003) trans-ferred Calyptahyla crucialis, H. marianae,H. pulchrilineata, H. vasta, and H. wilderi toOsteopilus that now includes eight species.Osteopilus is grouped together only on thebasis of the immunological distance results,as no discrete character data set supportingits monophyly has yet been published. Thespecies of Osteopilus available for our studywere O. crucialis, O. dominicensis, O. sep-tentrionalis, and O. vastus.
Phrynohyas: This genus was reviewed byDuellman (1971b). Although very distinctiveexternally, the only seeming synapomorphyin the diagnostic definition of Phrynohyasprovided by Duellman (1971b) is the exten-sively developed parotoid glands in the oc-cipital and scapular regions. Likely related tothis character state, the viscous, milky secre-tions of the species of this genus could alsobe considered synapomorphic. Lescure andMarty (2000) transferred Hyla hadroceps tothis genus; this was confirmed in a phylo-genetic analysis using the mitochondrial ri-bosomal gene 12S by Guillaume et al.(2001). Pombal et al. (2003) described P.lepida. See Osteocephalus for further com-ments. Phrynohyas currently contains sevenspecies: P. coriacea, P. hadroceps, P. imi-tatrix, P. lepida, P. mesophaea, P. resinific-trix, and P. venulosa. In our analysis we in-clude P. hadroceps, P. mesophaea, P. resi-nifictrix, and P. venulosa.
Tepuihyla: This genus was defined by Ay-arzaguena et al. (‘‘1992’’ [1993b]) for fivespecies of Osteocephalus previously consid-
40 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
ered to constitute the O. rodriguezi speciesgroup (Duellman and Hoogmoed, 1992; Ay-arzaguena et al., ‘‘1992’’ [1993a]). Ayarza-guena et al. (‘‘1992’’ [1993b]) differentiatedTepuihyla from Osteocephalus using the fol-lowing character states present in Tepuihyla:subgular vocal sac, absence or extreme re-duction of hand webbing, more reduced toewebbing, smaller size, absence of cranial co-ossification, large frontoparietal fontanelle,shorter nasals, and shorter frontoparietals. Itis unclear which, if any, of these characterstates are apparent synapomorphies of Te-puihyla. There are eight species currently in-cluded in this genus: T. aecii, T. celsae, T.edelcae, T. galani, T. luteolabris, T. rima-rum, T. talbergae, and T. rodriguezi. In thisanalysis we include only T. edelcae.
Trachycephalus: The relationships of thiscasque-headed taxon were discussedby Trueb (1970a) and Trueb and Duellman(1971). They diagnosed Trachycephalus fromArgenteohyla, Osteocephalus, and Phrynoh-yas for having heavily casqued and co-ossi-fied skulls, a medial ramus of pterygoid thatdoes not articulate with the prootic, and a par-asphenoid having odontoids. A likely syna-pomorphy of Trachycephalus is the presenceof exostosis on the alary process of the pre-maxillae (Trueb, 1970a). Trachycephalus con-tains three species: T. atlas, T. jordani, and T.nigromaculatus. In this analysis we include T.jordani and T. nigromaculatus.
Species and Species Groups of Hyla NotAssociated with Any Major Clade
Hyla aromatica Group: This group wasproposed by Ayarzaguena and Senaris(‘‘1993’’ [1994]) for two species from theVenezuelan Tepuis, H. aromatica and H. in-parquesi, which they could not associatewith any of the species groups known fromthe Guayanas. Ayarzaguena and Senaris(‘‘1993’’ [1994]) noticed that the H. aroma-tica group shares some characters with theH. larinopygion group; however, they pre-ferred to retain it as a separate group. Theyjustified this decision based on the smallersize of members of the H. aromatica group,different coloration pattern, supraorbital car-tilaginous process, vomerine odontophoressmoothly S-shaped and with more odonto-
phores, small nasals, and large prepollex.They included as well other character states,that actually, like some of these just men-tioned, are either shared with several neo-tropical groups (supraorbital cartilaginousprocess; Faivovich, personal obs.), or somespecies of the H. larinopygion group (vo-merine odontophores smoothly S-shaped;Duellman and Hillis, 1990: 5), or with the H.armata group (labial tooth row formula; seeCadle and Altig, 1991), or they support themonophyly of the H. aromatica group(adults with strong odor). Considering thelack of evidence of monophyly for the H.larinopygion group, Ayarzaguena and Sena-ris (‘‘1993’’ [1994]) cannot be questioned forrecognizing a separate species group.
Ongoing research by Faivovich andMcDiarmid suggests that Hyla loveridgeishould also be considered part of the H. aro-matica group. For this analysis, we includeH. inparquesi.
Hyla uruguaya Group: This group hasnever been mentioned as such in the litera-ture. However, clear similarities had beenshown by Langone (1990) between H. uru-guaya and H. pinima (these species being al-most undistinguishable). Possible synapo-morphies of the H. uruguaya group are thebicolored iris (also shared with Aplastodis-cus; see Garcia et al., 2001a), the presencein tadpoles of two small, keratinized platesbelow the lower jaw sheath, and a reductionin the size of the marginal papillae of theposterior margin of the oral disc relative tothe other papillae (Kolenc et al., ‘‘2003’’[2004]). From this apparent clade we includeH. uruguaya in our analysis.
Hyla chlorostea: Duellman at al. (1997)proposed the recognition of a species groupto include the enigmatic Hyla chlorostea, aspecies known only from its holotype (a sub-adult male), which could not be associatedwith any known group of Hyla after its de-scription (Reynolds and Foster, 1992). Un-fortunately, we were unable to include thistaxon in our analysis.
Hyla vigilans: Different perspectives con-cerning this enigmatic species were summa-rized by Suarez-Mayorga and Lynch(2001a). These authors rejected the possibil-ity of a relationship with Scinax (as suggest-ed by La Marca in Frost, 1985). Instead, they
2005 41FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
asserted that they suspected a possible rela-tionship with Sphaenorhynchus or with H.picta (from the H. godmani species group)based on ‘‘oral disc and mouth position’’ ofthe tadpoles. We could not obtain samples ofthis species for our analysis.
Hyla warreni: This species, known onlyfrom two adult females, was described byDuellman and Hoogmoed (1992), who didnot associate it with any other species or spe-cies group. Unfortunately, we could not ob-tain samples of this species for our analysis.
Other Genera
Aplastodiscus: The taxonomy and historyof this genus was recently reviewed thor-oughly by Garcia et al. (2001a). Accordingto these authors the monophyly of the genusis supported by four putative synapomor-phies: (1) the absence of webbing betweentoes I and II and basal webbing between theother toes; (2) bicolored iris; (3) females withunpigmented eggs; and (4) great develop-ment of internal metacarpal and metatarsaltubercles. Based on overall morphologicaland advertisement call similarities B. Lutz(1950) suggested a close relationship of thisgenus with Hyla albosignata. Garcia et al.(2001a) suggest that Aplastodiscus could berelated with the H. albofrenata and H. al-bosignata complexes of the H. albomargin-ata group, as defined by Cruz and Peixoto(1984), based on the presence of enlarged in-ternal metacarpal and metatarsal tubercles,and unpigmented eggs. Haddad et al. (2005)described the reproductive mode of A. per-viridis and noticed that it was the same asthat described by Haddad and Sawaya (2000)and Hartmann et al. (2004) in species in-cluded in H. albofrenata and H. albosignatacomplexes. Based on this, Haddad et al.(2005) suggested a possible relationship be-tween these two species complexes andAplastodiscus. Aplastodiscus is composed oftwo species, Aplastodiscus cochranae and A.perviridis; we include both in our analysis.
Nyctimantis: This monotypic Neotropicalgenus was considered a member of the Hem-iphractinae by Duellman (1970) and Trueb(1974). Duellman and Trueb (1976) reviewedthe taxon and placed it in Hylinae. Duellmanand Trueb (1976) considered Nyctimantis to
be related with Anotheca spinosa becauseboth share the medial ramus of the pterygoidthat is juxtaposed squarely against the an-terolateral corner of the ventral ledge of theotic capsule. Also, frogs of both genera areknown (Anotheca; Taylor, 1954; Jungfer,1996) or suspected (Nyctimantis; Duellmanand Trueb, 1976) to deposit their eggs in wa-ter-filled tree cavities. However, Duellman(2001) latter placed Anotheca in the MiddleAmerican/Holarctic clade, implicitly sug-gesting no relationship with Nyctimantis.Considering the uncertainty of the positionof Nyctimantis within hylines, at this stage itis difficult to interpret which character statesare autapomorphic. We include the singlespecies Nyctimantis rugiceps in this analysis.
Phyllodytes: The history of this genus wasreviewed by Bokermann (1966b). Possiblesynapomorphies of the taxon are the pres-ence of odontoids on the mandible and onthe cultriform process of the parasphenoid(Peters, ‘‘1872’’ [1873]), something uniquewithin the Hylinae. Peixoto and Cruz (1988)noticed that among the six species recog-nized at that time, four species (P. acumi-natus, P. brevirostris, P. luteolus, and P. tub-erculosus) share the presence of series of en-larged tubercles on the venter and an en-larged tubercle on each side at the origin ofthe thigh (Bokermann, 1966b: fig. 6). Theother two species, P. auratus and P. kautskyi,have uniform granulation on the venter andlack enlarged tubercles on the thighs, as alsoseems to be the case in P. melanomystax, aspecies described later (see Caramaschi et al.,1992). Peixoto et al. (2003) described twoadditional species, P. edelmoi and P. gyri-naethes; both have a tubercle on each side atthe origin of the thigh. Phyllodytes edelmoihas a series of indistinct tubercles on the ven-ter; in P. gyrinaethes they do not form series.Caramaschi and Peixoto (2004) added P.punctatus, which has two medial, poorly dis-tinct rows of tubercles. Caramaschi et al.(2004a) resurrected P. wuchereri. Peixoto etal. (2003) suggested three different speciesgroups based on color pattern, and Caramas-chi et al. (2004) further expanded the defi-nitions. The P. luteolus group is character-ized by a plain pattern with a variably de-fined dorsolateral dark brown to black lineon canthus rostralis and/or behind the corner
42 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
of eye. This group includes P. acuminatus,P. brevirostris, P. edelmoi, P. kautskyi, P.luteolus, and P. melanomystax. The P. tub-erculosus group has a pale brown dorsumwith scattered dark brown dots and includesP. punctatus and P. tuberculosus. The P. au-ratus group has a dorsal pattern of two dor-solateral, longitudinal white or yellowishstripes, with each stripe being bordered by adark brown or black line from posterior cor-ner of eye to groin. This group includes P.auratus and P. wuchereri. Finally, P. gyri-naethes is placed in its own group for havingred color on hidden surfaces of thighs and ahighly modified tadpole. It is unclear if anyof these groups is monophyletic. Tissueswere available for P. luteolus and an uniden-tified species, Phyllodytes sp., from Bahia,Brazil.
Lysapsus and Pseudis: The monophyly ofthe former subfamily Pseudinae has not beenhistorically controversial; it is supported bythe presence of a long, ossified intercalaryelement between the ultimate and penulti-mate phalanges. Haas (2003) added severalsynapomorphies from larval morphology,based on the study of larvae of two speciesof Pseudis. The limits and definitions ofPseudis and Lysapsus were reviewed by Sav-age and Carvalho (1953) and by Klappen-bach (1985). From their observations it is un-clear which character states support themonophyly of either genus. Savage and Car-valho (1953: 199) implicitly proposed theparaphyly of Pseudis, when they suggestedthat Lysapsus ‘‘seems to have arisen fromPseudis.’’ Garda et al. (2004) recently distin-guished both genera on the basis of spermmorphology. In Lysapsus laevis (the onlyspecies of Lysapsus available to them) thesubacrosomal cone is nearly absent, but it isclearly present in the four species of Pseudisthey studied. Regardless, the monophyly ofeither genus has not been satisfactorily doc-umented.
Morphological diversity within Pseudis in-cludes large species, several of which wereincluded in the past in the synonymy of P.paradoxa and were recently resurrected (Car-amaschi and Cruz, 1998), and smaller spe-cies with a double vocal sac, P. cardosoi andP. minuta (Klappenbach, 1985; Kwet, 2000).Lysapsus includes three species, L. caraya,
L. laevis and L. limellum; we include in ouranalysis the last two. Pseudis is composed ofsix species: P. bolbodactyla, P. cardosoi, P.fusca, P. minuta, P. paradoxa, and P. tocan-tins, of which we include in our analysis P.minuta and P. paradoxa.
Scarthyla: Duellman and de Sa (1988) andDuellman and Wiens (1992) suggested thatthis monotypic genus was sister to Scinax,but more recently, Darst and Cannatella(2003) presented evidence supporting a sistergroup relationship between Scarthyla and‘‘pseudids’’. The single species, Scarthylagoinorum, is included in our analysis.
Scinax: With roughly 86 recognized spe-cies, Scinax is the second largest genus with-in Hylinae. This genus includes the speciesformerly placed in the Hyla catharinae andH. rubra groups; a taxonomic history waspresented by Faivovich (2002). The relation-ships among the species of Scinax were re-cently addressed by Faivovich (2002), whoperformed a phylogenetic analysis using 38species representing the five species groupsthen recognized. Although he employedeight outgroups, the analysis is not a strongtest of the monophyly of Scinax nor of therelationships of Scinax with other hylines.Duellman and Wiens (1992) suggested thatScinax is the sister group of Scarthyla andthat this clade is sister to Sphaenorhynchus.Faivovich (2002) did not test this assertionbecause his selection of outgroups washeavily influenced by da Silva’s results(1998), which did not suggest a close rela-tionship between Scinax and these two gen-era. Taxon choice in the present study willtest more appropriately the hypothesis of(Scinax 1 Scarthyla) 1 Sphaenorhynchus.
Faivovich’s (2002) results suggested thatScinax contains two major clades: (1) a S.ruber clade composed of species that hadbeen previously grouped into the S. rostra-tus, S. ruber, and S. staufferi groups; and (2)a S. catharinae clade composed of the spe-cies that were included in the S. catharinaeand S. perpusillus groups. Faivovich (2002)continued recognition of these two speciesgroups within the S. catharinae clade, as wellas the S. rostratus group within the S. ruberclade, as the individual monophyly of the S.catharinae and S. rostratus groups were cor-roborated by his analysis. The S. perpusillus
2005 43FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
group is recognized because its monophylycould not be tested, and it still awaits a rig-orous test. All species previously included inthe nonmonophyletic groups of S. ruber andS. staufferi are included in the larger S. ruberclade, without being assigned to any group.For a list of the species currently included inScinax, see page 95.
In anticipation of a forthcoming study ofthe phylogeny of Scinax by Faivovich andassociates, we include only S. berthae and S.catharinae as exemplars of the S. catharinaeclade, and S. acuminatus, S. boulengeri, S.elaeochrous, S. staufferi, S. fuscovarius, S.ruber, S. squalirostris, and S. nasicus as ex-emplars of the S. ruber clade.
Sphaenorhynchus: More has been writtenabout nomenclatural confusion surroundingSphaenorhynchus than about its systematics(see Frost, 2004). This genus has been re-viewed by Caramaschi (1989). Duellman andWiens (1992) proposed the following syna-pomorphies for Sphaenorhynchus: posteriorramus of pterygoid absent; zygomatic ramusof squamosal absent or reduced to a smallknob; pars facialis of maxilla and alary pro-cess of premaxilla reduced; postorbital pro-cess of maxilla reduced, not in contact withquadratojugal; neopalatine reduced to a sliveror absent; pars externa plectri entering tym-panic ring posteriorly (rather than dorsally);pars externa plectri round; hyale curved me-dially; coracoids and clavicle elongated;transverse process of presacral vertebra IVelongate, oriented posteriorly; and prepollexossified, bladelike. The genus is composed of11 species: S. bromelicola, S. carneus, S.dorisae, S. lacteus, S. orophilus, S. palustris,S. pauloalvini, S. planicola, S. prasinus, S.platycephalus, and S. surdus. In our analysiswe include S. dorisae and S. lacteus.
Xenohyla: This genus was named by Iz-ecksohn (1996) for the bizarre frog Hylatruncata, which had previously been sug-gested to be related to Sphaenorhynchus byIzecksohn (1959, 1996) and Lutz (1973). Ac-cording to Izecksohn (1996), Xenohylashares with Sphaenorhynchus the reducednumber of maxillary teeth, a relatively shorturostyle, and the development of the trans-verse processes of presacral vertebra IV; fur-thermore, Xenohyla shares with Sphaenor-hynchus the quadratojugal not in contact with
the maxilla. Izecksohn (1996) suggested alsoa close relationship with Scinax based on thepresence in Xenohyla of a coracoid ridge andan internal, subgular vocal sac. While thecoracoid ridge is present in Scinax, it is alsopresent in several other hylines (e.g., see Fai-vovich, 2002). The internal, subgular vocalsac is not a synapomorphy of all Scinax, butonly of the S. catharinae clade. Caramaschi(1998) added X. eugenioi, a second speciesfor the genus. We include X. truncata in ourstudy.
CHARACTER SAMPLING
GENE SELECTION
Because this study involves the simulta-neous analysis of taxa of disparate levels ofdivergence, we assembled a large data set,including four mitochondrial and five nucleargenes, spanning a broad range of variation,from the fast-evolving cytochrome b (Gray-beal, 1993) to the much conserved nucleargenes such as 28S (Hillis and Dixon, 1991).
Ribosomal mitochondrial genes and cyto-chrome b have been employed recently inseveral phylogenetic studies of various an-uran groups at various levels of divergence(Read et al., 2001; Vences and Glaw, 2001;Cunningham, 2002; Salducci et al., 2002).Nuclear genes have been poorly explored fortheir use in anuran phylogenetics. The 28Sribosomal nuclear gene has been used in am-phibians by Hillis et al. (1993). The protein-coding genes rhodopsin, tyrosinase, RAG-1,and RAG-2 were used to study problems atdifferent levels by Bossuyt and Milinkovitch(2000), Biju and Bossuyt (2003), and Hoegget al. (2004). In this study we include 12S,tRNA valine, 16S, and fragments of cyto-chrome b, rhodopsin, tyrosinase, 28S, RAG-1, and seventh in absentia. The last gene isused here for the first time in amphibians.
DNA ISOLATION AND SEQUENCING
Whole cellular DNA was extracted fromfrozen and ethanol-preserved tissues (usuallyliver or muscle) using either phenol-chloro-form extraction methods or the DNeasy(QIAGEN) isolation kit. See table 2 for a listand sources of the primers employed.
Amplification was carried out in a 25-ml-
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TABLE 2Primers Used in this Study
Primer Sequence Reference
MVZ59 59-ATAGCACTGAAAAYGCTDAGATG-39 Graybeal (1997)12L1 59-AAAAAGCTTCAAACTGGGATTAGATACCCCACTAT-39 Feller and Hedges (1998)12SM 59-GGCAAGTCGTAACATGGTAAG-39 Darst and Cannatella (2004)MVZ50 59-TYTCGGTGTAAGYGARAKGCTT-39 Graybeal (1997)12sL13 59-TTAGAAGAGGCAAGTCGTAACATGGTA-39 Feller and Hedges (1998)16sTitus I 59-GGTGGCTGCTTTTAGGCC-39 Titus and Larson (1996)16sL2A 59-CCAAACGAGCCTAGTGATAGCTGGTT-39 Hedges (1994)16sH10 59-TGATTACGCTACCTTTGCACGGT-39 Hedges (1994)16sAR 59-CGCCTGTTTATCAAAAACAT-39 Palumbi et al. (1991)16sBR 59-CCGGTCTGAACTCAGATCACGT-39 Palumbi et al. (1991)16sWilk2 59-GACCTGGATTACTCCGGTCTGA-39 Wilkinson et al. (1996)MVZ15 59-GAACTAATGGCCCACACWWTACGNAA-39 Moritz et al. (1992)H15149(H) 59-AAACTGCAGCCCCTCAGAAATGATATTTGTCCTCA-39 Kocher at al. (1989)Rhod1A 59-ACCATGAACGGAACAGAAGGYCC-39 Bossuyt and Milinkovitch (2000)Rhod1C 59-CCAAGGGTAGCGAAGAARCCTTC-39 Bossuyt and Milinkovitch (2000)Rhod1Da 59-GTAGCGAAGAARCCTTCAAMGTA-39 Bossuyt and Milinkovitch (2000)R1-GFF 59-GAGAAGTCTACAAAAAVGGCAAAG-39 Taran Grant and Julian FaivovichR1-GFR 59-GAAGCGCCTGAACAGTTTATTAC-39 Taran Grant and Julian FaivovichTyr 1C 59-GGCAGAGGAWCRTGCCAAGATGT-39 Bossuyt and Milinkovitch (2000)Tyr 1G 59-TGCTGGGCRTCTCTCCARTCCCA-39 Bossuyt and Milinkovitch (2000)sia1b 59-TCGAGTGCCCCGTGTGYTTYGAYTA-39 Bonacum et al. (2001)sia2 59-GAAGTGGAAGCCGAAGCAGSWYTGCATCAT-39 Bonacum et al. (2001)28SV 59-AAGGTAGCCAAATGCCTCGTCATC-39 Hillis and Dixon (1991)28SJJ 59-AGTAGGGTAAAACTAACCT-39 Hillis and Dixon (1991)
a This primer, instead of Rhod1C, was used to amplify this gene in the 30-chromosome Hyla.b The primer pair sia1-2 was used together with the universal primers T3 and T7, as done by Bonacum et al. (2001).
volume reaction using either puRe TaqReady-To-Go PCR beads (Amersham Bio-sciences, Piscataway, NJ) or InvitrogenePCR SuperMix. For all the amplifications,the PCR program included an initial dena-turing step of 30 seconds at 948C, followedby 35 or 38 cycles of amplification (948C for30 seconds, 48–608C for 60 seconds, 728Cfor 60 seconds), with a final extension stepat 728C for 6 min.
Polymerase chain reaction (PCR)-ampli-fied products were cleaned either with aQIAquick PCR purification kit (QIAGEN,Valencia, CA) or with ARRAY-IT (Tele-Chem International, Sunnyvale, CA) and la-beled with fluorescent-dye labels terminators(ABI Prism Big Dye Terminators v. 3.0 cyclesequencing kits; Applied Biosystems, FosterCity, CA). Depending on whether thecleaned product was purified with QIAquickor Array-It, the sequencing reaction was car-ried out in either 10 ml or 8 ml volume re-action following standard protocols. The la-
beled PCR products were isopropanol-pre-cipitated following the manufacturer’s pro-tocol. The products were sequenced eitherwith an ABI 3700 or with an ABI Prism 377sequencer. Most samples were sequenced inboth directions.
Chromatograms obtained from the auto-mated sequencer were read and contigs madeusing the sequence editing software Se-quencher 3.0. (Gene Codes, Ann Arbor, MI).Complete sequences were edited with Bio-Edit (Hall, 1999).
MORPHOLOGY
Because the present study is mostly basedon molecular data, the failure to include athorough morphological data set doubtless isits weakest point. As trained morphologists,most of the authors of this paper think that aphylogenetic hypothesis that explains all theavailable data is the best hypothesis that wecan aspire to, and that no class of data is
2005 45FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
better than any other. Apart from da Silva’sunpublished dissertation, which is comment-ed upon below, published comparative stud-ies involving a diversity of hylid exemplarsare rare. Major exceptions are the thoroughosteological studies by Trueb (1970a) andthose on hand muscles of Pelodryadinae(Burton, 1996), distal extensor muscles ofanurans (Burton, 1998a), and foot muscles ofHylidae (Burton, 2004). Explicit characterdescriptions in the context of phylogeneticcomparisons include those by Duellman andTrueb (1983), Campbell and Smith (1992),Duellman and Campbell (1992), Duellmanand Wiens (1992), Fabrezi and Lavilla(1992), Kaplan (1994, 1999), Cocroft (1994),Burton (1996, 1998a, 2004), Haas (1996,2003), da Silva (1997), Duellman et al.(1997), Kaplan and Ruiz-Carranza (1997),Mendelson et al. (2000), Sheil et al. (2001),Faivovich (2002), and Alcalde and Rosset(‘‘2003’’ [2004]). Most of these studies weretargeted in general to very specific apparentclades or to very large clades using very fewterminals, which leaves particular sets ofcharacters known for very few terminals.Unfortunately, for the inclusion of the char-acters employed in these studies to be infor-mative, detailed anatomic work would be re-quired on a very large number of terminals(besides the potentially serious need to re-define several characters), a task that we findimpossible to pursue at this time. Much toour regret, we find that there are almost nopublished studies from which we could de-rive character scorings to enrich our data setwithout extensive work. The only data setthat we thought could be included, due to itsrelatively dense taxon sampling, is the oneresulting from the collection of observationspresented by Burton (2004). Although itssampling of nonhylid taxa that match oursampled taxa is particularly sparse, we con-sider Burton’s study to be an important ad-dition to this analysis. Characters are listedand discussed in appendix 3.
da Silva’s (1998) Dissertation
da Silva (1998) presented his Ph.D. dis-sertation on phylogeny of hylids with em-phasis on Hylinae. Although da Silva’s dis-sertation has not been published, some of its
results and conclusions were described andcommented in detail by Duellman (2001).Because the present paper deals specificallywith the phylogeny of Hylinae, we cannotavoid a few comments dealing with da Sil-va’s work. Considering the mostly coincidentscope of both da Silva’s dissertation and thispaper, it is evident that a thorough discussionand comparison of his results with ourswould almost amount to the publication ofhis chapter on Hylinae relationships. This isa situation with which we feel most uncom-fortable, because we think that this is a re-sponsibility that rests on Helio R. da Silva.
From a purely practical perspective, at thispoint the integration of da Silva’s data setwith ours is impracticable for two reasons:(1) The data matrix as printed in the disser-tation distributed by the University of Mich-igan is incomplete, as it lacks the scoringsfor 10 characters (chars. 110–120) for alltaxa. This is also the situation with the thesisthat is deposited at the Department of Her-petology library of the University of Kansas,Natural History Museum (Faivovich, person-al obs.). (2) A few scorings for groups thatwe are familiar with are not coincident withour observations on the same species, some-thing suggestive either of polymorphism inthose characters or mistaken scorings.21 Ifthis were the case, it would not be surprising,as scoring mistakes are to be expected insuch an impressive data set. The problemwith them is that once detected, they have tobe corrected and the analysis has to be re-done. It is evident that a revision of the dataset is necessary before any integration cantake place.
PHYLOGENETIC ANALYSIS
Our optimality criterion to choose amongtrees is parsimony. The logical basis of par-simony as an optimality criterion has beenpresented by Farris (1983). However, parsi-
21 For example, character 60 (anterior process of thehyale) is scored 0 (absent) in Aplastodiscus, where it ispresent in the material available to us (Faivovich, 2002;Garcia, personal obs.). Character 61 (anterolateral pro-cess of the hyoid plate) is scored 0 (absent) in Hylaalbofrenata, H. albomarginata, H. albopunctata, H. al-bosignata, H. faber, and H. multifasciata, whereas it ispresent in the specimens available to us (Garcia and Fai-vovich, personal obs.)
46 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
mony has repeatedly been attacked from dif-ferent perspectives, all of which tend to por-tray parsimony as inferior to such model-based approaches as maximum likelihood.Criticisms of parsimony have centered ontwo main topics: statistical inconsistency andthe notion that parsimony is an overpara-meterized likelihood model. As stated by Go-loboff (2003), the emphasis on statisticalconsistency decreased following severalstudies showing that: (1) maximum likeli-hood can be inconsistent even with minor vi-olations of the model when they were gen-erated with a mix of models (Chang, 1996);(2) given some evolutionary models, maxi-mum likelihood estimators could be incon-sistent (Steel et al., 1994; Farris, 1999); (3)parsimony can be consistent (Steel et al.,1993); (4) assuming likelihood as a more ac-curate method, inferences based on trees sub-optimal under the maximum likelihood couldbe less reliable than inferences made on treesoptimal under otherwise inferior but fastercriteria (Sanderson and Kim, 2000); and (5)at least under some conditions, parsimonymay be more likely than maximum likeli-hood to find the correct tree, given finiteamounts of data (Yang, 1997; Siddall; 1998;Pol and Siddall, 2001). Tuffley and Steel(1997) demonstrated that parsimony is amaximum likelihood estimator when eachsite has its own branch length. Farris (1999,2000) and Siddall and Kluge (1999) sug-gested that the results of Tuffley and Steel(1997) were an indication that the model im-plied by parsimony (‘‘no special model ofevolution’’ or ‘‘no common mechanism mod-el’’) was indeed more realistic. However,likelihood advocates (Steel and Penny, 2000;Lewis, 2001; Steel, 2002) countered thatmodels that assume constant probabilities ofchange across all sites are to be preferred onthe grounds of simplicity (i.e., as having few-er parameters to estimate). Goloboff (2003)demonstrated that parsimony could actuallybe derived from models that require evenfewer parameters than the commonly usedlikelihood models.
The use of Bayesian Markov chain MonteCarlo (BMCMC) techniques has becomequite popular among evolutionary biologists.However, for reasons outlined by Simmonset al. (2004), the posterior probability values
of the clades cannot be interpreted as valuesof truth or support. Furthermore, Kola-czkowski and Thornton (2004) demonstrat-ed, by using simulations in the presence ofheterogeneous data, that parsimony performsbetter than both maximum likelihood andBMCMC over a wide range of conditions.
We contend that all serious criticisms ofparsimony have been rebutted. We considerthat while the first point mentioned (incon-sistency of likelihood when the data are gen-erated with different models) could certainlyoccur in any analysis, it is particularly prob-lematic in the present one, because we arecombining morphology with both mitochon-drial and nuclear coding and non-codinggenes. Furthermore, for a data set of this size,maximum likelihood is quite impractical toapply for computational reasons.
For the phylogenetic analyses of the DNAsequence data, we used the method of DirectOptimization (Wheeler, 1996, 1998, 2002),as implemented in the program POY (Wheel-er et al., 2002), a heuristic approximation tothe optimal tree alignment methods of San-koff (1975) and Sankoff and Cedergren(1983). Sequence alignment and tree search-ing have traditionally been treated as two in-dependent steps in phylogenetic analyses: se-quences are first aligned, and a fixed or staticmultiple alignment is then treated as a stan-dard character matrix that is the basis for treesearching in the test of character congruence.However, there may be other equally defen-sible multiple sequence alignments thatwould require fewer hypothesized transfor-mations to explain the observed sequencevariation; an explanation that requires fewertransformations is more parsimonious and istherefore objectively preferred over expla-nations that require a greater number oftransformations (see De Laet [2005] for amuch more sophisticated approach to theproblems of constructing multiple alignmentsprior to tree searching). Direct Optimizationseeks the cladogram-alignment combination(i.e., the optimal tree alignment) that mini-mizes the total number of hypothesizedtransformation events required to explain theobservations. Within this framework, inser-tion/deletion events (indels, gaps) are histor-ical evidence that is taken into account whenhypothesizing common ancestry.
2005 47FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
The simplest minimization of transforma-tions is obtained when tree searches are con-ducted under equal weights for indels and allsubstitutions (1:1:1, this is the ratio of thecost of opening gap:extension gap:substitu-tions) (Frost et al., 2001). This weightingscheme implies that indels are as costly asthe number of nucleotides they span. This isnot a situation with which we are comfort-able, inasmuch as a single deletion eventcould entail more than a single nucleotideand hence necessarily require a lower costthan if all the nucleotides it includes werelost independently of each other. However,theoretical justifications for the selection ofdifferential costs for gap opening and gap ex-tension are not evident.
De Laet and Smets (1998) suggested thatparsimony analysis searches for the trees onwhich the highest number of compatible in-dependent pairwise similarities can be ac-commodated; that is, they described parsi-mony as a two-taxon analysis. When dealingwith static data sets, this approach and theminimization of transformations give thesame rank of tress. However, De Laet (2005)showed that when considering parsimony asa two-taxon analysis in the presence of in-applicable character states (e.g., unequal-length sequences), the minimization of trans-formations (as obtained under 1:1:1) does notmaximize the number of accommodatedcompatible independent pairwise a priorisimilarities. De Laet (2005) suggested thatsequence homology has two components, ho-mology of subsequences (the fragments ofsequences that are comparable across abranch) and base-to-base homology withinhomologous subsequences. When maximi-zation of homology is transformed into aproblem of minimization of changes, the op-timization of the two components that max-imizes the accommodated independent pair-wise similarities is obtained by summing upthe cost regimes that are involved for eachcomponent. The number of subsequences isquantified by counting the number of inser-tion/deletion events (independent of theirlength, and therefore represented each as awhole by a unit opening gap). Base-to-basehomology within homologous subsequencesis maximized when substitutions are weight-ed twice as much as unit gaps (Smith et al.,
1981). These result in a substitution cost of2, a gap opening cost of 2 1 1 (the samecost of a substitution plus the cost of the firstunit gap), and a gap extension cost of 1. Allthis development rests on the perspective ofparsimony as a two-taxon analysis (De Laetand Smets, 1998). The most immediately ap-pealing aspect of De Laet’s perspective isthat it offers a rationale for the use of gap-extension costs different from substitutioncosts, thus avoiding giving an insertion/de-letion event of n nucleotides the same weightof n substitutions.
We conducted our searches using equalweights for minimizing transformations. Inorder to examine the effect of the gap treat-ment in our results, and following De Laet’sdevelopment (2005), we also submitted ourfinal tree to a round of tree-bisection and re-connection branch swapping (TBR) by usinga weighting scheme of 2 for substitutions andmorphological transformations, 3 for a gapopening, and 1 for a gap extension.
This study is guided by the idea that a si-multaneous analysis of all available evidencemaximizes explanatory power (Kluge, 1989;Nixon and Carpenter, 1996). Consequently,we analyzed all molecular and available mor-phological evidence simultaneously. Theanalysis was performed using subclusters of60–100 processors of the American Museumof Natural History parallel computer cluster.
Heuristic algorithms applied to both treesearching and length calculation (i.e., align-ment cost) were employed throughout theanalysis. As with any heuristic solution, theoptimal solution from these analyses underDirect Optimization represents the upperbound, and more exhaustive searching couldresult in an improved solution. Consideringthe large size of our data set, we tried twodifferent approaches. The first strategy triesto collect many locally optimal trees frommany replications to input them into a finalround of tree fusing (Goloboff, 1999). Forthe second strategy, quick concensus esti-mates (Goloboff and Farris, 2001) are usedas constraints for additional tree searching,following the suggestion of Pablo Goloboff(personal commun.).
For maximizing the number of trees fortree fusing, we employed two different rou-tines:
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1. Three hundred fifty random addition se-quences were done in groups of 5 or 10,followed by a round of tree fusing, sendingthe best tree to 10–25 parsimony ratchet cy-cles (Nixon, 1999a) using TBR, reweightingbetween 15 and 35% of the fragments, keep-ing one tree per cycle, and by setting thecharacter weight multiplier between two andfive in different replicates, with a final roundof TBR branch swapping. Tree fusing wasalways done fusing sectors of at least threetaxa with two successive rounds of fusing.
2. One hundred fifty random addition sequenc-es were built in groups of 5, 7, or 10 bysubmitting the best of each group to 10–25ratchet cycles using TBR.
The 40 best trees resulting from theseanalyses where submitted to tree fusing ingroups of five, and the resulting eight treeswere subsequently fused. This final tree wassubmitted to 30 replicates of Ratchet usingthe same settings as above, with the resultingtrees being submitted to a final round of TBRbranch swapping.
Alternatively, we did 50 random additionsequences followed by a round of TBR andmade an 85% majority rule consensus, assuggested by Goloboff and Farris (2001) toquickly estimate the groups actually presentin the consensus of large data sets withouthaving to do intensive searches. The ap-proach of Goloboff and Farris (2001) as-sumes that groups that are present in all ormost independent searches are more likely tobe actually supported by the data. To speedup the searches for the estimation of thequick consensus, we treated the partial se-quences of the RAG-1, rhodopsin, SIA, andtyrosinase genes as prealigned. Once thequick consensus was estimated, it was input-ted in POY as a constraint file, with whichwe built 100 Wagner trees, each followed by10 ratchet replicates. All trees resulting fromthese constrained searches were fused ingroups of different size, and the final treeswere submitted to a round of TBR. The orig-inal constraint file was not used during thefusing and final TBR steps.
While all searches were done using stan-dard direct optimization, all were submittedto final rounds of TBR under the command‘‘iterative pass’’ (Wheeler, 2003a). This rou-tine does a three-dimensional optimization,taking into account the states of the three ad-
jacent nodes of the internal node of interest.Because any change in the reconstructed se-quence could potentially affect adjacentnodes, the procedure is done iteratively untilstabilization is achieved.
The large size of the data set imposes aheavy burden in computer times to estimatesupport measures. Bremer supports (Bremer,1988) were calculated using POY, withoutusing ‘‘iterative pass’’. Parsimony Jackknifevalues (Farris et al., 1996) were calculatedusing the implied alignment (Wheeler,2003b) of the best topology. In turn, this im-plies that the parsimony jackknife valuescould be overestimated. Parsimony Jackknifewas calculated in TNT (Goloboff et al.,2000); 1000 pseudoreplicates were per-formed. For each pseudoreplicate the best to-pology was searched for by using sectorialsearches and tree fusing, starting with twoWagner trees generated through random ad-dition sequences.
Final tree lengths under the 1:1:1 weight-ing scheme were checked with TNT. Lists ofsynapomorphies were generated with TNT;only unambiguous transformations commonto all most parsimonious trees were consid-ered.
For the analysis, the complete 12S-tRNAvaline-16S sequence was cut into 14 frag-ments and the partial 28S sequence was cutinto 4 fragments coincident with conservedregions (Giribet, 2001). Although this con-strains homology assessment, the universe ofalternative ancestral sequences that has to beexplored is a more tractable problem than us-ing long single fragments. The sequence filesas they were input into POY are availablefrom http://research.amnh.org/users/julian.Tree editing was done using WinClada (Nix-on, 1999b).
RESULTS
In total, we sequenced 256 terminals. Thecontiguous 12S, tRNA valine, and 16S geneswere sequenced for all but seven terminals.For these terminals we were unable to am-plify or sequence one or two of the overlap-ping PCR fragments. The partial cytochromeb fragment was sequenced for all but 12 ter-minals. The success with the nuclear locivaried from 232 terminals sequenced for the
2005 49FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
first exon of rhodopsin to as few as 166 se-quenced for 28S. See appendix 2 for a com-plete list of the loci sequenced for each tax-on, voucher specimens, locality data, andGenBank accessions. All sequences wereproduced for this project and for that of Fai-vovich et al. (2004) with the exception of 21sequences taken from GenBank that wereproduced by Bijou and Bossuyt (2003) andby Darst and Cannatella (2004). The fact thatthe morphological characters are not scoredfor 70% of the terminals led to several am-biguous optimizations. A list of nonambigu-ous morphological synapomorphies is pro-vided in appendix 3, many of which are men-tioned throughout the discussion and in thesection ‘‘Taxonomic Conclusions: A New Tax-onomy of Hylinae and Phyllomedusinae’’.
The phylogenetic analysis resulted in fourmost parsimonious trees of 65,717 steps. Oneof these trees resulted from one of the roundsof tree fusing of the trees resulting from theconstrained search, and the other three treeswere obtained after a round of TBR swap-ping of the first one. Parsimony Jackknifeand Bremer support values are generallyhigh. Most of the 272 nodes of the strict con-sensus (figs. 1–5) are well supported, with226 nodes having a Bremer support of $10and 162 nodes with a Bremer support of$20; additionally, 255 nodes have a jack-knife value of $75% and 245 nodes have ajackknife value of $90%.
All conflict among the trees is restricted totwo points: (1) the relationships among Hylacircumdata, H. hylax, and the undescribedspecies Hyla sp. 4 (fig. 3); and (2) the rela-tionships of H. femoralis with the H. versi-color and H. eximia groups (fig. 5).
When the best trees were submitted to around of TBR using a weighting scheme of3:1:2 as suggested by De Laet (in press) andmentioned earlier, the resulting tree differsfrom the original ones only in that (1) theclade composed of Cryptobatrachus and Ste-fania moves to be the sister group of Flec-tonotus and Gastrotheca (fig. 2), and (2) theclade composed of Lysapsus, Pseudis, andScarthyla moves from the sister taxon of Sci-nax to the sister taxon of the 30-chromosomeHyla groups, Sphaenorhynchus, and Xeno-hyla (fig. 4).
DISCUSSION
MAJOR PATTERNS OF RELATIONSHIPS OF
HYLIDAE AND OUTGROUPS
As in previous analyses (Ruvinsky andMaxson, 1996; Haas, 2003; Darst and Can-natella, 2004), our results do not recover Hy-lidae as a monophyletic taxon (figs. 1, 2).Hemiphractinae appears as only distantly re-lated to the Hylinae, Pelodryadinae, andPhyllomedusinae, each of which is mono-phyletic. For this reason, we exclude Hemi-phractinae from Hylidae, being thereby re-stricted to Hylinae, Pelodryadinae, and Phyl-lomedusinae. In the same way, Centroleni-dae, for a long time suspected to be relatedwith hylids, appears as a distantly relatedclade, as suggested by previous studies(Haas, 2003; Darst and Cannatella, 2004).
Hylidae, as understood here, excludes theHemiphractinae. Otherwise, the major cladeswithin the Hylidae are coincident with theremaining subfamilies currently recognized.The Pelodryadinae is the sister taxon ofPhyllomedusinae, corroborating the results ofDarst and Cannatella (2004); in turn, Pelo-dryadinae 1 Phyllomedusinae is the sistertaxon of Hylinae (figs. 1, 2).
Ranoids appear as monophyletic, with thetwo microhylid exemplars being the sistertaxon of the Astylosternidae 1 remainingranoids (fig. 2). Ranidae forms a paraphyleticmelange, with the exemplars of Hemisotidae,Mantellidae, and Rhacophoridae being nest-ed among the few ranid exemplars. Ranoidsare the sister taxon of all remaining terminals(fig. 2).
Within hyloids, as expected, Leptodacty-lidae is rampantly paraphyletic, with all otherincluded families nested within it (figs. 1, 2).Ceratophryinae, Eleutherodactylinae, Lepto-dactylinae, and Telmatobiinae are not mono-phyletic (fig. 2).
At the base of hyloids, the two exemplarsof Eleutherodactylus are the sister taxon of aclade composed of Hemiphractus helioi,Brachycephalus ephippium, and Phrynopussp. This situation renders Eleutherodactyli-nae and Hemiphractinae nonmonophyletic(fig. 2). The nonmonophyly of Hemiphrac-tinae is further given by the fact that in the1:1:1 analysis, Stefania 1 Cryptobatrachusand Gastrotheca 1 Flectonotus are not
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Fig. 1. A reduced image of the strict consensus of the four most parsimonious trees showing themajor patterns of relationships of the outgroups, hylid subfamilies, and the four major clades of Hylinaerecovered in the analysis.
monophyletic but occur as a grade leading tothe other hyloids (fig. 2); however, the groupis monophyletic in the 3:1:2 analysis. Mov-ing upward in the tree finds two large clades:one composed of Hylidae in the sense usedhere (i.e., excluding Hemiphractinae), andthe other composed of the remaining hyloidfamilies and subfamilies of Leptodactylidae.Leptodactylinae as defined by Laurent(1986) is not monophyletic in that Limno-medusa is only distantly related to the re-maining ‘‘Leptodactylinae’’, being the sistertaxon of Odontophrynus. Note that this ar-rangement is congruent with Leptodactylinaeas defined by Lynch (1971). Centrolenidaeobtains as monophyletic and as the sister tax-on of Allophrynidae. The only cycloram-phine exemplar, Crossodactylus schmidti, isthe sister taxon of the dendrobatid exemplars.
Telmatobiinae is not monophyletic forhaving one of the Ceratophryinae exemplars,Ceratophrys cranwelli, nested within it. Fur-thermore, the other two Telmatobiinae ex-emplars, Alsodes gargola and Euspsophuscalcaratus, form a clade with the Leptodac-tylinae exemplar Limnomedusa macroglossaand the other Ceratophryinae exemplarOdontophrynus americanus. This clade isalso the sister taxon of all Bufonidae exem-plars (fig. 2).
In general, most results concerning the re-lationships among outgroup taxa should beconsidered cautiously, because the taxonsampling of this analysis was not designedto address those specific questions. Some re-sults are nonetheless expected or at least sug-gestive. In the former group we include, forexample, the monophyly of Bufonidae, Den-drobatidae, Centrolenidae, and Ranoidea.The relationship between Crossodactylus anddendrobatids is consistent with the results ofHaas (2003).
The fact that hemiphractines are not relat-ed to the Hylidae, and are likely nonmono-phyletic, requires a change in how study ofthis group is approached. For instance, rela-tionships of Hemiphractinae as recovered
here are quite different from the results ofthe cladistic analysis based on morphologyand life-history data presented by Mendelsonet al. (2000). These authors found Hemi-phractus to be nested within Gastrotheca, aresult that Duellman (2001) considered im-plausible. Although we did not incorporatetheir data set into our analysis, the fact thatthey employed only hylid outgroups couldhave affected their results. With the impor-tant difference that we do not recover amonophyletic Hemipractinae, our results cor-roborate the sister taxon relationship betweenthe northern Andean Cryptobatrachus andthe Guayanan Stefania, as well as the sistergroup relationship between Flectonotus andGastrotheca, as suggested by Duellman andHoogmoed (1984) and Wassersug and Duell-man (1984). Regarding the monophyly andactual position of Hemiphractinae withinNeobatrachia, our results are inconclusive(similar to those of Darst and Cannatella,2004) most likely because of a lack of theappropriate taxon sampling to address theproblem. Their positions in the tree suggestthat a much denser taxon sampling of ‘‘Lep-todactylidae’’ and perhaps Eleutherodactyli-nae will be necessary to better understandtheir relationships.
PELODRYADINAE AND PHYLLOMEDUSINAE
Our results of a monophyletic Pelodryadi-nae 1 Phyllomedusinae corroborate earlysuggestions by Trewavas (1933), Duellman(1970), Bagnara and Ferris (1975), and morerecent results by Darst and Cannatella (2004)and Hoegg et al. (2004). The monophyly ofPelodryadinae and Phyllomedusinae was notrecovered in the analyses by Duellman(2001), Burton (2004), and Haas (2003). Dis-crepancies between the analyses of Burton(2004) and Haas (2003) and our’s may be theresult of different taxon sampling and as-sumptions. Duellman (2001) assumed thatHylidae, in the classical sense (includingHemiphractinae), was monophyletic, and
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Fig. 2. A partial view of the strict consensus showing the relationships of the outgroups, Pelodry-adinae, and Phyllomedusinae. Numbers above nodes are Bremer support values. Numbers below nodesare Parsimony Jackknife absolute frequencies; those with an asterisk (*) have a 100% frequency. Num-bers in boldfaced italic are node numbers for the list of morphological synapomorphies (see appendix3). Black circles denote nodes that are present in the quick consensus estimation. The arrow showsalternative placement of the (Cryptobatrachus 1 Stefania) clade when using the 3:1:2 weighting scheme(see text).
Burton (2004) assumed that Hylidae, Centro-lenidae, and Allophrynidae formed a mono-phyletic group. Our analysis did not includethe morphological characters employed byHaas (2003) in his analysis, nor does our pe-lodryadine taxon sampling match his. Forthis reason, our results are not directly com-parable to his, and we do not consider themonophyly of the Pelodryadinae a settled is-sue.
In our analysis, the presence of a tendonof the m. flexor ossis metatarsi II arising onlyfrom distal tarsal 2–3 is a synapomorphy ofPelodryadinae plus Phyllomedusinae. Fur-thermore, the presence of the pigment pter-orhodin (Bagnara and Ferris, 1975) may bea synapomorphy of this clade, although thedistribution of this character state requiresfurther elucidation. Both Pelodryadinae andPhyllomedusinae share the presence of sup-plementary elements of the m. intermandi-bularis. These elements are apical in Pelod-ryadinae and posterolateral in Phyllomedu-sinae (Tyler, 1971). Both character stateshave previously been considered nonhomo-logs (Tyler and Davies, 1978a) that separate-ly support the monophyly of each of thesegroups (Duellman, 2001). In the context ofour analysis, however, the sole presence ofsupplementary elements is more parsimoni-ously interpreted as a putative synapomorphyof this clade, while it is ambiguous which ofthe positions of the elements (apical or pos-terolateral) is the plesiomorphic state. Notethat this ambiguity is a potential challenge tothe only known morphological synapomor-phy of Pelodryadinae. Future anatomicalwork will corroborate whether these twomorphologies could be considered as statesof the same transformation series, as is ten-tatively being done here.
PELODRYADINAE
As stated previously, our analysis does notinclude enough of a comprehensive taxonsampling of Pelodryadinae to address its in-ternal relationships in a meaningful way.Nonetheless, our results corroborate thelong-held idea (see previous discussions) thatCyclorana and Nyctimystes are nested withinLitoria. For the reasons detailed earlier, ouranalysis is not an overly strong test of thepositions of the former two genera within Li-toria. Nevertheless, the single exemplar ofCyclorana is the sister taxon of L. aurea, anexemplar of the L. aurea group with whichCyclorana is supposed to be related based onvarious sources of evidence (King et al.,1979; Tyler, 1979; Tyler et al., 1981). Nyc-timystes is the sister taxon of L. infrafrenata,one of the groups of Litoria that Tyler andDavies (1979) considered as possibly relatedto Nyctimystes based on morphological sim-ilarities of its skull with that of N. zweifeli.The other groups they considered are mostlythe montane species of Litoria; from thesewe included a single exemplar, L. arfakiana,that is quite distant from Nyctimystes, beingthe sister taxon of L. meiriana (with which,incidentally, it also shares the presence of aflange in the medial surface of metacarpalIII; Tyler and Davies, 1978b). In our analy-sis, the fibrous origin of the m. extensorbrevis superficialis digiti III on the distal endof the fibulare is a synapomorphy of Pelod-ryadinae; we are skeptical, however, that thisoptimization will hold with better sampledoutgroups for muscular characters, becauseBurton (2004) found the same character statein several leptodactylids, none of which isincluded in our outgroup sample.
PHYLLOMEDUSINAE
Several authors (Funkhouser, 1957; Duell-man, 1970; Donnelly et al., 1987; Hoogmoed
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and Cadle, 1991) noticed the distinctivenessof Agalychnis calcarifer and its presumedsister taxon, A. craspedopus, from the otherspecies of Agalychnis. Corroborating the re-sults of Duellman (2001), we found no evi-dence for the monophyly of Agalychnis. Ourresults indicate that A. calcarifer is the sistergroup of the remaining Phyllomedusinae,and it has no close relationship with the otherexemplars of Agalychnis.
Phyllomedusa lemur, the only exemplar ofthe P. buckleyi group available for this anal-ysis, is recovered, although with low Bremersupport (3), as the sister group of Hyloman-tis, and it is only distantly related with theother exemplars of Phyllomedusa. This situ-ation corroborates previous suggestions(Funkhouser, 1957; Cannatella, 1980; Jung-fer and Weygoldt, 1994) that the P. buckleyigroup should not be included in Phyllome-dusa. Cruz (‘‘1988’’ [1989]) suggested, onthe basis of iris coloration, skin texture, poordevelopment of webbing, and slender body,that Hylomantis is related to two species ofthe P. bukleyi group, P. buckleyi and P. psi-lopygion. On the basis of the same characterstates, Cruz (1990) associated the P. buckleyigroup with both Hylomantis and Phasmahy-la. Our results support these ideas only inpart, because while our only exemplars ofHylomantis and the P. buckleyi group areeach monophyletic, Phasmahyla is moreclosely related to Phyllomedusa (excludingthe P. buckleyi group).
We had no clear idea regarding the posi-tion of Hylomantis and Phasmahyla. Mor-phologically, the evidence is conflicting inthat each one shares at least one differentpossible synapomorphy with the restrictedPhyllomedusa (Phyllomedusa excluding theP. buckleyi group). Phasmahyla has the sametype of nest where the eggs are wrapped ina leaf; nests are unknown in Hylomantis, butspecies of this genus share with Phyllome-dusa (excluding the P. buckleyi group) thepresence of the slip of the m. depressor man-dibulae that originates from the dorsal fasciaat the level of the m. dorsalis scapulae (Cruz,1990; Duellman et al., 1988b), a characterstate that is absent in all other Phyllomedu-sinae. Our analysis recovers Phasmahyla asthe sister group of the restricted Phyllome-
dusa, suggesting that the eggs wrapped in aleaf are a synapomorphy of this clade.
MAJOR PATTERNS OF RELATIONSHIPS
WITHIN HYLINAE
For purposes of discussion, we considerHylinae to be composed of four major clades(fig. 1), called here: (1) the South Americanclade I; (2) South American clade II (SA-II);(3) Middle American/Holarctic clade; and (4)South American/West Indian Casque-headedFrogs. These major sections and their sub-clades will be discussed in this order.
SOUTH AMERICAN CLADE I
This clade is composed of all GladiatorFrogs, the Andean stream-breeding Hyla, thegenus Aplastodiscus, and a Tepuian clade ofHyla. It contains five major clades (fig. 3).The first of these is called the Tepuian clade,and is composed solely of two exemplars ofthe H. aromatica and H. geographicagroups. The second clade is composed of allAndean stream-breeding Hyla. The third iscomposed of all the exemplars of the H. cir-cumdata, H. martinsi, and H. pseudopseudisgroups, from southeastern Brazil, and we arecalling it informally the Atlantic/Cerradoclade. The fourth is composed of the south-eastern Brazilian H. albosignata and H. al-bofrenata complexes of the larger, nonmon-ophyletic H. albomarginata group plus thetwo species of Aplastodiscus, and we arecalling it informally the Green clade. Thefifth clade is composed of all the remainingspecies groups (H. geographica, H. pulchel-la, H. boans, H. granosa, H. punctata, H.albomarginata complex of the H. albomar-ginata group) and unassigned species asso-ciated in the past with the Gladiator Frogs,and we are calling it informally the TGFclade (for True Gladiator Frogs.)
Six currently recognized species groupswithin the South American clade I are notmonophyletic. The Hyla albomarginatagroup is not monophyletic because its three‘‘complexes’’ defined by Cruz and Peixoto(‘‘1985’’ [1987]) are spread throughout theGreen clade and the TGF clade. The H. al-bomarginata complex is not monophyletic,with its species being related with differentgroups in the TGF clade (see below). The H.
2005 55FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Fig. 3. A partial view of the strict consensus showing the relationships of the South American Iclade and its correspondence with the currently recognized species groups. Numbers above nodes areBremer support values. Numbers below nodes are Parsimony Jackknife absolute frequencies; those withan asterisk (*) have a 100% frequency. Black circles denote nodes that are present in the quick consensusestimation.
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Fig. 4. A partial view of the strict consensus showing the relationships of the South American IIclade and its correspondence with the currently recognized species groups. Numbers above nodes areBremer support values. Numbers below nodes are Parsimony Jackknife absolute frequencies; those withan asterisk (*) have a 100% frequency. Numbers in boldfaced italic are node numbers for the list ofmorphological synapomorphies (see appendix 3). Black circles denote nodes that are present in the quickconsensus estimation. The arrow shows alternative placement of the (Scarthyla 1 (Lysapsus 1 Pseudis))clade when using the 3:1:2 weighting scheme (see text).
albosignata complex is monophyletic, as isthe H. albofrenata complex. These two com-plexes, however, do not form a monophyleticgroup, because Aplastodiscus is the sistergroup to the H. albosignata complex, andthis clade is sister to the H. albofrenata com-plex. Within the TGF Clade, the only groupthat is not represented by a single exemplar(the Hyla punctata group) that results asmonophyletic is the H. pulchella group. TheH. albomarginata complex, H. albopunctata,H. boans, H. geographica, and H. granosagroups are nonmonophyletic.
Hyla pellucens and H. rufitela are the sis-
ter group of the clade composed of H. heil-prini and the paraphyletic H. albopunctatagroup (see below); H. albomarginata is thesister taxon of a fragment of the H. boansgroup (see below). The H. albopunctatagroup is paraphyletic inasmuch as H. fasciataplus H. calcarata is nested within it. The H.boans group is polyphyletic because themostly southeastern Brazil/northeastern Ar-gentina exemplars (H. faber, H. lundii, H.pardalis, H. crepitans, and H. albomargina-ta) together with H. albomarginata are thesister taxon of the H. pulchella group and areonly distantly related to H. boans. Hyla
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boans is the sister taxon of H. geographicaplus H. semilineata. The H. geographicagroup is rampantly polyphyletic, with its ex-emplars partitioned into five different cladeswithin the South American clade I: (1) H.kanaima is the sister taxon of H. inparquesi,the single exemplar of the H. aromaticagroup; (2) H. roraima and H. microdermaform a monophyletic group with four Guay-anese and one Amazonian species; (3) Hylapicturata is related to the exemplars of theH. punctata and H. granosa groups; (4) Hylasemilineata 1 H. geographica are related toH. boans; and (5) H. fasciata 1 H. calcarataare nested within the H. albopunctata groupas detailed above. The H. granosa group isparaphyletic by having H. picturata and H.punctata nested within it.
Andean Stream-Breeding Hyla and the Te-puian Clade
The monophyly of the Andean stream-breeding Hyla is congruent with suggestionspresented by Duellman et al. (1997) and Mi-jares-Urrutia (1997), who noticed similaritiesin larval morphology of the H. bogotensisand H. larinopygion groups. Duellman et al.(1997) presented a phylogenetic analysis re-stricted to wholly or partially Andean speciesgroups of Hyla. In their most parsimonioustree, the H. armata, H. bogotensis, and H.larinopygion groups formed a monophyleticgroup supported by three transformations intadpole morphology: the enlarged, ventrallyoriented oral disc; the complete marginal pa-pillae; and a labial tooth row formula 4/6 orhigher.
Duellman et al. (1997) suggested a closerelationship between the H. armata and H.larinopygion groups based on the presencein males of a greatly enlarged prepollex lack-ing a projecting spine. While our results arecongruent with this, note that males of the H.bogotensis group also have a prepollex withthe same external morphology as those of theH. armata and H. larinopygion groups. Ki-zirian et al. (2003) suggested that the H. ar-mata group was nested in the H. larinopy-gion group. Our results do not support thissuggestion. However, this could be a conse-quence of the few exemplars of the H. lari-nopygion group available for our study.
Kizirian et al. (2003) had doubts about theplacement of Hyla tapichalaca. Faivovich etal. (2004) showed that this species is relatedto the H. armata–H. larinopygion groups (al-though they only included H. armata in theiranalysis). Our results go a step further, indi-cating a closer relationship with the H. lari-nopygion group.
Hyla inparquesi and H. kanaima formingthe sister taxon of all the remaining SouthAmerican clade I is an unexpected result. Onthe basis of morphology, we expected ouronly exemplar of the H. aromatica group, H.inparquesi22, to be related to the Andeanstream-breeding clade of Hyla, because bothshare the character states that Duellman et al.(1997) suggested as synapomorphies in sup-port of the monophyly of the Andean stream-breeding Hyla: (1) known larvae with ven-tral, enlarged oral discs; (2) complete mar-ginal papillae; and (3) with a minimum labialtooth row formula of 4/6. Furthermore,adults of the H. aromatica group share agreatly enlarged prepollex without a project-ing spine in males that, as we mentionedabove, is present in most species of Andeanstream-breeding Hyla (the only known ex-ception being H. tapichalaca; Kizirian et al.,2003).
This sister-group relationship between theTepuian clade and all the remaining groupsof the South American clade I has furtherimplications. Based on the available material,Duellman et al. (1997) considered the pre-pollex greatly enlarged without a projectingspine to be an intermediate state in an or-dered transformation series from prepollexnot greatly enlarged to prepollex greatly en-larged with a projecting spine. Our topologyimplies that the greatly enlarged prepollexwithout a projecting spine is a synapomorphyof the whole South American clade I (withsubsequent transformations, including thedevelopment of a projecting spine). Howev-er, H. kanaima does not have an enlargedprepollex as prominent as that found in theH. armata, H. aromatica, H. bogotensis, andH. larinopygion groups. In order to clarifythis situation, it would be necessary to (1)define the prepollex character states osteo-logically, and (2) include a denser sampling
22 The tadpole of Hyla kanaima is unknown.
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Fig. 5. A partial view of the strict consensus showing the relationships of the Middle American–Holarctic and South American/West Indian Casqued-headed frog clades and it correspondence with thecurrently recognized species groups. Numbers above nodes are Bremer support values. Numbers belownodes are Parsimony Jackknife absolute frequencies; those with an asterisk (*) have a 100% frequency.Numbers in boldfaced italic are node numbers for the list of morphological synapomorphies (see ap-pendix 3). Black circles denote nodes that are present in the quick consensus estimation.
of the H. aromatica group, to better under-stand its relationship with H. kanaima. (Per-haps the character state of H. kanaima couldbe interpreted as a reversal.)
Our topology implies an interesting sce-nario regarding the evolution of larval mor-phology in the South American clade I; thatis, that larval morphology of the Atlantic/Cerrado, Green, and TGF clades evolvedfrom an ancestor with the highly modifiedmorphology typical of stream larvae (includ-ing large numbers of labial tooth rows, alarge oral disc with complete marginal pa-pillae, and relatively low fins) that duringevolution, underwent a transformation ofthese character states (specifically, reductionin labial tooth row formulae, a reduced oraldisc, and formation of an anterior gap in themarginal papillae). These transformationswere coincident with distributional shiftsfrom high-elevation mountain streams (as inthe Tepuian and Andean stream-breedingclades) toward lower elevation forest moun-tain streams (the cases of the Atlantic/Cer-rado clade, the Green clade, and, and sometaxa of the TGF clade), and Amazonian low-lands and the Cerrado-Chaco (several taxa ofthe TGF clade).
Gladiator Frogs
Duellman et al. (1997) suggested the ex-istence of a clade composed of the Hyla al-bomarginata, H. albopunctata, H. boans, H.circumdata, H. geographica, and H. pul-chella groups. The synapomorphy supportingthis clade is, according to these authors, thepresence of ‘‘an enlarged prepollical spinelacking a quadrangular base’’. Faivovich etal. (2004), based on the analysis of mito-chondrial DNA sequences and on the obser-vation (Garcia and Faivovich, personal obs.)that H. punctata and the H. polytaenia groupshow the same morphology of the prepollicalspine, argued that these additional groups
also belong to that clade (these authors fur-ther included the H. polytaenia group withinthe H. pulchella group), which was calledearlier in this paper ‘‘Gladiator Frogs’’. Ourresults indicate that a clade with the com-position suggested by Duellman et al. (1997)and Faivovich et al. (2004) is paraphyleticbecause Aplastodiscus is nested within it;furthermore, H. kanaima of the H. geogra-phica group is only distantly related to thisclade.
Atlantic/Cerrado Clade
Our results corroborate the long-suspectedassociation of the Hyla circumdata groupwith the groups of H. pseudopseudis and H.martinsi (Bokermann, 1964a; Cardoso, 1983;Caramaschi and Feio, 1990; Pombal andCaramaschi, 1995), even though the mono-phyly of these two groups could not be testedby our analysis. We also consider as corrob-orated the suspected relationship of the Hylacircumdata and H. pseudopseudis groupswith H. alvarengai (Bokermann, 1964a;Duellman et al., 1997), because Hyla sp. 9(aff. H. alvarengai) is nested in this clade.
Unfortunately, due to the unavailability ofsamples, we could not test the relationships ofthe Hyla claresignata group with this clade.Considering the phylogenetic context of theAtlantic/Cerrado clade within the SouthAmerican clade I, we must revisit the appar-ent synapomorphies of the H. claresignatagroup mentioned earlier (oral disc completelysurrounded by marginal papillae, and 7/12–8/13 labial tooth rows). The presence and dis-tribution of these character states in the Te-puian and Andean stream-breeding Hylaclades is suggestive, not of a closer relation-ship of the H. claresignata group with any ofthem (these character states are plesiomor-phies in this context), but of the nature of itsrelationship with the Atlantic/Cerrado clade.Perhaps the H. claresignata group is not a
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member of the Atlantic/Cerrado clade, but isa basal group related either to the Tepuian orAndean stream-breeding clade, or perhaps itis even the sister taxon of the clade composedof the Atlantic/Cerrado, Green, and TGFclades. At this point we are not aware of ev-idence favoring any of these alternatives.
Green Clade
Lutz (1950) was the first to suggest thatAplastodiscus was related to species latter in-cluded in the Hyla albosignata complex, asdefined by Cruz and Peixoto (‘‘1985’’[1987]). This was supported by Garcia et al.(2001a) based on the presence of enlargedinternal metacarpal and metatarsal tuberclesand unpigmented eggs. More recently, Had-dad et al. (2005) described the reproductivemode of Aplastodiscus perviridis, which in-cludes egg deposition in a subterranean nestexcavated by the male, where exotrophic lar-vae spend the early stages of developmentuntil flooding releases them to a nearby waterbody. This mode is the same as that observedin one species of the H. albosignata com-plex, H. leucopygia (Haddad and Sawaya,2000), and for the undescribed species of theH. albofrenata complex included in our anal-ysis, Hyla sp.1 (aff. H. ehrhardti) (Hartmannet al., 2004); this mode is further suspectedto occur in all species of both the H. albo-signata and H. albofrenata complexes (Had-dad and Sawaya, 2000, Hartmann et al.,2004). Species included in the H. albomar-ginata complex instead lay their eggs on thewater film surface (Duellman, 1970). Basedon the shared reproductive mode, Haddad etal. (2005) suggested that Aplastodiscus couldbe related to the H. albofrenata and H. al-bosignata complexes. Our results support themonophyly of both the H. albofrenata andH. albosignata complexes and their close re-lationship with Aplastodiscus, that is the sis-ter taxon of the H. albosignata complex.
While we are not aware of nonmolecularsynapomorphies for several nodes supportedby molecular evidence, the few osteologicaldata available for the South American cladeI indicate that the Green clade and the TGFclade share the presence of transverse pro-cesses of the sacral diapophyses notably ex-panded distally, while species of the Atlantic/
Cerrado clade (Bokermann, 1964a; Garcia2003) and H. tapichalaca (Kizirian et al.,2003), the only Andean stream-breedingHyla with any described postcranial osteol-ogy, have the transverse processes poorly ex-panded or not expanded at all. However, thedistribution of this character state is in con-flict with the presence of a prepollical spinein the Atlantic/Cerrado clade and the TGFclade, which is absent in the Green clade,where there is a fairly enlarged prepollex butno spine. A detailed study of prepollex mor-phology in these taxa would help to betterdefine the relevant transformation series andit transformation sequences in the tree.
True Gladiator Frog Clade
The results imply the nonmonophyly of sev-eral species groups within this clade, as de-scribed earlier. This situation is not unexpectedconsidering the paucity of evidence of mono-phyly previously available for most of them.
The monophyly of the group composed ofthe two unassigned species from the Gua-yana Highlands, Hyla benitezi, H. lemai,Hyla sp. 2, two members of the H. geogra-phica group (H. microderma and H. rorai-ma), and Hyla sp. 8 is further supported bythe presence of a mental gland in males (Fai-vovich et al., in prep.)
In the DNA-based phylogenetic analysisof the Hyla pulchella group performed byFaivovich et al. (2004), H. punctata is thesister species of H. granosa, with this overallclade forming the sister taxon of a cladecomposed of the H. albopunctata, H. geo-graphica, H. albomarginata, H. boans, andH. pulchella groups. In our analysis, H.punctata, our only exemplar of the group, isthe sister taxon of H. granosa, and this taxonis at the apex of a pectinated series that in-cludes H. sibleszi and, curiously, H. pictur-ata. Prior to the analysis, we had no idea asto with which species group H. picturatawould be related, but certainly we did notexpect this colorful frog to be nested withina group of green species.
Our results lend only partial support for arelationship between Hyla heilprini and theH. albomarginata group, as tentatively sug-gested by Duellman (1974) and Trueb andTyler (1974) based on overall pigmentation
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and the white peritoneum, because H. heil-prini is the sister taxon of the H. albopunc-tata group (including a fragment of the H.geographica group), with H. heilprini plusthis unit being the sister taxon of a fragmentof the H. albomarginata group. The non-monophyly of the H. albopunctata groupcorroborates comments advanced by de Sa(1995, 1996) as to the lack of evidence forits monophyly.
Hyla crepitans, H. faber, H. lundii, and H.pardalis form, together with H. albomargin-ata, a monophyletic group only distantly re-lated to H. boans, the species that gives thename to the former group. Within this clade,the nest builders23 H. faber, H. lundii, and H.pardalis, are monophyletic.
The polyphyly of the Hyla boans grouphas implications for the evolution of repro-ductive modes, in that it implies independentorigins of nest-building behavior by males.Although theoretically possible, certainly noauthor had ever suggested that such a char-acteristic behavior as the nest building couldbe a homoplastic feature.24 However, the mo-lecular evidence points that way, and furtherevidence indicates that this or a similar re-productive mode also occurs in at least somespecies of the H. circumdata group (Pombaland Haddad, 1993, Pombal and Gordo,2004), implying then at least three indepen-dent occurrences within the South Americanclade I. In a wider context, nest building wasreported as well in Pelodryadinae in malesof Litoria jungguy (Richards, 1993; using thename L. lesueuri, see Donnellan and Mahony[2004]), thus implying a fourth instance ofhomoplasy within Hylidae.
Our topology for the Hyla pulchella groupis identical to that of Faivovich et al. (2004),including the exemplars of the former H. po-lytaenia group nested within it. Following
23 Caldwell (1992) referred to the facultative nature ofnest building in males of Hyla crepitans on specimensfrom Venezuela, far away from the range of H. crepitansin Brazil. Other authors (Lynch and Suarez-Mayorga,2001), expressed doubts regarding the taxonomic statusof northwestern South American H. crepitans, and un-published molecular data from Faivovich and Haddadindicate that more than one species is involved.
24 Our surprise with these results led us to sequencean additional sample of each Hyla boans and H. faberto check for the possibility of cross-contaminations; bothwere identical with the sequences we already had.
Faivovich et al. (2004), we continue to rec-ognize a H. polytaenia clade within the H.pulchella group. These authors stated that thelack of any pattern on the hidden surfaces ofthighs was one of two possible morphologicalsynapomorphies of this clade (with the otherbeing the mostly striped dorsal pattern). Thisobservation is mistaken, because the samecharacter state occurs in H. ericae (Caramas-chi and Cruz, 2000), H. joaquini, H. margin-ata, H. melanopleura, H. palaestes, and H.semiguttata (Duellman et al., 1997; Garcia etal., 2001b), suggesting that it actually may bea synapomorphy of a more inclusive cladewhose contents are still undefined.
SOUTH AMERICAN II CLADE
The South American II clade (fig. 4) iscomposed of the 30-chromosome Hyla, theHyla uruguaya group, Lysapsus, Pseudis,Scarthyla, Scinax, Sphaenorhynchus, and Xe-nohyla. It contains two main clades: onecomposed of Lysapsus, Pseudis, Scarthyla,and Scinax (including the H. uruguayagroup), and the other composed of Sphae-norhynchus, Xenohyla, and all the exemplarsof 30-chromosome Hyla species groups.
Within this clade, Scinax and the Hyla mi-crocephala group are not monophyletic. Sci-nax has H. uruguaya, an exemplar of the H.uruguaya group, nested within it. The H. mi-crocephala group is paraphyletic with re-spect to the available exemplars of the H.decipiens and H. rubicundula groups.
Relationships of Scinax
Several hypotheses have been advanced onthe relationships of Scinax. Aparasphenodonwas considered by Trueb (1970a) to be closelyrelated to Corythomantis and, in turn, she con-sidered these two genera to be nested withinthe (then) Hyla rubra group (currently the ge-nus Scinax), based on overall similarities incranial morphology. Scarthyla was consideredto be related to Scinax by Duellman and de Sa(1988). Duellman and Wiens (1992) extendedthis to suggest that Scarthyla is the sister taxonof Scinax, which together form the sister taxonof Sphaenorhynchus. According to Duellmanand Wiens (1992), character states supportingthe monophyly of these three genera are nar-row sacral diapophyses, anteriorly inclined ala-
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ry processes of the premaxillae, and tadpoleswith large, laterally placed eyes. Duellman andWiens (1992) suggested that possible morpho-logical synapomorphies of Scinax and Scarthy-la are reduced webbing on the hand and thepresence of an anterior process of the hyale.Tepuihyla was suggested to be closely relatedwith Scinax, this being supported by the ab-sence or extreme reduction of webbing be-tween toes I and II, adhesive discs wider thanlong, and the presence of double-tailed sperm(Ayarzaguena et al., ‘‘1992’’ [1993b]). This as-sociation was questioned by Duellman andYoshpa (1996) on the grounds that the absenceor extreme reduction of webbing between toesI and II was homoplastic among hylids (al-though Duellman and Wiens [1992] suggestedthis same character state to be a synapomorphyof Scinax). These authors suggested that theonly evidence uniting Tepuihyla with Scinaxcould be the double-tailed sperm reported byAyarzaguena et al. (‘‘1992’’ [1993b]) for Te-puihyla and by Fouquette and Delahoussaye(1977) for Scinax.25
Mijares-Urrutia et al. (1999) again suggesteda close relationship of Tepuihyla with Scinax,but also with Scarthyla and Sphaenorhynchus–Scinax relatives, as suggested by Duellman and
25 The interpretation of the double-tailed sperm as aputative synapomorphy is problematic for two practicalreasons: (1) Taboga and Dolder (1998), Kuramoto(1998), and Costa et al. (2004) suggested that previousreports of double-tailed spermatozoa in several Anurabased on optical microscopy are in error, because scan-ning electron microscopy and transmission electron mi-croscopy of ultrathin serial sections show that actuallythere is a single axoneme/paraxonemal rod and the axialfiber. This suggests that there could be a problem ofhomology between the structures present in Scinax andthose in Tepuihyla. (2) Even if we would assume thatthe problem is only about the correct interpretation oftwo different states in the optical microscopy (i.e.,whether the ‘‘double tail’’ is actually a double flagellumor an axoneme/paraxonemal rod and the axial fiber), wefind that the studied hylid taxa using optical microscopyare not numerous. Although Fouquette and Delahous-saye (1977) mentioned that they studied several hylines,an exhaustive list of those taxa was not given, and pub-lished records only include Acris (Delahoussaye, 1966),10 species of Hyla (Delahoussaye, 1966; Pyburn, 1993;Kuramoto, 1998; Taboga and Dolder, 1998; Costa et al.,2004), 1 species of Pseudacris (Delahoussaye, 1966),Pseudis and Lysapsus (Garda et. al., 2004), Scarthylagoinorum (Duellman and de Sa, 1988), several speciesof Scinax (Fouquette and Delahoussaye, 1977; Tabogaand Dolder, 1998; Costa et al., 2004), and Sphaenorhyn-chus lacteus (Fouquette and Delahoussaye, 1977).
Wiens (1992). Mijares-Urrutia et al. (1999)also noted that Tepuihyla has rounded sacraldiapophyses as found in these three genera(Duellman and Wiens, 1992).
Our results do not support a relationshipof Scinax with Aparasphenodon, Corythom-antis, or Tepuihyla because these three gen-era are nested within the South American/West Indies Casque-headed Frog clade. Fur-thermore, Scinax is not the sister group ofScarthyla but of a clade composed of Scar-thyla plus Lysapsus and Pseudis (in the 1:1:1 analysis) or of all the remaining genera in-cluded in the South American II clade (in the3:1:2 analysis).
Scinax is also paraphyletic with respect tothe Hyla uruguaya group, for which Boker-mann and Sazima (1973a) and Langone(1990) could not suggest affinities with anyother hylids. Most recently, Kolenc et al.(‘‘2003’’ [2004]) observed in the larvae ofH. uruguaya and H. pinima the morpholog-ical synapomorphies of the larvae of the Sci-nax ruber clade that were reported by Fai-vovich (2002), and they suggested a possiblerelationship between the H. uruguaya groupand the Scinax ruber clade. Adults of the H.uruguaya group are quite characteristic mor-phologically, and perhaps as a consequencethis group was never associated with anyspecies of Scinax prior to Kolenc et al.(‘‘2003’’ [2004]).
Our results reveal that none of the out-groups employed by Faivovich (2002) in thephylogenetic analysis of Scinax is particular-ly close to Scinax; instead, all other compo-nents of the South American II clade aremuch more suitable to establish character-state polarities in this genus. Consequently,exemplars of these closer neighbors of Sci-nax need to be added, and the synapomor-phies of Scinax resulting from that analysisneed to be reevaluated.
Lysapsus, Pseudis, and Scarthyla
The sister-group relationship betweenScarthyla goinorum and ‘‘pseudids’’ (Lysap-sus 1 Pseudis), and this group being nestedwithin Hylinae, corroborates recent findingsby Darst and Cannatella (2004) and Haas(2003). Burton (2004) reported a likely syn-apomorphy for the Scarthyla plus the ‘‘pseu-
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did’’ clade that is corroborated in the presentanalysis; that is, the m. transversus metatar-sus II oblique, with a narrow, proximal con-nection to metatarsus II, and a broad, distalconnection to metatarsus III. Another char-acter state described by Burton (2004), theundivided tendon of the m. flexor digitorumbrevis superficialis, optimizes in this analysisas a synapomorphy of this clade plus Scinax.
Besides the molecular data, the monophy-ly of Lysapsus plus Pseudis is further sup-ported by the tendo superficialis pro digiti IIIarising from the m. flexor digitorum brevissuperficialis, with no contribution from theaponeurosis plantaris; the origin of m. flexorossis metatarsi IV and the joint tendon of or-igin of mm. flexores ossum metatarsorum IIand III crossing each other; the m. flexor os-sis metatarsi IV very short, inserting on theproximal two-thirds of metatarsal IV or less;absence of a tendon from the m. flexor dig-itorum brevis superficialis to the medial slipof the medial m. lumbricalis brevis digiti V;and m. transversus metatarsus III oblique,with a narrow, proximal connection ontometatarsal III, and a broad, distal connectionto metatarsal IV. Another likely morpholog-ical synapomorphy is the elongated interca-lary elements.
Vera Candioti (2004) noticed that Lysap-sus limellum and most of the 30-chromosomeHyla (H. nana and H. microcephala) studiedby her and Haas (2003) share two of the syn-apomorphies that Haas (2003) reported forPseudis paradoxa and P. minuta: insertion ofthe m. levator mandibulae lateralis in the na-sal sac, and a distinct gap in the m. subar-cualis rectus II–IV. Haas (2003) observeddifferent character states for H. ebraccata(the m. levator mandibulae lateralis inserts intissue close to posterodorsal process of su-prarostral cartilage or adrostral tissue; contin-uous m. subarcualis rectus II–IV), suggestingthe need for additional studies on its taxo-nomic distribution within the 30-chromo-some Hyla and in the other genera of theSouth American II clade.
Sphaenorhynchus, Xenohyla and the 30-Chromosome Hyla
Izecksohn (1959, 1996) suggested possiblerelationships of Xenohyla with Sphaenorhyn-
chus and Scinax (Izecksohn, 1996). Theseideas are partially corroborated by our 1:1:1results, with the exception that they also sug-gest that Xenohyla is the sister group of the30-chromosome Hyla. The karyotype is stillunknown in Xenohyla, and this poses an ob-stacle to our understanding of the limits ofthe 30-chromosome Hyla. Interestingly,while both Sphaenorhynchus and Xenohylado have a quadratojugal, in both cases it doesnot articulate with the maxilla (Duellman andWiens, 1992; Izecksohn, 1996), which couldbe seen as an intermediate step before theextreme reductions of the quadratojugal seenin the 30-chromosome Hyla (Duellman andTrueb, 1983). Tadpoles of Xenohyla and sev-eral species of 30-chromosome Hyla sharethe presence of the tail tip extended into aflagellum, as well as the presence of highcaudal fins (e.g., see Bokermann, 1963; Ken-ny, 1969; Gomes and Peixoto, 1991a, 1991b;Izecksohn, 1996; Peixoto and Gomes, 1999).
Phylogenetic hypotheses of the 30-chro-mosome Hyla species groups using morpho-logical characters were presented by Duell-man and Trueb (1983), Duellman et al.(1997), Kaplan (1991, 1994), and Kaplanand Ruız (1997); none of these tested themonophyly of the contained species groups.A summary of their proposals and the sup-porting evidence are depicted in figure 6.
Chek et al. (2001) presented a phyloge-netic analysis using partial 16S and cyto-chrome b sequences of the Hyla leucophyl-lata group, including exemplars of other 30-chromosome Hyla species groups. Becausethey did not include non-30-chromosome hy-lids, they did not test the monophyly of thisclade.
The distribution of certain characters inseveral species associated with the currentlyrecognized species groups suggests problemsin our phylogenetic understanding of thesefrogs. The monophyly of a group composedof the Hyla leucophyllata, H. marmorata, H.microcephala, and H. parviceps groups iscurrently supported by the absence of labialtooth rows in their larvae (Duellman andTrueb, 1983). However, within the H. parv-iceps group, H. microps (Santos et al., 1998)and H. giesleri (Bokermann, 1963; Santos etal., 1998) have at least one labial tooth row.Similarly, Gomes and Peixoto (1991a) and
64 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 6. Current state of phylogenetic knowledge of the 30-chromosome Hyla. Redrawn from Duell-man (2001: 857) with the addition of Kaplan’s (2000) suggestion regarding the relationships of Hylapraestans with the H. garagoensis group. Numbered synapomorphies, as textually described by theseauthors, are: 1, 30 chromosomes; 2, reduced quadratojugal; 3, 1/2 labial tooth rows; 4, nuptial excres-cences absent; 5, tadpole tail xiphicercal; 6, tadpole mouth terminal; 7, 0/1 labial tooth rows; 8, 0/0labial tooth rows; 9, one ventral row of small labial papillae in tadpoles; 10, extensive axillary mem-brane; 11, longitudinal stripes on hindlimbs of tadpoles; 12, one ventral row of large labial papillae intadpoles; 13, tadpole body violin-shaped in dorsal view; 14, body of tadpole depressed; 15, labialpapillae absent in tadpoles; 16, internal surface of the arytenoids with a small medial depression.
Peixoto and Gomes (1999) noticed in the H.marmorata group the presence of one labialtooth row in the larvae of H. nahdereri, H.senicula, and H. soaresi. Based on thesefacts and similarities in tail depth, tail color,general body shape, and predatory habits,they suggested that the H. marmorata groupcould instead be more closely related to H.minuta than to the groups suggested byDuellman and Trueb (1983). Gomes andPeixoto (1991b) pointed out the presence ofa labial tooth row in the larva of H. elegans.Wild (1992) further noted the absence ofmarginal papillae (an apparent synapomor-phy of the H. microcephala group) in the lar-va of H. allenorum (a species of the H. parv-iceps group).
The reproductive modes of the differentspecies are also informative. According toDuellman and Crump (1974), Hyla parvicepsdeposits its eggs directly in the water, as doesH. microps (Bokermann, 1963), whereas H.bokermanni and H. brevifrons oviposit onleaves overhanging ponds; upon hatching,
the tadpoles drop into the water where theycomplete development. Hyla ruschii ovipos-its on leaves overhanging streams (Weygoldtand Peixoto, 1987). The oviposition onleaves occurs in most species of the H. leu-cophyllata group, whereas both reproductivemodes occur in the H. microcephala group.
Our results recover the 30-chromosomeHyla species as monophyletic; however, ourtopology differs from previous hypotheses.In our topology, the root is placed betweenthe H. marmorata group and the other ex-emplars, instead of between the H. labialisgroup and the other exemplars, as was as-sumed in previous analyses (Duellman andTrueb, 1983; Kaplan, 1991, 1994, 1999;Chek et al., 2001).
Topological differences from previous hy-potheses are not due merely to a re-rootingof the previously accepted tree; the relation-ships obtained by our analysis are quite dif-ferent from previous proposals. Our analysisdoes not recover as monophyletic the ex-emplars of the three species groups once
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thought to be monophyletic on the basis oflacking labial tooth rows, that is, the Hylaleucophyllata, H. microcephala, and H.parviceps groups. Instead, the H. microce-phala group (including the taxa imbeddedwithin it) is the sister taxon of a clade com-posed of H. anceps and the exemplars of theH. leucophyllata group. The exemplars of theH. parviceps group are the sister taxon of aclade composed of the exemplars of the H.columbiana and H. labialis groups. Thisshows that the scenario of labial tooth rowevolution is more complex than previouslythought, because it implies several transfor-mations in both directions between presenceand absence of labial teeth within the clade.
Observations by Wassersug (1980), Spi-randeli Cruz (1991), and Kaplan and Ruiz-Carranza (1997) on the internal oral featuresof larvae of representatives of the Hyla leu-cophyllata (H. ebraccata and H. sarayacuen-sis), H. microcephala (H. microcephala, H.nana, H. phlebodes, and H. sanborni), andH. garagoensis (H. padreluna and H. viro-linensis) groups revealed a reduction of in-ternal oral structures (including reduction ofmost internal papillation, reduction of bran-chial baskets, reduction or absence of secre-tory ridges and secretory pits) that is mostextreme in the representatives of the H. mi-crocephala group. Hyla minuta does notshow the reductions seen in these speciesgroups (Spirandeli Cruz, 1991). This speciesalso shares with representatives of the H. leu-cophyllata group described by Wassersug(1980) a reduction in the density of the filtermesh of the branchial baskets in comparisonwith other hylid tadpoles. It is clear that thestudy of internal oral features will provideseveral additional characters relevant for thestudy of the 30-chromosome species of Hyla.
The exemplars of the Hyla parvicepsgroup obtain as monophyletic. However, weincluded only 3 of the 15 species currentlyincluded in this problematic group. We arenot confident that the monophyly of the H.parviceps group will be maintained as moretaxa are added. Regarding the exemplars ofthe H. leucophyllata group, their relation-ships are equivalent to those obtained byChek et al. (2001).
The sister-group relationship of Hyla an-ceps and the H. leucophyllata group corrob-
orates early suggestions by Lutz (1948,1973) that these could be related on the basisof sharing a large axilar membrane and flashcoloration.
While we could not test the monophyly ofthe Hyla minima and H. minuta groups, ourexemplars of these groups are sister taxa, andthey are only distantly related with the ex-emplars of the H. parviceps group. This po-sition does not support Duellman’s (2001)tentative suggestion that the species of the H.minima group should be included in the H.parviceps group.
The paraphyly of the Hyla microcephalagroup with respect to the H. rubicundulagroup is an expected result, as historically itsspecies were associated with H. nana and H.sanborni (Lutz, 1973). Nevertheless, the as-sociation of the H. microcephala and H. rub-icundula groups were reinforced by Puglieseet al. (2001), who described the larva of H.rubicundula and noted similarities (like thelack of marginal papillae) with the larvae ofmembers of the H. microcephala group. Inparticular, these authors noticed similaritieswith H. nana and H. sanborni, with the latterbeing the sister taxon of H. rubicundula inour analysis.
Carvalho e Silva et al. (2003) segregatedthe Hyla decipiens group from the H. micro-cephala group on the basis that the larvae ofthese species lack the possible morphologicalsynapomorphies currently diagnostic of theH. microcephala group (body of tadpole de-pressed, labial papillae absent in tadpoles)and the putative clade composed of the H.leucophyllata, H. microcephala, and H.parviceps groups (absence of labial toothrows). However, as mentioned above, our re-sults imply a complex scenario for labialtooth row transformations and place the H.decipiens group within the H. microcephalagroup. The available taxon sampling did notallow testing the monophyly of the H. deci-piens group. The fact that its known speciesshare the oviposition on leaves above the wa-ter, and the reversals in larval morphologythat led Carvalho e Silva et al. (2003) to con-sider them unrelated to the H. microcephalagroup, probably indicates that, even if nestedinside this group, the species assigned to theH. decipiens group could be a monophyleticunit.
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The relationships of the Hyla garagoensisgroup, from which no exemplar was avail-able for this study, were discussed by Kaplanand Ruiz-Carranza (1997). Based on the ab-sence of labial tooth rows, they placed the H.garagoensis group in a polytomy togetherwith the H. marmorata group and the cladecomposed of the H. microcephala, H. parv-iceps, and H. leucophyllata groups. Duell-man et al. (1997) presented a cladogram formost of the 30-chromosome Hyla groups,where the H. garagoensis and H. marmoratagroups appear together as a clade supportedby the presence of one ventral row of smallmarginal papillae in larvae. This characterstate needs further assessment, as indicatedby Gomes and Peixoto (1991a) and Peixotoand Gomes (1999), because tadpoles of theH. marmorata group have either one (H.nahdereri) or two rows of marginal papillae(H. senicula; H. soaresi); the tadpoles of H.padreluna, a species of the H. garagoensisgroup, also has a double row (Kaplan andRuiz-Carranza, 1997). Considering this andearlier comments, we do not see evidencethat associates the H. garagoensis group withthe H. marmorata group more than with anyother group within the 30-chromosome Hylaclade.
MIDDLE AMERICAN/HOLARCTIC CLADE
This clade is composed of most of theMiddle American/Holarctic genera and spe-cies groups of treefrogs (fig. 5). For the pur-poses of discussion, we divide it into fourlarge clades. The first of these includes Acrisand Pseudacris. The second includes Plec-trohyla, the Hyla bistincta group, the H. sum-ichrasti group, and various elements of theH. miotympanum group. The third clade in-cludes Duellmanohyla, Ptychohyla, H. mili-aria, H. bromeliacia (the sole exemplar ofthe H. bromeliacia group), and one elementof the H. miotympanum group. The fourthclade includes Smilisca, Triprion, Anotheca,and the exemplars of the H. arborea, H. ci-nerea, H. eximia, H. godmani, H. mixoma-culata, H. pictipes, H. pseudopuma, H. tae-niopus, and H. versicolor groups.
Within the Middle American/Holarcticclade, the genera Ptychohyla and Smilisca, aswell as the Hyla arborea, H. cinerea, H. ex-
imia, H. miotympanum, H. tuberculosa, andH. versicolor groups, are not monophyletic.The H. miotympanum group is polyphyletic;its exemplars split among three differentclades: H. miotympanum is the sister taxonof one of the exemplars of the H. tuberculosagroup, H. miliaria; H. arborescandens andH. cyclada are related to an undescribed spe-cies close to H. thorectes, and together arerelated to exemplars of the H. bistinctagroup; H. melanomma and H. perkinsi are atthe base of the H. sumichrasti group. Smilis-ca is not monophyletic, having Pternohylafodiens nested within it. Ptychohyla is para-phyletic with respect to H. dendrophasma(H. tuberculosa group). The H. arboreagroup is polyphyletic, with H. japonica nest-ed within the H. eximia group. The H. ci-nerea group is not monophyletic, with H. fe-moralis being more closely related to mem-bers of the H. eximia and H. versicolorgroups than to H. cinerea, H. gratiosa, andH. squirella. The H. versicolor group is notmonophyletic because H. andersonii is moreclosely related to the H. eximia group.
The monophyly of all genera and speciesgroups of Hyla contained in this clade wasmaintained by Duellman (1970, 2001) basedmostly on biogeographic grounds, becausemorphological evidence of monophyly waslacking. Duellman (2001) further presented adiagram depicting ‘‘suggested possible evo-lutionary relationships’’ among Middle andNorth American Hylinae, using as terminalsthe species groups of Hyla and the differentgenera (redrawn here as fig. 7). Duellman(2001) envisioned a North American basallineage being the sister taxon of what hecalled the Middle American basal lineage.This Middle American basal lineage is fur-ther divided into a lower Central Americanlineage (itself divided into an isthmian high-land lineage and a lowland lineage) and theMexican-Nuclear Central American lineage(in turn divided into a Mexican-Nuclear Cen-tral American highland lineage and a low-lands lineage).
While the basal position of Acris andPseudacris in this clade is consistent withDuellman’s (2001) intuitive suggestion of aNorth American basal lineage, it differs inthat the Holarctic species groups of Hyla areonly distantly related to them.
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Fig. 7. Relationships of Middle American and North American Hylinae as envisioned by Duellman(2001). Broken lines are tentative placements.
We found no evidence supporting Duell-man’s (2001) exclusively Mexican-NuclearCentral American lineage, nor of his Mexi-can-Nuclear Central American highland lin-eage. The latter has nested within it the Mex-ican-Nuclear Central American lowlandclade (the H. godmani group), a clade remi-niscent of Duellman’s (2001) isthmian high-land-lowlands lineage, and also all speciesgroups of Holarctic Hyla. Biogeographic im-plications of the discordant nature of our re-sults with Duellman’s suggested relationshipsbetween Middle- and North American Hyli-nae will be dealt with from a biogeographicperspective later in this paper.
The nonmonophyly of North American
Hylinae does not agree with previous anal-yses (Hedges, 1986; Cocroft, 1994; da Silva,1997; Moriarty and Cannatella, 2004) be-cause those analyses assumed implicitly thatAcris, the Holarctic Hyla, and Pseudacris aremonophyletic. In spite of this, the internalrelationships of Pseudacris recovered in thisanalysis are consistent to those obtained byMoriarty and Cannatella (2004).
Previous analyses either did not find evi-dence that Acris was particularly close to anygroup of North American hylids (Cocroft,1994), or else suggested relationships withdifferent species groups of North AmericanHyla (Hedges, 1986; da Silva, 1997). Twomorphological synapomorphies of this Acris
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1 Pseudacris clade could be the spherical orovoid testes (as opposed to elongate testes)and the presence of dark pigmentation in theperitoneum surrounding the testes (Ralin,1970, as cited by Hedges, 1986).
The nonmonophyly of the Hyla miotym-panum group is not surprising because, asdiscussed in the section on taxon sampling,it did not appear that any of its putative syn-apomorphies could withstand a test with abroader taxon sampling. Duellman (2001)merged the formerly recognized H. pinorumgroup (Duellman, 1970) with the H. miotym-panum group. With the notable exception ofH. miotympanum, our results are close to re-covering both groups as originally envi-sioned by Duellman (1970), because H. cy-clada and H. arborescandens (H. miotym-panum group) are recovered as monophylet-ic, and the former H. pinorum group isrecovered as a paraphyletic assemblage thatincludes the H. sumichrasti group nestedwithin it. In his analysis (Duellman, 2001:912), the two species included by Duellman(1970) in the H. pinorum group (H. melan-omma and H. pinorum), plus the two speciesthat were later associated with this group (H.perkinsi and H. juanitae), form a monophy-letic group supported by a single synapo-morphy, the presence of an extensive (equalto or more than one-half length of upper arm)axillary membrane.
The dubious monophyly of the 17 speciesassigned to the Hyla bistincta group was notseriously tested in our analysis, because only2 species were available. These two exem-plars form the sister group of two taxa pre-viously associated with the H. miotympanumgroup, and together they form the sister tax-on of Plectrohyla. According to Duellmanand Campbell (1992), character states sup-porting a monophyletic H. bistincta groupplus Plectrohyla are: medial ramus of pter-ygoid long, in contact with otic capsule; dor-sal skin thick (but see Mendelson and Toal[1995] and Duellman [2001] for discussionsof this character); complete marginal papillaeof the oral disc; and presence of at least onerow of submarginal papillae (called by theseauthors accessory labial papillae) on the pos-terior labium (but see Wilson et al. [1994a]for discussion of this character state). Fromthese, the complete marginal papillae of the
oral disc occur in all known larvae of theclade containing Plectrohyla, and the H. bis-tincta, H. sumichrasti, and fragments of theH. miotympanum group, as well as in larvaeof several other nearby clades (Ptychohyla,Duellmanohyla, the H. mixomaculata, and H.taeniopus groups; see Duellman 1970, 2001).Furthermore, the row of submarginal papillaein the larval oral disc presents a fair amountof variation in the extent and distribution ofthe papillae, within which could probably besubsumed the morphology seen in all knownlarvae of the clade mentioned above (see il-lustrations of all these oral discs in Duell-man, 1970, 2001). Besides discussions pro-vided by Mendelson and Toal (1995) andDuellman (2001) regarding the definition ofthe character state ‘‘thick skin’’, it does notoccur in the following species currently as-signed to the H. bistincta group: H. calvi-collina, H. charadricola, H. chryses, H. la-bedactyla, and H. sabrina (Duellman, 2001).Considering that 15 of 17 species currentlyincluded in the H. bistincta group and 15 of18 included in Plectrohyla could not be in-cluded in the analysis, we do not considerour results a strong test of their intrarelation-ships, particularly when several species ofthe H. bistincta group that present suspiciouscharacter state combinations, like the onesmentioned above, were not available. Our re-sults relating H. arborescandens with speciesof the H. bistincta group corroborate earliersuggestions by Caldwell (1974) that relatethis species to species currently placed in theH. bistincta group (H. mykter, H. robertso-rum, and H. siopela.). Mendelson and Toal(1996) also suggested affinities of H. arbo-rescandens and H. hazalae with the H. bis-tincta group on the basis of unpublished os-teological data.
Duellman (2001) noted the lack of evi-dence for the monophyly of the Hyla tuber-culosa group. Although poor, our taxon sam-pling does not recover it as monophyletic,because H. dendrophasma is nested withinPtychohyla, and H. miliaria is the sister tax-on of H. miotympanum. Duellman (2001)also referred to the possibility advanced byda Silva (1997) of a relationship of the H.tuberculosa group with the Gladiator Frogs;at this point the evidence presented hereindoes not support this idea, but in case a dens-
2005 69FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
er sampling of the group still corroborates itspolyphyly, we would not be surprised ifsome of its elements (particularly H. tuber-culosa26) are shown to be related with theTGF clade.
Hyla miotympanum has repeatedly beenconsidered a generalized Middle Americanhyline (Duellman, 1963, 1970; Campbell andSmith, 1992; Duellman, 2001), largely be-cause the larva of H. miotympanum exhibitsa labial tooth row formula of 2/3 and a rel-atively small oral disc with an anterior gapin the marginal papillae. We are not aware ofany morphological synapomorphy support-ing its relationship with H. miliaria, althoughour molecular data firmly place it there. Ac-cording with our results, there is a morpho-logical synapomorphy supporting the mono-phyly of the clade composed of these twospecies plus Duellmanohyla, the H. brome-liacia group, and Ptychohyla (including H.dendrophasma): the tendo superficialis hal-lucis that tapers from an expanded corner ofthe aponeurosis plantaris, with fibers of them. transversus plantae distalis originating ondistal tarsal 2–3 that insert on the lateral sideof the tendon.
Considering its overall external appear-ance, we are surprised by the position of thepoorly known Hyla dendrophasma. This spe-cies was originally considered to be a mem-ber of the H. tuberculosa group (Campbellet al., 2000) based on its large snout–ventlength and extensive hand webbing, althoughwith the caveat that it lacks dermal fringes,the only character state shared by all otherspecies placed in the H. tuberculosa group.DNA was isolated and sequenced twice fromtissues of the female holotype, the onlyknown specimen (Campbell et al., 2000). In-asmuch as most previous notions of relation-ships among species of Ptychohyla derivefrom adult male morphology and tadpoles,the discovery of at least one male specimenof H. dendrophasma could hopefully allow
26 The other South American species of the Hyla tu-berculosa group, H. phantasmagoria, is known onlyfrom the holotype. It was considered a junior synonymof H. miliaria by Duellman (1970), who later resurrectedit (Duellman, 2001). Besides a few comments by thisauthor, no morphological comparisons with other speciesof the group are available.
us to better understand its relationships with-in Ptychohyla.
Campbell and Smith (1992) and Duellman(2001) suggested five morphological syna-pomorphies for Ptychohyla. One of these isapparently unique to Ptychohyla (pars pala-tina of the premaxilla with well-developedlingual flange), while the other four show amore extensive taxonomic distribution. (1)The cluster of ventrolateral mucous glands inbreeding males is present in Duellmanohylachamulae, D. ignicolor, and D. schmidtorum(Campbell and Smith, 1992; see also Thomaset al., 1993). (2) The presence in the ventro-lateral edge of forearm of tubercles coalescedinto a ridge (as opposed to the absence oftubercles) was reported for D. lythrodes, D.salvavida, D. schmidtorum, and D. soralia(Duellman, 1970; 2001); H. bromeliacia hasan indistinct row of tubercles that do not co-alesce into a ridge (absent in H. dendroscar-ta) (Duellman, 1970). (3) The double row ofmarginal papillae is present as well in larvaeof the H. bromeliacia group (Duellman,1970). Finally, (4) larvae of the H. bromelia-cia group have a labial tooth row formula of2/4 or 2/5, and all known larvae of Duell-manohyla have a labial tooth row formula of3/3; the minimum known for a species ofPtychohyla is 3/5 (P. legleri and P. salva-dorensis) (Duellman, 1970, 2001; Campbelland Smith, 1992).
The monophyly of the group composed ofPtychohyla euthysanota, P. hypomykter, P.leonhardschultzei, and P. zophodes is con-gruent with the results of Duellman (2001),who supported the monophyly of these taxabased on the presence of a thick, roundedtarsal fold. These species further share thepresence of hypertrophied ventrolateralglands in breeding males with two specieswe could not include in our analysis: P. ma-crotympanum and P. panchoi. Furthermore,all these species also share with P. spinipol-lex the presence of the nuptial excrescencescomposed of enlarged individual spines. Thestates of these characters are unknown inPtychohyla sp. and Hyla dendrophasma be-cause the only available specimens are fe-males. The nonmonophyly of P. hypomykterplus P. spinipollex is most surprising, con-sidering that both were considered to be a
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single species (Wilson and McCranie, 1989;see also McCranie and Wilson, 1993).
Although we included several species ofPtychohyla in our analysis, we do not thinkthat we have apprehended a good represen-tation of the morphological diversity of thegroup, and the absence of species like P. er-ythromma, P. legleri, and P. sanctaecruciscertainly weakens the test of monophyly ofPtychohyla. This is more so considering thefact that several of the putative morphologi-cal synapomorphies of Ptychohyla are actu-ally shared with some species of its sistertaxon, as discussed earlier, and that themonophyly of our exemplars of Ptychohylais weakly supported. The low Bremer sup-port (3) for Ptychohyla also suggests that theevidence for its monophyly deserves furtherattention.
The Hyla bromeliacia group was tenta-tively associated with the polyphyletic H.miotympanum group by Duellman (2001:779). Other than this, we are not aware of itbeing associated with any other group. Be-sides the molecular evidence, we are awareof at least one likely morphological synapo-morphy supporting the monophyly of Duell-manohyla plus the Hyla bromeliacia group:the presence of pointed serrations of the lar-val jaw sheaths (Campbell and Smith, 1992;Duellman, 1970; 2001). These are apparentlylonger in some species of Duellmanohylathan in the H. bromeliacia group, but bothseem to be notably more pointed than in Pty-chohyla (see descriptions and illustrations inDuellman, 1970).
We share with Duellman (2001) and Men-delson and Campbell (1999) doubts regard-ing the monophyly of the Hyla taeniopusgroup. Nevertheless, our two exemplars arerecovered as monophyletic in the analysis.Duellman (2001) examined the possibility ofa relationship between this group and the H.bistincta group, based on the fact that bothhave large stream-adapted tadpoles withsmall, ventral oral discs with complete mar-ginal papillae and bear a labial tooth row for-mula of 2/3 (but noting that the tooth-rowformula is slightly higher for H. nephila andH. trux). Our results suggest instead that theH. taeniopus group is the sister taxon of aclade composed of H. mixe (the only avail-able exemplar of the H. mixomaculata group)
plus the clade composed of the HolarcticHyla groups, the H. godmani, H. pictipes,and H. pseudopuma groups, and Anotheca,Smilisca (including Pternohyla), and Tri-prion. Furthermore, in the context of our re-sults, the ventral oral disc with completemarginal papillae seems to be a synapomor-phy of the whole Middle American clade,with subsequent transformations in the cladejust mentioned and in other points of the tree.
We were unable to test the monophyly ofthe Hyla mixomaculata group, because H.mixe was the only taxon available. Regard-less, and until a rigorous test is possible, themonophyly of this group could be reasonablyassumed based on the presence of the en-larged oral disc with 7/10 or 11 labial toothrows. At this point it should be stressed thatthe sequenced sample comes from a tadpolethat was assigned to the H. mixomaculatagroup based that on that characteristic, andthat it was tentatively assigned to H. mixe forbeing the only species of the group knownfrom the region where the larva was collect-ed; thus, considering the uncertainty in itsdetermination, its position in the tree shouldbe viewed cautiously.
Duellman (2001) included the species ofthe former Hyla picta group in the H. god-mani group. Unfortunately, the two exem-plars available to us for this analysis are onlythe two members of the former H. pictagroup and none of the restricted H. godmanigroup; therefore, this is not a satisfactory testof the monophyly of the H. godmani group(sensu lato).
The Lower Central American Lineage
The monophyly of the included exemplarsof the Hyla pictipes and H. pseudopumagroups, and its relationship with a clade com-posed of Anotheca, Smilisca, Triprion, andPternohyla, is quite consistent (in the sensethat it contains almost the same groups) withDuellman’s intuitive proposal of a lowerCentral American clade that contains an Isth-mian Highland lineage and a Lowland line-age (fig. 7), with the only exception beingthat he tentatively considered the H. miliariagroup related to Anotheca.
The monophyly of a group composed ofHyla pseudopuma and H. rivularis, the only
2005 71FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
two exemplars available from the H. pseu-dopuma and H. pictipes groups, could besuggestive of a lineage of highland isthmianHylinae as suggested by Duellman (2001).Although the monophyly of each of thesetwo groups has not been tested here, the po-sition of the undescribed Mexican speciesHyla sp. 5 (aff. H. thorectes) deserves somecomments.
Duellman (1970) recognized the Hyla ha-zelae group in which he included the nomi-nal species and H. thorectes. Reasons for rec-ognizing this group were ‘‘the combinationof large hands with vestigial webbing, halfwebbed feet . . . and presence of a tympa-num are external features which separatethese species from other small stream-breed-ing Mexican Hyla. Furthermore both specieshave small, relatively narrow tongues andlarge tubercles below the anal opening. Thenature of the nasals and sphenethmoid areunique among northern Middle Americanhylids’’ (Duellman, 1970: 384). It is unclearif any of these character states could havebeen considered as evidence of monophylyof the group. It is also unclear on what basisDuellman (2001) dismantled the group andplaced H. hazelae in the H. miotympanumgroup, while H. thorectes was transferred tothe H. pictipes group. Wilson et al. (1994b)suggested that H. thorectes could be relatedto H. insolita and H. calypsa (under the nameH. lancasteri; see Lips, 1996) because theyshare oviposition on leaves overhangingstreams and have dark ventral pigmentation;these character states were not included inDuellman’s (2001) analysis of the group. Al-though we do not have data to take a positionregarding these actions, the fact that Hyla sp.5 (aff. H. thorectes) is unrelated to H. rivu-laris is here taken as evidence that H. tho-rectes should not be included in the H. pic-tipes group. Furthermore, all character statesadvanced by Duellman (2001) as shared byH. thorectes and the species of the H. picti-pes group are also shared by H. thorectes andthe taxa to which Hyla sp. 5 (aff. H. thorec-tes) appears to be closely related in our anal-ysis. Because we were unable to include H.calypsa, H. insolita, or H. lancasteri, we donot have elements to test the hypothesis ofWilson et al. (1994b) regarding the close re-
lationship of H. calypsa, H. insolita, and H.thorectes.
In order to better understand the relation-ships of the Hyla pictipes group, it would beimportant to add to this analysis exemplarsof the former H. zeteki and H. lancasterigroups, because together with the original H.pictipes and H. rivularis groups (as definedby Duellman, 1970) they represent the threemain morphological extremes of the group.The fact that so few exemplars of these twogroups were available is one of the weakerpoints of our analysis.
The paraphyly of Smilisca is partly con-sistent with Duellman’s (2001) phylogeneticanalysis of this genus in that Pternohyla isnested within it. However, we did not recoverTriprion nested within Smilisca, as didDuellman (2001).
A possible relationship between Triprionand Anotheca was first advanced by Lutz(1968) because she considered them to be theextreme of one specialization consisting ‘‘inexcessive ossification of the head, accompa-nied at some stages by extra dentition’’(Lutz, 1968: 10). Within this same line, sheincluded all casque-headed frogs, includingtogether South American and Middle Amer-ican forms. Duellman and Trueb (1976) sug-gested a possible link of Anotheca with Nyc-timantis (discussed below). Duellman (2001:332) proposed a tentative relation of Anoth-eca with the Hyla miliaria group based onthe oophagous tadpoles that develop in bro-meliads or tree-holes (known for the onlyspecies of the H. tuberculosa group with aknown tadpole, H. salvaje; see Wilson et al.,1985). While our results support a sister-group relationship of Anotheca and Triprionas suggested by Lutz (1968), it occurs withinthe Holarctic/Middle American clade and notwithin a group composed of all casque-head-ed frogs as she suggested. In the context ofthis analysis, both Triprion and Anothecashare the posterior expansion of the fronto-parietals that cover almost all the otoccipitaldorsally (see figures in Duellman, 1970).
Our analysis indicates that the insertion ofm. extensor digitorum comunis longus onmetatarsal II is a synapomorphy of a groupcomposed of Anotheca, Triprion, and theparaphyletic Smilisca. Furthermore, Smilisca(including Pternohyla) and Triprion share
72 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
the type I septomaxillary (see Trueb, 1970a)and bifurcated cavum principale of the olfac-tory capsule (Trueb, 1970a); the distributionof these character states should be studied inAnotheca and nearby groups to determine thelevel of inclusiveness of these possible syn-apomorphies.
Real Hyla
Our results concerning the relationshipsbetween the North American and Eurasiaticspecies groups of Hyla differ from previousanalyses, in part likely because of the pre-vious assumption of monophyly of NorthAmerican/Holarctic Hylinae. In the firstplace, the molecular evidence supports aclade containing all North American and Eu-rasiatic species groups of Hyla, a result thatdiffers from previous analyses where rela-tionships either were unresolved (Cocroft,1994) or were paraphyletic with respect toAcris and/or Pseudacris (Hedges, 1986; daSilva, 1997). The polyphyly of the Hyla ar-borea group and the paraphyly of the H. ex-imia group corroborate previous ideas byAnderson (1991) and Borkin (1999) regard-ing their nonmonophyly and the closer rela-tionship of H. japonica with the H. eximiaand H. versicolor groups. A likely synapo-morphy of the H. eximia and H. versicolorgroups, including H. japonica and H. ander-sonii, is the nucleolar organizer region(NOR) present in chromosome 6 instead ofchromosome 10 (Anderson, 1991).
SOUTH AMERICAN/WEST INDIAN CASQUE-HEADED FROGS
This clade (fig. 5) is composed of Phyl-lodytes, Phrynohyas, Nyctimantis, and allSouth American/West Indian casque-headedfrogs: Argenteohyla, Aparasphenodon, Cor-ythomantis, Osteopilus, Osteocephalus, Tra-chycephalus, and Tepuihyla. It is divided ba-sally in a group composed of the two ex-emplars of Phyllodytes, and another groupcomposed of all casque-headed frog genera,including Phrynohyas and Nyctimantis.
Within this clade, other than those generathat are not monotypic (Argenteohyla, Nyc-timantis, Corythomantis) or represented inthis analysis by a single species (Aparas-phenodon, Tepuihyla), Phyllodytes and Os-
teopilus are monophyletic, and Osteocephal-us, Phrynohyas, and Trachycephalus are notmonophyletic. Osteocephalus is not mono-phyletic because O. langsdorffii (the onlyspecies of the genus distributed in the Atlan-tic forest) is not related to the remaining ex-emplars of Osteocephalus, which form amonophyletic group that is the sister taxonof Tepuihyla. Phrynohyas is not monophy-letic, having Trachycephalus nigromaculatusnested within it, and Trachycephalus is notmonophyletic, with T. jordani forming thesister taxon of ‘‘Phrynohyas’’ 1 T. nigro-maculatus.
With respect to Phyllodytes, we were un-able to find any published hypothesis regard-ing its relationships, and considering thescant information available on its morphol-ogy, we had no previous clue as to othergroups of Hylidae with which it might berelated. The only morphological characterstate of which we are aware that Phyllodytesshares with several members of the SouthAmerican/West Indian Casque-headed Frogsclade is the presence of at least four posteriorlabial tooth rows in the tadpole oral disc (seebelow, ‘‘Taxonomic Conclusions: A NewTaxonomy of Hylinae and Phyllomedusi-nae’’, for further details).
The polyphyly of Osteocephalus was notunexpected considering the lack of any evi-dence of its monophyly. This polyphyly re-sults because of the position of O. langs-dorffii. This is the only species of the genuspresent in the Atlantic forest and a speciesthat had been particularly poorly discussed inthe context of the systematics of Osteoce-phalus (Duellman, 1974). While we do nottest the monophyly of the bromeliad-breed-ing/single vocal sac species (here representedby a O. oophagus), our results show that ourexemplars with lateral vocal sacs are para-phyletic with respect to O. oophagus.
The monophyly of Tepuihyla was not test-ed in this analysis. Its sister-group relation-ship with Osteocephalus (excluding O.langsdorffii) is supported by our data, insteadof with Scinax as first suggested (Ayarza-guena et al., ‘‘1992’’ [1993b]; see earlier dis-cussion on the relationships of Scinax). Thissituation requires changes in the original in-terpretation of two character states that pro-vided evidence of the relationships of Te-
2005 73FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
puihyla with other hylids. The presence ofspicules in the dorsum of males is more par-simoniously interpreted as a putative syna-pomorphy of Tepuihyla plus Osteocephalus,instead of a homoplasy, as advanced by Ay-arzaguena et al. (‘‘1992’’ [1993b]). Similarly,the reduction of webbing between toes I andII is more parsimoniously interpreted as a pu-tative synapomorphy of Tepuihyla (homo-plastic with Scinax) instead of a synapomor-phy of Scinax 1 Tepuihyla.
The monophyly of the four exemplars ofOsteopilus corroborates in a much broadertaxonomic context the results of Maxson(1992), Hedges (1996), and Hass et al.(2001), based on albumin immunologicaldistances and still unpublished sequence dataregarding its monophyly, and the recent tax-onomic changes summarized by Powell andHenderson (2003b). Unfortunately, we couldnot include in our analysis O. marianae, O.pulchrilineatus, and O. wilderi, and we arenot aware of any possible morphologicalsynapomorphy supporting their monophyly.In the absence of other evidence, it could besuggested that the oviposition and develop-ment in bromeliads, which occurs in thesethree species and O. brunneus, is a possiblesynapomorphy uniting these with O. cruci-alis, which apparently also has this repro-ductive mode (Hedges, 1987).
The presence of paired lateral vocal sacsand biogeographic considerations led Trueb(1970b) and Trueb and Duellman (1971) tosuggest the collective monophyly of Argen-teohyla, Osteocephalus, Phrynohyas, andTrachycephalus (at that time the species ofOsteocephalus having a single subgular vo-cal sac were still unknown). Furthermore,these authors considered Trachycephalus andPhrynohyas to be a monophyletic group onthe basis of sharing vocal sacs that are morelateral and protrude posteriorly to the anglesof the jaws when inflated. Trueb and Tyler(1974) suggested that the West Indian Osteo-pilus and the former Calyptahyla crucialis(now Osteopilus crucialis) were also relatedto this clade, although they exhibit a single,subgular vocal sac. Our results corroboratethe monophyly of Phrynohyas plus Trachy-cephalus (see below), but they also suggesta more complex situation where the casque-headed frogs with double vocal sacs are par-
aphyletic, with all of the genera of casque-headed frogs that have a single subgular sacbeing nested within them.
Duellman and Trueb (1976) suggested thatNyctimantis was related to Anotheca, be-cause both share the medial ramus of thepterygoid being juxtaposed squarely againstthe anterolateral corner of the ventral ledgeof the otic capsule. Also, frogs of both gen-era are known (Anotheca; Taylor, 1954;Jungfer, 1996) or suspected (Nyctimantis;Duellman and Trueb, 1976) to deposit theireggs in water-filled tree cavities. Our resultssuggest a radically different picture, withNyctimantis nested within the South Ameri-can/West Indian Casque-headed Frogs, whileAnotheca is nested within the Middle Amer-ican/Holarctic clade, being the sister taxon ofTriprion.
The topology has some discrepancies withprevious suggestions as to the relationshipsof Argenteohyla, Aparasphenodon, and Cor-ythomantis (for the latter two genera, see alsocomments for Scinax above). Argenteohylasiemersi was segregated by Trueb (1970b)from Trachycephalus, where it had beenplaced by Klappenbach (1961), because itlacks the diagnostic character states of Tra-chycephalus established by Trueb (1970a).Further, she suggested that Argenteohyla is aclose ally of Osteocephalus based on thepresence of paired lateral vocal sacs. Al-though Trueb (1970a) suggested that Apar-asphenodon and Corythomantis are sistertaxa, our evidence suggests that both Argen-teohyla and Nyctimantis are closer to Apar-asphenodon than to Corythomantis.
Most species in the South American/WestIndies Casque-headed Frog clade frequentlylive in or seek refuge in bromeliads or tree-holes. This has been reported for Aparas-phenodon (Paolillo and Cerda, 1981; Teix-eira et al., 2002), Argenteohyla (Barrio andLutz, 1966; Cespedez, 2000), Corythomantis(Jared et al., 1999), Nyctimantis (Duellmanand Trueb, 1976), Osteocephalus langsdorffii(Haddad, personal obs.), Phrynohyas (Goel-di, 1907; Prado et al., 2003), Tepuihyla (Ay-arzaguena et al., ‘‘1992’’ [1993b]), and Tra-chycephalus (Lutz, 1954; Bokermann,1966c). Furthermore, all species of Phyllod-ytes (Peixoto et al., 2003), some species ofOsteocephalus (Jungfer and Schiesari, 1995;
74 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Jungfer and Weygoldt, 1999; Jungfer andLehr, 2001), some species of Osteopilus(Hedges, 1987; Lannoo et al., 1987), and atleast two species of Phrynohyas (Goeldi,1907; Lescure and Marty, 2000) even laytheir eggs in phytotelmata or treeholes wheretheir exotrophic larvae develop. While bro-meliads and treeholes are used as refuges orfor reproduction in other groups of hylids(e.g., Anotheca spinosa, the Hyla bromelia-cia group, the two bromeliad breeding frogsof the H. pictipes group, H. astartea, the H.tuberculosa group, Scinax alter, the Scinaxperpusillus group of the S. catharinae clade),the South American/West Indian Casque-headed Frog clade seems to be the largestclade of hylids that consistently makes useof bromeliads or treeholes.
The phylogenetic structure of the SouthAmerican/West Indies Casque-headed Frogclade implies a minimum of one instance ofreversal from presence of heavily exostosedand co-ossified skulls to normal looking, al-beit heavily built skulls (the case of Phry-nohyas), and at least a possible second andthird instance involving reversals from ex-ostosed skulls (the cases of Tepuihyla andOsteopilus vastus).
From a morphological perspective, theparaphyly of Phrynohyas with respect toTrachycephalus nigromaculatus and the con-comitant nonmonophyly of Trachycephalusare most interesting and surprising. Herpe-tologists have been noticing for years that T.nigromaculatus and Phrynohyas mesophaeaproduce hybrids throughout their overlappingranges of distribution (Haddad, personalobs.; Pombal, personal commun.; Ramos andGasparini, 2004). The possibility of hybrid-ization leads us to think that perhaps the in-trogression of P. mesophaea mitochondriacould actually be the reason for the recoveredparaphyly of Trachycephalus. However, phy-logenetic analyses using either tyrosinase orrhodopsin alone (the only two nuclear genesthat were successfully sequenced in T. nigro-maculatus) still recover a paraphyletic Tra-chycephalus (results not shown).
TAXONOMIC CONCLUSIONS: A NEWTAXONOMY OF HYLINAE AND
PHYLLOMEDUSINAEBelow we present a new taxonomic ar-
rangement of Hylinae and Phyllomedusinae,
based on the results discussed above. Whileour data clearly point to the fact that wecould hardly have a more paraphyletic anduninformative hylid taxonomy as the currentone, we foresee some resistance to this newmonophyletic taxonomy, due mostly to thelack of morphological evidence in the anal-ysis with consequent few morphological syn-apomorphies in the diagnoses (for many ofthe groups, only the molecular evidence pre-sented here provides the evidence of mono-phyly) or to insufficient numbers of exem-plars. The lack of a complete, well-re-searched nonmolecular data set is admittedlya weakness of this project. However, ourstudy represents the largest amount of evi-dence for the largest number of terminalsever put together and analyzed in a consistentway to address the phylogenetic relationshipsof hylids. Until a nonmolecular data set isassembled, we are left only with the evidenceprovided by our analysis. The alternatives areevident: either we ignore the present resultsand stick to the traditional, grossly uninfor-mative taxonomy, or we dare to present anew monophyletic taxonomy based on theevidence provided here. The latter option isfar closer to the goals of phylogenetic sys-tematics than is the former one. The new tax-onomy is the result of our attempt to recon-cile the need to recognize only monophyleticgroups and to minimize changes to the ex-isting taxonomy while keeping it informative(e.g.: we could have included all currentlyrecognized genera of Hylinae in the synon-ymy of Hyla; this would have resulted in aperfectly monophyletic though utterly unin-formative taxonomy).
We have not tested the monophyly of sev-eral genera and species groups either becausewe could not sample them at all (e. g. thecases of Phrynomedusa and the Hyla clare-signata and H. garagoensis group) or be-cause of insufficiency in sampling (e.g., theH. pictipes, H. pseudopuma, and H. mixo-maculata groups). There are, as well, sevenspecies that we could not associate with anygroup (see ‘‘Incertae Sedis and Nomina Du-bia’’ below and appendix 4). Nevertheless,we are being bold in the recognition ofgroups. We recognize all groups that werepreviously recognized but for which we havenot sampled sufficiently to test their mono-
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phyly (i.e., only one species was available).These are noted in text. We also assume thatthe transformation series supporting themonophyly of the exemplars of any givenclade are correctly extrapolated as being ev-idence of the monophyly of the whole group.In the worse case scenario, we will be shownto be wrong; in the best case our hypotheseswill withstand further testing. By default, wehope our arrangement will stimulate furtherresearch.
The former subfamily Hemiphractinae isnow tentatively considered to be part of theparaphyletic Leptodactylidae, pending fur-ther research on this vast nonmonophyleticconglomerate of hyloids.
The total number of DNA transformationssupporting the monophyly of each relevantclade is informed in the respective diagnoses,with the exception of species groups whosemonophyly has not been tested in our anal-ysis. See figure 8 for a summary of the newtaxonomy and figures 9–12 for the strict con-sensus of our analysis updated with the newtaxonomy. See appendix 5 for details regard-ing the number of transitions, transversions,and inferred insertion/deletion events as wellas the specific positions involved for eachgene.
HYLINAE RAFINESQUE, 1815
SYNONYMS: See sections for tribes.DIAGNOSIS: This subfamily is diagnosed by
32 transformations in nuclear and mitochon-drial proteins and ribosomal genes. See ap-pendix 5 for a complete list of these trans-formations. Possible morphological synapo-morphies are the tendo superficialis digiti V(manus) with an additional tendon that arisesventrally from m. palmaris longus (da Silva,1998, as cited by Duellman, 2001).
COMMENTS: The 2n 5 24 chromosomesmay be another putative synapomorphy ofthis clade, but it will be necessary to betterunderstand its distribution in the more basalmembers of the different tribes.
COPHOMANTINI HOFFMANN, 1878
Cophomantina Hoffmann, 1878. Type genus: Co-phomantis Peters, 1870.
DIAGNOSIS: This tribe is diagnosed by 65transformations in nuclear and mitochondrial
proteins and ribosomal genes. See appendix5 for a complete list of these transformations.Possible morphological synapomorphies in-clude a ventral oral disc, and complete mar-ginal papillae in larvae, (these characterstates subsequently transform in more inclu-sive groups within this clade).
COMMENTS: This tribe includes the generaAplastodiscus, Bokermannohyla new genus,Hyloscirtus, Hypsiboas, and Myersiohylanew genus.
The increase in the number of labial toothrows is likely another synapomorphy of Co-phomantini, because all known larvae of Hy-loscirtus and Myersiohyla new genus colec-tively have a minimum of 6/7 labial toothrows. However, at this time the minimumnumber of labial tooth rows that is synapo-morphic for Cophomantini is ambiguous, be-cause the tadpole is still unknown in H. kan-aima.
An enlarged prepollex is present in allspecies of Hyloscirtus and in most speciesof Myersiohyla, new genus. (Unlike speciesof the H. aromatica group, in H. kanaima,the prepollex is not enlarged.) This charac-teristic could be a synapomorphy of Co-phomantini as an intermediate state leadingto the enlarged prepollex with a projectingspine, as proposed by Duellman et al.(1997). In order to understand whether thischaracter state is a synapomorphy of Copho-mantini, further research is required, includ-ing (1) more osteological work to define thecharacter states involved, and (2) additionalstudies on the phylogenetic relationshipswithin Myersiohyla, new genus, to under-stand whether the character state present inthe former H. kanaima could be interpretedas a reversal.
Burton (2004) suggested that the tendo su-perficialis hallucis tapering from an expand-ed corner of the aponeurosis plantaris, withfibers of the m. transversus plantae distalisoriginating on distal tarsal 2–3 inserting onthe lateral side of the tendon, provides evi-dence of monophyly of a group composed ofthe H. albomarginata, H. albopunctata, H.boans, H. geographica, and H. pulchellagroups. The lack of information on the tax-onomic distribution of this character statewithin several terminals of Cophomantinirenders its optimization ambiguous in all our
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→
Fig. 8. A schematic summary of the new taxonomy of Hylidae proposed here, as indicated by thephylogenetic relationships of the genera of Hylinae, Pelodryadinae, and Phyllomedusinae. The genusPhrynomedusa was unavailable for this study and is not included.
most parsimonious trees. While it is clearthat it is a synapomorphy of some compo-nent of Cophomantini, at this point we do notknow its level of inclusiveness. (Burtonpoints out its presence in H. phyllognatha,the only member of the H. bogotensis groupthat he studied.) The same point holds for thepresence of an accessory tendon of the m.lumbricalis longus digiti III, which Burton(2004) considered characteristic of thosesame species groups.
There are other character states that wereobserved in exemplars of this tribe whosetaxonomic distribution needs to be assessedin its most basal taxa in order to know withmore precision the limits of the clade orclades they diagnose. One of these is thepoint of insertion of the tendon of the m.extensor brevis medius digiti IV that Fai-vovich (2002) found to insert in the medialproximal margin of phalanx 2 in the ex-emplars of this tribe that he studied (Aplas-todiscus perviridis, Hyla albopunctata, H.faber, and H. raniceps). In other hyloidsthis tendon is known to insert in the ante-rior medial margin of metacarpal IV(Burton, 1996, 1998b; Faivovich, 2002).Subsequently, this character state was ob-served in other species available for studies(H. albomarginata, H. andina, H. circum-data, H. clepsydra, H. geographica, H.granosa, H. multifasciata, and H. polytaen-ia; Faivovich, personal obs.).
There are at least two other characterstates whose taxonomic distribution withinthis tribe deserve further scrutiny. The firstof these is the presence in the dorsal surfaceof the larval oral cavity of an anteromedialloop of the prenarial wall into the prenarialarena. Wassersug (1980) described and re-ported it in Hyla rufitela, Spirandeli Cruz(1991) in Aplastodiscus perviridis, H. faber,H. lundii, and H. prasina, and D’Heursel andde Sa (1999) in H. geographica and H. semi-lineata. The second character state is thepresence of one (most frequently) or more (afew species of the H. bogotensis group; Mi-
jares-Urrutia, 1992b) fleshy projections ofvariable shape (triangular, round, or elliptic;sometimes called papillae) in the inner mar-gin of the nostrils of the larvae, which vari-ous authors (Kenny, 1969; Peixoto, 1981;Peixoto and Cruz, 1983; Lavilla, 1984; Mi-jares-Urrutia, 1992b; Wild, 1992; Ayarza-guena and Senaris, ‘‘1993’’ [1994]; de Sa,1995, 1996; Duellman et al., 1997; Faivov-ich, 2002; Gomes and Peixoto, 2002; Fai-vovich, personal obs.) noticed in several spe-cies: Aplastodiscus perviridis, Hyla albofren-ata, H. albomarginata, H. albopunctata, H.albosignata, H. alemani, H. andina, H. aro-matica, H. charazani, H. balzani, H. carval-hoi, H. circumdata, H. faber, H. fasciata, H.granosa, H. inparquesi, H. jahni, H. leuco-pygia, H. multifasciata, H. palaestes, H. pla-tydactyla, H. punctata, H. raniceps, H. sib-leszi, and an undescribed species of the H.aromatica group. Although not explicitlymentioned in the descriptions, this characterstate seems evident in illustrations of otherlarvae: H. goiana, H. joaquini, H. marginata,H. polytaenia, and H. pulchella (Eterovick etal., 2002; Gallardo, 1964; Garcia et al.,2001b, 2003).
Aplastodiscus A. Lutz in B. Lutz, 1950
TYPE SPECIES: Aplastodiscus perviridis A.Lutz in B. Lutz, 1950, by original designa-tion.
DIAGNOSIS: This genus is diagnosed by 72transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these molecular syn-apomorphies. Other apparent synapomor-phies of this clade are the particular repro-ductive modes, where the male constructs asubterranean nest in the muddy side ofstreams and ponds, and where larvae spendearly stages of development; subsequent toflooding, the exotrophic larvae live in pondsor streams (Haddad and Sawaya, 2000; Hart-mann et al., 2004, Haddad et al., 2005). Thepresence of proportionally very developed
2005 77FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
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Fig. 9. A partial view of the strict consensus, updated with the new taxonomy for Phyllomedusinaeproposed here.
metacarpal and metatarsal tubercles is a pos-sible morphological synapomorphy of thegenus (Garcia et al., 2001).
COMMENTS: Our results imply a clade com-posed of Aplastodiscus and two complexesof the former Hyla albomarginata group, asdefined by Cruz and Peixoto (‘‘1985’’[1987]): the H. albofrenata and H. albosig-nata complexes, which are here included inAplastodiscus.
Garcia et al. (2001) suggested four syna-pomorphies for Aplastodiscus as then under-stood (that is, containing only A. cochranaeand A. perviridis): (1) lack of webbing be-tween toes I and II, and very reduced web-bing in the remaining toes, (2) bicolored iris,(3) females with unpigmented eggs, and (4)highly developed inner metacarpal and meta-tarsal tubercles. The lack of webbing be-tween toes I and II, the reduction of webbingamong the remaining toes, and the bicolorediris occur only in the two species originallycontained in Aplastodiscus. These species, A.cochranae and A. perviridis, are here includ-ed in the A. perviridis group, and thereforethese two character states are possibly syna-
pomorphic only of this group, not of Aplas-todiscus as redefined here. The very devel-oped inner metacarpal and metatarsal tuber-cles are also present in all species of the Hylaalbofrenata and H. albosignata complexes(Cruz and Peixoto ‘‘1984’’ [1985], ‘‘1985’’[1987]; Cruz et al., 2003), and we considerthis feature as a putative synapomorphy ofAplastodiscus as redefined here. The pres-ence of unpigmented eggs is known to occurin all species of the former H. albofrenataand H. albosignata complexes with knowneggs (Haddad and Sawaya, 2000; Garcia etal., 2001; Hartmann et al., 2004; Haddad etal., 2005). However, the taxonomic distribu-tion of egg pigmentation within Cophoman-tini is not well known. It is possible that un-pigmented eggs are actually a synapomorphyof a more inclusive clade, as they are knownto occur in at least some species of Hylo-scirtus (H. jahni, H. larinopygion, H. pal-meri, and H. platydactyla; La Marca, 1985,and Faivovich, personal obs.), Hypsiboas (H.lemai, Duellman [1997], and the undescribedspecies here called Hyla sp. 2), and Myers-iohyla new genus (Hyla inparquesi; Faivo-
2005 79FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Fig. 10. A partial view of the strict consensus, updated with the new taxonomy for the tribe Co-phomantini.
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Fig. 11. A partial view of the strict consensus, updated with the new taxonomy for the tribe Den-dropsophini.
vich, Myers, and McDiarmid, in prep.; eggsof M. kanaima are pigmented, Duellman andHoogmoed, 1992); the only known eggs ofspecies of Bokermannohyla, new genus havea pigmented animal pole (Sazima and Bok-ermann, 1977; Eterovick and Brandao,2001).
Most species of Aplastodiscus, as rede-fined here, possess a white parietal perito-neum (Garcia and Faivovich, personal obs.),as it occurs in some other Cophomantini(Hyla bogotensis, H. granosa, and H. punc-tata groups, H. marginata; Ruiz-Carranzaand Lynch [1991: 4]; Garcia [2003]; Faivo-vich, personal obs.). While this could be apossible synapomorphy of Aplastodiscus, the
taxonomic distribution of this character stateis still poorly known in various componentsof the Cophomantini, so we prefer to awaitfurther research on the issue, before hypoth-esizing polarities.
Bokermann (1967c) pointed out the over-all similarity among advertisement calls ofthe Hyla albofrenata and H. albosignatacomplexes and Aplastodiscus perviridis. Fu-ture research will define whether any char-acter state related to the advertisement callscould be considered as a synapomorphy ofAplastodiscus as redefined here, or of any ofits internal clades.
CONTENTS: Fourteen species included inthree species groups.
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Fig. 12. A partial view of the strict consensus, updated with the new taxonomy for the tribes Hyliniand Lophiohylini.
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Aplastodiscus albofrenatus Group
DIAGNOSIS: This species group is diag-nosed by 114 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesetransformations. We are not aware of anymorphological synapomorphies for thisgroup.
CONTENTS: Six species. Aplastodiscus al-bofrenatus (A. Lutz, 1924), new comb.;Aplastodiscus arildae (Cruz and Peixoto,‘‘1985’’ [1987]), new comb.; Aplastodiscusehrhardti (Muller, 1924), new comb.; Aplas-todiscus musicus (B. Lutz, 1948), newcomb.; Aplastodiscus weygoldti (Cruz andPeixoto, ‘‘1985’’ [1987]), new comb.
Aplastodiscus albosignatus Group
DIAGNOSIS: This species group is diag-nosed by 42 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. A possible mor-phological synapomorphy of this group is thepresence of elaborate tubercles and ornamen-tation around the cloacal region (Cruz andPeixoto, ‘‘1985’’ [1987]).
CONTENTS: Seven species. Aplastodiscusalbosignatus (A. Lutz and B. Lutz, 1938),new comb.; Aplastodiscus callipygius (Cruzand Peixoto, ‘‘1984’’ [1985]), new comb.;Aplastodiscus cavicola (Cruz and Peixoto,‘‘1984’’ [1985]), new comb.; Aplastodiscusflumineus (Cruz and Peixoto, ‘‘1984’’[1985]), new comb.; Aplastodiscus ibirapi-tanga (Cruz, Pimenta, and Silvano, 2003),new comb.; Aplastodiscus leucopygius (Cruzand Peixoto, ‘‘1984’’ [1985]), new comb.;Aplastodiscus sibilatus (Cruz, Pimenta, andSilvano, 2003), new comb.
Aplastodiscus perviridis Group
DIAGNOSIS: This species group is diag-nosed by 58 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. Apparent mor-phological synapomorphies of this group in-clude the bicolored iris and the absence ofwebbing between toes I and II (known in-stances of homoplasy within hylids occur in
some Scinax and in various groups of Lo-phiohylini) and reduction of webbing be-tween the other toes (Garcia et al., 2001).
CONTENTS: Two species. Aplastodiscuscochranae (Mertens, 1952); Aplastodiscusperviridis A. Lutz in B. Lutz, 1950.
Bokermannohyla, new genus
TYPE SPECIES: Hyla circumdata Cope,‘‘1870’’ [1871].
DIAGNOSIS: This genus is diagnosed by 65transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy.
ETYMOLOGY: This genus is dedicated toWerner Carlos Augusto Bokermann (1929–1995), as homage to his contribution to theknowledge of Brazilian anurans. He also de-scribed several species now included in thenew genus. The name derives from Boker-mann 1 connecting -o 1 Hyla. We are adopt-ing the ending -hyla for several of the newgenera described here, most of which containspecies groups formerly placed in Hyla. Thegender is feminine.
COMMENTS: Bokermannohyla includes allspecies previously allocated in the Hyla cir-cumdata, H. martinsi, and H. pseudopseudisgroups. We include tentatively the H. clare-signata group pending the inclusion of itsspecies in the analysis, because it was asso-ciated to the H. circumdata group by Bok-ermann (1972) and Jim and Caramaschi(1979). Hyla alvarengai is also included be-cause our analysis shows that Hyla sp. 9 (aff.H. alvarengai) is nested within this new ge-nus. These species groups should be main-tained within Bokermannohyla until theirmonophyly is rigorously tested.
CONTENTS: Twenty-three species, placed infour species groups.
Bokermannohyla circumdata Group
DIAGNOSIS: This species group is diag-nosed by 52 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. A putative mor-phological synapomorphy of this group is thepresence of (usually thin) dark vertical
2005 83FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
stripes on the posterior surface of the thigh(Heyer, 1985).
CONTENTS: Fifteen species. Bokermanno-hyla ahenea (Napoli and Caramaschi, 2004),new comb.; Bokermannohyla astartea (Bok-ermann, 1967), new comb.; Bokermannohylacaramaschii (Napoli, 2005) new comb.; Bok-ermannohyla carvalhoi (Peixoto, 1981), newcomb.; Bokermannohyla circumdata (Cope,‘‘1870’’ [1871]), new comb.; Bokermanno-hyla feioi (Napoli and Caramaschi, 2004),new comb.; Bokermannohyla gouveai (Peix-oto and Cruz, 1992), new comb.; Bokerman-nohyla hylax (Heyer, 1985), new comb.; Bok-ermannohyla ibitipoca (Caramaschi andFeio, 1990), new comb.; Bokermannohyla iz-eckshoni (Jim and Caramaschi, 1979), newcomb.; Bokermannohyla lucianae (Napoliand Pimenta, 2003), new comb.; Bokerman-nohyla luctuosa (Pombal and Haddad, 1993),new comb.; Bokermannohyla nanuzae (Bok-ermann and Sazima, 1973), new comb.; Bok-ermannohyla ravida (Caramaschi, Napoliand Bernardes, 2001), new comb.; Boker-mannohyla sazimai (Cardoso and Andrade,‘‘1982’’ [1983]), new comb.
Bokermannohyla claresignata Group
DIAGNOSIS: We are not aware of any syn-apomorphy supporting the monophyly of thisgroup; see below.
COMMENTS: We did not include any ex-emplar of this group, and as such we did nottest its monophyly. See the earlier discussionregarding its position in the South AmericanI clade. Considering the topology of Co-phomantini, synapomorphies suggested forthe Bokermannohyla claresignata group canonly be maintained if assumed to be reversals(i.e., a enlarged larval oral disc, completemarginal papillae, and large number of labialtooth rows are also present in Myersiohylanew genus and the Hyloscirtus armatusgroup; complete marginal papillae and largenumber of labial tooth rows are also presentin the H. bogotensis and H. larinopygiongroups; the marginal papillae are also com-plete or with an extremely reduced gap inother species of Bokermannohyla). Thiswould be unproblematic if it were a result ofthe analysis, but we prefer not to assume ita priori. While these character states cannot
be considered at this stage to support themonophyly of the B. claresignata group,considering that its two species are barelydistinguishable from each other, we think itsnonmonophyly is unlikely and we continueto recognize the group as a hypothesis to betested.
CONTENTS: Two species. Bokermannohylaclaresignata (A. Lutz and B. Lutz, 1939),new comb.; Bokermannohyla clepsydra (A.Lutz, 1925), new comb.
Bokermannohyla martinsi Group
DIAGNOSIS: Apparent morphological syna-pomorphies of this group are the develop-ment of the humeral crest into a hook-likeprojection, and a bifid prepollex (Boker-mann, 1965b).
COMMENTS: We included a single exemplarof this group, and as such we did not test itsmonophyly. We continue recognizing it onthe basis of the morphological evidence not-ed above.
CONTENTS: Two species. Bokermannohylalangei (Bokermann, 1965), new comb.; Bok-ermannohyla martinsi (Bokermann, 1964),new comb.
Bokermannohyla pseudopseudis Group
DIAGNOSIS: This group is diagnosed by 48transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy for this group.
COMMENTS: Considering our results, whichshow the undescribed species of the Hylapseudopseudis group (Hyla sp. 6) to be thesister taxon of an undescribed species similarwith H. alvarengai (Hyla sp. 9), we are ten-tatively including the H. alvarengai in thisspecies group. Eterovick and Brandao (2001)characterized this group on the basis of thepresence of short, lateral irregular tooth rowsand for having more tooth rows (between sixand eight rows) in the oral discs of the larvaethan do those of the Bokermannohyla cir-cumdata group. However, the tadpole of H.ibitiguara, included in this group by Cara-maschi et al. (2001), has a labial tooth for-mula of 2/4 (Cardoso, 1983) and seems tolack the short, lateral irregular tooth rows, as
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do tadpoles of H. alvarengai (Sazima andBokermann, 1977).
CONTENTS: Four species. Bokermannohylaalvarengai (Bokermann, 1956), new comb.;Bokermannohyla ibitiguara (Cardoso, 1983),new comb.; Bokermannohyla pseudopseudis(Miranda-Ribeiro, 1937), new comb.; Bok-ermannohyla saxicola (Bokermann, 1964),new comb.
Hyloscirtus Peters, 1882
TYPE SPECIES: Hyloscirtus bogotensis Pe-ters, 1882.
Hylonomus Peters, 1882. Type species: Hylono-mus bogotensis Peters, 1882, by monotypy. Pri-mary homonym of Hylonomus Dawson, 1860.
DIAGNOSIS: This genus is diagnosed by 56transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. The only putative morphologi-cal synapomorphy that we are aware of forthis genus is the wide dermal fringes on fin-gers and toes.
COMMENTS: This genus contains all speciesincluded in the Hyla armata, H. bogotensis,and H. larinopygion species groups. Thegroups are maintained unchanged within Hy-loscirtus until the monophyly of each ofthem is properly tested with denser taxonsampling.
While the wide dermal fringes in fingersand toes are present in the three speciesgroups, in the H. armatus group they aremore obvious in the first manual digit. In theH. bogotensis and H. larinopygion groupsthe fringes look even wider, apparently dueto a combination of proportionally smallerdiscs and wider fringe, which gives the fingeror toe the appearance of being almost aswide as the disc.
CONTENTS: Twenty-eight species placed inthree species groups.
Hyloscirtus armatus Group
DIAGNOSIS: This species group is diag-nosed by 103 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. Duellman et al.(1997) suggested four synapomorphies of the
H. armatus group: the presence of keratin-covered bony spines on the proximal ventralsurface of the humerus, on the expanded dis-tal element of the prepollex, and on the firstmetacarpal; forearms hypertrophied; tadpoletail long with low fins and bluntly roundedtip; and the presence of a ‘‘shelf’’ on the lar-val upper jaw sheath.
COMMENTS: Our observations of breedingmales of the two species of this group indi-cate the presence of darkly pigmented, ke-ratinized spicules in the dorsum, head (par-ticularly lips), forelimbs, undersides of fore-limbs, and pectoral and abdominal region. Asthe breeding biology of this and the othertwo species groups of the genus becomesbetter known, it will be possible to under-stand if the presence of these spicules are aputative synapomorphy of the H. armatusgroup.
CONTENTS: Two species. Hyloscirtus ar-matus (Boulenger, 1902), new comb.; Hylos-cirtus charazani (Vellard, 1970), new comb.
Hyloscirtus bogotensis Group
DIAGNOSIS: This species group is diag-nosed by 95 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesetransformations. The only morphologicalsynapomorphy that has been proposed forthis group is the presence of a mental glandin adult males (Duellman, 1972b). See ap-pendix 4 for a justification of the inclusionof ‘‘Hyalinobatrachium’’ estevesi (Rivero,1968) in this species group.
COMMENTS: Species of the Hyloscirtus bo-gotensis group have a white parietal perito-neum (Ruiz-Carranza and Lynch, 1991: 4),as do some other Cophomantini (see com-ments for Aplastodiscus). Further research onthe taxonomic distribution of this characterstate in the other species groups of Hyloscir-tus, and in the other genera of Cophomantini,would clarify which group or groups are di-agnosed by this synapomorphy.
CONTENTS: Sixteen species. Hyloscirtus al-bopunctulatus (Boulenger, 1882), newcomb.; Hyloscirtus alytolylax (Duellman,1972), new comb.; Hyloscirtus bogotensisPeters, 1882; Hyloscirtus callipeza (Duell-man, 1989), new comb.; Hyloscirtus colymba
2005 85FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
(Dunn, 1931), new comb.; Hyloscirtus den-ticulentus (Duellman, 1972), new comb.; Hy-loscirtus estevesi (Rivero, 1968), new comb.;Hyloscirtus jahni (Rivero, 1961), new comb.;Hyloscirtus lascinius (Rivero, 1969), newcomb.; Hyloscirtus lynchi (Ruiz-Carranzaand Ardila-Robayo, 1991), new comb.; Hy-loscirtus palmeri (Boulenger, 1908), newcomb.; Hyloscirtus phyllognathus (Melin,1941), new comb.; Hyloscirtus piceigularis(Ruiz-Carranza and Lynch, 1982), newcomb.; Hyloscirtus platydactylus (Boulenger,1905), new comb.; Hyloscirtus simmonsi(Duellman, 1989), new comb.; Hyloscirtustorrenticola (Duellman and Altig, 1978),new comb.
Hyloscirtus larinopygion Group
DIAGNOSIS: This species group is diag-nosed by 32 transformations in mitochondri-al protein and ribosomal genes. See appendix5 for a complete list of these molecular syn-apomorphies. We are not aware of any mor-phological synapomorphy for this group.
CONTENTS: Ten species. Hyloscirtus cau-canus (Ardila-Robayo, Ruiz-Carranza, andRoa-Trujillo, 1993), new comb.; Hyloscirtuslarinopygion (Duellman, 1973), new comb.;Hyloscirtus lindae (Duellman and Altig,1978), new comb.; Hyloscirtus pacha (Duell-man and Hillis, 1990), new comb.; Hyloscir-tus pantostictus (Duellman and Berger,1982), new comb.; Hyloscirtus psarolaimus(Duellman and Hillis, 1990), new comb.; Hy-loscirtus ptychodactylus (Duellman and Hil-lis, 1990) new comb.; Hyloscirtus saram-piona (Ruiz-Carranza and Lynch, 1982) newcomb.; Hyloscirtus staufferorum (Duellmanand Coloma, 1993), new comb.; Hyloscirtustapichalaca (Kizirian, Coloma, and Paredes-Recalde, 2003), new comb.
Hypsiboas Wagler, 1830
TYPE SPECIES: Hyla palmata Daudin, 1802(5 Rana boans Linnaeus, 1758), by subse-quent designation by implication of Duell-man, 1977 (not monotypy as stated by Duell-man, 1977: 24).
Boana Gray, 1825. Type species: Rana boans Lin-naeus, 1758, by monotypy. Coined as a syno-nym of Hyla and never subsequently validatedas available under article 11.6.1 (ICZN, 1999).
Auletris Wagler, 1830. Type species: Rana boansLinnaeus, 1758, by subsequent designation ofStejneger (1907).
Lobipes Fitzinger, 1843. Type species: Hyla pal-mata Daudin, 1801 (5 Rana boans Linnaeus,1758), by original designation.
Phyllobius Fitzinger, 1843. Type species: Hyla al-bomarginata Spix, 1824, by original designa-tion. Primary homonym of Phyllobius Schoen-herr, 1824.
Centrotelma Burmeister, 1856. Type species: Hylainfulata Wied-Neuwied, 1824 (5 H. albomar-ginata Spix, 1824), by subsequent implicationby Duellman (1977) (not monotypy as stated byDuellman, 1977: 24).
Hylomedusa Burmeister, 1856. Type species: Hylacrepitans Wied-Neuwied, 1825, by subsequentdesignation by implication of Duellman (1977)(not monotypy as stated by Duellman, 1977:24).
Cinclidium Cope, 1867. Type species: Cinclidiumgranulatum Cope, 1867 (5 Rana boans Lin-naeus, 1758), by monotypy. Primary homonymof Cinclidium Blyth, 1842.
Cophomantis Peters, 1870. Type species: Co-phomantis punctillata Peters, 1870 (5 Hylasemilineata Spix, 1824) by monotypy.
Cincliscopus Cope, ‘‘1870’’ [1871]. Replacementname for Cinclidium Cope, 1867.
DIAGNOSIS: This genus is diagnosed by 33transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy for this genus.
COMMENTS: This genus is resurrected forall species formerly included in the Hyla al-bopunctata, H. boans, H. geographica, H.granosa, H. pulchella, and H. punctata spe-cies groups, the H. albomarginata complex,and several species previously unassigned toany group. Most of the former species groupsare retained using the new combinations andits contents are redefined in accordance withour results to avoid paraphyly.
It is tempting to suggest that the presenceof a prepollical spine is a possible synapo-morphy of this group. However, as discussedearlier, further anatomical studies are neces-sary to determine whether this character stateis homologous with that present in Boker-mannohyla.
Duellman (2001) and Savage (2002b) sug-gested that the name Boana Gray, 1825could be applied to a clade of Gladiator
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Frogs. Gray (1825) suggested the nameBoana as a synonym of Hyla, and Stejneger(1907) subsequently designated Hyla boansas its type species. Unfortunately, as far aswe can determine from literature research,the name Boana has never been used in com-bination with an active species name, there-fore failing to fulfill the criterion establishedby article 11.6.1 (ICZN, 1999) for the avail-ability of names originally proposed as syn-onyms. Duellman (2001) further stated thatif Hyla punctata were included within thisgroup, ‘‘the generic Hylaplesia Boie wouldbe included in the synonymy of Boana.’’However, this is not the case since, as dis-cussed by Dubois (1982), the type species ofHylaplesia is Rana tinctoria Cuvier, 1797 (5Dendrobates tinctorius), by subsequent des-ignation of Dumeril and Bibron (1841).
Wagler (1830) did not combine Hypsiboaswith any of the species included in this ge-nus. Subsequent authors (e.g., Tschudi, 1838;Fitzinger, 1843, Cope, 1862) considered itmasculine, as we are doing here.
CONTENTS: Seventy species placed in sev-en species groups, plus two species unas-signed to group.
Hypsiboas albopunctatus Group
DIAGNOSIS: This species group is diag-nosed by 43 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. We are not awareof any morphological synapomorphy.
COMMENTS: To avoid paraphyly, we in-clude Hyla fasciata and H. calcarata in thegroup, and to avoid the creation of a speciesgroup for a single species, we also includethe sister taxon of the former H. albopunc-tata group, H. heilprini. Larvae so far knownof the group (H. albopunctata, H. calcarata,H. fasciata, H. multifasciata, H. raniceps),with the exception of H. heilprini and H. lan-ciformis are reported to have the mediodistalportion of the internal wall of the spiracleseparated from the body wall (de Sa, 1995;1996; Faivovich, 2002, personal obs.; Peix-oto and Cruz, 1983; Wild, 1992). Peixoto andCruz (1983) reported the same character statefor larvae of H. albomarginata. Additionalstudies on the taxonomic distribution of this
peculiar character state are needed to knowthe limits of the clade or clades it diagnoses.Wild (1992) noticed that larvae of H. cal-carata, H. fasciata, and H. lanciformis sharethe presence of pigmented caudal verticalbands that interconnect laterally along themusculature, with this pattern occurring aswell in H. raniceps (Faivovich, personalobs.).
Hyla dentei was originally associated withboth H. raniceps and the former H. geogra-phica group. We are tentatively including itin the Hypsiboas albopunctatus group inview of its overall similarities with Hyla cal-carata and H. fasciata.
CONTENTS: Nine species. Hypsiboas albo-punctatus (Spix, 1824), new comb.; Hypsi-boas calcaratus (Troschel, 1848), newcomb.; Hypsiboas dentei (Bokermann,1967), new comb.; Hypsiboas fasciatus(Gunther, 1858), new comb.; Hypsiboas heil-prini (Noble, 1923), new comb.; Hypsiboaslanciformis Cope, 1871; Hypsiboas leucoch-eilus (Caramaschi and Niemeyer, 2003), newcomb.; Hypsiboas multifasciatus (Gunther,1859), new comb.; Hypsiboas ranicepsCope, 1862.
Hypsiboas benitezi Group
DIAGNOSIS: This species group is diag-nosed by 30 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. A putative syn-apomorphy of this group is the presence ofa flat mental gland in males (Faivovich et al.,in prep.) (known instance of homoplasy inHyla granosa).
COMMENTS: This new species group in-cludes the clade composed of three speciesfrom the Guayana Highlands and three spe-cies from the northwestern Amazon Basin.Because one of the Guayanan and one of theAmazonian species are still undescribed,they are not further considered. Two species,Hyla microderma and H. roraima, are a frag-ment of the former H. geographica group.We are also including tentatively H. hutch-insi and H. rhythmicus, based on their overallsimilarity with H. benitezi. See appendix 4for a justification of the inclusion of Hylapulidoi (Rivero, 1968) in this species group.
2005 87FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
CONTENTS: Seven species. Hypsiboas ben-itezi (Rivero, 1961), new comb.; Hypsiboashutchinsi (Pyburn and Hall, 1984), newcomb.; Hypsiboas lemai (Rivero, 1971), newcomb.; Hypsiboas microderma (Pyburn,1977), new comb.; Hypsiboas pulidoi (Riv-ero, 1968), new comb.; Hypsiboas rhythmi-cus (Senaris and Ayarzaguena. 2002), newcomb.; Hypsiboas roraima (Duellman andHoogmoed, 1992), new comb.
Hypsiboas faber Group
DIAGNOSIS: This species group is diag-nosed by 28 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. We are not awareof any morphological synapomorphy for thisgroup.
COMMENTS: We are including in this groupa clade of some species resulting from thefragmentation of the Hyla boans group. Ourresults indicate that H. crepitans, H. faber,H. lundii, and H. pardalis form, togetherwith H. albomarginata (a component of theHyla albomarginata group), a monophyleticgroup only distantly related to H. boans, thespecies that gives the name to the formergroup. For this reason, we recognize thisclade as the Hypsiboas faber species group.We tentatively include Hyla exastis in thisgroup because Caramaschi and Rodrigues(2003) related it to H. lundii and H. pardalison the basis of the lichenous color patternand the rugose dorsal skin texture.
A likely behavioral synapomorphy of mostspecies of this group, with the exception ofHyla albomarginata, is the construction ofnests by males (with two instances of ho-moplasy within Hylinae, some species of thenow called Hypsiboas semilineatus group(see p. 88), and the Bokermannohyla circum-data group.) The inclusion of Hyla pugnaxand H. rosenbergi is tentative, based on thefact that males construct nests, but they lackthe reticulated palpebral membrane, a likelysynapomorphy present in most species of thenow called Hypsiboas semilineatus group(see below). Future research will test thisbold hypothesis.
CONTENTS: Eight species. Hypsiboas al-bomarginatus (Spix, 1824), new comb.; Hyp-
siboas crepitans (Wied-Neuwied, 1824), newcomb.; Hypsiboas exastis (Caramaschi andRodrigues, 2003), new comb.; Hypsiboas fa-ber (Wied-Neuwied, 1821), new comb.; Hyp-siboas lundii (Burmeister, 1856), new comb.;Hypsiboas pardalis (Spix, 1824), new comb.;Hypsiboas pugnax (O. Schmidt, 1857), newcomb.; Hypsiboas rosenbergi (Boulenger,1898), new comb.
Hypsiboas pellucens Group
DIAGNOSIS: This species group is diag-nosed by 115 transformations in mitochon-drial ribosomal genes. See appendix 5 for acomplete list of these transformations. Weare not aware of any morphological syna-pomorphy for the group.
COMMENTS: We recognize this new speciesgroup to include the clade composed of thefragment of the former Hyla albomarginatacomplex that includes H. pellucens and H.rufitela. The inclusion of H. rubracyla is ten-tative, based on its previous association withH. pellucens.
CONTENTS: Three species. Hypsiboas pel-lucens (Werner, 1901), new comb.; Hypsi-boas rubracylus (Cochran and Goin, 1970),new comb.; Hypsiboas rufitelus (Fouquette,1958), new comb.
Hypsiboas pulchellus Group
DIAGNOSIS: This species group is diag-nosed by 55 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. Observations byFaivovich and Garcia (unpubl.) suggest thatthe absence of the slip of the m. depressormandibulae that originates on the dorsal fas-cia at the level of the m. dorsalis scapularis(present in all other exemplars so far studiedof Hypsiboas, and also of Aplastodiscus, Hy-loscirtus, and Bokermannohyla) is a possiblesynapomorphy of the group.
COMMENTS: We continue to recognizewithin this species group a Hypsiboas poly-taenius clade that includes Hyla beckeri, H.buriti, H. cipoensis, H. goiana, H. latistriata,H. leptolineata, H. phaeopleura, H. poly-taenia, and H. stenocephala. Besides molec-ular data, a likely morphological synapomor-phy that supports this clade is the dorsally
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striped pattern (homoplastic in Hyla bischoffiwhere a striped pattern occurs on some in-dividuals).
CONTENTS: Thirty species. Hypsiboas al-boniger (Nieden, 1923), new comb.; Hypsi-boas andinus (Muller, 1926), new comb.;Hypsiboas balzani (Boulenger, 1898), newcomb.; Hypsiboas beckeri (Caramaschi andCruz, 2004), new comb.; Hypsiboas bischoffi(Boulenger, 1887), new comb.; Hypsiboasburiti (Caramaschi and Cruz, 1999), newcomb.; Hypsiboas caingua (Carrizo, ‘‘1990’’[1991]), new comb.; Hypsiboas callipleura(Boulenger, 1902), new comb.; Hypsiboas ci-poensis (B. Lutz, 1968), new comb.; Hypsi-boas cordobae (Barrio, 1965), new comb.;Hypsiboas cymbalum (Bokermann, 1963),new comb.; Hypsiboas ericae (Caramaschiand Cruz, 2000), new comb.; Hypsiboas frei-canecae (Carnaval and Peixoto, 2004), newcomb.; Hypsiboas goianus (B. Lutz, 1968),new comb.; Hypsiboas guentheri (Boulenger,1886), new comb.; Hypsiboas joaquini (B.Lutz, 1968), new comb.; Hypsiboas latistria-tus (Caramaschi and Cruz, 2004), newcomb.; Hypsiboas leptolineatus (P. Braunand C. Braun, 1977), new comb.; Hypsiboasmarginatus (Boulenger, 1887), new comb.;Hypsiboas marianitae (Carrizo, 1992), newcomb.; Hypsiboas melanopleura (Boulenger,1912), new comb.; Hypsiboas palaestes(Duellman, De la Riva, and Wild, 1997), newcomb.; Hypsiboas phaeopleura (Caramaschiand Cruz, 2000), new comb.; Hypsiboas po-lytaenius (Cope, 1870), new comb.; Hypsi-boas prasinus (Burmeister, 1856), newcomb.; Hypsiboas pulchellus (Dumeril andBibron, 1841), new comb.; Hypsiboas rio-janus (Koslowsky, 1895), new comb.; Hyp-siboas secedens (B. Lutz, 1963), new comb.;Hypsiboas semiguttatus (A. Lutz, 1925), newcomb.; Hypsiboas stenocephalus (Caramas-chi and Cruz, 1999), new comb.
Hypsiboas punctatus Group
DIAGNOSIS: This species group is diag-nosed by 30 transformations in mitochondri-al protein and ribosomal genes. See appendix5 for a complete list of these molecular syn-apomorphies. We are not aware of any mor-phological synapomorphy for the group.
COMMENTS: We do not see any reason to
keep the former Hyla granosa and H. punc-tata groups separated, as our analysis showsthat the two nominal species form a mono-phyletic group and they are phenotypicallysimilar, so we include all the species in theHypsiboas punctatus group. The inclusion ofHyla alemani, H. atlantica, H. hobbsi, andH. ornatissima is tentative, based on theirprevious association with the former H. gra-nosa and H. punctata groups. The inclusionof Hyla picturata is based on our analysis.
CONTENTS: Eight species. Hypsiboas ale-mani (Rivero, 1964), new comb.; Hypsiboasatlanticus (Caramaschi and Velosa, 1996),new comb.; Hypsiboas granosus (Boulenger,1882), new comb.; Hypsiboas hobbsi (Coch-ran and Goin, 1970), new comb.; Hypsiboasornatissimus (Noble, 1923), new comb.;Hypsiboas picturatus (Boulenger, 1882), newcomb.; Hypsiboas punctatus (Schneider,1799), new comb.; Hypsiboas sibleszi (Riv-ero, 1971), new comb.
Hypsiboas semilineatus Group
DIAGNOSIS: This species group is diag-nosed by 128 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. A possible mor-phological synapomorphy of this speciesgroup is the presence of a reticulated palpe-bral membrane (with instances of homoplasyin other species of Hypsiboas: H. hutchinsiand H. microderma).
COMMENTS: We include in this group thefragments of the former Hyla boans and H.geographica groups that form a monophy-letic group. We prefer to call it the Hypsiboassemilineatus group because the use of eitherthe H. boans or H. geographicus groupswould only cause confusion regarding itscontents. The inclusion of Hyla wavrini isbased on the combination of the same repro-ductive mode of H. boans (eggs deposited ina basin built by the male) and a reticulatedpalpebral membrane (Hoogmoed, 1990).Hyla pombali is tentatively included basedon comments by Caramaschi et al (2004a)stressing its similarities with H. semilineata,but with the caveat that it lacks the reticu-lated palpebral membrane. Schooling behav-ior has been reported for tadpoles of H. geo-
2005 89FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
graphica (Caldwell, 1989) and H. semilinea-ta (D’Heursel and Haddad, 2002), and thismay be a synapomorphy for at least thesetwo species.
CONTENTS: Six species. Hypsiboas boans(Linnaeus, 1758), new comb.; Hypsiboasgeographicus (Spix, 1824), new comb.; Hyp-siboas pombali (Caramaschi, Silva, and Feio,2004); Hypsiboas semilineatus (Spix, 1824),new comb.; Hypsiboas wavrini (Parker,1936), new comb.
Species of Hypsiboas Unassigned to Group
There are two species of Hypsiboas thatwe do not assign to any group because wedo not have evidence favoring a relationshipwith any of the species groups that we arerecognizing for the genus. These species areHypsiboas fuentei (Goin and Goin, 1968),new comb., and Hypsiboas varelae (Carrizo,1992), new comb.
Myersiohyla, new genus
TYPE SPECIES: Hyla inparquesi Ayarza-guena and Senaris (‘‘1993’’ [1994]).
DIAGNOSIS: This genus is diagnosed by 48transformations in mitochondrial protein andribosomal genes. See appendix 5 for com-plete list of these molecular synapomorphies.We are not aware of any morphological syn-apomorphy for this group.
ETYMOLOGY: Dedicated to Charles W. My-ers in recognition of his contributions to her-petology, particularly to the herpetofauna ofthe Guayana Highlands. The name derivesfrom Myersius (latinized Myers) 1 connect-ing -o 1 Hyla. The gender is feminine (My-ers and Stothers, MS).
COMMENTS: This new genus includes thespecies of Hyla aromatica group and H. kan-aima, a former member of the H. geogra-phica group. Ayarzaguena and Senaris(‘‘1993’’ [1994]) included the presence of astrong odor in the definition of the Hyla aro-matica group; this could be a possible syn-apomorphy of Myersiohyla. The presence ofa strong odor has yet to be recorded in H.kanaima. It should be noted that the samplewe included of H. inparquesi was not col-lected in the type locality, but in Cerro de laNeblina, ca. 300 km southward.
CONTENTS: Four species. Myersiohyla aro-
matica (Ayarzaguena and Senaris, ‘‘1993’’[1994]), new comb.; Myersiohyla inparquesi(Ayarzaguena and Senaris, ‘‘1993’’ [1994]),new comb.; Myersiohyla loveridgei (Rivero,1961), new comb.; Myersiohyla kanaima(Goin and Wodley, 1961), new comb.
DENDROPSOPHINI FITZINGER, 1843
Dendropsophi Fitzinger, 1843. Type genus: Den-dropsophus Fitzinger, 1843.
DIAGNOSIS: This tribe is diagnosed by 23transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. Apparent morphological syna-pomorphies of this tribe are the absence oflingual papillae in the larvae (known instanc-es of reversal in Lysapsus and Pseudis) andthe absence of nuptial excrescences (with in-stances of homoplasy in some species ofSphaenorhynchus and several Cophomanti-ni).
COMMENTS: This tribe contains the generaDendropsophus, Lysapsus, Pseudis, Scarthy-la, Scinax, Sphaenorhynchus, and Xenohyla.The absence of lingual papillae in the larvaeis the condition reported in all species ofDendropsophus, Scarthyla, and Scinax,whose larvae have been studied (Wassersug,1980; Duellman and de Sa, 1988; Echeverria,1997; Faivovich, 2002; Vera Candioti et al.,2004); a reversal occurs in Lysapsus andPseudis (de Sa and Lavilla, 1997; Vera Can-dioti, 2004). This character state is still un-known in Sphaenorhynchus and Xenohyla.Another possible morphological synapomor-phy is the absence of keratinized nuptial ex-crescences. Duellman et al. (1997) andDuellman (2001) suggested that the absenceof nuptial excrescences was a synapomorphyof the 30-chromosome Hyla. Nuptial excres-cences are also absent in Lysapsus, Pseudis,Scarthyla, Scinax, some species of Sphae-norhynchus, and Xenohyla (Caramaschi,1989; Duellman and Wiens, 1992; Faivovich,personal obs.; Rodriguez and Duellman,1994). Note that, while pigmented kerati-nized structures are absent in all thesegroups, nuptial pads are present at least insome species of Dendropsophus, Scarthyla,and Scinax (Faivovich, personal obs.).
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Dendropsophus Fitzinger, 1843
TYPE SPECIES: Hyla frontalis Daudin, 1800(5 Rana leucophyllata Beireis, 1783), byoriginal designation.
Lophopus Tschudi, 1838. Type species: Hyla mar-morata Daudin (5 Bufo marmoratus Laurenti,1768), by monotypy. Primary homonym of Lo-phopus Dumeril, 1837.
Hylella Reinhardt and Lutken, ‘‘1861’’ [1862].Type species: Hylella tenera Reinhardt and Lut-ken, 1862 (5 Hyla bipunctata Spix, 1824), bysubsequent designation of Smith and Taylor(1948).
Guntheria Miranda-Ribeiro, 1926. Type species:Hyla dasynota Gunther, 1869 (5 Hyla seniculaCope, 1868), by monotypy.
DIAGNOSIS: This genus is diagnosed by 33transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. Karyological evidence is thepresence of 30 chromosomes. Morphologicalsynapomorphies of this clade are possibly theextreme reduction in the quadratojugal (alsooccurs in some Cophomantini and Hylini)and a 1/2 labial tooth row formula (knowninstance of homoplasy in Hyla anceps; sub-sequent reductions in the formula in someclades) (Duellman and Trueb, 1983; Wogelet al., 2000).
COMMENTS: This genus contains all speciesformerly placed in Hyla that are known orsuspected to have 30 chromosomes. How-ever, the fact that the karyotype of its sistertaxon, Xenohyla, is still unknown, precludesthe 30-chromosome condition to be consid-ered a synapomorphy of Dendropsophus, be-cause it could be a synapomorphy of Den-dropsophus 1 Xenohyla. A similar situationoccurs with two muscle characters. Burton(2004) suggested that the m. contrahentishallucis reduced or absent and the presenceof m. flexor teres hallucis are synapomor-phies of this group. Unfortunately, bothtransformations optimize ambiguously be-cause corresponding character states are stillunknown in Xenohyla.
While we consider the extreme reductionof the quadratojugal to be a possible mor-phological synapomorphy of Dendropso-phus, we warn that the condition requiresfurther study, because the quadratojugal is
reduced as well in Sphaenorhynchus and Xe-nohyla (Caramaschi, 1989; Duellman andWiens, 1992; Izecksohn, 1996), although ap-parently not to the level seen in Dendrop-sophus.
Bogart (1973), Gruber (2002), Skuk andLangone (1991), and Kaiser et al. (1996) de-scribed variation in chromosome morpholo-gy for several species of Dendropsophus.
CONTENTS: Eighty-eight species, most ofthem placed in nine species groups, and sev-en unassigned to group.
Dendropsophus columbianus Group
DIAGNOSIS: The only morphological syna-pomorphy suggested for this group is thepresence of two close, triangular lateral spac-es between the cricoid and arytenoids at theposterior part of the larynx (Kaplan, 1999).
COMMENTS: We included a single exemplarof this group, and as such we did not test itsmonophyly, but following Kaplan (1999) werecognize it on the basis of the evidencementioned above.
CONTENTS: Three species. Dendropsophusbogerti (Cochran and Goin, 1970), newcomb.; Dendropsophus carnifex (Duellman,1969), new comb.; Dendropsophus colum-bianus (Boettger, 1892), new comb.
Dendropsophus garagoensis Group
DIAGNOSIS: A possible morphological syn-apomorphy of this group is the internal sur-face of the arytenoids with a small medialdepression (Kaplan, 1999).
COMMENTS: We did not include any ex-emplar of this group in the analysis. We rec-ognize it following Kaplan (1999), who con-sidered Hyla praestans to be the sister taxonof the H. garagoensis group on the basis ofthem sharing the aforementioned putativesynapomorphy. We find it more informativeat this stage to include it in the group thanto consider it as a species unassigned to anygroup.
CONTENTS: Four species. Dendropsophusgaragoensis (Kaplan, 1991), new comb.;Dendropsophus padreluna (Kaplan andRuiz-Carranza, 1997), new comb.; Dendrop-sophus praestans (Duellman and Trueb,1983), new comb.; Dendropsophus viroli-
2005 91FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
nensis (Kaplan and Ruiz-Carranza, 1997),new comb.
Dendropsophus labialis Group
DIAGNOSIS: We are not aware of any syn-apomorphy for this group.
COMMENTS: We included a single exemplarof this group in the analysis, and as such wedid not test its monophyly. Following Duell-man and Trueb (1983) and Duellman (1989),we continue to recognize the group pendinga rigorous test of its monophyly.
CONTENTS: Three species. Dendropsophuslabialis (Peters, 1863), new comb.; Dendrop-sophus meridensis (Rivero, 1961) newcomb.; Dendropsophus pelidna (Duellman,1989), new comb.
Dendropsophus leucophyllatus Group
DIAGNOSIS: This species group is diag-nosed by 35 transformations in mitochondri-al ribosomal genes. See appendix 5 for acomplete list of these molecular synapomor-phies.
COMMENTS: On the basis of our molecularresults, we are including Hyla anceps in thisgroup. With the exception of this species, allother members of the group share the pres-ence of pectoral glands in males and females(Duellman, 1970).
CONTENTS: Eight species. Dendropsophusanceps (A. Lutz, 1929), new comb.; Den-dropsophus bifurcus (Andersson, 1945), newcomb.; Dendropsophus ebraccatus (Cope,1874), new comb.; Dendropsophus elegans(Wied-Neuwied, 1824), new comb.; Den-dropsophus leucophyllatus (Beireis, 1783),new comb.; Dendropsophus rossalleni(Goin, 1959), new comb.; Dendropsophussarayacuensis (Shreve, 1935), new comb.;Dendropsophus triangulum (Gunther,‘‘1868’’ [1869]), new comb.
Dendropsophus marmoratus Group
DIAGNOSIS: This species group is diag-nosed by 73 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. Possible mor-phological synapomorphies of this group arethe warty skin around the margin of the low-
er lip, the crenulated margin of limbs, andthe dorsal marbled pattern (Bokermann,1964b) (instances of homoplasy in other Hy-linae). Furthermore, as it is discernible fromthe illustrations presented by Gomes andPeixoto (1991b) and Peixoto and Gomes(1999), and confirmed by Peixoto (personalcommun. cited in Altig and McDiarmid,1999a), known larvae of this species groupshare the presence of a thick sheath of tissuein the basal portion of the tail muscle andadjacent fins, another likely morphologicalsynapomorphy.
COMMENTS: Bokermann (1964b) diag-nosed this group as having large vocal sacs.While this could be a synapomorphy, we arehesitant to consider it as such until more an-atomical and comparative studies are done inDendropsophus. In connection with the largevocal sacs of the species of this group, Tyler(1971) mentioned that in Hyla marmoratathe pectoral lymphatic septum is modified ina way that permits the inflated sac to intrudeinto sub-humeral spaces.
CONTENTS: Eight species. Dendropsophusacreanus (Bokermann, 1964), new comb.;Dendropsophus dutrai (Gomes and Peixoto,1996), new comb.; Dendropsophus marmor-atus (Laurenti, 1768), new comb.; Dendrop-sophus melanargyreus (Cope, 1887), newcomb.; Dendropsophus nahdereri (B. Lutzand Bokermann, 1963), new comb.; Den-dropsophus novaisi (Bokermann, 1968) newcomb.; Dendropsophus seniculus (Cope,1868), new comb.; Dendropsophus soaresi(Caramaschi and Jim, 1983), new comb.
Dendropsophus microcephalus Group
DIAGNOSIS: This species group is diag-nosed by 42 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. Morphologicalsynapomorphies include the lack of labialtooth rows and marginal papillae (Duellmanand Trueb, 1983) (a reversal occurs in theDendropsophus decipiens clade).
COMMENTS: This group now includes allspecies from the Hyla decipiens, H. micro-cephala, and H. rubicundula groups. We didnot test the monophyly of the H. decipiensor H. rubicundula groups. We continue rec-
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ognition of the Dendropsophus microcepha-lus group and within it, pending a rigoroustest, a D. decipiens clade (including H. ber-thalutzae, H. decipiens, H. haddadi, and H.oliveirai), and a D. rubicundulus clade (in-cluding H. anataliasiasi, H. araguaya, H.cachimbo, H. cerradensis, H. elianeae, H.jimi, H. rhea, H. rubicundula, and H. tri-taeniata). Putative synapomorphies of the D.decipiens clade are the oviposition on leavesoverhanging water (homoplastic with the D.leucophyllatus group and some species of thenow D. parviceps group) and the presence ofa posterior row of marginal papillae (a re-versal). A putative synapomorphy of the D.rubicundulus clade is the green dorsum inlife that changes to pinkish or violet whenpreserved (Napoli and Caramaschi, 1998).
It seems likely that additional synapomor-phies for at least some species of Dendrop-sophus microcephalus group will be hy-pothesized as larval anatomy is carefullystudied. For example, the four species of thegroup studied by Spirandeli Cruz (1991) andWassersug (1980) (H. microcephala, H.nana, H. phlebodes, H. sanborni) showknob-like vestiges of the filter rows in larvae.It also remains to be seen whether the pe-culiarities of the mannicoto glandulare de-scribed by Lajmanovich et al. (2000) for H.nana are common to other larvae of thegroup.
CONTENTS: Thirty-three species. Dendrop-sophus anataliasiasi (Bokermann, 1972),new comb.; Dendropsophus araguaya (Na-poli and Caramaschi, 1998), new comb.;Dendropsophus berthalutzae (Bokermann,1962), new comb.; Dendropsophus bipunc-tatus (Spix, 1824), new comb.; Dendropso-phus branneri (Cochran, 1948), new comb.;Dendropsophus decipiens (A. Lutz, 1925),new comb.; Dendropsophus cachimbo (Na-poli and Caramaschi, 1999), new comb.;Dendropsophus cerradensis (Napoli andCaramaschi, 1998) new comb.; Dendropso-phus cruzi (Pombal and Bastos, 1998), newcomb.; Dendropsophus elianeae (Napoli andCaramaschi, 2000), new comb.; Dendropso-phus gryllatus (Duellman, 1973), new comb.;Dendropsophus haddadi (Bastos and Pom-bal, 1996), new comb.; Dendropsophus jimi(Napoli and Caramaschi, 1999), new comb.;Dendropsophus joannae (Kohler and Lotters,
2001), new comb.; Dendropsophus leali(Bokermann, 1964), new comb.; Dendrop-sophus mathiassoni (Cochran and Goin,1970), new comb.; Dendropsophus meridi-anus (B. Lutz, 1954), new comb.; Dendrop-sophus microcephalus (Cope, 1886), newcomb.; Dendropsophus minusculus (Rivero,1971), new comb.; Dendropsophus nanus(Boulenger, 1889), new comb.; Dendropso-phus oliveirai (Bokermann, 1963), newcomb.; Dendropsophus phlebodes (Stejneger,1906), new comb.; Dendropsophus pseudom-eridianus (Cruz et al., 2000), new comb.;Dendropsophus rhea (Napoli and Caramas-chi, 1999), new comb.; Dendropsophus rho-dopeplus (Gunther, 1858), new comb.; Den-dropsophus robertmertensi (Taylor, 1937),new comb.; Dendropsophus rubicundulus(Reinhardt and Lutken, ‘‘1861’’ [1862]), newcomb.; Dendropsophus sanborni (Schmidt,1944), new comb.; Dendropsophus sartori(Smith, 1951), new comb.; Dendropsophusstuderae (Carvalho e Silva, Carvalho e Silva,and Izecksohn, 2003), new comb.; Dendrop-sophus tritaeniatus (Bokermann, 1965), newcomb.; Dendropsophus walfordi (Boker-mann, 1962), new comb.; Dendropsophuswerneri (Cochran, 1952), new comb.
Dendropsophus minimus Group
DIAGNOSIS: No synapomorphy is knownfor this group.
COMMENTS: We included a single speciesof this group in the analisis, and as such wedid not test its monophyly and we are notaware of any evidence supporting it. Follow-ing Duellman (1982), we continue to recog-nize it pending a rigorous test of its mono-phyly.
CONTENTS: Four species. Dendropsophusaperomeus (Duellman, 1982), new comb.;Dendropsophus minimus (Ahl, 1933), newcomb.; Dendropsophus miyatai (Vigle andGoberdhan-Vigle, 1990), new comb.; Den-dropsophus riveroi (Cochran and Goin,1970), new comb.
Dendropsophus minutus Group
DIAGNOSIS: No synapomorphy is knownfor this group
COMMENTS: We included a single speciesof this group in the analysis, and as such we
2005 93FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
did not test its monophyly. Following Mar-tins and Cardoso (1987), we continue to rec-ognize the species group pending a rigoroustest of its monophyly. Considering similari-ties between Hyla minuta and H. limai (Had-dad, personal obs.), we tentatively includethe latter in the group.
CONTENTS: Four species. Dendropsophusdelarivai (Kohler and Lotters, 2001), newcomb.; Dendropsophus limai (Bokermann,1962), new comb.; Dendropsophus minutus(Peters, 1872), new comb.; Dendropsophusxapuriensis (Martins and Cardoso, 1987),new comb.
Dendropsophus parviceps Group
DIAGNOSIS: This species group is diag-nosed by 27 transformations in nuclear andmitochondrial protein and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. We are not awareof any morphological synapomorphy sup-porting the monophyly of this group.
COMMENTS: We expressed our scepticismregarding the monophyly of this group, ascurrently defined, but found no evidence toreject monophyly. We recognize the grouppending a rigorous test of its monophyly.
CONTENTS: Fifteen species. Dendropso-phus allenorum (Duellman and Trueb, 1989),new comb.; Dendropsophus bokermanni(Goin, 1960), new comb.; Dendropsophusbrevifrons (Duellman and Crump, 1974),new comb.; Dendropsophus gaucheri (Les-cure and Marty, 2001), new comb.; Den-dropsophus giesleri (Mertens, 1950), newcomb.; Dendropsophus grandisonae (Goin,1966), new comb.; Dendropsophus koechlini(Duellman and Trueb, 1989), new comb.;Dendropsophus luteoocellatus (Roux, 1927),new comb.; Dendropsophus microps (Peters,1872), new comb.; Dendropsophus parviceps(Boulenger, 1882), new comb.; Dendropso-phus pauiniensis (Heyer, 1977), new comb.;Dendropsophus ruschii (Weygoldt and Peix-oto, 1987), new comb.; Dendropsophusschubarti (Bokermann, 1963), new comb.;Dendropsophus subocularis (Dunn, 1934),new comb.; Dendropsophus timbeba (Mar-tins and Cardoso, 1987), new comb.
Species of Dendropsophus Unassigned toGroup
There are several species of Dendropso-phus that have not been associated with anygroup. These are: Dendropsophus amicorum(Mijares-Urrutia, 1998), new comb.; Den-dropsophus battersbyi (Rivero, 1961), newcomb.; Dendropsophus haraldschultzi (Bok-ermann, 1962), new comb.; Dendropsophusstingi (Kaplan, 1994), new comb.; Dendrop-sophus tintinnabulum (Melin, 1941), newcomb.; Dendropsophus yaracuyanus (Mija-res-Urrutia and Rivero, 2000), new comb.
Lysapsus Cope, 1862
TYPE SPECIES: Lysapsus limellum Cope,1862, by monotypy.
Podonectes Steindachner, 1864. Type species: Po-donectes palmatus Fitzinger, 1864 (5 Lysapsuslimellum Cope, 1862), by monotypy.
DIAGNOSIS: This genus is diagnosed by 47transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these transformations.A possible morphological synapomorphy ofthis genus is the near absence of subacroso-mal cone in the sperm (Garda et al., 2004).
CONTENTS: Three species. Lysapsus carayaGallardo, 1964; Lysapsus laevis Parker,1935; Lysapsus limellum Cope, 1862.
Pseudis Wagler, 1830
TYPE SPECIES: Rana paradoxa Linnaeus,1758, by monotypy.
Batrachychthys Pizarro, 1876. Type species: notdesignated; based on larvae of Pseudis para-doxa (Linnaeus, 1758), according to Caramas-chi and Cruz (1998).
DIAGNOSIS: This genus is diagnosed by 28transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these transformations.We are not aware of any morphological syn-apomorphy for this genus.
COMMENTS: Garda et al. (2004) distin-guished Lysapsus and Pseudis on the basisof the ultrastructure of the sperm acrosomecomplex, but they stated that the morphologypresent in Lysapsus (near absence of suba-crosomal cone) is the apomorphic condition,
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with only the plesiomorphic condition beingfound in Pseudis and therefore not providingevidence of its monophyly.
CONTENTS: Six species. Pseudis bolbod-actyla A. Lutz, 1925; Pseudis cardosoiKwet, 2000; Pseudis fusca Garman, 1883;Pseudis minuta Gunther, 1858; Pseudis par-adoxa (Linnaeus, 1758); Pseudis tocantinsCaramaschi and Cruz, 1998.
Scarthyla Duellman and de Sa, 1988
TYPE SPECIES: Scarthyla ostinodactyla (5Hyla goinorum Bokermann, 1962), by orig-inal designation.
DIAGNOSIS: Molecular autapomorphies in-clude 227 transformations in nuclear and mi-tochondrial protein and ribosomal genes. Seeappendix 5 for a complete list of these mo-lecular autapomorphies. Apparent morpho-logical autapomorphies include the ability ofits tadpoles to propel themselves out of thewater, elongated tadpoles (Duellman andWiens, 1992) and the presence of a labialarm on the oral disc (McDiarmid and Altig,1990).
COMMENTS: The oral structure known asthe labial arm has also been reported for theScinax rostratus group (McDiarmid and Al-tig, 1990; Faivovich, 2002) and for four oth-er species of Scinax (Heyer et al., 1990; Al-ves and Carvalho e Silva, 2002; Alves et al.,2004). Suarez Mayorga and Lynch (2001b)recently reported a similar structure in thelarvae of Sphaenorhynchus dorisae, addingthat as yet unpublished studies suggest thatthe structures present in Scinax, Scarthyla,and Sphaenorhynchus are not homologs.
CONTENTS: Monotypic. Scarthyla goino-rum (Bokermann, 1962)
Scinax Wagler, 1830
TYPE SPECIES: Hyla aurata Wied-Neuwied1821, by subsequent designation of Stejneger(1907).
Ololygon Fitzinger, 1843. Type species: Hyla stri-gilata, Spix, 1824, by original designation.
Garbeana Miranda-Ribeiro, 1926. Type species:Garbeana garbei Miranda-Ribeiro, 1926, bymonotypy.
DIAGNOSIS: This genus is diagnosed by 83transformations in nuclear and mitochondrial
protein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. Morphological synapomorphiesinclude webbing between toes I and II thatdoes not extend beyond the subarticular tu-bercle of toe I, ability to bend finger I andtoe I, origin of the m. pectoralis abdominalisthrough well-defined tendons, and m. pecto-ralis abdominalis overlapping m. obliquusexternus (da Silva, 1998; Faivovich, 2002).
COMMENTS: Besides the first five characterstates mentioned above, Faivovich (2002)considered as synapomorphies of Scinax theround or poorly expanded sacral diapophy-ses, the occluded frontoparietal fontanelle,single origin of the m. extensor brevis su-perficialis digiti III from the ulnare, and thepresence of the m. lumbricalis longus digitiV that originates from the lateral corner ofthe aponeurosis palmaris. As mentioned ear-lier, the position of Scinax within Hylinaesuggests that outgroups employed by Faivo-vich (2002) are phylogenetically distant fromScinax. Because of this, we contend that thetaxonomic distribution of the aforementionedcharacter states needs to be reassessed, atleast among the other Dendropsophini, be-fore considering them synapomorphies ofScinax. For example, it seems evident thatthe round or poorly expanded sacral diapoph-yses are not a synapomorphy of Scinax, butof a more inclusive group (also present, atleast, in Scarthyla and Sphaenorhynchus;Duellman and Wiens, 1992), whose limitsare still unclear. In the same way, the absenceof the lingual papillae, as mentioned earlier,might be a synapomorphy of Dendropsophi-ni. The truncated discs of the digits wereconsidered a synapomorphy of Scinax byDuellman and Wiens (1992) and Faivovich(2002). The phylogenetic position of the sin-gle exemplar of the H. uruguaya group in theanalysis, as sister group of the S. ruber clade,complicates this interpretation. Discs in thetwo species of the group, H. uruguaya andH. pinima, are proportionally reduced in sizewith respect to most species of Scinax, can-not be considered truncated, and thereforedetermine an ambiguous optimization of thischaracter state.
Burton (2004) considered that m. flexorossis metatarsus IV with insertions on bothmetatarsi IV and V was a synapomorphy of
2005 95FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Scinax. Because this character state is stillunknown in our two exemplars of the S. ca-tharinae clade, and in the exemplar of Hylauruguaya group, it optimizes ambiguously inour analysis; it is unclear if it is a synapo-morphy of Scinax or of a more exclusiveclade.
The Hyla uruguaya group is being includ-ed in Scinax to avoid rendering Scinax para-phyletic. Larvae of the two species of the H.uruguaya group share with members of theS. ruber clade some synapomorphies (theproctodeal tube not reaching the free marginof the lower fin, and the presence of kerati-nized spurs behind the lower jaw sheath andover the infralabial papillae [Kolenc et al.,‘‘2003’’ [2004]]). However, preliminary ob-servations on H. uruguaya indicate thatadults show at least one conflicting characterstate, the m. depressor mandibulae withoutan origin from the dorsal fascia at the levelof the m. dorsalis scapulae (Faivovich, per-sonal obs.)—a character state that optimizedas a synapomorphy of the S. catharinae cladein the analysis of Faivovich (2002). Thiscontroversy may be resolved when all theconflicting evidence is analyzed, including amuch denser sampling of Scinax. In themeantime, since the molecular evidence in-dicates affinities with the S. ruber clade, wetentatively include the two species of the H.uruguaya group in this clade, where they arerecognized as a separate group.
CONTENTS: Eighty-eight species placed intwo major clades.
Scinax catharinae Clade
DIAGNOSIS: This clade is diagnosed by 90transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these molecular syn-apomorphies. Morphological synapomor-phies suggested for this clade by Faivovich(2002) are absence of the anterior process ofthe suprascapula, internal vocal sac, distal di-vision of the middle branch of the m. exten-sor digitorum comunis longus, and insertionof the medial side of this branch on the ten-don of the m. extensor brevis medius digitiIV.
COMMENTS: Regardless of problems im-posed by the present results to interpretation
of the possible synapomorphies of Scinax re-sulting from Faivovich’s (2002) analysis, thesparse available knowledge on the taxonomicdistribution of the transformations supportingthe monophyly of the very distinctive S. ca-tharinae clade suggests that most of themstill hold in the present analysis. An excep-tion is the m. depressor mandibulae withoutan origin from the dorsal fascia at the levelof the m. dorsalis scapulae, which also oc-curs in Hyla uruguaya, rendering its opti-mization ambiguous in our analysis.
Scinax catharinae Group
DIAGNOSIS: Because we only included twospecies of the Scinax catharinae group as ex-emplars of the S. catharinae clade, the mo-lecular transformations that diagnose thisgroup are redundant with those diagnosingthe S. catharinae clade. Presumed morpho-logical synapomorphies of this group includethe posterior part of the cricoid ring exten-sively elongated and curved, the partial min-eralization of intercalary elements betweenultimate and penultimate phalanges, and thelaterodistal origin of the m. extensor brevisdistalis digiti III (Faivovich, 2002).
CONTENTS: Twenty-seven species. Scinaxagilis (Cruz and Peixoto, 1983); Scinax al-bicans (Bokermann, 1967); Scinax angrensis(B. Lutz, 1973); Scinax argyreornatus (Mi-randa-Ribeiro, 1926); Scinax ariadne (Bok-ermann, 1967); Scinax aromothyella Faivov-ich, 2005; Scinax berthae (Barrio, 1962);Scinax brieni (De Witte, 1927); Scinax can-astrensis (Cardoso and Haddad, 1982); Sci-nax carnevalli (Caramaschi and Kisteumach-er, 1989); Scinax catharinae (Boulenger,1888); Scinax centralis (Pombal and Bastos,1996); Scinax flavoguttatus (A. Lutz and B.Lutz, 1939); Scinax heyeri (Peixoto andWeygoldt, 1986); Scinax hiemalis (Haddadand Pombal, 1987); Scinax humilis (B. Lutz,1954); Scinax jureia (Pombal and Gordo,1991); Scinax kautskyi (Carvalho e Silva andPeixoto, 1991); Scinax littoralis (Pombal andGordo, 1991); Scinax longilineus (B. Lutz,1968); Scinax luizotavioi (Caramaschi andKisteumacher, 1989); Scinax machadoi(Bokermann and Sazima, 1973); Scinax ob-triangulatus (B. Lutz, 1973); Scinax ranki(Andrade and Cardoso, 1987); Scinax rizi-
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bilis (Bokermann, 1964); Scinax strigilatus(Spix, 1824); and Scinax trapicheiroi (B.Lutz, 1954).
Scinax perpusillus Group
DIAGNOSIS: Presumed synapomorphies ofthis group are the oviposition in bromeliadsand the extreme reduction of webbing be-tween toes II and III (Peixoto, 1987; Faivov-ich, 2002).
COMMENTS: The monophyly of this groupwas not tested by Faivovich (2002) becauseonly one species of the group was availablefor his analysis, where it obtained as the sis-ter taxon of all exemplars of the S. cathari-nae clade. For these two reasons, we contin-ue recognizing this group until its monophy-ly is rigorously tested.
CONTENTS: Seven species. Scinax alcatraz(B. Lutz, 1973); Scinax arduous Peixoto,2002; Scinax atratus (Peixoto, 1989); Scinaxlittoreus (Peixoto, 1988); Scinax melloi(Peixoto, 1989); Scinax perpusillus (A. Lutzand B. Lutz, 1939); Scinax v-signatus (B.Lutz, 1968).
Scinax ruber Clade
DIAGNOSIS: This clade is supported by 53transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. A morphological synapomorphysuggested for this clade by Faivovich (2002)is the proctodeal tube positioned above themargin of the lower fin.
COMMENTS: Faivovich (2002) was skepti-cal about the monophyly of the S. ruberclade; however, the present analysis recoversit as monophyletic, with a considerable num-ber of transformations supporting its mono-phyly.
As in the case of several of the synapo-morphies suggested by Faivovich’s (2002)analysis for Scinax, we are unsure as towhether the suggested morphological syna-pomorphies are optimized identically in ouranalysis. In particular, we do not know thetaxonomic distribution within Dendropsophi-ni for two other synapomorphies proposedfor this clade (Faivovich, 2002): the aryte-noids with a dorsal prominence developedover the pharyngeal margin, and absence of
the lateral m. extensor brevis distalis digiti V(pes). Preliminary observation on the larvaeof some species of Sphaenorhynchus (Sphae-norhynchus bromelicola, S. orophilus, S.pauloalvini, and S. prasinus; Faivovich, per-sonal obs.) indicate that their proctodealtubes are attached to the free margin of thelower fin, similar to the S. catharinae clade,instead of having the characteristic positionseen in larvae of the S. ruber clade.
Scinax megapodius and S. trachythoraxare considered here to be junior synonyms ofS. fuscovarius for reasons discussed in ap-pendix 4. There are two species, Hyla dolloiand H. karenanneae, that upon examinationof their type series we consider to be speciesof Scinax (see appendix 4 for further com-ments on them).
CONTENTS: Fifty-six species. Eleven as-signed to two groups, 43 unassigned to anygroup.
Scinax rostratus Group
DIAGNOSIS: Putative morphological syna-pomorphies of this group include the juxta-posed inner margins of the vomers; overlapof the otic plate of the crista parotica due toa broad otic plate; nonfenestration of the car-tilaginous plate of the squamosal with theoblique cartilage; pointed tubercle on heel;absence of the m. extensor brevis distalisdigiti II; presence of m. extensor brevis dis-talis digiti I (pes); discontinuity of lateralmargins with the posterior portion of the oraldisc; third posterior labial tooth row placedon a labial arm; reduction of the third pos-terior labial tooth to one-quarter the lengthof the second row; absence of keratinizedspurs behind the lower jaw sheath; and head-down calling position (Faivovich, 2002).
CONTENTS: Nine species. Scinax boulen-geri (Cope, 1887); Scinax garbei (Miranda-Ribeiro, 1926); Scinax jolyi (Lescure andMarty, 2001); Scinax kennedyi (Pyburn,1973); Scinax nebulosus (Spix, 1824); Scinaxpedromedinae (Henle, 1991); Scinax probos-cideus (Brongersma, 1933); Scinax rostratus(Peters, 1870); Scinax sugillatus (Duellman,1973).
Scinax uruguayus Group
DIAGNOSIS: Putative morphological syna-pomorphies of this group include the bicol-
2005 97FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
ored iris and the presence of two keratinizedand pigmented plates on the sides of the low-er jaw sheath (Kolenc et al., ‘‘2003’’ [2004]).
COMMENTS: The marginal papillae of theposterior margin of the oral disc being largerthan those of the lateral margins (Kolenc etal., ‘‘2003’’ [2004]) and the reduction in toewebbing could be other synapomorphies ofthe group.
CONTENTS: Two species. Scinax pinima(Bokermann and Sazima, 1973) new comb.;Scinax uruguayus (Schmidt, 1944) newcomb.
Species of the Scinax ruber Clade Unas-signed to a Species Group
We follow Faivovich (2002) in not rec-ognizing the former Scinax ruber and S.staufferi groups, as both were not monophy-letic on his analysis. We are considering allspecies formerly included in these groups asmembers of the S. ruber clade, although weconsider them as unassigned to any group.These species are Scinax acuminatus (Cope,1862); Scinax altae (Dunn, 1933); Scinax al-ter (B. Lutz, 1973); Scinax auratus (Wied-Neuwied, 1821); Scinax baumgardneri (Riv-ero, 1961); Scinax blairi (Fouquette and Py-burn, 1972); Scinax boesemani (Goin, 1966);Scinax caldarum (B. Lutz, 1968); Scinaxcardosoi (Carvalho e Silva and Peixoto,1991); Scinax castroviejoi De la Riva, 1993;Scinax chiquitanus (De la Riva, 1990); Sci-nax crospedospilus (A. Lutz, 1925); Scinaxcruentommus (Duellman, 1972); Scinax cur-icica Pugliese, Pombal, and Sazima, 2004;Scinax cuspidatus (A. Lutz, 1925); Scinaxdanae (Duellman, 1986); Scinax dolloi (Wer-ner, 1898) new comb.; Scinax duartei (B.Lutz, 1951); Scinax elaeochrous (Cope,1875); Scinax eurydice (Bokermann, 1964);Scinax exiguus (Duellman, 1986); Scinaxflavidus La Marca, 2004; Scinax funereus(Cope, 1874); Scinax fuscomarginatus (A.Lutz, 1925); Scinax fuscovarius (A. Lutz,1925); Scinax granulatus (Peters, 1871); Sci-nax hayii (Barbour, 1909); Scinax ictericusDuellman and Wiens, 1993; Scinax karen-anneae (Pyburn, 1992) comb. nov.; Scinaxlindsayi Pyburn, 1992; Scinax manriqueiBarrio-Amoros, Orellana, and Chacon, 2004;Scinax maracaya (Cardoso and Sazima,
1980); Scinax nasicus (Cope, 1862); Scinaxoreites Duellman and Wiens, 1993; Scinaxpachycrus (Miranda-Ribeiro, 1937); Scinaxparkeri (Gaige, 1926); Scinax perereca Pom-bal, Haddad, and Kasahara, 1995; Scinaxquinquefasciatus (Fowler, 1913); Scinax rub-er (Laurenti, 1768); Scinax similis (Cochran,1952); Scinax squalirostris (A. Lutz, 1925);Scinax staufferi (Cope, 1865); Scinax trili-neatus (Hoogmoed and Gorzula, 1977); Sci-nax wandae (Pyburn and Fouquette, 1971);Scinax x-signatus (Spix, 1824).
Sphaenorhynchus Tschudi, 1838
TYPE SPECIES: Hyla lactea Daudin, 1801,by original designation.
Dryomelictes Fitzinger, 1843. Type species: Hylalactea Daudin, 1802, by original designation.
Dryomelictes Cope, 1865. Type species: Hyla au-rantiaca Daudin, 1802, by original designation.Junior homonym of Dryomelictes Fitzinger,1843.
Hylopsis Werner, 1894. Type species: Hylopsisplatycephalus Werner, 1894, by monotypy.
Sphoenohyla Lutz and Lutz, 1938. Substitutename (explicit subgenus of Hyla) for Sphae-norhynchus thought erroneously to be preoc-cupied by Sphenorhynchus Lichtenstein, 1823.
DIAGNOSIS: This genus is diagnosed by157 transformations in nuclear and mito-chondrial protein and ribosomal genes. Seeappendix 5 for a complete list of these mo-lecular synapomorphies. Duellman andWiens (1992) proposed the following syna-pomorphies for Sphaenorhynchus: posteriorramus of pterygoid absent; zygomatic ramusof squamosal absent or reduced to a smallknob; pars facialis of maxilla and alary pro-cess of premaxilla reduced; postorbital pro-cess of maxilla reduced, not in contact withquadratojugal; neopalatine reduced to a sliveror absent; pars externa plectri entering tym-panic ring posteriorly (rather than dorsally);pars externa plectri round; hyale curved me-dially; coracoids and clavicle elongated; andprepollex ossified, bladelike. Other likelysynapomorphies include the differentiationof the m. intermandibularis into a small api-cal supplementary element, and the extremedevelopment of the m. interhyoideus (Tyler,1971).
COMMENTS: Duellman and Wiens (1992)
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considered that the transverse process of pre-sacral vertebra IV elongate, oriented poste-riorly is a synapomorphy of Sphaenorhyn-chus. The presence of this character state inXenohyla (Izecksohn, 1996) suggests that inthe context of our topology, its optimizationis ambiguous. There are also some larval fea-tures that could be considered synapomor-phies of at least some species of Sphaeno-rhynchus, such as the morphology and po-sition of the nostrils and the presence ofsome notably large marginal papillae (seeKenny, 1969; Bokermann, 1973; Cruz, 1973;Cruz and Peixoto, 1980; Suarez-Mayorgaand Lynch, 2001b). The presence of a whiteperitoneum in five species (S. carneus, S.lacteus, S. planicola, S. prasinus, and S. sur-dus; Haddad and Faivovich, personal obs.)may be another synapomorphy of this genus(with several instances of homoplasy withinHylinae). Observations on six species (S.carneus, S. dorisae, S. lacteus, S. planicola,S. prasinus, and S. surdus) suggest that theyare ant specialists (Duellman, 1978; Rodri-guez and Duellman, 1994; Parmalee, 1999;Haddad, personal obs.), another likely syna-pomorphy whose taxonomic distributionwithin the group deserves additional study.
CONTENTS: Eleven species. Sphaenorhyn-chus bromelicola Bokermann, 1966; Sphae-norhynchus carneus (Cope, 1868); Sphae-norhynchus dorisae (Goin, 1967); Sphaeno-rhynchus lacteus (Daudin, 1801); Sphaeno-rhynchus orophilus (A. Lutz and B. Lutz,1938); Sphaenorhynchus palustris Boker-mann, 1966; Sphaenorhynchus pauloalviniBokermann, 1973; Sphaenorhynchus plani-cola (A. Lutz and B. Lutz, 1938); Sphaeno-rhynchus platycephalus (Werner, 1894);Sphaenorhynchus prasinus Bokermann,1973; Sphaenorhynchus surdus (Cochran,1953).
Xenohyla Izecksohn, 1996
TYPE SPECIES: Hyla truncata Izecksohn,1959, by original designation.
DIAGNOSIS: For the purposes of this paper,we consider that the 128 transformations inmitochondrial protein and ribosomal genesautapomorphic of Xenohyla truncata are syn-apomorphies of this genus. See appendix 5for a complete list of these molecular syna-
pomorphies. Although species of Xenohylaare very distinctive, we are aware of onlythree putative morphological synapomor-phies: the retention in adults of the scars ofthe windows of forelimbs emergence (but seeComments below); the presence of a small,transverse process in the urostyle; and fru-givorous habits (reported for X. truncata byda Silva et al. [1989] and Izecksohn [1996];unknown in X. eugenioi).
COMMENTS: We included a single speciesof this genus in the analysis, and as such wedid not test its monophyly, but consider itvery likely on the basis of the evidence notedabove and its unique external aspect. Izeck-sohn (1996) and Caramaschi (1998) noticedthat adults of Xenohyla retain scars of thelarge windows of forelimb emergence thatare evident in recently metamorphosed indi-viduals. Each of these scars actually corre-sponds to a thick pectoral patch of glandsthat is macroscopically evident upon super-ficial dissection (Faivovich, personal obs.).
CONTENTS: Two species. Xenohyla eugen-ioi Caramaschi, 2001; Xenohyla truncata (Iz-ecksohn, 1959).
HYLINI RAFINESQUE, 1815
Hylarinia Rafinesque, 1815. Type genus: HylariaRafinesque, 1814 (an unjustified emendation ofHyla Laurenti, 1768).
Hylina Gray, 1825. Type genus: Hyla Laurenti,1768.
Dryophytae Fitzinger, 1843. Type genus: Dry-ophytes Fitzinger, 1843.
Acridina Mivart, 1869. Type genus: Acris Du-meril and Bibron, 1841.
Triprioninae Miranda-Ribeiro, 1926. Type genus:Triprion Cope, 1866.
DIAGNOSIS: This tribe is diagnosed by 107transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. The only known morphologicalsynapomorphy is the undivided tendon of them. flexor digitorum brevis superficialis (thereare several instances of homoplasy withinHylidae including at least Scinax, Scarthyla1 Pseudis, and a reversal within Hylini).
COMMENTS: The tribe Hylini is proposedfor the clade of Middle American/Holarctichylids. It includes Acris, Anotheca, Duell-manohyla, Exerodonta, Hyla, Pseudacris,
2005 99FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Ptychohyla, Smilisca (including Pternohyla),Triprion, and six new genera, Bromeliohylanew gen., Charadrahyla new gen., Ecnom-iohyla new gen., Isthmohyla new gen., Me-gastomatohyla new gen., and Tlalocohylanew gen. Morescalchi (1973) recognized thetribe Hylini in which he included most gen-era currently placed in Hylinae. Subsequentauthors have not used Hylini in the sense thatMorescalchi (1973) used it.
Acris Dumeril and Bibron, 1841
TYPE SPECIES: Rana gryllus LeConte,1825, by subsequent designation of Fitzinger(1843).
DIAGNOSIS: This genus is diagnosed by138 transformations in nuclear and mito-chondrial proteins and ribosomal genes. Seeappendix 5 for a complete list of these mo-lecular synapomorphies. Other apparent syn-apomorphies include the differentiation ofthe m. intermandibularis into an apical sup-plementary element (Tyler, 1971), and dip-loid chromosome number of 22 (Bushnell etal., 1939; Cole, 1966; Duellman, 1970).
CONTENTS: Two species. Acris crepitansBaird, 1854; Acris gryllus (LeConte, 1825).
Anotheca Smith, 1939
TYPE SPECIES: Gastrotheca coronata Ste-jneger, 1911 (5 Hyla spinosa Steindachner,1864), by original designation.
DIAGNOSIS: This monotypic genus is di-agnosed by 219 transformations of nuclearand mitochondrial protein and ribosomalgenes. See appendix 5 for a complete list ofthese transformations. Morphological auta-pomorphies include the tendo superficialishallucis that tapers from an expanded cornerof the aponeurosis plantaris; with fibers ofthe m. transversus plantae distalis originatingon distal tarsal 2–3 inserting on the lateralside of the tendon (several of homoplasy, seeappendix 1); the unique skull ornamentationcomposed of sharp, dorsally pointed spinesin the margins of frontoparietal, maxilla, na-sal (including canthal ridge), and squamosal,and character states that result in its repro-ductive mode, including maternal provision-ing of trophic eggs to tadpoles (see Jungfer,1996).
CONTENTS: Monotypic. Anotheca spinosa(Steindachner, 1864).
Bromeliohyla, new genus
TYPE SPECIES: Hyla bromeliacia Schmidt,1933.
DIAGNOSIS: For the purposes of this paperwe consider that the 141 transformations innuclear and mitochondrial protein and ribo-somal genes autapomorphic of Bromeliohylabromeliacia are synapomorphies of this ge-nus. See appendix 5 for a complete list ofthese molecular synapomorphies. Possiblenonmolecular synapomorphies of this genusare the reproductive mode, where eggs arelaid in water accumulated in bromeliads(several instances of homoplasy, e.g., twospecies of Isthmohyla, Phyllodytes, somespecies Osteopilus, and the Scinax perpusil-lus group), and tadpoles with dorsoventrallyflattened bodies and elongated tails.
ETYMOLOGY: From Bromelia 1 Hyla, inreference to the bromeliad breeding habits ofits species. The gender is feminine.
COMMENTS: We included a single speciesof this genus, and as such we did not test itsmonophyly. We consider it likely based onthe evidence noted above.
CONTENTS: Two species. Bromeliohylabromeliacia (Schmidt, 1933), new comb.;Bromeliohyla dendroscarta (Taylor, 1940),new comb.
Charadrahyla, new genus
TYPE SPECIES: Hyla taeniopus Gunther,1901.
DIAGNOSIS: This genus is diagnosed by 56transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these molecular syn-apomorphies. We are not aware of any mor-phological synapomorphy supporting this ge-nus.
ETYMOLOGY: Derived from the Greek wordcharadra- (ravine) 1 Hyla. In reference tothe habits of these frogs. The gender is fem-inine.
COMMENTS: This new genus includes thespecies formerly placed in the Hyla taenio-pus group.
CONTENTS: Five species. Charadrahyla al-tipotens (Duellman, 1968), new comb.;
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Charadrahyla chaneque (Duellman, 1961),new comb.; Charadrahyla nephila (Mendel-son and Campbell, 1999), new comb.; Char-adrahyla taeniopus (Gunther, 1901), newcomb.; Charadrahyla trux (Adler and Denis,1972), new comb.
Duellmanohyla Campbell and Smith, 1992
TYPE SPECIES: Hyla uranochroa Cope,1876, by original designation.
DIAGNOSIS: This genus is diagnosed by 48transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. Likely morphological synapo-morphies of this group are the red iris, thelabial stripe expanded below orbit, the lackof nuptial excrescences, the ventrally orient-ed funnel-shaped oral disc in larvae, labialtooth rows reduced in length, and absence oflateral processes on upper jaw sheath (Duell-man, 2001).
CONTENTS: Eight species. Duellmanohylachamulae (Duellman, 1961); Duellmanohylaignicolor (Duellman, 1961); Duellmanohylalythrodes (Savage, 1968); Duellmanohyla ru-fioculis (Taylor, 1952); Duellmanohyla sal-vavida (McCranie and Wilson, 1986); Duell-manohyla schmidtorum (Stuart, 1954);Duellmanohyla soralia (Wilson and Mc-Cranie, 1985); Duellmanohyla uranochroa(Cope, 1876).
Ecnomiohyla, new genus
TYPE SPECIES: Hypsiboas miliarius Cope,1886.
DIAGNOSIS: This genus is diagnosed by 37transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy supporting this ge-nus.
ETYMOLOGY: From the Greek, ecnomios,meaning marvelous, unusual; an obvious ref-erence to the incredible frogs of the Hylatuberculosa group. The gender is feminine.
COMMENTS: This new genus contains theHyla tuberculosa group, excluding H. den-drophasma, and including one species of theH. miotympanum group as well. Erecting anew genus for this clade is the only way of
being consistent with the new monophyletictaxonomy that is proposed for hylids. Al-though naming the former H. tuberculosagroup as a genus constitutes a testable claimof monophyly, we expect that it will ulti-mately be found to be two or three differentclades, with one of these being the onenamed here.
CONTENTS: Ten species. Ecnomiohylaechinata (Duellman, 1962), new comb.; Ec-nomiohyla fimbrimembra (Taylor, 1948), newcomb.; Ecnomiohyla miliaria (Cope, 1886),new comb.; Ecnomiohyla minera (Wilson,McCranie, and Williams, 1985), new comb.;Ecnomiohyla miotympanum (Cope, 1863),new comb.; Ecnomiohyla phantasmagoria(Dunn, 1943); Ecnomiohyla salvaje (Wilson,McCranie, and Williams, 1985), new comb.;Ecnomiohyla thysanota (Duellman, 1966),new comb.; Ecnomiohyla tuberculosa (Bou-lenger, 1882), new comb.; Ecnomiohyla va-lancifer (Firschein and Smith, 1956), newcomb.
Exerodonta Brocchi, 1879
TYPE SPECIES: Exerodonta sumichrastiBrocchi, 1879, by monotypy.
DIAGNOSIS: This genus is diagnosed by 80transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these molecular syn-apomorphies. We are not aware of any mor-phological synapomorphy supporting this ge-nus.
COMMENTS: Exerodonta is resurrected forthe species previously placed in the Hylasumichrasti group and a fragment of the for-mer H. miotympanum group as defined byDuellman (2001) that corresponds to the tra-ditionally recognized H. pinorum group(Duellman 1970). Although we did not in-clude the type species, E. sumichrasti, in theanalysis, but only H. chimalapa and H. xera,we consider that these two species and H.sumichrasti and H. smaragdina are so similarthat we are not hesitant to consider themclosely related. Although we are not awareof any synapomorphy supporting the mono-phyly of the former H. pinorum group(Duellman, 1970), and our results suggest itis paraphyletic with respect to the H. sumi-chrasti group, we are tentatively including
2005 101FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
the other species associated with it, H. ab-divita, H. bivocata, H. catracha, and H.juanitae by Snyder (1972), Porras and Wil-son (1987), and Campbell and Duellman(2000), in this resurrected genus.
CONTENTS: Eleven species, four placed inone species group, seven unassigned togroup.
Exerodonta sumichrasti Group
DIAGNOSIS: This species group is diag-nosed by 76 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. Putative mor-phological synapomorphies of this group arethe massive nasals (Duellman, 1970) and, inthe species with a known tadpole, the en-larged larval oral disc (homoplastic with theformer Hyla mixomaculata group), and the3/6 to 7 labial tooth row formula (Canseco-Marquez et al., 2003; Duellman, 1970).
COMMENTS: The illustrations of the oraldiscs of Hyla smaragdina, H. sumichrasti(Duellman, 1970), and H. xera (Canseco-Marquez et al., 2003) show that they sharethe multiple interruption of the last posteriorlabial tooth row into shorter rows, possiblyanother synapomorphy.
CONTENTS: Four species. Exerodonta chi-malapa (Mendelson and Campbell, 1994),new comb.; Exerodonta smaragdina (Taylor,1940), new comb.; Exerodonta sumichrastiBrocchi, 1879; Exerodonta xera (Mendelsonand Campbell, 1994), new comb.
Species of Exerodonta Unassigned toGroup
Considering that in our analysis Hyla me-lanomma and H. perkinsi are a grade leadingto the E. sumichrasti group, we are not as-signing to any group these and the other spe-cies associated with the former Hyla pinorumgroup (Duellman, 1970; Snyder, 1972; Porrasand Wilson, 1987; Campbell and Duellman,2000). These species are: Exerodonta abdi-vita (Campbell and Duellman, 2000), newcomb.; Exerodonta bivocata (Duellman andHoyt, 1961), new comb.; Exerodonta catra-cha (Porras and Wilson, 1987), new comb.;Exerodonta juanitae (Snyder, 1972), newcomb.; Exerodonta melanomma (Taylor,
1940), new comb.; Exerodonta perkinsi(Campbell and Brodie, 1992), new comb.;Exerodonta pinorum (Taylor, 1937), newcomb.
Hyla Laurenti, 1768
TYPE SPECIES: Hyla viridis Laurenti, 1768(5 Rana arborea Linnaeus, 1758), by sub-sequent designation of Stejneger (1907).
Calamita Schneider, 1799. Type species: Rana ar-borea Linnaeus, 1758, by subsequent designa-tion of Stejneger (1907).
Hylaria Rafinesque, 1814. Unjustified emendationfor Hyla.
Hyas Wagler, 1830. Type species: Rana arboreaLinnaeus, 1758, by monotypy. Junior homonymof Hyas Leach, 1815.
Dendrohyas Wagler, 1830. Substitute name forHyas Wagler, 1830.
Dryophytes Fitzinger, 1843. Type species: Hylaversicolor LeConte, 1825, by original designa-tion.
Epedaphus Cope, 1885. Type species: Hyla gra-tiosa LeConte, 1856, by monotypy.
DIAGNOSIS: This genus is diagnosed by 25transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy for the genus.
COMMENTS: Hyla is restricted to all speciespreviously placed in the H. arborea, H. ci-nerea, H. eximia, and H. versicolor groups,which are redefined herein.
CONTENTS: Thirty-two species, with 31placed in four species groups and one speciesunassigned to group.
Hyla arborea Group
DIAGNOSIS: This species group is diag-nosed by 37 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. We are not awareof any morphological synapomorphy sup-porting this group.
COMMENTS: The contents of the Hyla ar-borea group are restricted to avoid its para-phyly. The inclusion of the species that werenot included in the present analysis and donot show the NOR in chromosome 6 is ten-tative, because no evidence, other than the
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molecular data presented here, is known tosupport its monophyly.
CONTENTS: Fourteen species. Hyla annec-tans (Jerdon, 1870); Hyla arborea (Linnaeus,1758); Hyla chinensis Gunther, 1858; Hylahallowellii Thomson, 1912; Hyla immacula-ta Boettger, 1888; Hyla intermedia Boulen-ger, 1882; Hyla meridionalis Boettger, 1874;Hyla sanchiangensis Pope, 1929; Hyla sarda(De Betta, 1853); Hyla savignyi Audouin,1827; Hyla simplex Boettger, 1901; Hylatsinlingensis Liu and Hu, 1966; Hyla ussur-iensis Nikolsky, 1918; Hyla zhaopingensisTang and Zhang, 1984.
Hyla cinerea Group
DIAGNOSIS: This species group is diag-nosed by 35 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. We are not awareof any morphological synapomorphy sup-porting this group.
COMMENTS: Hyla femoralis is excludedfrom the H. cinerea group to avoid the par-aphyly of the group.
CONTENTS: Three species. Hyla cinerea(Schneider, 1799); Hyla gratiosa LeConte,‘‘1856’’ [1857]; Hyla squirella Bosc, 1800.
Hyla eximia Group
DIAGNOSIS: This species group is diag-nosed by 17 transformations in mitochondri-al proteins and ribosomal genes. See appen-dix 5 for a complete list of these molecularsynapomorphies. We are not aware of anymorphological synapomorphy supportingthis group.
COMMENTS: The inclusion of Hyla suweo-nensis is tentative, based on the fact that An-derson (1991) reported a NOR in chromo-some 6, a character state shared by H. fe-moralis and the H. eximia and H. versicolorgroups.
CONTENTS: Eleven species. Hyla anderson-ii Baird, 1854; Hyla arboricola Taylor, 1941;Hyla arenicolor Cope, 1886; Hyla bocourti(Mocquard, 1889); Hyla euphorbiacea Gun-ther, 1859; Hyla eximia Baird, 1854; Hylajaponica Gunther, ‘‘1858’’ [1859]; Hyla pli-cata Brocchi, 1877; Hyla suweonensis Ku-
ramoto, 1980; Hyla walkeri Stuart, 1954;Hyla wrightorum Taylor, 1939.
Hyla versicolor Group
DIAGNOSIS: This species group is diag-nosed by 51 transformations in nuclear andmitochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesemolecular synapomorphies. We are not awareof any morphological synapomorphy sup-porting this group.
COMMENTS: Hyla andersonii is transferredto the H. eximia group to avoid the paraphylyof the H. versicolor group.
CONTENTS: Three species. Hyla avivocaViosca, 1928; Hyla chrysoscelis Cope, 1880;Hyla versicolor LeConte, 1825.
Species of Hyla Unassigned to Group
Considering that relationships of Hyla fe-moralis Bosc, 1800 with the H. versicolorand H. eximia groups are unresolved, we pre-fer to keep this species unassigned as a morestable alternative to merging the H. versicol-or and the H. eximia groups into a singlegroup.
Isthmohyla, new genus
TYPE SPECIES: Hyla pseudopuma Gunther,1901.
DIAGNOSIS: This genus is diagnosed by 42transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy for the genus.
ETYMOLOGY: From Isthmo, Greek, in ref-erence to the mostly isthmian distribution ofthese frogs (the only exception is Hyla in-solita) 1 Hyla. The gender is feminine.
COMMENTS: This new genus includes allspecies of the Hyla pseudopuma and H. pic-tipes groups, as defined by Duellman (2001),with the exception of H. thorectes, which istransferred to Plectrohyla. Our taxon sam-pling of the relevant species groups was toosparse to result in a test of their respectivemonophyly. We tentatively recognize thesespecies groups as reviewed by Duellman(2001), with the exception that H. thorectesis excluded from the former H. pictipesgroup and not included in Isthmohyla.
2005 103FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
CONTENTS: Fourteen species placed in twospecies groups.
Isthmohyla pictipes Group
DIAGNOSIS: We are not aware of any syn-apomorphy supporting the monophyly of thisgroup.
COMMENTS: We included a single exemplarof this group, and as such we did not test itsmonophyly. It is being tentatively recognizedfollowing Duellman (2001) until a rigoroustest is performed. This species group isformed by the former Hyla lancasteri, H.pictipes, H. rivularis, and H. zeteki groups(Duellman, 1970, 2001). The monophyly ofthe former H. zeteki group does not seem tobe controversial, as its two species sharemassive temporal musculature, bromeliaddwelling larvae, a terminal oral disc, and alabial tooth row formula of 1/1. The mono-phyly of the group composed of the formerH. rivularis and H. pictipes groups (as de-fined by Duellman, 1970) is supported by thepresence of an enlarged oral disc (Duellman,2001) with a broad band of conic submar-ginal papillae on the posterior part of thedisc, about three rows on the anterior part,and an M-shaped upper jaw sheath27 (Fai-vovich, personal obs.; see also illustrations inDuellman, 2001). The monophyly of the for-mer H. lancasteri group seems to be sup-ported by the presence of granular dorsalskin (Duellman, 2001; known homoplasticinstance in H. debilis), a short snout, and thepresence of dark ventral pigmentation (Wil-son et al., 1994b, known homoplastic in-stance in H. thorectes.)
CONTENTS: Ten species. Isthmohyla calyp-sa (Lips, 1996), new comb.; Isthmohyla de-bilis (Taylor, 1952), new comb.; Isthmohylainsolita (McCranie, Wilson, and Williams,1993), new comb.; Isthmohyla lancasteri(Barbour, 1928), new comb.; Isthmohyla pi-cadoi (Dunn, 1937), new comb.; Isthmohylapictipes (Cope, 1876), new comb.; Isthmo-hyla rivularis (Taylor, 1952), new comb.;Isthmohyla tica (Starret, 1966), new comb.;Isthmohyla xanthosticta (Duellman, 1968),
27 The description and illustrations of the tadpole ofHyla debilis by Duellman (1970) do not show thesecharacter states.
new comb.; Isthmohyla zeteki (Gaige, 1929),new comb.
Isthmohyla pseudopuma Group
DIAGNOSIS: We are not aware of any syn-apomorphy supporting the monophyly of thisgroup.
COMMENTS: We included a single exemplarof this group, and as such we did not test itsmonophyly. It is being tentatively recognizedfollowing Duellman (2001) until a rigoroustest is performed.
CONTENTS: Four species. Isthmohyla an-gustilineata (Taylor, 1952), new comb.; Isth-mohyla graceae (Myers and Duellman,1982), new comb.; Isthmohyla infucata(Duellman, 1968), new comb.; Isthmohylapseudopuma (Gunther, 1901), new comb.
Megastomatohyla, new genus
TYPE SPECIES: Hyla mixe Duellman, 1965.DIAGNOSIS: For the purposes of this paper,
we consider that the 209 transformations innuclear and mitochondrial protein and ribo-somal genes autapomorphic of Hyla mixe aresynapomorphies of this genus. See appendix5 for a complete list of these molecular syn-apomorphies. A possible morphological syn-apomorphy of this genus is the greatly en-larged oral disc of the known larvae bearing7–10 anterior rows and 10–11 posteriorrows.
ETYMOLOGY: From the Greek, mega, large,plus the stem of the genitive stomatos,mouth, in reference to the enlarged oral discof the larvae 1 Hyla. The gender is feminine.
COMMENTS: We included a single speciesof this genus, and as such we did not test itsmonophyly, but consider it very likely on thebasis of the evidence noted above. As men-tioned earlier, the sequenced sample comesfrom a tadpole that was assigned to the Hylamixomaculata group based on the enlargedoral disc and the labial tooth row formulaand tentatively assigned to H. mixe for beingthe only species of the group known fromthe region where it was collected. Consider-ing the uncertainty in its determination, itsposition in the tree should be viewed cau-tiously. This is not a situation we feel mostcomfortable with, but for a matter of beingconsistent with the general approach of this
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contribution, we consider that it is better todescribe a new genus for the H. mixomacu-lata group than to leave it as incerta sedis.Although we did not test the monophyly ofthe group, as stated earlier, we consider itbased on the morphological synapomorphyfor larvae mentioned above. Another possi-ble synapomorphy of this group could be thelack of vocal slits (Duellman, 1970), but thisis contingent on the internal relationships ofthe nearby Charadrahyla; C. chaneque is theonly species of that genus known to lack vo-cal slits (Duellman, 2001). If future studiesshow it to be the sister group of the remain-ing species of Charadrahyla, it could renderthe optimization of the lack of vocal slits asambiguous for both Charadrahyla and Me-gastomatohyla. Males of species included inMegastomatohyla lack nuptial excrescenceson the thumb (Duellman, 1970). The polarityof this character state is unclear because italso occurs in Charadrahyla altipotens andin Hyla godmani and H. loquax (Duellman,1970).
CONTENTS: Four species. Megastomatohylamixe (Duellman, 1965), new comb.; Megas-tomatohyla mixomaculata (Taylor, 1950),new comb.; Megastomatohyla nubicola(Duellman, 1964), new comb.; Megastoma-tohyla pellita (Duellman, 1968), new comb.
Plectrohyla Brocchi, 1877
TYPE SPECIES: Plectrohyla guatemalensisBrocchi, 1877, by original designation.
Cauphias Brocchi, 1877. Replacement name forPlectrohyla Brocchi, 1877.
DIAGNOSIS: This genus is supported by 43transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy supporting this ge-nus as redefined here.
COMMENTS: We are including in Plectro-hyla all species formerly placed in the Hylabistincta group and some of the members ofthe former H. miotympanum (H. cyclada andH. arborescandens) and H. pictipes (H. tho-rectes) groups. H. thorectes is being tenta-tively included because a still undescribedspecies, very similar to H. thorectes (Hylasp. 5) is nested within this clade. Hyla ha-
zelae is tentatively included because of itssimilarities with H. thorectes. Technicallyour results are certainly compatible with therecognition of a separate genus for the mem-bers of the H. bistincta group and the fewspecies from other groups associated withthem. However, we are particularly con-cerned that the present, clean separation be-tween Plectrohyla and these exemplars prob-ably will not hold when more species of thetwo clades, particularly from the H. bistinctagroup, are added. The facts that no apparentmorphological synapomorphies are knownfor the H. bistincta group and that some au-thors raised doubts regarding the limits be-tween it and Plectrohyla support the conser-vative stance of including all these species inPlectrohyla. We preserve a Plectrohyla gua-temalensis group for all the species of Plec-trohyla as defined in the past and tentativelyrecognize a group that contains all membersof the H. bistincta group plus the species ofother groups shown to be related with it inthis analysis.
The reasons why we are not consideringsome of the characters states shared by Plec-trohyla and the Hyla bistincta group thatwere advanced by Duellman (2001) as syn-apomorphies of the redefined Plectrohylawere discussed earlier in this paper (p. 68).The only character state that seems to be in-clusive of Plectrohyla and the H. bistinctagroup is the long medial ramus of the pter-ygoid in contact with the otic capsule. How-ever, both H. arborescandens and H. cycladawere reported by Duellman (2001) to have ashort medial ramus that does not contact theprootic. In a more densely sampled context,this character state could probably be inter-preted as a reversal; however, in the presentcontext it optimized ambiguously, so we donot consider it a morphological synapomor-phy of the redefined Plectrohyla.
CONTENTS: Thirty-nine species placed intwo species groups.
Plectrohyla bistincta Group
DIAGNOSIS: Exemplars of this speciesgroup in our analysis are diagnosed by 16transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these transformations.
2005 105FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
We are not aware of any morphological syn-apomorphy supporting this group.
CONTENTS: Twenty-one species. Plectro-hyla ameibothalame (Canseco-Marquez,Mendelson, and Gutierrez-Mayen et al.,2002), new comb.; Plectrohyla arborescan-dens (Taylor, ‘‘1938’’[1939]), new comb.;Plectrohyla bistincta (Cope, 1877), newcomb.; Plectrohyla calthula (Ustach, Men-delson, McDiarmid, and Campbell, 2000),new comb.; Plectrohyla calvicollina (Toal,1994), new comb.; Plectrohyla celata (Toaland Mendelson, 1995), new comb.; Plectro-hyla cembra (Caldwell, 1974), new comb.;Plectrohyla charadricola (Duellman, 1964),new comb.; Plectrohyla chryses (Adler,1965), new comb.; Plectrohyla crassa (Broc-chi, 1877), new comb.; Plectrohyla cyanom-ma (Caldwell, 1974), new comb.; Plectro-hyla cyclada (Campbell and Duellman,2000) new comb.; Plectrohyla hazelae (Tay-lor, 1940), new comb.; Plectrohyla labedac-tyla (Mendelson and Toal, 1996), new comb.;Plectrohyla mykter (Adler and Dennis,1972), new comb.; Plectrohyla pachyderma,(Taylor, 1942), new comb.; Plectrohyla pen-theter (Adler, 1965), new comb.; Plectrohylapsarosema (Campbell and Duellman, 2000),new comb.; Plectrohyla robertsorum (Taylor,1940), new comb.; Plectrohyla sabrina(Caldwell, 1974), new comb.; Plectrohylasiopela, (Duellman, 1968), new comb.; Plec-trohyla thorectes (Adler, 1965), new comb.
Plectrohyla guatemalensis Group
DIAGNOSIS: This group is diagnosed by 34transformations in nuclear and mitochondrialproteins and ribosomal genes. See appendix5 for a complete list of these transformations.Possible morphological synapomorphies ofthis species group are bifurcate alary processof the premaxilla; sphenethmoid with ante-rior part ossified; frontoparietals abbutingposteriorly, exposing only small part of thefrontoparietal fontanelle; humerus havingwell-developed flanges; hypertrophied fore-arm; prepollex enlarged and ossified in bothsexes; prepollex truncate; and absence of lat-eral labial folds in larvae (Duellman andCampbell, 1992; Duellman, 2001).
CONTENTS: Eighteen species. Plectrohylaacanthodes Duellman and Campbell, 1992;
Plectrohyla avia Stuart, 1952; Plectrohylachrysopleura Wilson, McCranie, and Cruz-Dıaz, 1994; Plectrohyla dasypus McCranieand Wilson, 1981; Plectrohyla exquisitiaMcCranie and Wilson, 1998; Plectrohylaglandulosa (Boulenger, 1883); Plectrohylaguatemalensis Brocchi, 1877; Plectrohylahartwegi Duellman, 1968; Plectrohyla ixilStuart, 1942; Plectrohyla lacertosa Bumhan-zen and Smith, 1954; Plectrohyla matudaiHartweg, 1941; Plectrohyla pokomchi Duell-man and Campbell, 1984; Plectrohyla psi-loderma McCranie and Wilson, 1992; Plec-trohyla pycnochila Rabb, 1959; Plectrohylaquecchi Stuart, 1942; Plectrohyla sagorumHartweg, 1941; Plectrohyla tecunumaniDuellman and Campbell, 1984; Plectrohylateuchestes Duellman and Campbell, 1992.
Pseudacris Fitzinger, 1843
TYPE SPECIES: Rana nigrita LeConte,1825, by monotypy.
Chorophilus Baird, 1854. Type species: Rana ni-grita LeConte, 1825, by original designation.
Helocaetes Baird, 1854. Type species: Hyla tris-eriata Wied-Neuwied, 1839, by subsequentdesignation of Schmidt (1953).
Hyliola Mocquard, 1899. Type species: Hyla re-gilla Baird and Girard, 1852, by subsequentdesignation of Stejneger (1907).
Limnaoedus Mittleman and List, 1953. Type spe-cies: Hyla ocularis Bosc and Daudin, 1801, byoriginal designation.
Parapseudacris Hardy and Borrough, 1986. Typespecies: Hyla crucifer Wied-Neuwied, 1838, byoriginal designation.
DIAGNOSIS: This genus is diagnosed by 37transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. The m. transversus metatarsus IIbroad, occupying the entire length of meta-tarsus II optimizes in this analysis as a mor-phological synapomorphy of this genus.
CONTENTS: Fourteen species placed in fourclades.
Pseudacris crucifer Clade
DIAGNOSIS: This clade is diagnosed by mo-lecular data presented by Moriarty and Can-natella (2004).
CONTENTS: Two species. Pseudacris cru-
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cifer (Wied-Neuwied, 1838); Pseudacrisocularis (Bosc and Daudin, 1801).
Pseudacris ornata Clade
DIAGNOSIS: This clade is diagnosed by mo-lecular data presented by Moriarty and Can-natella (2004).
CONTENTS: Three species. Pseudacris illi-noensis Smith, 1951; Pseudacris ornata(Holbrook, 1836); Pseudacris streckeri A.A.Wright and A.H. Wright, 1933.
Pseudacris nigrita Clade
DIAGNOSIS: This clade is diagnosed by mo-lecular data presented by Moriarty and Can-natella (2004).
COMMENTS: According to Moriarty andCannatella (2004), this clade contains a moreexclusive clade that contains all species butP. brimleyi and P. brachyphona.
CONTENTS: Seven species. Pseudacris bra-chyphona (Cope, 1889); Pseudacris brimleyiBrandt and Walker, 1933; Pseudacris clarkii(Baird, 1854); Pseudacris feriarum (Baird,1854); Pseudacris maculata (Agassiz, 1850);Pseudacris nigrita (LeConte, 1825); Pseu-dacris triseriata (Wied-Neuwied, 1838).
Pseudacris regilla Clade
DIAGNOSIS: This clade is diagnosed by mo-lecular data presented by Moriarty and Can-natella (2004).
CONTENTS: Two species. Pseudacris ca-daverina (Cope, 1866); Pseudacris regilla(Baird and Girard, 1852).
Ptychohyla Taylor, 1944
TYPE SPECIES: Ptychohyla adipoventrisTaylor, 1944 (5 Hyla leonhardschultzei Ahl,1934), by original designation.
DIAGNOSIS: This genus is diagnosed by 11transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. An apparent morphological syn-apomorphy of this group is the well-devel-oped lingual flange of the pars palatina ofpremaxillar (Campbell and Smith, 1992).
COMMENTS: To avoid paraphyly, we are in-cluding Hyla dendrophasma in Ptychohyla.As mentioned earlier in the discussion, other
synapomorphies of Ptychohyla proposed byCampbell and Smith (1992) and Duellman(2001) are also present in some species of itssister taxon (Bromeliohyla 1 Duellmanohy-la), so we do not recognize them as syna-pomorphies until a phylogenetic analysis in-cluding that evidence is performed. Knownmales of the exemplars of Ptychohyla in-cluded in the analysis share the presence ofenlarged individual nuptial spines and hy-pertrophied ventrolateral glands (Duellman,2001), as do males of P. macrotympanumand P. panchoi. Discovery of males of H.dendrophasma will confirm whether thischaracter state is an apparent synapomorphyof these species or of a less inclusive clade.
CONTENTS: Thirteen species. Ptychohylaacrochorda Campbell and Duellman, 2000;Ptychohyla dendrophasma (Campbell,Smith, and Acevedo, 2000), new comb.; Pty-chohyla erythromma (Taylor, 1937); Pty-chohyla euthysanota Kellogg, 1928; Pty-chohyla hypomykter McCranie and Wilson,1993; Ptychohyla legleri (Taylor, 1958); Pty-chohyla leonhardschultzei (Ahl, 1934); Pty-chohyla macrotympanum (Tanner, 1957);Ptychohyla panchoi Duellman and Camp-bell, 1982; Ptychohyla salvadorensis (Mer-tens, 1952); Ptychohyla sanctaecrucis Camp-bell and Smith, 1992; Ptychohyla spinipollex(Schmidt, 1936); Ptychohyla zophodesCampbell and Duellman, 2000.
Smilisca Cope, 1865
TYPE SPECIES: Smilisca daulinia Cope,1865 (5 Hyla baudinii Dumeril and Bibron,1841), by monotypy.
Pternohyla Boulenger, 1882. Type species Pter-nohyla fodiens Boulenger, 1882, by monotypy.NEW SYNONYMY.
DIAGNOSIS: This genus is diagnosed by 38transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy.
COMMENTS: Pternohyla is included in thesynonymy of Smilisca to avoid paraphyly.
CONTENTS: Eight species. Smilisca baudi-nii (Dumeril and Bibron, 1841); Smilisca cy-anosticta (Smith, 1953); Smilisca dentata(Smith, 1957), new comb.; Smilisca fodiens
2005 107FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
(Boulenger, 1882), new comb.; Smiliscaphaeota (Cope, 1862); Smilisca puma (Cope,1885); Smilisca sila (Duellman and Trueb,1966); Smilisca sordida (Peters, 1863).
Tlalocohyla, new genus
TYPE SPECIES: Hyla smithii Boulenger,1902.
DIAGNOSIS: This genus is diagnosed by 92transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy.
ETYMOLOGY: From Tlaloc, the Olmec Godof the rain, 1 connecting -o 1 Hyla. Thegender is feminine.
COMMENTS: The inclusion of Hyla god-mani and H. loquax is tentative and based onits association with H. picta and H. smithiiin the former H. godmani group by Duellman(2001). The larvae of H. loquax and H. smi-thii share a reduction in the length of thethird posterior tooth row (Caldwell, 1986;Lee, 1996). This feature is not present in thelarvae of H. godmani as described by Duell-man (1970).
CONTENTS: Four species. Tlalocohyla god-mani (Gunther, 1901), new comb.; Tlalocoh-yla loquax (Gaige and Stuart, 1934), newcomb.; Tlalocohyla picta (Gunther, 1901),new comb.; Tlalocohyla smithii (Boulenger,1902), new comb.
Triprion Cope, 1866
TYPE SPECIES: Pharyngodon petasatusCope, 1865, by monotypy.
Pharyngodon Cope, 1865. Junior homonym ofPharyngodon Diesing, 1861. Type species:Pharyngodon petasatus Cope, 1865, by mono-typy.
Diaglena Cope, 1887. Type species: Triprion spa-tulatus Gunther, 1882, by monotypy.
DIAGNOSIS: For the purposes of this paperwe consider that the 125 transformations innuclear and mitochondrial protein and ribo-somal genes autapomorphic of Triprion pe-tasatus are synapomorphies of this genus.See appendix 5 for a complete list of thesemolecular synapomorphies. In Duellman’s(2001) phylogenetic analysis of Pternohyla,
Smilisca, and Triprion, the monophyly ofTriprion is supported by three synapomor-phies28: maxilla greatly expanded laterally;prenasal bone present (known homoplasticinstance in Aparasphenodon); and presenceof parasphenoid odontoids.
COMMENTS: We included a single speciesof this genus, and as such we did not test itsmonophyly, but we do not consider it con-troversial on the basis of the morphologicalevidence mentioned above.
CONTENTS: Two species. Triprion petasa-tus (Cope, 1865); Triprion spatulatus Gun-ther, 1882.
LOPHIOHYLINI MIRANDA-RIBEIRO, 1926
Lophiohylinae Miranda-Ribeiro, 1926.Type genus: Lophyohyla Miranda-Ribeiro,1926.
Trachycephalinae B. Lutz, 1969. Type genus: Tra-chycephalus Tschudi, 1838.
DIAGNOSIS: This tribe is diagnosed by 63transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. A putative morphological syn-apomorphy of this tribe is the presence of atleast four posterior labial tooth rows in thelarval oral disc (e.g., Bokermann, 1966b;Duellman, 1974; de Sa, 1983; Lannoo et al.,1987; McDiarmid and Altig, 1990; Schiesariet al., 1996; da Silva in Altig and Mc-Diarmid, 1999b; Wogel et al., 2000) (rever-sals in Osteopilus marianae, O. crucialis, O.wilderi [Dunn, 1926] and in Osteocephalusoophagus [Jungfer and Schiesari, 1995]).
COMMENTS: This tribe contains all SouthAmerican and West Indian casque-headedfrogs and related groups. It includes Apar-asphenodon, Argenteohyla, Corythomantis,Osteopilus, Phyllodytes, Tepuihyla, a newmonotypic genus, and the genera Osteoce-phalus and Trachycephalus as redefinedhere.
Recently, Kasahara et al. (2003) noticedthat Aparasphenodon brunoi, Corythomantis
28 Note that on his preferred tree (fig. 410) one ofthese character transformations is numbered 18, whichseems to be a typographical error for 12, the only othercharacter that supports this clade but that is not shownin the tree.
108 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
greeningi, and Osteocephalus langsdorffiishare similar chromosome morphology,where there is a clear discontinuity in thechromosome lengths of the first five pairsand the remaining seven pairs. Furthermore,they share the presence of a secondary con-striction in pair 10. Available information onkaryotypes of other casque-headed frogs ofthis clade suggests that the discontinuity inchromosome lengths occurs as well in Ar-genteohyla (apparent from plates publishedby Morand and Hernando, 1996), Phrynoh-yas venulosa (apparent from plates publishedby Bogart, 1973), and some species of Os-teopilus (O. brunneus, O. dominicensis, O.marianae, O. septentrionalis), but not in Os-teocephalus taurinus, the only species of thegenus Osteocephalus, as redefined here,whose karyotype was studied (Anderson,1996). The position of the secondary con-striction also varies, having been observed inchromosome 4 in Argenteohyla (Morand andHernando, 1996), chromosome 9 in Osteo-pilus brunneus, O. dominicensis, O. septen-trionalis, and O. wilderi (Anderson, 1996),chromosome 10 in Phrynohyas venulosa (ap-parent from plates published by Bogart,1973), and in chromosome 12 in Osteoce-phalus taurinus. The taxonomic distributionof these character states needs further studyto define the inclusiveness of the clades theysupport.
Delfino et al. (2002) noticed that serousskin glands of Osteopilus septentrionalis andPhrynohyas venulosa produce secretorygranules with a dense cortex and a pale me-dulla; they observed the same in a photo-graph of a section of skin of Corythomantisgreeningi published by Toledo and Jared(1995). Very few hylid taxa were studied forserous gland histology, and these include afew species of Phyllomedusa, HolarcticHyla, Scinax, and Pseudis paradoxa (seeDelfino et al., 2001, 2002). The taxonomicdistribution of these peculiar secretory gran-ules requires additional study to assess itslevel of generality and the clade or cladesthat it diagnoses.
Aparasphenodon Miranda-Ribeiro, 1920
TYPE SPECIES: Aparasphenodon brunoiMiranda-Ribeiro, 1920, by monotypy.
DIAGNOSIS: For the purposes of this paperwe consider that the 83 transformations innuclear and mitochondrial protein and ribo-somal genes autapomorphic of Aparasphen-odon brunoi are synapomorphies of this ge-nus. See appendix 5 for a complete list ofthese molecular synapomorphies. A possiblemorphological synapomorphy of this genusis the presence of a prenasal bone (Trueb,1970a.)
COMMENTS: We included a single speciesof this genus, and as such we did not test itsmonophyly, but we consider it possible basedon the morphological evidence mentionedabove.
CONTENTS: Three species. Aparaspheno-don bokermanni Pombal, 1993; Aparasphen-odon brunoi Miranda-Ribeiro, 1920; Apara-sphenodon venezolanus (Mertens, 1950).
Argenteohyla Trueb, 1970
TYPE SPECIES: Hyla siemersi Mertens,1937, by original designation.
DIAGNOSIS: Molecular autapomorphies in-clude 102 transformations in nuclear and mi-tochondrial protein and ribosomal genes. Seeappendix 5 for a complete list of these mo-lecular synapomorphies. Apparent morpho-logical autapomorphies of this taxon includethe articulation of the zygomatic ramus of thesquamosal with the pars fascialis of the max-illary, and the noticeable reduction in the sizeof discs of fingers and toes (Trueb, 1970b.)
CONTENTS: Monotypic. Argenteohyla sie-mersi (Mertens, 1937).
Corythomantis Boulenger, 1896
TYPE SPECIES: Corythomantis greeningiBoulenger, 1896, by monotypy.
DIAGNOSIS: Molecular autapomorphies in-clude 132 transformations in nuclear and mi-tochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesetransformations. Morphological autapomor-phies of this monotypic genus include the ab-sence of palatines, and nasals that conceal thealary processes of premaxillaries (Trueb,1970a).
CONTENTS: Monotypic. Corythomantisgreeningi Boulenger, 1896.
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Itapotihyla, new genus
TYPE SPECIES: Hyla langsdorffii Dumeriland Bibron, 1841.
DIAGNOSIS: Molecular autapomorphies in-clude 122 transformations in nuclear and mi-tochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesetransformations. A possible morphologicalautapomorphy is the presence of a prominentsubcloacal flap.
ETYMOLOGY: From Itapoti 1 -Hyla. Thegeneric name is an allusion to the resem-blance of the unique known species of thisgenus with lichens and mosses. Itapoti is aTupi-Guarani term, a composition of ‘‘ita’’(5 rock) with ‘‘poti’’ (5 flower or to flour-ish), which means lichen or moss.
CONTENTS: Monotypic. Itapotihyla langs-dorffii (Dumeril and Bibron, 1841), newcomb.
Nyctimantis Boulenger, 1882
TYPE SPECIES: Nyctimantis rugiceps Bou-lenger, 1882, by monotypy.
DIAGNOSIS: Molecular autapomorphies in-clude 139 transformations in mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. Possible morphological autapo-morphies are the development of an irregularorbital flange in the frontoparietal, and thesphenethmoid almost completely concealeddorsally by the frontoparietals and nasals(Duellman and Trueb, 1976).
CONTENTS: Monotypic. Nyctimantis rugi-ceps Boulenger, 1882.
Osteocephalus Steindachner, 1862
TYPE SPECIES: Osteocephalus taurinusSteindachner, 1862, by subsequent designa-tion of Kellogg (1932).
DIAGNOSIS: This genus is diagnosed by 34transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. We are not aware of any mor-phological synapomorphy supporting this ge-nus.
CONTENTS: Seventeen species. Osteoce-phalus buckleyi (Boulenger, 1882); Osteoce-phalus cabrerai (Cochran and Goin, 1970);
Osteocephalus deridens Jungfer, Ron, Seipp,and Almendariz, 2000; Osteocephalus elke-jungingerae (Henle, 1981); Osteocephalusexophthalmus Smith and Noonan, 2001; Os-teocephalus fuscifacies Jungfer, Ron, Seipp,and Almendariz, 2000; Osteocephalus heyeriLynch, 2002; Osteocephalus leoniae Jungferand Lehr, 2001; Osteocephalus leprieurii(Dumeril and Bibron, 1841); Osteocephalusmutabor Jungfer and Hodl, 2002; Osteoce-phalus oophagus Jungfer and Schiesari,1995; Osteocephalus pearsoni (Gaige,1929); Osteocephalus planiceps Cope, 1874;Osteocephalus subtilis Martins and Cardoso,1987; Osteocephalus taurinus Steindachner,1862; Osteocephalus verruciger (Werner,1901); Osteocephalus yasuni Ron and Pra-muk, 1999.
Osteopilus Fitzinger, 1843
TYPE SPECIES: Trachycephalus marmora-tus Dumeril and Bibron, 1841 (5 Hyla sep-tentrionalis Dumeril and Bibron, 1841).
Calyptahyla Trueb and Tyler, 1974. Type species:Trachycephalus lichenatus Gosse, 1851 (5Hyla crucialis Harlan, 1826), by original des-ignation.
DIAGNOSIS: This genus is diagnosed by 43transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. No morphological synapomor-phies are known for this genus.
CONTENTS: Eight species. Osteopilus brun-neus (Gosse, 1851); Osteopilus crucialis(Harlan, 1826); Osteopilus dominicensis(Tschudi, 1838); Osteopilus marianae(Dunn, 1926); Osteopilus pulchrilineatus(Cope ‘‘1869’’ [1870]); Osteopilus septen-trionalis (Dumeril and Bibron, 1841); Osteo-pilus vastus (Cope, 1871); Osteopilus wilderi(Dunn, 1925).
Phyllodytes Wagler, 1830
TYPE SPECIES: Hyla luteola Wied-Neu-wied, 1824, by monotypy.
Amphodus Peters, ‘‘1872’’ [1873]. Type species:Amphodus wuchereri Peters, ‘‘1872’’ [1873],by original designation.
Lophyohyla Miranda-Ribeiro, 1923. Type species:Lophyohyla piperata Miranda-Ribeiro, 1923 (5
110 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Hyla luteola Wied-Neuwied, 1824), by originaldesignation.
DIAGNOSIS: This genus is diagnosed by174 transformations in nuclear and mito-chondrial protein and ribosomal genes. Seeappendix 5 for a complete list of these mo-lecular synapomorphies. Two morphologicalsynapomorphies of this taxon are the pres-ence of odontoids on the mandible and onthe cultriform process of the parasphenoid(Noble, 1931).
CONTENTS: Eleven species placed in fourspecies groups (Caramaschi et al., 2004a),one of which is monotypic.
Phyllodytes auratus Group
DIAGNOSIS: We are not aware of any pos-sible synapomorphy for this group.
COMMENTS: We did not include any ex-emplar of this group, but we continue to rec-ognize it following Caramaschi et al. (2004b)pending a rigorous test. Caramaschi et al.(2004b) diagnosed the different speciesgroups based on color patterns; it is unclearif any of these patterns could be consideredsynapomorphic.
CONTENTS: Two species. Phyllodytes au-ratus (Boulenger, 1917); Phyllodytes wuch-ereri (Peters, ‘‘1872’’ [1873]).
Phyllodytes luteolus Group
DIAGNOSIS: We are not aware of any pos-sible synapomorphy for this group.
COMMENTS: We included a single exemplarof this group and thus did not test its mono-phyly, but we continue to recognize it fol-lowing Caramaschi et al. (2004b) pending arigorous test. See comments for the P. au-ratus group.
CONTENTS: Six species. Phyllodytes acu-minatus Bokermann, 1966; Phyllodytes bre-virostris Peixoto and Cruz, 1988; Phyllodytesedelmoi Peixoto, Caramaschi, and Freire,2003; Phyllodytes kautskyi Peixoto and Cruz,1988; Phyllodytes luteolus (Wied-Neuwied,1824); Phyllodytes melanomystax Caramas-chi, Silva, and Britto-Pereira, 1992.
Phyllodytes tuberculosus Group
DIAGNOSIS: We are not aware of any pos-sible synapomorphy for this group.
COMMENTS: We did not include any ex-emplar of this group, but we continue to rec-ognize it following Caramaschi et al. (2004b)pending a rigorous test. See comments forthe P. auratus group.
CONTENTS: Two species. Phyllodytes punc-tatus Caramaschi and Peixoto, 2004; Phyl-lodytes tuberculosus Bokermann, 1966.
Species of Phyllodytes Unassigned toGroup
Peixoto et al. (2003) assigned Phyllodytesgyrinaethes Peixoto, Caramaschi, and Freire,2003 to its own species group. As stated ear-lier in this paper, we consider that monotypicspecies groups are not informative.
Tepuihyla Ayarzaguena and Senaris,‘‘1992’’ [1993]
TYPE SPECIES: Hyla rodriguezi Rivero,1968, by original designation.
DIAGNOSIS: For the purposes of this paperwe consider that the 90 transformations innuclear and mitochondrial protein and ribo-somal genes autapomorphic of Tepuihylaedelcae are synapomorphies of this genus.See appendix 5 for a complete list of thesemolecular synapomorphies. In the context ofour results, the reduction of webbing be-tween toes I and II is a putative morpholog-ical synapomorphy of this genus (Ayarza-guena et al., ‘‘1992’’ [1993b]; several in-stances of homoplasy within Lophiohylini, inPhyllodytes, and in the clade composed ofCorythomantis, Argenteohyla, Aparaspheno-don, and Nyctimantis).
COMMENTS: We included a single speciesof Tepuihyla, and as such we did not test itsmonophyly. We continue to recognize it fol-lowing Ayarzaguena and Senaris (‘‘1992’’[1993b]) until its monophyly is rigorouslytested.
CONTENTS: Eight species. Tepuihyla aecii(Ayarzaguena, Senaris, and Gorzula, ‘‘1992’’[1993]); Tepuihyla celsae Mijares-Urrutia,Manzanilla-Pupo, and La Marca, 1999; Te-puihyla edelcae (Ayarzaguena, Senaris, andGorzula, ‘‘1992’’ [1993]); Tepuihyla galani(Ayarzaguena, Senaris, and Gorzula, ‘‘1992’’[1993]); Tepuihyla luteolabris (Ayarzaguena,Senaris, and Gorzula, ‘‘1992’’ [1993]); Te-puihyla rimarum (Ayarzaguena, Senaris, and
2005 111FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Gorzula, ‘‘1992’’ [1993]); Tepuihyla rodri-guezi (Rivero, 1968); Tepuihyla talbergaeDuellman and Yoshpa, 1996.
Trachycephalus Tschudi, 1838
TYPE SPECIES: Trachycephalus nigroma-culatus Tschudi, 1838, by monotypy.
Phrynohyas Fitzinger, 1843. Type species: Hylazonata Spix, 1824 (5 Rana venulosa Laurenti,1768). NEW SYNONYMY.
Acrodytes Fitzinger, 1843. Type species: Hylavenulosa Daudin, 1802 (5 Rana venulosa Lau-renti, 1768), by original designation.
Scytopis Cope, 1862. Type species: Scytopis hebesCope, 1862, by monotypy.
Tetraprion Stejneger and Test, 1891. Type spe-cies: Tetraprion jordani Stejneger and Test,1891, by original designation.
DIAGNOSIS: This genus is diagnosed by 37transformations in nuclear and mitochondrialprotein and ribosomal genes. See appendix 5for a complete list of these molecular syna-pomorphies. The only possible morphologi-cal synapomorphy that we are aware of forthis genus is the presence of paired vocalsacs protruding posterior to the angles of thejaws when inflated (Trueb and Duellman,1971; see also Tyler, 1971).
COMMENTS: We are including Phrynohyasin the synonymy of Trachycephalus to avoidthe nonmonophyly of the two genera. Thereare other alternatives to resolve this situation,such as restricting Trachycephalus to thesoutheastern Brazilian taxa, including P. me-sophaea, while retaining Phrynohyas for theremaining species currently placed in that ge-nus, and resurrecting Tetraprion to accom-modate T. jordani. We consider that the ac-tion taken here is the most conservative.
CONTENTS: Ten species. Trachycephalusatlas Bokermann, 1966; Trachycephalus cor-iaceus (Peters, 1867), new comb.; Trachy-cephalus hadroceps (Duellman and Hoog-moed, 1992), new comb.; Trachycephalusimitatrix (Miranda-Ribeiro, 1926), newcomb.; Trachycephalus lepidus (Pombal,Haddad, and Cruz, 2003), new comb.; Tra-chycephalus mesophaeus (Hensel, 1867),new comb.; Trachycephalus nigromaculatusTschudi, 1838; Trachycephalus resinifictrix(Goeldi, 1907), new comb.; Trachycephalusvenulosus (Laurenti, 1768), new comb.
Incertae Sedis and Nomina Dubia
The taxonomic scheme introduced abovecomprises most of the valid species of Hy-linae. However, there are a number of speciesof former Hyla whose position in this newtaxonomy is uncertain. There are two likelyreasons for this. (1) The species have knowntype material and/or are known from multi-ple specimens, but the available informationis not sufficient to allow even the tentativeassignment to any of the taxonomic groups,so they are here considered as incerta sedis.(2) The species are known mostly from theiroriginal descriptions or type materials are re-ported to be lost or lack clear locality data.These are considered nomina dubia. See ap-pendix 4 for additional comments on someof these species. Within the first category fallHyla alboguttata Boulenger, 1882, Hylachlorostea Reynolds and Foster, 1992, Hylahelenae Ruthven, 1919, Hyla imitator (Bar-bour and Dunn, 1921), Hyla inframaculataBoulenger, 1882, Hyla vigilans Solano, 1971,and Hyla warreni Duellman and Hoogmoed,1992. In the second category we include(those with extant type material are followedby an asterisk) Calamita melanorabdotusSchneider, 1799, Calamita quadrilineatusSchneider, 1799, Hyla auraria* Peters, 1873,Hyla fusca Laurenti, 1768, Hypsiboas hyp-selops Cope, 1871, Hyla molitor* Schmidt,1857, Hyla palliata Cope, 1863, Hylaroeschmanni De Grys, 1938, Hyla surina-mensis Daudin, 1802, and Litoria ameri-cana* Dumeril and Bibron, 1841.
PHYLLOMEDUSINAE GUNTHER, 1858
Phyllomedusidae Gunther, 1858. Type genus:Phyllomedusa Wagler, 1830.
Pithecopinae B. Lutz, 1969. Type genus: Pithe-copus Cope, 1866.
DIAGNOSIS: The monophyly of this sub-family is supported by 95 transformations innuclear and mitochondrial protein and ribo-somal genes. See appendix 5 for a completelist of these molecular synapomorphies. Apossible morphological synapomorphy is thepupil constricting to vertical ellipse (Duell-man, 2001; known instance of homoplasy inNyctimystes). There are several larval char-acter states that may be synapomorphies,
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such as: the ventrolateral position of the spi-racle; arcus subocularis of larval chondrocra-nium with distinct lateral processes; ultralowsuspensorium; secondary fenestrae parieta-les; and absence of a passage between cera-tohyal and ceratobranchial I (Haas, 2003).
COMMENTS: Duellman (2001) consideredthe presence of a process on the medial sur-face of metacarpal II a synapomorphy ofPhyllomedusinae, with a known instance ofhomoplasy in Centrolenidae. However, be-cause Phyllomedusinae appears to be the sis-ter taxon of Pelodryadinae, the situation ismore complex. As noticed by Tyler and Da-vies (1978b), this character state is also pre-sent in some species groups of Litoria, so theinternal topology of Pelodryadinae will de-termine whether this character state is indeeda synapomorphy of Phyllomedusinae, withhomoplastic instances in Pelodryadinae, or ifit is a synapomorphy of Phyllomedusinae 1Pelodryadinae, with subsequent reversals inthe latter taxon. The supplementary postero-lateral elements of the m. intermandibularishave been considered a synapomorphy ofPhyllomedusinae (Duellman, 2001; Tyler,1971). As mentioned earlier, because it ismore parsimonious to interpret the sole pres-ence of supplementary elements of the m. in-termandibularis as a synapomorphy of Pelod-ryadinae 1 Phyllomedusinae, at this point itis ambiguous which of the positions (apicalas present in Pelodryadinae or posterolateralas in Phyllomedusinae) is the plesiomorphicstate of this clade.
The absence of the slip of the m. depressormandibulae that originates from the dorsalfascia at the level of the m. dorsalis scapulae(which subsequently reverses in Hylomantisand Phyllomedusa, see below) could also bea synapomorphy of Phyllomedusinae; how-ever, its taxonomic distribution among non-phyllomedusines needs to be assessed. Thisis most needed in Pelodryadinae, where asfar as we are aware, all observations on thismuscle are limited to Starrett’s (1968) un-published dissertation where she commentedon its morphology in 2 of the 172 knownvalid species of the subfamily. Ovipositionon leaves out of water could also be anothersynapomorphy of Phyllomedusinae, but thisis dependent on the position of Phrynome-dusa within Phyllomedusinae (species of this
genus do not oviposit on leaves but on rockcrevices or fallen trunks) and on the topologyof Pelodryadinae (however, only two speciesof Pelodryadinae, Litoria iris and L. longi-rostris, are known to lay eggs out of water,and not necessarily on leaves; Tyler, 1963;McDonald and Storch, 1993).
Several transformations that resulted assynapomorphies of Phyllomedusinae inBurton’s (2004) analysis optimize ambigu-ously in our trees because their distributionis unknown in Cruziohyla new genus. Con-sequently, it is unclear which transformationsare synapomorphic of the subfamily andwhich ones support the monophyly of inter-nal clades. These transformations are: two in-sertions of the m. flexor digitorum brevis su-perficialis; the tendon of the m. flexor digi-torum brevis superficialis divided along itslength into a medial tendon, from which arisetendo superficialis IV and m. lumbricalis lon-gus digiti V, and a lateral tendon from whicharise tendo superficialis V and m. lumbricalislongus digiti IV; tendo superficialis pro digitiII arising from a deep, triangular muscle,which originates on the distal tarsal 2–3; ten-do superficialis pro digiti III arising entirelyfrom the margin of the aponeurosis plantaris;two tendons of insertion of m. lumbricalislongus digiti V arising from two equal mus-cle slips; pennate insertion of the lateral slipof the medial m. lumbricalis brevis digiti V;m. transversus metatarsus II broad, occupy-ing the entire length of metatarsal II; m.transversus metatarsus III broad, occupyingmore than 75% of the length of metatarsalIII; m. extensor brevis superficialis digiti IIIwith two insertions, a flat tendon onto basalphalanx III and a pennate insertion on meta-tarsus III; and finally the m. extensor brevissuperficialis digiti IV with a single originwith belly undivided. The presence of m.flexor teres hallucis is shared with Pelodry-adinae; however, Burton (2004) stressed thatin that subfamily, presence or absence of thismuscle is subject to great intraspecific vari-ation, without providing information as tothe states present in the particular specimenshe studied, so the character was scored asmissing data in our matrix.
There are several other character systemsthat will likely provide additional synapo-morphies for this group of frogs. Manzano
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and Lavilla (1995b) and Manzano (1997) de-scribed several unique character states frommusculature, whose taxonomic distributionacross all Phyllomedusinae needs to be as-sessed. Tyler and Davies (1978a) mentionedthat Phyllomedusinae are the only hylidswhere the mandibular branch of the trigem-inal nerve subdivides into two twigs after tra-versing the mandible. Various authors (e.g.,Kenny; 1969; Cruz, 1982; Lescure et al.,1995) noticed that larvae of several speciesof Phyllomedusinae are usually suspended inwater in an oblique or even vertical positionrelative to the water surface. Bagnara (1974)observed a light-sensitive tail-darkening re-action in larvae of two phyllomedusines (Pa-chymedusa dacnicolor and Phyllomedusatrinitatis), and we observed a similar reactionin tadpoles of Phyllomedusa tetraploidea(Faivovich, pers. obs.). Further research willdetermine how inclusive is the clade orclades supported by these synapomorphies.
The presence of multiple bioactive pep-tides has been suggested as a distinctivecharacter of Phyllomedusinae (Cei, 1985).Since the beginning of the biochemical pros-pecting, it has become evident that Phyllo-medusinae have several different classes ofbioactive peptides (Erspamer, 1994), someunique (e.g., sauvagine, deltorphins), somenot (e.g., bombesins, caeruleins), as do thePelodryadinae (Apponyi et al., 2004). Be-cause there are multiple bioactive peptides,it seems reasonable to consider the differentpeptide families individually as potentialsynapomorphies of Phyllomedusinae, Pelod-ryadinae, or Phyllomedusinae 1 Pelodryadi-nae. More work needs to be done to betterunderstand the taxonomic distribution of thedifferent classes of peptides.
Agalychnis Cope, 1864
TYPE SPECIES: Agalychnis callidryas Cope,1862, by original designation.
DIAGNOSIS: The monophyly of this groupis supported by 23 transformations in nuclearand mitochondrial protein and ribosomalgenes. See appendix 5 for a complete list ofthese molecular synapomorphies. Agalychnishas extensively developed webbing on handsand feet in relationship with Pachymedusa,Hylomantis, Cruziohyla new genus, Phas-
mahyla, and Phyllomedusa. Also, with theexception of A. annae, which has a yellowiris, the other species have either a red or adark red iris.
COMMENTS: Considering the lack ofknowledge regarding the internal structure ofPelodryadinae, where some species have ex-tensive hand and foot webbing (e.g., Tyler,1968), it is still unknown if these characterstates are plesiomorphic for Phyllomedusi-nae. Consequently, at this stage we do notknow exactly in which point of the topologyof Phyllomedusinae they are homoplastic(both hands and foot webbing are developedin Cruziohyla new genus and, somewhat lessextensively, in Phrynomedusa).
CONTENTS: Six species. Agalychnis annae(Duellman, 1963); Agalychnis callidryasCope, 1862; Agalychnis litodryas (Duellmanand Trueb, 1967); Agalychnis moreletii (Du-meril, 1853); Agalychnis saltator (Taylor,1955); Agalychnis spurrelli (Boulenger,‘‘1913’’ [1914].)
Cruziohyla, new genus
TYPE SPECIES: Agalychnis calcarifer Bou-lenger, 1902.
DIAGNOSIS: For the purposes of this paperwe consider that the 171 transformations innuclear and mitochondrial protein and ribo-somal genes autapomorphic of Cruziohylacalcarifer are synapomorphies of this genus.See appendix 5 for a complete list of thesemolecular synapomorphies. Possible mor-phological synapomorphies include the ex-tensive hand and foot webbing (but see com-ments for Agalychnis) and the developmentof tadpoles in water-filled depressions onfallen trees. See comments below.
ETYMOLOGY: The name comes from theLatinization of Cruz, Cruzius 1 connecting-o 1 Hyla. We dedicate this new genus toour colleague and friend Carlos Alberto Gon-calves da Cruz, in recognition of his variouscontributions to our knowledge of Phyllo-medusinae.
COMMENTS: Phrynomedusa, the only genusof Phyllomedusinae missing from our anal-ysis, shares with Cruziohyla a bicolored iris,developed foot webbing (although more ex-tensively developed in Cruziohyla), and oraldisc with complete marginal papillae in the
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larvae. However, they differ in that eggs ofPhrynomedusa are laid in rock crevices (A.Lutz and B. Lutz, 1939; Weygoldt, 1991) orfallen trunk cavities above streams, fromwhere tadpoles drop and develop. The larvaeof Cruziohyla, unlike those of most otherknown Phyllomedusinae, develop in water-filed depressions of fallen trees (Donnelly etal., 1987; Hoogmoed and Cadle, 1991; Cald-well, 1994; Block et al., 2003). Hoogmoedand Cadle (1991) reported two situationswhere tadpoles associated with Agalychniscraspedopus were found in small pools in theforest, without a clear indication of where theeggs were laid. This could be interpreted ei-ther as a polymorphic reproductive trait or asan indication that more than one species isinvolved.
The oral disc with marginal papillae as amorphological synapomorphy of Cruziohyla1 Phrynomedusa should be taken cautiouslybecause of our general ignorance of the in-ternal topology of Pelodryadinae. Some Pe-lodryadinae also have an oral disc with com-plete marginal papillae (see Anstis, 2002),and further analysis could show that this isactually a plesiomorphy for Phyllomedusi-nae. The same problem holds for the pres-ence of foot webbing.
Instead of creating Cruziohyla to includeAgalychnis calcarifer and A. craspedopus,we could place both species in Phrynome-dusa. Both alternatives imply taxonomicrisks (in particular, that Cruziohyla could beshown to be nested within Phrynomedusa).Taking into account our almost complete ig-norance of the relationships of Pelodryadi-nae, and therefore character-state polarities atits base, and that Phrynomedusa could notbe included in this analysis, we consider thatat this stage it is more appropriate to createCruziohyla than to enlarge Phrynomedusa,without being certain about character polar-ities at the base of Phyllomedusinae.
Agalychnis craspedopus could not be in-cluded in the analysis, but the close relation-ship between A. craspedopus and A. calcar-ifer seems uncontroversial, as both have beenrepeatedly associated by some authors(Duellman, 1970; Hoogmoed and Cadle,1991; Duellman, 2001).
CONTENTS: Two species. Cruziohyla cal-carifer (Boulenger, 1902), new comb., Cru-
ziohyla craspedopus (Funkhouser, 1957),new comb.
Hylomantis Peters, ‘‘1872’’ [1873]
TYPE SPECIES: Hylomantis aspera Peters,‘‘1872’’ [1873], by monotypy.
DIAGNOSIS: The monophyly of this groupis supported by 38 transformations in mito-chondrial protein and ribosomal genes. Seeappendix 5 for a complete list of these mo-lecular synapomorphies. We are not aware ofany morphological synapomorphy support-ing this genus.
COMMENTS: Only Phyllomedusa lemur ofthe P. buckleyi group was included in theanalysis, and it obtains as the sister group ofour exemplar of Hylomantis, H. granulosa,but with a Bremer support value of 3. Whileit is evident that this group should be ex-cluded from Phyllomedusa, the possible tax-onomic actions (whether to create a new ge-nus or to include it in Hylomantis) deservefurther discussion. From the definition of thegroup given by Cannatella (1980), the onlycharacter state that could be considered asynapomorphy is the bright orange flanks inlife. The other character states included byCannatella (1980) are either likely symple-siomorphies (absence of the slip of the m.depressor mandibulae originating from thedorsal fascia at the level of the m. dorsalisscapulae; hands and feet less than one-fourthwebbed; parotoid gland not differentiated;palpebrum unpigmented; frontoparietal fon-tanelle exposed, large, and oval; oral discs oflarvae lacking marginal papillae anteriorly)or character states whose taxonomic distri-bution in Phyllomedusinae makes their po-larity unclear (lack of spots or pattern onflanks; cream or white iris; size; dorsum uni-formly green by day; presence or absence ofcalcars). Like the P. buckleyi group, the twospecies included in Hylomantis by Cruz(1990) also lack spots or pattern on flanks,which are light yellow (instead of bright or-ange). At this point, we have no evidenceregarding the polarity of these two characterstates; consequently, we consider that themorphological evidence of monophyly of theP. buckleyi group is weak. Considering thatwe could not test the monophyly of the P.buckleyi group, and that available morpho-
2005 115FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
logical evidence for its monophyly is notcompelling, provisionally, and with the ca-veat that the molecular support for thisgrouping is rather weak, we prefer to includeall species of this group in Hylomantis,where we recognize them as a separate spe-cies group, pending a rigorous test of itsmonophyly when a denser taxon samplingbecomes available.
CONTENTS: Eight species placed in twospecies groups.
Hylomantis aspera Group
DIAGNOSIS: Possible morphological syna-pomorphies of this group are the lanceolatediscs and presence of the slip of the m. de-pressor mandibulae originating from the dor-sal fascia at the level of the m. dorsalis scap-ulae (known homoplastic instance in Phyl-lomedusa and several other anurans).
COMMENTS: We included a single speciesof this group, and as such we did not we didnot test its monophyly, but we recognize itbased on the aforementioned evidence.
CONTENTS: Two species. Hylomantis as-pera Peters, ‘‘1872’’ [1873]; Hylomantisgranulosa (Cruz, ‘‘1988’’ [1989]).
Hylomantis buckleyi Group
DIAGNOSIS: The only apparent morpholog-ical synapomorphy of this group is the pos-session of bright orange flanks in life (Can-natella, 1980).
COMMENTS: We included a single speciesof this group, and as such we did not we didnot test its monophyly. We recognize it fol-lowing Cannatella (1980), pending a rigoroustest of its monophyly. Ruiz-Carranza et al.(1988) tentatively included Phyllomedusadanieli in the P. buckleyi group because ofthe reduced webbing, absence of parotoidglands, toe I shorter than toe II, presence ofa calcar, and unpigmented palpebrum. As theauthors noted, these characteristics are alsoshared with Phasmahyla and Hylomantis(they refer to these genera using the formerspecies groups of Phyllomedusa), but somealso with Phrynomedusa. One differencethey noticed was the golden iris colorationinstead of white; however, in the present sce-nario the polarity of this state is unclear (awhite iris is present in Phasmahyla and Hy-
lomantis, as redefined here). A difference isthe large snout–vent length (SVL) of P. dan-ieli compared with the species of Hylomantis(the only reported specimen of P. danieli, afemale, is 81 mm SVL; females of the otherspecies reach a maximum of 57 mm, accord-ing to Cannatella [1980]). Phyllomedusadanieli shares with the Hylomantis buckleyigroup its only apparent morphological syn-apomorphy (but see comments above), thebright orange flanks in life. Because of this,we tentatively include P. danieli in Hylo-mantis.
CONTENTS: Six species. Hylomantis buck-leyi (Boulenger, 1882), new comb.; Hylo-mantis danieli (Ruiz-Carranza, Hernandez-Camacho, and Rueda-Almonacid, 1988),new comb.; Hylomantis hulli (Duellman andMedelson, 1995), new comb.; Hylomantis le-mur (Boulenger, 1882), new comb.; Hylo-mantis medinai (Funkhouser, 1962), newcomb.; Hylomantis psilopygion (Cannatella,1980), new comb.
Pachymedusa Duellman, 1968
TYPE SPECIES: Phyllomedusa dacnicolorCope, 1864.
DIAGNOSIS: Molecular autapomorphies in-clude 105 transformations in nuclear and mi-tochondrial proteins and ribosomal genes.See appendix 5 for a complete list of thesetransformations. Possible morphological au-tapomorphies are the first toe opposable toothers, reticulated palpebral membrane (ho-moplastic with some species of Phyllome-dusa; Duellman et al., 1988b), and the irisreticulation (Duellman, 2001).
COMMENTS: Duellman (2001) also includ-ed the toes about one-fourth webbed as anautapomorphy of Pachymedusa. In the con-text of our results, this is probably not anautapomorphy, as the webbing is also equallyor more reduced in Hylomantis (as redefinedhere), Phasmahyla, and Phyllomedusa.
CONTENTS: Monotypic. Pachymedusa dac-nicolor (Cope, 1864).
Phasmahyla Cruz, 1990
TYPE SPECIES: Phyllomedusa guttata A.Lutz, 1924, by original designation.
DIAGNOSIS: The monophyly of this genusis supported by 94 transformations in mito-
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chondrial protein and ribosomal genes. Seeappendix 5 for a complete list of these mo-lecular synapomorphies. Possible morpho-logical synapomorphies of this genus are theabsence of a vocal sac, and the modificationof the larval oral disc into an anterodorsalfunnel-shaped structure (Cruz, 1990).
COMMENTS: Cruz (1990) mentioned the ab-sense of parotoid glands in Phasmahyla butstressed the presence of a pair of latero-dor-sal glands. While these glands could be cos-idered as possible synapomorphies of Phas-mahyla, additional work is needed in orderto determine if they could be considered ashomologous to the parotoid glands present inPhyllomedusa.
CONTENTS: Four species. Phasmahylacochranae (Bokermann, 1966); Phasmahylaexilis (Cruz, 1980); Phasmahyla guttata (A.Lutz, 1924); Phasmahyla jandaia (Boker-mann and Sazima, 1978).
Phrynomedusa Miranda-Ribeiro, 1923
TYPE SPECIES: Phrynomedusa fimbriataMiranda-Ribeiro, 1923, by subsequent des-ignation of Miranda-Ribeiro (1926).
DIAGNOSIS: A likely synapomorphy of thistaxon is the oviposition in rock crevices orfallen trunks overhanging streams (A. Lutzand B. Lutz, 1939; Weygoldt, 1991).
COMMENTS: We did not include any speciesof this genus in our analysis. Besides theplace of oviposition, we are not aware of anyother possible synapomorphy of Phrynome-dusa. This is not a strong support for itsmonophyly, particularly if we consider thatwith the exception of Weygoldt’s (1991)studies in captivity of Phrynomedusa mar-ginata, reports on oviposition of Phrynome-dusa are mostly anecdotal.
The most obvious difference betweenPhrynomedusa and Cruziohyla is the impres-sive SVL difference. Although the reductionin SVL could actually be a synapomorphy ofPhrynomedusa, considering how rudimenta-ry is our knowledge of the topology of Pe-lodryadinae, and considering its taxonomicdistribution in Phyllomedusinae (Phasmahy-la, Hylomantis, and Cruziohyla also have aproportionally smaller SVL, as do some spe-cies of Phyllomedusa), the polarity of SVLas a character, if definable at all, is far from
clear. Phrynomedusa could either be the sis-ter group of Cruziohyla or the remainingPhyllomedusinae.
CONTENTS: Five species. Phrynomedusaappendiculata (A. Lutz, 1925); Phrynome-dusa bokermanni Cruz, 1991; Phrynomedusafimbriata Miranda-Ribeiro, 1923; Phryno-medusa marginata (Izecksohn and Cruz,1976); Phrynomedusa vanzolinii Cruz, 1991.
Phyllomedusa Wagler, 1830
TYPE SPECIES: Rana bicolor Boddaert,1772 by monotypy.
Pithecopus Cope, 1866. Type species: Phyllome-dusa azurea Cope, 1862.
Bradymedusa Miranda-Ribeiro, 1926. Type spe-cies: Hyla hypochondrialis Daudin, 1800, bysubsequent designation of Vellard (1948).
DIAGNOSIS: The monophyly of this taxonis supported by 49 transformations in nuclearand mitochondrial protein and ribosomalgenes. See appendix 5 for a complete list ofthese transformations. Apparent morpholog-ical synapomorphies of Phyllomedusa are thepresence of parotoid glands, toe I longer thantoe II, and presence of the slip of the m. de-pressor mandibulae originating from the dor-sal fascia at the level of the m. dorsalis scap-ulae (known instance of homoplasy in theHylomantis granulosa group, and severalother anurans) (Duellman et al., 1988b).
COMMENTS: The transformation from pres-ence to absence of the m. abductor brevisplantae hallucis optimizes ambiguously inour analysis because the state of this char-acter is unknown in Phasmahyla.
Blaylock et al. (1976) described the pe-culiar wiping behavior in P. boliviana (as P.pailona), P. hypochondrialis, P. sauvagii,and P. tetraploidea (as P. iheringii). This be-havior was subsequently reported in P. dis-tincta, P. tarsius (Castanho and De Luca,2001), and P. iheringii (Langone et al.,1985). Castanho and De Luca (2001) furthernoticed a peculiar daily molting behavior.Further research on the taxonomic distribu-tion of these behaviors in Phyllomedusa willdetermine the limits of the group(s) they sup-port. The presence of the so-called lipidglands has been so far been reported in thefive species of Phyllomedusa that were stud-ied (P. bicolor, P. boliviana, P. hypochon-
2005 117FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
drialis, P. sauvagii, and P. tetraploidea;Blaylock et al., 1976; Delfino et al., 1998;Lacombe et al., 2000) and were noticed tobe unique to the genus by Delfino et al.(1998), so they could likely be another syn-apomorphy. As noticed by Cruz (1982), andcorroborated by most larval descriptions ofPhyllomedusinae, the larvae of most speciesof Phyllomedusa,29 as redefined here, havethe third posterior row of labial teeth reducedin relation to the first and second posteriorrows.
CONTENTS: Twenty-six species, some ofthem included in four species groups.
Phyllomedusa burmeisteri Group
DIAGNOSIS: We are not aware of any syn-apomorphy of this group.
COMMENTS: We included only a single spe-cies of this group in our analysis, and as suchwe did not test its monophyly, but we rec-ognize it following Pombal and Haddad(1992), pending a rigorous test of its mono-phyly.
CONTENTS: Four species. Phyllomedusaburmeisteri Boulenger, 1882; Phyllomedusadistincta B. Lutz, 1950; Phyllomedusa iher-ingii Boulenger, 1885; Phyllomedusa tetra-ploidea Pombal and Haddad, 1992.
Phyllomedusa hypochondrialis Group
DIAGNOSIS: We are not aware of any syn-apomorphy of this group.
COMMENTS: We included a single speciesof this group, and as such we did not test itsmonophyly, but we recognize it followingBrandao (2002), pending a rigorous test ofits monophyly. Manzano and Lavilla (1995b)described the muscle epicoracoideus in Phyl-lomedusa hypochondrialis, and Manzano(1997) noticed its absence in other speciesthat she studied (P. atelopoides, P. boliviana,and P. sauvagii). Our observations on theonly other species of the group available tous, P. rohdei (AMNH A-20263), indicatethat it also has the m. epicoracoideus, so weconsider the presence of this muscle a pos-sible synapomorphy of the group. All species
29 The only exception we are aware of is the larvaeof Phyllomedusa vaillanti, where P-3 almost equals P-2(Caramaschi and Jim, 1983).
of this group lack vomerine teeth, as doPhasmahyla, Phyllomedusa palliata andsome species of Phrynomedusa (Brandao,2002; Cruz, 1990). The taxonomic distribu-tion of other myological peculiarities de-scribed by Manzano and Lavilla (1995b) inP. hypochondrialis, such as the presence ofthin and/or shortened muscles, and unusualinsertions of some of them, needs to be as-sessed in other Phyllomedusinae.
CONTENTS: Six species. Phyllomedusa ay-eaye (B. Lutz, 1966); Phyllomedusa centralisBokermann, 1965; Phyllomedusa hypochon-drialis (Daudin, 1800); Phyllomedusa me-gacephala (Miranda-Ribeiro, 1926); Phyllo-medusa oreades Brandao, 2002; Phyllome-dusa rohdei Mertens, 1926.
Phyllomedusa perinesos Group
DIAGNOSIS: A possible synapomorphy ofthis group is the purple coloration on thehands, feet, flanks, and concealed surfaces,as well as the purple venter with white gran-ules (Cannatella, 1982).
COMMENTS: We did not include any ex-emplar of this group in the analysis. Itsmonophyly is tentatively assumed followingCannatella (1982) and is based on the evi-dence mentioned above.
CONTENTS: Four species. Phyllomedusabaltea Duellman and Toft, 1979; Phyllome-dusa duellmani Cannatella, 1982; Phyllome-dusa ecuatoriana Cannatella, 1982; Phyllo-medusa perinesos Duellman, 1973.
Phyllomedusa tarsius Group
DIAGNOSIS: We are not aware of any syn-apomorphy supporting the monophyly of thisgroup.
COMMENTS: We included a single speciesof this group, and as such we did not test itsmonophyly, but we continue to recognize itfollowing De la Riva (1999) until its mono-phyly is rigorously tested.
CONTENTS: Four species. Phyllomedusaboliviana Boulenger, 1902; Phyllomedusacamba De la Riva, 2000; Phyllomedusa sau-vagii Boulenger, 1882; Phyllomedusa tarsius(Cope, 1868).
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Species of Phyllomedusa Unassigned toGroup
There are several species that are currentlynot assigned to any group. These are: Phyl-lomedusa atelopoides Duellman, Cadle, andCannatella, 1988; Phyllomedusa bicolor(Boddaert, 1772); Phyllomedusa coelestis(Cope, 1874); Phyllomedusa palliata Peters,‘‘1872’’ [1873]; Phyllomedusa tomopterna(Cope, 1868); Phyllomedusa trinitatis Mer-tens, 1926; Phyllomedusa vaillanti Boulen-ger, 1882; and Phyllomedusa venusta Duell-man and Trueb, 1967.
BIOGEOGRAPHICAL COMMENTARYOur objective here is to comment on pat-
terns of distribution among the major bio-geographic/tectonic units and not to providea detailed biogeographic analysis. The distri-bution/biogeographic units of our discussionare (1) Australia plus New Guinea, (2) con-tinental South America, (3) Middle America(in the sense of being composed of tropicalMexico, the Chortis Block of Central Amer-ica, and the Panamanian Isthmus), and (4)the temperate Holarctic. Clearly all of theseregions have histories that provide clues asto movements and diversifications withinthese areas. Because of the enormity of thetopic of biogeography for the entire Hylidae,our comments will be truncated, limited ei-ther by our taxonomic sampling, knowledgeof earth history, or phylogenetic resolution.Nevertheless, there are obvious geographicpatterns that warrant our attention.
The distribution of the Hylidae stronglysuggests a southern-continent origin of thetaxon, a conclusion in accord with sugges-tions based on different lines of evidence ad-vanced over the last 80 years (Metcalf,1923a, 1923b, 1928; Duellman, 1970, 2001;Savage, 2002a) and supported by the obser-vation that all the major groups of hylidshave their centers of diversity in southerncontinents, with only phylogenetically sec-ondary centers of diversification existing inMiddle America, North America, and evenmore attenuated areas of radiation in Eurasia.
RELATIONSHIPS BETWEEN AUSTRALIA AND
SOUTH AMERICA
The relationship between Australian andSouth American taxa has been previously
noted for the Hylidae (Darst and Cannatella,2004; Hoegg et al., 2004) and in several oth-er groups (see Sanmartın and Ronquist[2004] for a review). In our results (fig. 13),the Australopapuan Pelodryadinae forms thesister taxon of the predominantly SouthAmerican Phyllomedusinae, a distributionwhich we think speaks to one of the earliestpatterns in the entire Hylidae, that of an Aus-tralia–Antarctica–South American connec-tion. We cannot address any other topics ofpelodryadine biogeography due to our lim-ited sampling.
RELATIONSHIPS BETWEEN SOUTH AND
MIDDLE AMERICA
Having the sister taxon of the Phyllome-dusinae in Australia strongly suggests asouthern (South American) origin of thePhyllomedusinae, although distribution ofthe most basal taxon, Cruziohyla calcarifer,is in Choco/lower Middle America, with theremaining taxa found from northwesternMexico to southern Brazil. This distributionsuggests that the phyllomedusine biogeo-graphic pattern is not recent. Assuming aconnection between Australia and SouthAmerica was by way of Antarctica, onewould be driven to the conclusion that SouthAmerica is the home of the phyllomedusines.The fact that we could not include any ex-emplar of Phrynomedusa, an Atlantic Forestgenus possibly related with Cruziohyla,could possibly cloud the general picture.
Apart from the Hylini, there are severalinstances of members of the other three tribesof Hylinae having a Middle American distri-bution (figs. 14–16), corroborating the sug-gestion of Duellman (2001) regarding the ex-istence of several independent vicariance ordispersal events with hylids between SouthAmerica and Middle America. Our resultsimply eight independent events of dispersalfrom South America into Middle America:(1) Dendropsophus ebraccatus, (2) D. micro-cephalus, (3) ancestor of Hylini, (4) Hypsi-boas boans, (5) H. rufitelus, (6) Scinax bou-lengeri, (7) S. elaeochrous and S. staufferi,and (8) Trachycephalus venulosus. However,this number is a clear underestimation be-cause we did not include other terminals thatalso have a Middle American distribution
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(e.g., Dendropsophus phlebodes, D. robert-mertensi, D. sartori, D. subocularis, Hypsi-boas pugnax, H. rosenbergi, Scinax altae,and S. rostratus) and which may representadditional entries into Middle America fromSouth America. For some of these species(e.g., Scinax altae and S. rostratus) we con-sider it likely that they are related to otherMiddle American members of their respec-tive phylogenetic nearest relatives (hence notadding to the number of independent biogeo-graphic events). Other species (Dendropso-phus subocularis) might imply additionalevents because they are probably nestedwithin mostly South American clades. Theuncertain position of the other species (e.g.,Dendropsophus phlebodes, D. robertmerten-si, D. sartori, Hypsiboas pugnax, H. rosen-bergi) in our phylogenetic hypothesis doesnot allow us to suggest that their presence inMiddle America either represents indepen-dent events or that they are contained withinother groups of species whose ancestorsmoved into Middle America.
Considering the Hylinae with a MiddleAmerican origin, the results imply two bio-geographic events to explain the presence ofthese lineages in South America: the cases ofScinax elaeochrous (fig. 15) and Smiliscaphaeota (fig. 16). Once again, this is a min-imal number of events. The monophyly ofEcnomiohyla could be in error, because mostspecies were unavailable for study and atleast E. tuberculosa (not studied) is possiblyunrelated to the Middle American fringe-limbed treefrogs. Duellman (2001) suggestedthat Smilisca sila and S. sordida together aremonophyletic (apparent synapomorphies:ventral oral disc in the larvae and small innermetatarsal tubercle) and together are the sis-ter taxon of S. puma. If this hypothesis with-stands further testing, it would represent athird independent biogeographic event in-volving a Middle American lineage presentin northern South America.
SOUTH AMERICA
GUAYANA HIGHLANDS–ANDES–ATLANTIC FOREST
Within Cophomantini (fig. 14), the firstfour genera contain elements from threecharacteristic formations, quite distant geo-
graphically from each other: Myersiohyla iscomposed solely of Guayana Highlands spe-cies; Hyloscirtus is composed exclusively ofAndean species; Bokermannohyla and Aplas-todiscus are composed almost exclusively ofspecies from the southeastern Brazilian At-lantic Forest and Rocky fields associatedwith this formation and to the Cerrado. Weare not aware of any similar biogeographicpattern in any other animal group.
GUAYANA HIGHLANDS
Our analysis included 6 of the 19 hylidendemics (updated from Duellman’s [1999]list by adding Hypsiboas rhythmicus) of theGuayana Highlands, plus two undescribedspecies. The topology suggests a minimumof four independent occurrences of endemichylines in the Guayana Highlands (figs. 14,16): (1) the Hypsiboas benitezi group (thisgroup also contains three species from west-ern Amazonia: H. hutchinsi, H. microderma,and Hypsiboas sp. 2), (2) Hypsiboas sibleszi,(3) Myersiohyla, and (4) Tepuihyla. In the H.benitezi group, it is ambiguous whether thereis an origin in the Guayana Highlands witha subsequent dispersal/vicariance event intonorthwestern Amazonia, or two independentevents that led to the presence of these spe-cies in the highlands.
Considering the 13 taxa from the GuayanaHighlands that were unavailable for thisstudy, all but two species are members ofgroups represented in the analysis. Seven arespecies of Tepuihyla, three are species ofMyersiohyla, two are species of Scinax (S.danae, and S. exiguus), one is a species ten-tatively associated with the Hypsiboas beni-tezi group (H. rhythmicus), and one is incertasedis (‘‘Hyla warreni’’). Phylogenetic rela-tionships of the two species of Scinax withother species of the genus are still unknown,as is the position of ‘‘Hyla warreni’’. Whenconsidering relationships of the GuayanaHighlands lineages with the other Hylinae,current evidence suggests they are related toelements from the Amazon Basin (Osteoce-phalus, Hypsiboas microderma, Hypsiboassp. 2) and the Choco (Hypsiboas picturatus).
IMPACT OF THE ANDES IN HYLINE EVOLUTION
The uplift of the Andes and subsequentclimatic changes in the Quaternary have an
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Fig. 13. A partial view of the strict consensus showing major biogeographic patterns among out-groups, Pelodryadinae and Phyllomedusinae, and the geographic distribution of the exemplars of Phyl-lomedusinae. Distributions are taken from Duellman (1999) and Frost (2002). Only collective groupsreferred in ‘‘Biogeographic Commentary’’ are shown. An asterisk (*) indicates the distribution of Phyl-lomedusa hypochondrialis that ranges from the Chaco/Cerrado through the Amazon Basin and Guayanalowlands up to the Llanos.
impressive correlation with large radiationsof anuran groups, such as certain Bufonidae,Centrolenidae, Dendrobatidae, Hemiphracti-nae, and Eleutherodactylinae (e.g., Lynch,1986; Coloma, 1995; Lynch and Duellman,1997; Lynch et al., 1997; Lynch, 1998;Duellman, 1999). Previous knowledge of hy-lid distribution, as well as our results, sug-gests a much more limited impact of the An-des in hylid radiation and speciation, withthree hyline radiations in the Andes (figs. 14,16): Hyloscirtus, the Andean clade of theHypsiboas pulchellus group, and the Den-dropsophus columbianus 1 D. labialisgroups clade. If we consider the taxa thatwere not included in this analysis, there are41 with an Andean distribution: Dendrop-sophus aperomeus, D. battersbyi, ‘‘Hylachlorostea’’, D. delarivai, D. praestans, D.stingi, D. yaracuyanus, ‘‘Hyla vigilans’’, Os-teocephalus elkejungingerae, O. leoniae, O.pearsoni, Scinax fuscovarius, S. castroviejoi,S. manriquei, S. oreites, the four species ofthe Dendropsophus garagoensis group, plus23 additional members of Hyloscirtus, theAndean clade of the Hypsiboas pulchellusgroup, and the Dendropsophus columbianus1 D. labialis groups clade (Duellman, 1999;Mijares-Urrutia and Rivero, 2000; Jungferand Lehr, 2001; Kohler and Lotters, 2001a;Barrio-Amoros et al., 2004). If the D. gara-goensis group is not related to the Dendrop-sophus columbianus 1 D. labialis groupsclade, then it would represent a fourth An-dean radiation of hylids. Relationships of theAndean D. aperomeus, D. battersbyi, D. de-larivai, D. stingi, and D. yaracuyanus withinDendropsophus are unknown, so they mayrepresent as many as five additional eventsleading to the presence of hylids in the An-des. A similar situation occurs with ‘‘Hylachlorostea’’, ‘‘Hyla vigilans’’, Scinax man-riquei, S. oreites, and the species of Osteo-cephalus. Scinax castroviejoi and S. fusco-
varius are sister species (Faivovich, unpubl.data), so they are considered another inde-pendent entrance into the Andes. Adding upall these species and clades gives a maximumtotal of 17 independent biogeographicevents, of which 5 subsequently radiated and13 are single taxa. If the Andean species ofOsteocephalus were monophyletic, then thefigure could decrease to 14 independentevents, 6 of which subsequently radiated.
Considering the information outlinedabove, and returning to the beginning of thissection, whereas now we have an upper anda lower limit for the number of independentradiations of hylids in the Andes, we have noidea as to the number of independent radia-tions of Bufonidae, Centrolenidae, Dendro-batidae, and Hemiphractinae in the Andes.
A question that might arise is why there issuch poor diversification of hylids in the An-des (note that its complement, Why are thereso many species in extra-Andean areas?, isequally valid). There are several scenariosthat could answer this question. The presencein several areas of the Andes of anurangroups with obligate aquatic life-historystages dependent on either ponds or streams(Centrolenidae, Bufonidae) appears to be astrong argument against a hypothesis of lackof appropriate habitats. The fact that hylidsare found in fairly high altitudes in the Andes(e.g., species in the Andean stream-breedingclade reach up to 2400 m; Duellman et al.,1997; species of the D. labialis group reachup to 3500 m; Luddecke and Sanchez, 2002)and other places (e.g., several Hylini livingbetween 2000 and 3000 m; see Duellman,2001) could indicate that there may be fewphysiological constraints limiting the exploi-tation of higher areas.30
30 We understand that this argument is weak; perhapsthe hylid groups that are not physiologically constrainedare precisely those that could colonize and diversify inthe highlands.
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Fig. 14. A partial view of the strict consensus showing the geographic distribution of the componentsof Cophomantini. Distributions, in general, are taken from Duellman (1999). Only collective groupsreferred in ‘‘Biogeographic Commentary’’ are shown. An asterisk (*) indicates the distribution of Hyp-siboas punctatus that ranges from the Chaco/Cerrado through the Amazon Basin and Guayana lowlandsup to the Caribbean lowlands. Two asterisks (**) indicate the geographic distribution of Hypsiboascordobae that is restricted to the Sierras of Central Argentina.
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ATLANTIC FOREST
Of the several instances of hyline taxa pre-sent in the Atlantic Forest of Brazil, eight aresingle terminals and six are clades (figs. 14–16). The terminals are (1) Aparasphenodonbrunoi, (2) Dendropsophus anceps, (3) D.giesleri, (4) D. minutus, (5) D. seniculus, (6)Hypsiboas albopunctatus, (7) Scinax urugua-yus, and (8) Xenohyla truncata. The cladesare (1) Aplastodiscus, (2) the Hypsiboas fa-ber group plus the H. pulchellus group clade,(3) Trachycephalus nigromaculatus plus T.mesophaeus, (4) Phyllodytes, (5) the Scinaxcatharinae clade, and (6) Bokermannohyla.Including the approximately 134 hylid spe-cies that were unavailable for this study witha distribution in eastern Brazil would cer-tainly increase the number of clades and ter-minals in an unpredictable way.
Faivovich (2002) observed that Scinaxwas divided in two clades, one endemic tothe Atlantic Forest (the S. catharinae clade)and another that was widespread in the Neo-tropics (the S. ruber clade). Our results implyan ambiguous situation. The position of S.uruguayus as the sister taxon of the remain-ing species of the S. ruber clade suggests thatScinax could have as well originated insoutheastern Brazil and colonized other areasof the Neotropics in subsequent events. Adenser taxon sampling of Scinax would allowa test of this hypothesis.
In Lophiohylini, nearly all species of Phyl-lodytes are from the Atlantic Forest (the onlyexception being P. auratus from Trinidad),as is also true for Itapotihyla langsdorffii andseveral other species of the tribe. However,the situation here is equivocal because itwould be equally parsimonious to postulatetwo independent events leading to the pres-ence of I. langsdorffii and Phyllodytes in theAtlantic Forest.
Within Bokermannohyla, the B. circum-data species group, mostly from forested re-gions, is nested within a clade composed ofspecies and species groups (B. pseudopseudisand B. martinsi groups) restricted to thehighland formations of rocky fields (Boker-mann and Sazima, 1973b; Eterovick andBrandao, 2001; Lugli and Haddad, in prep.).The facts that the other species included inthe B. martinsi group, B. langei, is from for-
ested areas (Bokermann, 1964a) and that twospecies of the B. circumdata group, B. na-nuzae and B. sazimai are from rocky fields(Bokermann and Sazima, 1973b; Cardosoand Andrade ‘‘1982’’ [1983]) suggest that adenser taxon sampling of Bokermannohyla isnecessary to better understand whether thesefrogs are an original element from the rockyfields that secondarily radiated in the forestedareas or vice versa.
The clade composed of the Hypsiboas fa-ber and H. pulchellus groups could be an ex-ample of an Atlantic Forest (or at least aneastern Brazilian31) origin with subsequentradiations into other regions. Faivovich et al.(2004) found that within the H. pulchellusgroup, an Andean clade was nested within anAtlantic Forest clade. Exactly the same pat-tern for the group is corroborated by ouranalysis (fig. 14).
ORIGIN OF WEST INDIAN HYLIDS
Our results support assertions made byHass et al. (2001) and Hedges (1996), basedon immunological distances and unpublishedsequence data, regarding a diphyletic originof West Indian hylids. Osteopilus is the sistergroup of a clade composed of Osteocephalusand the montane Tepuihyla (fig. 16). Whilethe incomplete taxon sampling precludes acareful assessment of the distribution of themost basal taxa of these genera, our resultsare compatible with a northern South Amer-ican origin for Osteopilus; Tepuihyla is re-stricted to highlands in Venezuela and Guy-ana (Ayarzaguena et al., ‘‘1992’’ [1993b],Duellman and Yoshpa, 1996; Mijares-Urrutiaet al., 1999), and Osteocephalus is wide-spread in the Amazon Basin and surroundingregions, from Venezuela and French Guianato Bolivia (e.g., De la Riva et al., 2000; Les-cure and Marty, 2000; Jungfer and Lehr,2001; Smith and Noonan, 2001).
31 Hypsiboas crepitans in Brazil occurs in some areasof the Atlantic forest and mostly in adjacent CerradoCaatinga and Cerrado formations; however, we stillthink the observed pattern is meaningful, and the addi-tion of the remaining species of the group will help tobetter define it. Hypsiboas crepitans also has wider dis-tribution, including Panama, Colombia, Venezuela, andthe Guayanas; however, the status of those populationsneeds to be reassessed (Lynch and Suarez-Mayorga,2001), as do their relationships with the H. faber group.
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Regarding the other West Indian hylid spe-cies, Hypsiboas heilprini and its sister-grouprelationship with the H. albopunctatus group(fig. 14) is notable in that the basal taxon ofthe group, H. raniceps, has a broad distri-bution through open areas of South America,spanning about 3500 km, from French Gui-ana (Lescure and Marty, 2000) to easterncentral Argentina (Basso, 1995). The factthat both hylid lineages present in the WestIndies clearly have a northern South Ameri-can origin is coincident with the suggestionsmade by Hedges (1996).
MIDDLE AMERICA AND THE HOLARCTIC
The molecular evidence for Hylini doesnot support the idea of a basal split betweenan Isthmian–Lowlands clade and a Mexican–Nuclear Central American clade as was sug-gested by Duellman (2001). Instead, it sug-gests the existence of a clade composed pri-marily of, but not limited to, Mexican high-lands-Nuclear Central American elements(fig. 15). Nested within this clade, there is alineage composed of two lowland clades(Tlalocohyla, and the group composed of An-otheca, Triprion, and Smilisca), an IsthmianHighlands clade (Isthmohyla) and one NorthAmerican–Eurasiatic clade (Hyla).
Within the Mexican Highlands–NuclearCentral American clade, and besides the ma-jor lineage that diversified outside this region(Hyla, Isthmohyla), there are two other in-stances of terminals distributed in lower Cen-tral America (fig. 15), Duellmanohyla rufi-oculis and Ecnomiohyla miliaria. There arealso at least three more cases that were un-available for this study: two species of Duell-manohyla (D. lythrodes, D. uranochroa), thelower Central American species of Ecnom-iohyla (E. fimbrimembra, E. thysanota), andPtychohyla (P. legleri). The two instancesimplied by our results are a lower limit; theaddition of further exemplars of Duellma-nohyla, Ecnomiohyla, and Ptychohyla willdetermine if there were other events as well.
Hylini is nested within a South Americanclade, supporting a South American originfor this group. It is interesting, however, tosee a likely Mexican highlands–Nuclear Cen-tral American origin for the lowland lineages(Anotheca, Tlalocohyla, Smilisca, Triprion),
and that the Isthmian Highlands Isthmohylais nested within all these lineages. We seethese situations as being at least partiallycompatible with the current paleogeographicscenario for Middle America (see Iturralde-Vinent and McPhee, 1999; Savage, 2002a).
The topology of the Mexican Highlands–Nuclear Central American clades also showssome fine-grained patterns, like the possiblesister-taxon relationship between a cladefrom the Mexican highlands (Plectrohylabistincta group) and one from the NuclearCentral American highlands (Plectrohylaguatemalensis group). The problem is thatconsidering the scarce taxon sampling ofthese two species groups in our analysis, andthe very likely possibility of further rear-rangements upon addition of more exemplarsof these groups (see comments in earlier dis-cussion), it seems risky to hypothesize aboutthe recovered pattern.
Most authors who have discussed the ori-gin of Eurasiatic Hyla assumed its originfrom western North American Hyla and itsdispersion to Eurasia presumably throughBeringia (Anderson, 1991; Borkin, 1999;Duellman, 2001; Kuramoto, 1980). The to-pology of Hyla (fig. 15) shows two Eura-siatic taxa. One of these, the H. arboreagroup, forms the sister taxon of the remain-ing Hyla. The other, H. japonica, is imbed-ded within the H. eximia group, which inturn is nested within a grade of eastern NorthAmerican species. This situation implies twoindependent biogeographic events to explainthe origin of Eurasiatic Hyla, as suggested byAnderson (1991) and Borkin (1999). Whilethis pattern is partially compatible with theidea of a western North American origin ofat least some Eurasiatic Hyla (those nestedwithin the H. eximia group) claimed by pre-vious authors, it remains at least equivocalfor the clade that contains most exemplars ofthe H. arborea group. The only way of main-taining a western North American origin forthis clade is to invoke a very important his-torical shift in the distribution of the cur-rently eastern North American speciesgroups or to accept an eastern North Amer-ican–European (North Atlantic) vicariance ordispersal event. The position of the H. eximiagroup nested within eastern North Americanspecies groups requires a dispersal/vicariance
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Fig. 15. Partial view of the strict consensus showing the geographic distribution of the componentsof Dendropsophini. Distributions, in general, are taken from Duellman (1999). Only collective groupsreferred in ‘‘Biogeographic Commentary’’ are shown. An asterisk (*) indicates the geographic distri-bution of Scinax squalirostris that includes the Atlantic forest, Cerrado, Chaco, and Pampean grasslands.Two asterisks (**) indicate the geographic distribution of Scinax ruber that includes the Amazonian andGuayana lowlands, Caribbean lowlands, and Llanos. Three asterisks (***) indicate the geographic dis-tribution of Pseudis paradoxa that extends from the Chaco region to the Amazonian and Guayanalowlands, the Llanos, and the Caribbean lowlands.
event southward to explain the distributionof this lineage, as suggested by Duellman(2001).
A ROUGH TEMPORAL FRAMEWORK: CLUES
FROM THE HYLID FOSSIL RECORD
The hylid fossil record is remarkably scant(Sanchiz, 1998a), and hylid remains, on mostoccasions, are represented by disarticulatedilia. The oldest fossil record tentatively as-signed to Hylidae is remains of an ilium and
a humerus from the Maastrichtian of Naskal,India (Prasad and Rage, 1995) tentatively as-signed to Hylidae, although Sanchiz (1998a)considered the identification dubious. Hylidswere mentioned from the Paleocene of Ita-boraı (Brazil) by Estes (1970), Estes andReig (1973), Estes and Baez (1985), andBaez (2001), but the material—also iliac re-mains—is stills unstudied.
The earliest known record for NorthAmerica is Hyla swanstoni, from the Late
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Fig. 16. Partial view of the strict consensus showing the geographic distribution of the componentsof Hylini and Lophiohylini. Distributions were taken from Campbell (1999) and Duellman (1970, 2001).Only collective groups referred in ‘‘Biogeographic Commentary’’ are shown. An asterisk (*) indicatesthe distribution of Anotheca spinosa that is present in the Mexican, Nuclear Central American, andIsthmian highlands. Two asterisks (**) indicate the geographic distribution of Trachycephalus venulosus,which extends from central eastern Argentina to Southern Mexico.
Eocene of Cypress Hill Formation (Saskatch-ewan, Canada), described by Holman (1968)based on eroded iliac remains; Sanchiz(1998a) also considered this taxonomic as-signment to be dubious. Other iliac remainsfrom the mid-Miocene of North America(mostly from the Hemingfordian NorthAmerican Land Mammal Age) were also re-ferred to hylids (i.e., Acris barbouri, Hylamiocenica, H. miofloridiana, Proacris min-toni, Pseudacris nordensis), as well as sev-eral Pleistocene remains that were assignedto extant species (Holman, 2003).
The oldest known hylid record for Europewas reported by Sanchiz (1998b) from earlyMiocene lignite deposits of Austria. The re-mains consist of a fragmentary sacral verte-bra and a scapula, which Sanchiz (1998b)found to be similar to Hyla arborea and H.meridionalis, and assigned to Hyla sp. Otherremains from the late Miocene, as well asfrom the Pliocene to Holocene, were referredto Hyla sp., H. arborea, and the most recentones to some extant species (see Sanchiz[1998a] for review).
The oldest fossil record of hylids in Aus-tralia dates to the Lower to Middle Miocene,where some species of Litoria were de-scribed on the basis of iliac remains (e.g.,Tyler, 1991). Other Pleistocene and Holoceneremains were assigned to extant species (seeSanchiz [1998a] for a review).
We are not inclined to construct far-reach-ing hypotheses based on such a sparse fossilrecord. Without taking into account Hylaswanstoni, already questioned by Sanchiz(1998a), we must consider the possibility thatat least one of the North American iliac re-mains from the Miocene assigned to hylidsis actually related to the extant groups ofHolarctic hylids. If this were the case, itwould imply a minimum age of approxi-mately 15 mybp of hylid presence in NorthAmerica. If we consider with the same le-
niency the Miocene remains from Europe,especially the approximately 17 mybp Hylasp. reported by Sanchiz (1998b), we couldconsider an even earlier, though still unspec-ified, minimum age of hylid presence inNorth America. All the evidence regardinghylid fossil record and its minimum ages hasbeen presented.
How this information could be used de-pends on how much we are willing to assumeabout the role of known geophysical data indating cladogenetic events. In recent studiesaddressing phylogenetic studies of some froggroups, geophysical information was usedfor molecular clock calibrations (e.g., Bos-suyt and Milinkovitch, 2000; Vences et al.,2003c). Leaving aside particular objectionsabout molecular clock estimations, the use ofclade-independent information as calibrationpoints entails assumptions about the evolu-tionary process and about the primacy andprecision of paleogeographic reconstructionsthat should be taken cautiously (Page andLydeard, 1994). It has been this clade-inde-pendent information that has classically beenused to date one of the important events ofthe hylid radiation: the origin of the Hylini.
Both Savage (1966, 1982, 2002a) andDuellman (1970, 2001) suggested an earlycolonization event of Middle America byHylinae and Phyllomedusinae stocks in thelate Cretaceous–Eocene; from these stocks,all remaining Middle American and NorthAmerican elements differentiated due to sev-eral vicariant and dispersal events throughthe Tertiary. Considering that the evidencefor the existence of a late mid-Miocene land-bridge between North and South America(necessarily involving Central America) isambiguous (Iturralde-Vinent and McPhee,1999), a ‘‘classical’’ assumption of nonover-water dispersal would imply that actually theminimum age of colonization of MiddleAmerica must have been at least about 75
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mybp, during the existence of the so-calledpost-Cretaceous Arc landbridge (Iturralde-Vinent and McPhee, 1999). While this seemslike a logical deduction, we find it to be un-supported by the available data pertaining tohylids. We only know (or better, assume) thathylids were present in North America 15mybp and possibly in Europe 17 mybp. Ac-cordingly, we are still uncomfortable in as-signing 75 mybp as the minimum age of Hy-lini. Instead, this age should be the result ofeither a fossil record (still unknown) or a dat-ing exercise.
FUTURE DIRECTIONS
In order to increase our knowledge of hy-lid phylogenetics, we see several lines of in-quiry stemming from this project whose pur-suit would be fruitful. These are:
1. Nonmolecular data set: As stated elsewherein this paper, an extensive, well-researchednonmolecular data set is a major gap in ouranalysis. Every effort should be made tocomplete one and to combine it with theavailable molecular data.
2. Phylogeny of Pelodryadinae: By far thegreatest deficiency in our knowledge of hy-lid relationships that still need to be solvedis the relationships among Pelodryadinae.The combination of a densely sampled dataset of Pelodryadinae with ours will be help-ful for understanding Phyllomedusinae re-lationships.
3. Inclusion of unrepresented groups and dens-er taxon sampling of poorly representedgroups of Hylinae: No exemplars of theBokermannohyla claresignata and the Den-dropsophus garagoensis groups were avail-able for this analysis. Furthermore, severalgroups have been poorly represented; suchis the case for Ecnomiohyla, Exerodonta,Isthmohyla, Megastomatohyla, Plectrohyla,and Tlalocohyla. A major task in the futurewill be to rigorously test all the tentative as-sociations of species not included in theanalysis with the various clades that weidentified.
4. Relationships within smaller taxonomicunits: Our results provide a general frame-work for the study of relationships withinalmost any of the genera or species groupsof the species-rich genera of Hylidae. In par-ticular, we think that the study of relation-ships through an increased taxon samplingwithin Bokermannohyla, Dendropsophus,
Ecnomiohyla, Hyloscirtus, Hypsiboas, andPlectrohyla would result in importantchanges in our understanding of thosegroups.
5. Resolution of the taxa herein considered in-sertae sedis: A careful study of availablematerial of the taxa that we could not as-sociate with any of clades that we identifyand, if available, the inclusion of moleculardata derived from them would hopefullypermit their inclusion in the phylogeneticcontext of hylids.
ACKNOWLEDGMENTS
First and foremost, we thank William E.Duellman for his major contribution to ourknowledge of hylid frogs. His lifework wasthe foundation of this project and a perma-nent source of inspiration. Several of hisideas were corroborated here, while we wereunable to find evidence supporting a few oth-ers. These discrepancies are immaterial inour appreciation of his monumental contri-bution.
For the loan of tissue samples employedin this study, voucher specimens, help in thefield, or critical material, we thank (see ap-pendix 4 for institutional abreviations):Raoul Bain, Charles J. Cole, Linda S. Ford,and Charles W. Myers (AMNH); AngeliqueCorthals (AMCC-AMNH); Cornelya Klutsch(ZFMK); Diego F. Cisneros-Heredia (DFCH-USFQ); Eric Smith and Paul Ustach (UTA);Boris L. Blotto, Andres Sehinkman, MartinRamirez, Santiago Nenda, and Gustavo R.Carrizo (MACN); Oscar Flores-Villela andAdrian Nieto Montes de Oca (MZFC);Dwight Lawson (Zoo Atlanta), David Wake,and Meredith Mahoney (MVZ); Jose P. Pom-bal Jr. (MNRJ); Marcio Martins, Miguel T.Rodrigues, and Renata Cecilia Amaro Ghi-lardi (Universidade de Sao Paulo); HussamZaher, Paulo Nuin, and Patricia Narvaes(MZUSP); Frederick H. Sheldon and DonnaDittmann (LSU); W. Ronald Heyer and RoyW. McDiarmid (USNM); Karen Lips(SIUC); Luis Coloma (QCAZ); Jiri Moravec(NMP6V); Steve Lougheed (Queen’s Uni-versity); Ignacio De la Riva (MNCN); DiegoBaldo (Universidad Nacional de Misiones);George Lenglet (IRSNB); James Hanken(MCZ); Frank Glaw (ZSM); Elizabeth Scot(TMSA); Rainer Gunther (ZMB); Barry
2005 129FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Clarke (BMNH); Arnold Kluge and GregSchnider (UMMZ); Robert Murphy (ROM);Steve Donnellan (SAMA); Jorge Williams(MLP); Goran Nilson (NHMG); Janalee P.Caldwell (Sam Noble Oklahoma Museum ofNatural History, University of Oklahoma;samples collected under National ScienceFundation grants DEB-9200779 and DEB-9505518); Rafael de Sa (University of Rich-mond); Renoud Boistel (Universite ParisSud); Luis Canseco-Marquez (UniversidadNacional Autonoma de Mexico); Esteban O.Lavilla (Fundacion Miguel Lillo); MaureenA. Donnelly (Florida International Universi-ty); John D. Lynch (Instituto de Ciencias Na-turales, Universidad Nacional de Colombia);Magno Segalla (Museu de Historia NaturalCapao da Imbuia); Helio da Silva (Univer-sidade Federal Rural do Rio de Janeiro); AnaCarolina de Queiroz Carnaval (University ofChicago); Paul Moler (Florida Fish andWildlife Conservation Commission); RobertN. Fisher (U.S. Geological Survey, San Di-ego, CA); Valerie McKenzie (University ofCalifornia, Santa Barbara); Aaron Bauer(Villanova University); Marcos Vaira (Mu-seo de Ciencias Naturales, Universidad Na-cional de Salta); Claude Gascon (Conserva-tion International); Joseph Mendelsohn (UtahState University); Axel Kwet (StaatlichesMuseum fur Naturkunde, Stuttgart); RogerioP. Bastos (Universidade Federal de Goias);Joao Luis Gasparini (Universidade Federaldo Espırito Santo); Marcelo Gordo (Univer-sidade Federal do Amazonas); Sonia Cechin(Universidade Federal de Santa Maria); IvanSazima (Universidade Estadual de Campi-nas); Marılia T.A. Hartmann, Paulo Hart-mann, Luciana Lugli, Ariovaldo P. CruzNeto, Cynthia P.A. Prado, Luız O. M. Gias-son, and Luıs F. Toledo (Universidade Estad-ual Paulista–Rio Claro); Giovanni Vincipro-va (Universidade Federal do Rio Grande doSul); Phillipe Gaucher (Mission Pour la Cre-ation du Parc de la Guyane); Karl-HeinzJungfer; Sylvain Dubey; A.P. Almeida; PedroCacivio; Martin Henzel; and Pierrino andSandy Mascarino. Their generosity was in-strumental in the completion of this project.
Tissues from Bolivian AMNH specimenswere collected during an expedition support-ed by the Center for Biodiversity and Con-servation at the American Museum of Nat-
ural History and the Center for Environmen-tal Research and Conservation at ColumbiaUniversity, New York, in collaboration withthe Museo de Historia Natural Noel-KempffMercado, Santa Cruz, Bolivia, and ColeccionBoliviana de la Fauna, La Paz. Tissues fromAMNH Vietnamese specimens were collect-ed under National Science Foundation DEB-9870232 to the Center for Biosiversity andConservation at the AMNH. Exportation per-mits of Brazilian samples were issued by Au-torizacao de Acesso e de Remessa de Amos-tra de Componente do Patrimonio Geneticon8 037/2004; permits for collection were is-sued by IBAMA/RAN, licenca 057/03, pro-cesso 02001.002792/98–03.
For discussions on hylid systematics and/or nomenclatural problems, we thank Jose A.Langone, Jose P. Pombal, Jr., Charles W. My-ers, Esteban O. Lavilla, and Roy McDiarmid.Tom Burton answered many questions andrequests of clarifications regarding his im-portant contribution (Burton, 2004). Jan DeLaet and Pablo Goloboff made useful sug-gestions regarding the analyses. Maureen A.Donnelly, Jose P. Pombal, Jr., Charles W.Myers, Jose L. Langone, Taran Grant, Nor-berto P. Giannini, Raoul Bain, Diego Pol, andW. Leo Smith kindly read partially or com-pletely the manuscript and provided usefulcomments and ideas.
Julian Faivovich acknowledges the Amer-ican Museum of Natural History, E3B/Co-lumbia University, NASA/AMNH computa-tional phylogenetic fellowship, AMNH Roo-sevelt Grant, and National Science Founda-tion grant no. DEB-0407632 for financialsupport. Furthermore, he thanks Diego Pol,W. Leo Smith, Taran Grant, Rafael de Sa,Charles J. Cole, Fernando Lobo, and EstebanLavilla for their early encouragment to pur-sue this project.
Celio F.B. Haddad thanks programa BIO-TA/FAPESP (proc. 01/13341–3) and CNPqfor financial support.
Jonathan A. Campbell acknowledges Na-tional Science Foundation grant no. DEB-0102383 for financial support.
Darrel R. Frost and Ward C. Wheeler ac-knowledge support from NASA grantsNAG5–8443 and NAG5–12333. They alsoespecially thank Lisa Gugenheim and Mer-
130 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
rily Sterns for their efforts in support of mo-lecular research at the AMNH.
REFERENCES
Alcalde, L., and S.D. Rosset. ‘‘2003’’ [2004].Descripcion y comparacion del condrocraneoen larvas de Hyla raniceps (Cope, 1862), Sci-nax granulatus (Peters, 1871) y Scinax squali-rostris (A. Lutz, 1925) (Anura: Hylidae). Cuad-ernos de Herpetologıa 17: 33–49.
Almeida, C.G., and A.J. Cardoso. 1985. Variabi-lidade em medidas dos espermatozoides deHyla fuscovaria (Amphibia, Anura) e seu sig-nificado taxonomico. Revista Brasileira deBiologia 45: 387–391.
Altig, R., and R.W. McDiarmid. 1999a. Bodyplan. Development and morphology. In R.W.McDiarmid and R. Altig (editors), Tadpoles.The biology of anuran larvae: 24–51. Chicago:The University of Chicago Press.
Altig, R., and R.W. McDiarmid. 1999b. Diversity.Familial and generic characterizations. In R.W.McDiarmid and R. Altig (editors), Tadpoles.The biology of anuran larvae: 295–337. Chi-cago: The University of Chicago Press.
Alves, A.C.R., and S.P. Carvalho e Silva. 2002.Descriptions of the tadpoles of Scinax alter andScinax cuspidatus. Journal of Herpetology 36:133–137.
Alves, A.C.R., M.d.R. Gomes, and S.P. Carvalhoe Silva. 2004. Description of the tadpole of Sci-nax auratus (Wied-Neuwied) (Anura, Hylidae).Revista Brasileira de Zoologia 21: 315–317.
Anderson, K. 1991. Chromosome evolution inHolarctic Hyla treefrogs. In D.M. Green andS.K. Sessions (editors), Amphibian cytogenet-ics and evolution: 299–331. New York: Aca-demic Press.
Anderson, K. 1996. A karyological perspective onthe monophyly of the hylid genus Osteopilus.In R. Powell and R.W. Henderson (editors),Contributions to West Indian herpetology: atribute to Albert Schwartz: 157–168. Ithaca,NY: Society for the Study of Amphibians andReptiles.
Anstis, M. 2002. Tadpoles of south-eastern Aus-tralia. A guide with keys. Sydney: Reed NewHolland.
Apponyi, M.A., T.L. Pukala, C.S. Brinkworth,V.M. Maselli, J.H. Bowie, M.J. Tyler, G.W.Booker, J.C. Wallace, J.A. Carver, F. Separovic,J.J. Doyle, and L.E. Llewellyn. 2004. Host-de-fense peptides of Australian anurans: structure,mechanisms of action and evolutionary signif-icance. Peptides 25: 1035–1054.
Austin, J.D., S.C. Lougheed, K. Tanner, A.A.Chek, J.P. Bogart, and P.T. Boag. 2002. A mo-
lecular perspective on the evolutionary affini-ties of an enigmatic Neotropical frog, Allophry-ne ruthveni. Zoological Journal of the LinneanSociety 134: 335–346.
Ayarzaguena, J. 1992. Los centrolenidos de laGuayana Venezolana. Publicaciones de la Aso-ciacion de Amigos de Donana: 1–46.
Ayarzaguena, J., and J.C. Senaris. ‘‘1993’’[1994]. Dos nuevas especies de Hyla (Anura;Hylidae) para las Cumbres Tepuyanas del Es-tado Amazonas, Venezuela. Memoria de la So-ciedad de Ciencias Naturales La Salle 53: 127–146.
Ayarzaguena, J., J.C. Senaris, and S. Gorzula.‘‘1992’’ [1993a]. El grupo Osteocephalus rod-riguezi de las tierras altas de la guayana vene-zolana: descripcion de cinco nuevas especies.Memoria Sociedad de Ciencias Naturales LaSalle 52: 133–142.
Ayarzaguena, J., J.C. Senaris, and S. Gorzula.‘‘1992’’ [1993b]. Un nuevo genero para las es-pecies del ‘‘grupo Osteocephalus rodriguezi’’(Anura: Hylidae). Memoria de la Sociedad deCiencias Naturales La Salle 52: 213–221.
Baez, A.M. 2001. Tertiary anurans from SouthAmerica. In H. Heatwole and R.L. Carrol (ed-itors), Amphibian biology, vol. 4: 1388–1401.Sidney: Surrey Beatty and Sons.
Bagnara, J.T. 1974. The tail-darkening reaction ofphyllomedusine tadpoles. Journal of Experi-mental Zoology 187: 149–154.
Bagnara, J.T., and W. Ferris. 1975. The presenceof phyllomedusine melanosomes and pigmentsin Australian hylids. Copeia 1975: 592–595.
Barrio, A., and B. Lutz. 1966. Notas Batracolo-gicas. I. Sobre la supuesta presencia del generoFlectonotus Miranda-Ribeiro en la Argentina.II. Observaciones etoecologicas sobre Trachy-cephalus siemersi (Mertens) (Anura, Hylidae).Physis 26: 107–108.
Barrio, A., and P. Rinaldi de Chieri. 1971. Con-tribucion al esclarecimiento de la posicion taxo-filetica de algunos batracios patagonicos de lafamilia Leptodactylidae mediante el analisiscariotipico. Physis 30: 673–685.
Barrio-Amoros, C.L., and O. Fuentes. 2003. Anew species of Stefania (Anura: Hylidae:Hemiphractinae) from the summit of Cerro Au-tana, Estado Amazonas, Venezuela. Herpetolo-gica 59: 504–512.
Barrio-Amoros, C.L., A. Orellana, and A. Cha-con. 2004. A new species of Scinax (Anura:Hylidae) from the Andes of Venezuela. Journalof Herpetology 38: 105–112.
Basso, N.G. 1995. Hyla raniceps. Geographic dis-tribution. Herpetological Review 26: 105.
Bastos, R.P., and J.P. Pombal, Jr. 1996. A new
2005 131FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
species of Hyla (Anura: Hylidae) from easternBrazil. Amphibia–Reptilia 17: 325–331.
Biju, S.D., and F. Bossuyt. 2003. New frog fromIndia reveals an ancient biogeographical linkwith the Seychelles. Nature 425: 711–714.
Blair, W.F. 1958. Call structure and species groupsin U.S. treefrogs (Hyla). Southwestern Natural-ist 3: 77–89.
Blaylock, L.A., R. Ruibal, and K. Platt-Aloia.1976. Skin structure and wiping behavior ofPhyllomedusinae frogs. Copeia 1976: 283–295.
Block, J.E., S.L. Unser, J.K. Mooney, and E.R.Wild. 2003. Agalychnis craspedopus (Amazonleaf frog). Reproduction. Herpetological Re-view 34: 134–135.
Bogart, J.E. 1973. Evolution of anuran karyo-types. In J.L. Vial (editor), Evolutionary biol-ogy of the anurans. Contemporary research onmajor problems: 329–349. Columbia, MO:University of Missouri Press.
Bokermann, W.C.A. 1962. Cuatro nuevos hylidosdel Brasil. Neotropica 8: 81–91.
Bokermann, W.C.A. 1963. Girinos de anfıbiosbrasileiros—I (Amphibia–Salientia). Anais daAcademia Brasileira de Ciencias 35: 465–474.
Bokermann, W.C.A. 1964a. Dos nuevas especiesde Hyla de Minas Gerais y notas sobre Hylaalvarengai bok (Amphibia, Salientia, Hylidae).Neotropica 10: 67–76.
Bokermann, W.C.A. 1964b. Notes on treefrogs ofthe Hyla marmorata group with description ofa new species (Amphibia, Hylidae). Sencken-bergiana Biologica 45: 243–254.
Bokermann, W.C.A. 1965a. Tres novos batraquiosda regiao central de Mato Grosso, Brasil. Re-vista Brasileira de Biologia 25: 257–264.
Bokermann, W.C.A. 1965b. Hyla langei, a newfrog from Parana, southern Brasil. Journal ofthe Ohio Herpetological Society 5: 49–51.
Bokermann, W.C.A. 1966a. Notas sobre Hylidaedo Espirito Santo (Amphibia, Salientia). Revis-ta Brasileira de Biologia 26: 29–37.
Bokermann, W.C.A. 1966b. O genero PhyllodytesWagler, 1830 (Anura, Hylidae). Anais da Aca-demia Brasileira de Ciencias 38: 335–344.
Bokermann, W.C.A. 1966c. Una nueva especie deTrachycephalus de Bahia, Brasil (Amphibia,Hylidae). Neotropica 12: 120–124.
Bokermann, W.C.A. 1967a. Hyla astartea, novaespecie da Serra do Mar em Sao Paulo (Am-phibia, Hylidae). Revista Brasileira de Biologia27: 157–158.
Bokermann, W.C.A. 1967b. Nova especie de Hylado Amapa (Amphibia, Hylidae). Revista Bras-ileira de Biologia 27: 109–112.
Bokermann, W.C.A. 1967c. Notas sobre cantosnupciais de anfıbios brasileiros. I. (Anura). An-
ais da Academia Brasileira de Ciencias 39:441–443.
Bokermann, W.C.A. 1968. Notas sobre alguns an-fibios Brasileiros descritos por Reinhardt e Lut-ken em 1862 (Amphibia). Revista Brasileira deBiologia 28: 327–329.
Bokermann, W.C.A. 1972. Notas sobre Hyla clep-sydra A. Lutz (Anura, Hylidae). Revista Bras-ileira de Biologia 32: 291–295.
Bokermann, W.C.A. 1973. Duas novas especiesde Sphaenorynchus da Bahia (Anura, Hylidae).Revista Brasileira de Biologia 33: 589–594.
Bokermann, W.C.A., and I. Sazima. 1973a. An-fıbios da Serra do Cipo, Minas Gerais, Brasil.1: Duas especies novas de Hyla (Anura, Hyli-dae). Revista Brasileira de Biologia 33: 521–528.
Bokermann, W.C.A. and I. Sazima. 1973b. Anfı-bios da Serra do Cipo, Minas Gerais, Brasil. 1:Especies novas de ‘‘Hyla’’ (Anura, Hylidae).Revista Brasileira de Biologia 33: 329–336.
Bokermann, W.C.A., and I. Sazima. 1978. Anfı-bios da Serra do Cipo, Minas Gerais, Brasil. 4:Descricao de Phyllomedusa jandaia sp. n. (An-ura, Hylidae). Revista Brasileira de Biologia38: 927–930.
Bonacum, J., R. DeSalle, P. O’Grady, D. Olivera,J. Wintermute, and M. Zilversmith. 2001. Newnuclear and mitochondrial primers for system-atics and comparative genomics in Drosophili-dae. Drosophila Information Service 84: 201–204.
Borkin, L.J. 1999. Distribution of amphibians inNorth Africa, Europe, western Asia, and theformer Soviet Union. In W.E. Duellman (edi-tor), Patterns of distribution of amphibians. Aglobal perspective: 329–420. Baltimore: TheJohns Hopkins University Press.
Bossuyt, F., and M.C. Milinkovitch. 2000. Con-vergent adaptive radiations in Madagascan andAsian ranid frogs reveal covariation betweenlarval and adult traits. Proceedings of the Na-tional Academy of Sciences USA 97: 6585–6590.
Boulenger, G.A. 1882. Catalogue of the BatrachiaSalientia s. Ecaudata in the collection of theBritish Museum, 2nd ed. London: Taylor andFrancis.
Brandao, R.A. 2002. A new species of Phyllo-medusa Wagler, 1830 (Anura: Hylidae) fromcentral Brazil. Journal of Herpetology 36: 571–578.
Bremer, K. 1988. The limits of amino acid se-quence data in angiosperm phylogenetic recon-struction. Evolution 42: 795–803.
Burton, T.C. 1996. Adaptations and evolution inthe hand muscles of Australo-Papuan hylid
132 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
frogs (Anura: Hylidae: Pelodryadinae). Austra-lian Journal of Zoology 44: 611–623.
Burton, T.C. 1998a. Are the distal extensor mus-cles of the fingers of anurans an adaptation toarboreality? Journal of Herpetology 32: 611–617.
Burton, T.C. 1998b. Variation in the hand and su-perficial throat musculature of Neotropical lep-todactylid frogs. Herpetologica 54: 53–72.
Burton, T.C. 2004. Muscles of the pes of hylidfrogs. Journal of Morphology 260: 209–233.
Bushnell, R.J., E.P. Bushnell, and M.V. Parker.1939. A chromosome study of five members ofthe family Hylidae. Journal of the TennesseeAcademy of Science 14: 209–215.
Cadle, J.E., and R. Altig. 1991. Two lotic tadpolesfrom the Andes of southern Peru: Hyla armataand Bufo veraguensis, with notes on the call ofHyla armata (Amphibia: Anura: Hylidae andBufonidae). Studies on Neotropical Fauna andEnvironment 26: 45–53.
Caldwell, J.P. 1974. A re-evaluation of the Hylabistincta species group, with descriptions ofthree new species (Anura: Hylidae). OccasionalPapers of the Museum of Natural History, TheUniversity of Kansas 28: 1–37.
Caldwell, J.P. 1986. A description of the tadpoleof Hyla smithii with comments on tail colora-tion. Copeia 1986: 1004–1006.
Caldwell, J.P. 1989. Structure and behavior ofHyla geographica tadpole schools, with com-ments on classification of group behavior intadpoles. Copeia 1989: 938–950.
Caldwell, J.P. 1992. Diversity of reproductivemodes in anurans: facultative nest constructionin gladiator frogs. In W.C. Hamlett (editor), Re-productive biology of South American verte-brates: 85–97. New York: Springer-Verlag.
Caldwell, J.P. 1994. Natural history and survivalof eggs and early larval stages of Agalychniscalcarifer (Anura: Hylidae). HerpetologicalNatural History 2: 57–66.
Campbell, J.A. 1999. Distribution patterns of Am-phibians in Middle America. In W.E. Duellman(editor), Patterns of distribution of amphibians.A global perspective: 111–210. Baltimore: TheJohns Hopkins University Press.
Campbell, J.A., and W.E. Duellman. 2000. Newspecies of stream-breeding hylid frogs from thenorthern versant of the highlands of Oaxaca,Mexico. Scientific Papers of the Natural His-tory Museum, The University of Kansas 16: 1–28.
Campbell, J.A., and E.N. Smith. 1992. A new frogof the genus Ptychohyla (Hylidae) from the Si-erra de Santa Cruz, Guatemala, and descriptionof a new genus of Middle American stream-breeding treefrogs. Herpetologica 48: 153–167.
Campbell, J.A., E.N. Smith, and M. Acevedo.2000. A new species of fringe-limbed treefrog(Hylidae) from the Sierra de los Cuchumatanesof northwestern Guatemala. Herpetologica 56:250–256.
Cannatella, D.C. 1980. A review of the Phyllo-medusa buckleyi group (Anura: Hylidae). Oc-casional Papers of the Museum of Natural His-tory, The University of Kansas 87: 1–40.
Cannatella, D.C. 1982. Leaf-frogs of the Phyllo-medusa perinesos group (Anura: Hylidae).Copeia 1982: 501–513.
Canseco-Marquez, L., J.R. Mendelson III, and G.Gutierrez-Mayen. 2002. A new species of Hyla(Anura: Hylidae) from the Mixteca Alta, Oa-xaca, Mexico. Herpetologica 58: 260–269.
Canseco-Marquez, L., G. Gutierrez-Mayen, andJ.R. Mendelson III. 2003. Distribution and nat-ural history of the hylid frog Hyla xera in theTehuacan-Cuicatlan valley, Mexico, with a de-scription of the tadpole. The Southwestern Nat-uralist 48: 670–675.
Caramaschi, U. 1989. Revisao do genero Sphae-norhynchus Tschudi, 1838: Composicao, anal-ise cladıstica e biogeografia (Anura, Hylidae).Ph.D. dissertation, Instituto de Biociencias,Universidade de Sao Paulo, Sao Paulo
Caramaschi, U. 1998. Description of a secondspecies of the genus Xenohyla (Anura: Hyli-dae). Amphibia-Reptilia 19: 377–384.
Caramaschi, U., and C.A.G. Cruz. 1998. Notastaxonomicas sobre Pseudis fusca Garman e P.bolboactyla A. Lutz, com a descricao de umanova especie correlata (Anura, Pseudidae). Re-vista Brasileira de Biologia 15: 929–944.
Caramaschi, U., and C.A.G. Cruz. 2000. Duas es-pecies novas de Hyla Laurenti, 1768 do estadode Goias, Brasil (Amphibia, Anura, Hylidae).Boletim do Museu Nacional, Nova Serie, Zool-ogia 422: 1–12.
Caramaschi, U., and R.N. Feio. 1990. A new spe-cies of Hyla (Anura, Hylidae) from southernMinas Gerais, Brazil. Copeia 1990: 542–546.
Caramaschi, U., and J. Jim. 1983. Observacoessobre habitos e desenvolvimento dos girinos dePhyllomedusa vaillanti Boulenger, 1882 (Am-phibia, Anura, Hylidae). Revista Brasileira deBiologia 43: 261–268.
Caramaschi, U. and M.F. Napoli. 2004. The no-menclatural status of the synonyms of Hylapardalis Spix, 1824, and the taxonomic posi-tion of Hyla biobeba Bokerman and Sazima,1973. Journal of Herpetology 38: 501–509.
Caramaschi, U., and H. Niemeyer. 2003. Newspecies of the Hyla albopunctata group fromcentral Brazil (Amphibia, Anura, Hylidae).Boletim do Museu Nacional, Nova Serie, Zool-ogia 504: 1–8.
2005 133FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Caramaschi, U., and M.T. Rodrigues. 2003. Anew large treefrog species, genus Hyla Lauren-ti, 1768, from southern Bahia, Brazil (Amphib-ia, Anura, Hylidae). Arquivos do Museu Na-cional 61: 255–260.
Caramaschi, U., and O.L. Peixoto. 2004. A newspecies of Phyllodytes (Anura: Hylidae) fromthe state of Sergipe, northeastern Brazil. Am-phibia-Reptilia 25: 1–7.
Caramaschi, U., and A. Velosa. 1996. Nova es-pecie de Hyla Laurenti, 1768 do leste brasileiro(Amphibia, Anura, Hylidae). Boletim do Mu-seu Nacional, Nova Serie, Zoologia 365: 1–7.
Caramaschi, U., H.R. da Silva, and M.C.d. Britto–Pereira. 1992. A new species of Phyllodytes(Anura, Hylidae) from southern Bahia, Brazil.Copeia 1992: 187–191.
Caramaschi, U., M.F. Napoli, and A.T. Bernardes.2001. Nova especie do grupo de Hyla circum-data (Cope, 1870) do Estado de Minas Gerais,Brasil (Amphibia, Anura, Hylidae). Boletim doMuseu Nacional, Nova Serie, Zoologia 457: 1–11.
Caramaschi, U., B.V.S. Pimenta, and R.N. Feio.2004a. Nova especie do grupo de Hyla geogra-phica Spix, 1824 da Floresta Atlantica, Brasil(Amphibia, Anura, Hylidae). Boletim do Mu-seu Nacional, Nova Serie, Zoologia 518: 1–14.
Caramaschi, U., O.L. Peixoto, and M.T. Ro-drigues. 2004b. Revalidation and redescriptionof Phyllodytes wuchereri (Peters, 1873) (Am-phibia, Anura, Hylidae). Arquivos do MuseuNacional 62: 185–191.
Cardoso, A.J. 1983. Descricao e biologia de umanova especie de Hyla Laurenti, 1768 (Amphib-ia, Anura, Hylidae). Iheringia, Zoologia 62: 37–45.
Cardoso, A.J., and G.V. Andrade. ‘‘1982’’ [1983].Nova especie de Hyla do Parque Nacional Se-rra da Canastra (Anura, Hylidae). Revista Bras-ileira de Biologia 42: 589–593.
Carnaval, A.C.O.Q., and O.L. Peixoto. 2004. Anew species of Hyla from northeastern Brazil(Amphibia, Anura, Hylidae). Herpetologica 60:387–395.
Carrizo, G.R. 1992. Cuatro especies nuevas de an-uros (Bufonidae: Bufo e Hylidae: Hyla) del nor-te de la Argentina. Cuadernos de Herpetologıa7: 14–23.
Carvalho e Silva, S.P., A.M.P.T. Carvalho e Silva,and E. Izecksohn. 2003. Nova especie de HylaLaurenti do grupo de H. microcephala Cope(Amphibia, Anura, Hylidae) do nordeste doBrasil. Revista Brasileira de Zoologia 20: 553–558.
Castanho, L.M., and I.M.S. De Luca. 2001.Moulting behavior in leaf-frogs of the genus
Phyllomedusa (Anura: Phyllomedusinae). Zool-ogischer Anzeiger 240: 3–6.
Cei, J.M. 1985. Taxonomic and evolutionary sig-nificance of peptides in amphibian skin. Pep-tides 6(suppl. 3): 13–16.
Cei, J.M. 1987. Additional notes to ‘‘Amphibiansof Argentina’’: An update 1980–1986. Moni-tore Zoologico Italiano (N. S.) 21: 209–272.
Cespedez, J.A. 2000. Historia natural de la ranade Pedersen, Argenteohyla siemersi pederseni(Anura: Hylidae). Boletin de la AsociacionHerpetologica Espanola 11: 75–80.
Chang, J. 1996. Inconsistency of evolutionary treetopology reconstruction methods when substi-tution rates vary across characters. Mathemat-ical Biosciences 134: 189–215.
Chek, A.A., S.C. Lougheed, J.P. Bogart, and P.T.Boag. 2001. Perception and history: molecularphylogeny of a diverse group of Neotropicalfrogs, the 30-chromosomes Hyla (Anura: Hy-lidae). Molecular Phylogenetics and Evolution18: 370–385.
Cochran, D.M. 1955. Frogs of southeastern Bra-zil. Bulletin of the United States National Mu-seum 206: 1–423.
Cochran, D.M. 1961. Type specimens of reptilesand amphibians in the United States NationalMuseum. Bulletin of the United States NationalMuseum 220: 1–291.
Cochran, D.M., and C.J. Goin. 1970. Frogs of Co-lombia. Bulletin of the United States NationalMuseum 288: 1–655.
Cocroft, R.B. 1994. A cladistic analysis of chorusfrog phylogeny (Hylidae: Pseudacris). Herpe-tologica 50: 420–437.
Cole, C.J. 1966. The chromosomes of Acris cre-pitans blanchardi Harper (Anura: Hylidae).Copeia 1966: 578–580.
Coloma, L.A. 1995. Ecuadorian frogs of the ge-nus Colostethus (Anura: Dendrobatidae). Mis-cellaneous Publications of the Museum of Nat-ural History, University of Kansas 87: 1–72.
Cope, E.D. 1862. Catalogues of the reptiles ob-tained during the explorations of the Parana,Paraguay, Vermejo and Uruguay Rivers, byCapt. Thos. J. Page, U.S.N.; and of those pro-cured by Lieut. N. Michler, U.S. Top. Eng.,Commander of the expedition conducting thesurvey of the Atrato River. Proceedings of theAcademy Natural Sciences of Philadelphia 14:346–359.
Cope, E.D. 1863. On Trachycephalus, Scaphiopusand other Batrachia. Proceedings of the Acad-emy of Natural Sciences of Philadelphia 15:43–54.
Costa, G.C., G.H.C. Vieira, R.D. Teixeira, A.A.Garda, G.R. Colli, and S.N. Bao. 2004. An ul-trastructural comparative study of the sperm of
134 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Hyla pseudopseudis, Scinax rostratus, and S.squalirostris (Amphibia: Anura: Hylidae).Zoomorphology 123: 191–197.
Cruz, C.A.G. 1973. Observacoes sobre o girinode Sphaenorhynchus planicola (Lutz and Lutz,1938) (Amphibia, Anura, Hylidae). Arquivosde Universidade Federal Rural do Rio de Ja-neiro 3: 83–86.
Cruz, C.A.G. 1982. Conceituacao de grupos deespecies de Phyllomedusinae brasileiras combase em carcteres larvarios (Amphibia, Anura,Hylidae). Arquivos de Universidade FederalRural do Rio de Janeiro 5: 147–171.
Cruz, C.A.G. ‘‘1988’’ [1989]. Sobre Phyllome-dusa aspera e a descricao de uma especie novadesse genero (Amphibia, Anura, Hylidae). Ar-quivos de Universidade Federal Rural do Riode Janeiro 11: 39–44.
Cruz, C.A.G. 1990. Sobre as relacoes intergener-icas de Phyllomedusinae da Floresta Atlantica(Amphibia, Anura, Hylidae). Revista Brasileirade Biologia 50: 709–726.
Cruz, C.A.G., and U. Caramaschi. 1998. Defini-cao, composicao e distribucao geografica dogrupo de Hyla polytaenia Cope, 1870 (Am-phibia, Anura, Hylidae). Boletim do Museu Na-cional, Nova Serie, Zoologia 392: 1–19.
Cruz, C.A.G., and A.G. Dias. 1991. Girinos dogrupo ‘‘microcephala’’ do estado do Rio de Ja-neiro (Amphibia, Anura, Hylidae). RevistaBrasileira de Zoologia 7: 679–683.
Cruz, C.A.G., and O.L. Peixoto. 1980. Notas so-bre o girino de Sphaenorhynchus orophilus(Lutz and Lutz, 1938) (Amphibia, Anura, Hy-lidae). Revista Brasileira de Biologia 40: 383–386.
Cruz, C.A.G., and O.L. Peixoto. ‘‘1984’’ [1985].Especies verdes de Hyla: o complexo ‘‘albosig-nata’’ (Amphibia, Anura, Hylidae). Arquivosde Universidade Federal Rural do Rio de Ja-neiro 7: 31–47.
Cruz, C.A.G., and O.L. Peixoto. ‘‘1985’’ [1987].Especies verdes de Hyla: o complexo ‘‘albof-renata’’ (Amphibia, Anura, Hylidae). Arquivosde Universidade Federal Rural do Rio de Ja-neiro 8: 59–70.
Cruz, C.A.G., U. Caramaschi, and A.G. Dias.2000. Especie nova de Hyla Laurenti, 1768 doEstado do Rio de Janeiro, Brasil (Amphibia,Anura, Hylidae). Boletim do Museu Nacional,Nova Serie, Zoologia 434: 1–8.
Cruz, C.A.G., B.V.S. Pimenta, and D.L. Silvano.2003. Duas novas especies pertenecentes aocomplexo de Hyla albosignata Lutz and Lutz,1938, do leste do Brasil (Amphibia, Anura, Hy-lidae). Boletim do Museu Nacional, Nova Ser-ie, Zoologia 503: 1–13.
Cunningham, M. 2002. Identification and evolu-
tion of Australian torrent treefrogs (Anura: Hy-lidae: Litoria nannotis group). Memoirs of theQueensland Museum 48: 93–102.
Darst, C.R., and D.C. Cannatella. 2004. Novel re-lationships among hyloid frogs inferred from12S and 16S mitochondrial DNA sequences.Molecular Phylogenetics and Evolution 31:462–475.
da Silva, H.R. 1997. Two character states new forhylines and the taxonomy of the genus Pseu-dacris. Journal of Herpetology 31: 609–613.
da Silva, H.R. 1998. Phylogenetic relationships ofthe family Hylidae with emphasis on the rela-tionships within the subfamily Hylinae (Am-phibia: Anura). Ph.D. dissertation, Departmentof Systematics and Ecology, University of Kan-sas.
da Silva, H.R., M.C. De Britto-Pereira, and U.Caramaschi. 1989. Frugivory and seed dispers-al by Hyla truncata, a Neotropical tree-frog.Copeia 1989: 781–783.
De Laet, J. 2005. Parsimony and the problem ofinapplicables in sequence data. In V.A. Albert(editor), Parsimony, phylogeny, and genomics:81–116. Oxford University Press.
De Laet, J., and E. Smets. 1998. On the three-taxon approach to parsimony analysis. Cladis-tics 14: 363–381.
Delahoussaye, A.J. 1966. The comparative spermmorphology of the Louisiana Hylidae (Am-phibia: Anura). Proceedings of the LouisianaAcademy of Sciences 29: 140–152.
De la Riva, I., 1999. A new Phyllomedusa fromsouthwestern Amazonia (Amphibia: Anura:Hylidae). Revista Espanola de Herpetologıa 13:123–131.
De la Riva, I., and W.E. Duellman. 1997. Theidentity and distribution of Hyla rossalleniGoin. Amphibia-Reptilia 18: 433–436.
De la Riva, I., J. Kohler, S. Lotters, and S. Rei-chle. 2000. Ten years of research on Bolivianamphibians: updated checklist, distribution,taxonomic problems, literature and iconogra-phy. Revista Espanola de Herpetologıa 14: 19–164.
Delfino, G., B.B. Alvarez, R. Brizzi, and J.A. Ces-pedez. 1998. Serous cutaneous glands of Ar-gentine Phyllomedusa Wagler 1830 (Anura Hy-lidae): secretory polymorphism and adaptiveplasticity. Tropical Zoology 11: 333–351.
Delfino, G., D. Nosi, and F. Giachi. 2001. Secre-tory granule-cytoplasm relationships in serousglands of anurans: ultrastructural evidence andpossible functional role. Toxicon 39: 1161–1171.
Delfino, G., R. Brizzi, D. Nosi, and A. Terreni.2002. Serous cutaneous glands in New Worldhylid frogs: an ultrastructural study on skin poi-
2005 135FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
sons confirms phylogenetic relationships be-tween Osteopilus septentrionalis and Phrynoh-yas venulosa. Journal of Morpholgy 253: 176–186.
de Sa, R.O. 1983. Descripcion de la larva de Ar-genteohyla siemersi (Mertens, 1937), (Anura:Hylidae). Resumenes y Comunicaciones de lasJornadas de Ciencias Naturales, Montevideo 3:40–41.
de Sa, R.O. 1995. Hyla albopunctata. Catalogue ofAmerican Amphibians and Reptiles 602: 1–5.
de Sa, R.O. 1996. Hyla multifasciata. Catalogue ofAmerican Amphibians and Reptiles 624: 1–4.
de Sa, R.O., and E.O. Lavilla. 1997. The tadpoleof Pseudis minuta (Anura: Pseudidae), an ap-parent case of heterochrony. Amphibia-Reptilia18: 229–240.
D’Heursel, A., and R.O. de Sa. 1999. Comparingthe tadpoles of Hyla geographica and Hylasemilineata. Journal of Herpetology 33: 353–361.
D’Heursel, A., and C.F.B. Haddad. 2002. School-ing and swimming behaviors of Hyla semili-neata tadpoles (Anura, Hylidae). Iheringia,Zoologia 92: 99–104.
Donnellan, S.C., and M.J. Mahony. 2004. Allo-zyme, chromosomal and morphological vari-ability in the Litoria lesueuri species group(Anura: Hylidae), including a description of anew species. Australian Journal of Zoology 52:1–28.
Donnelly, M.A., and C.W. Myers. 1991. Herpe-tological results of the 1990 Venezuelan expe-dition to the summit of Cerro Guaiquinima,with new tepui reptiles. American MuseumNovitates 3017: 1–54.
Donnelly, M.A., C. Guyer, D.M. Krempels, andH.E. Braker. 1987. The tadpole of Agalychniscalcarifer (Anura: Hylidae). Copeia 1987: 247–250.
Dubois, A. 1982. Dendrobates Wagler, 1830 andDendrobatidae Cope, 1865 (Amphibia, Anura):proposed conservation. Bulletin of ZoologicalNomenclature 39: 267–278.
Dubois, A. 1987. Miscellanea taxinomica batrach-ologica (I). Alytes 5: 7–95.
Duellman, W.E. 1963. A review of the MiddleAmerican tree frogs of the genus Ptychohyla.University of Kansas Publications, Museum ofNatural History 15: 297–349.
Duellman, W.E. 1966. Taxonomic notes on someMexican and Central American Hylid frogs.University of Kansas Publications, Museum ofNatural History 17: 263–279.
Duellman, W.E. 1967. Additional studies of chro-mosomes of anuran amphibians. SystematicZoology 16: 38–43.
Duellman, W.E. 1968a. The genera of phyllome-
dusine frogs (Anura: Hylidae). University ofKansas Publications, Museum of Natural His-tory 18: 1–10.
Duellman, W.E. 1968b. The taxonomic status ofsome American hylid frogs. Herpetologica 24:194–209.
Duellman, W.E. 1969. Phyllomedusa buckleyiBoulenger: variation, distribution and synony-my. Herpetologica 25: 134–140.
Duellman, W.E. 1970. Hylid frogs of MiddleAmerica. Monographs of the Museum of Nat-ural History, University of Kansas 1–2: 1–753.
Duellman, W.E. 1971a. The nomenclatural statusof the names Hyla boans (Linnaeus) and Hylamaxima (Laurenti) (Anura: Hylidae). Herpeto-logica 27: 397–405.
Duellman, W.E. 1971b. A taxonomic review ofSouth American hylid frogs, genus Phrynoh-yas. Occasional Papers of the Museum of Nat-ural History, The University of Kansas 4: 1–21.
Duellman, W.E. 1972a. The systematic status andlife history of Hyla rhodopepla Gunther. Her-petologica 28: 369–375.
Duellman, W.E. 1972b. A review of the Neotrop-ical frogs of the Hyla bogotensis group. Occa-sional Papers of the Museum of Natural His-tory, University of Kansas 11: 1–31.
Duellman, W.E. 1973a. Frogs of the Hyla geogra-phica group. Copeia 1973: 515–533.
Duellman, W.E. 1973b. Descriptions of new hylidfrogs from Colombia and Ecuador. Herpetolo-gica 29: 219–227.
Duellman, W.E. 1974. A reassessment of the tax-onomic status of some Neotropical hylid frogs.Occasional Papers of the Museum of NaturalHistory, University of Kansas 27: 1–27.
Duellman, W.E. 1977. Liste der rezenten Amphi-bien und Reptilien. Hylidae, Centrolenidae,Pseudidae. Das Tierreich 95: 1–225.
Duellman, W.E. 1978. The biology of an Equa-torial herpetofauna in Amazonian Ecuador.Miscellaneous Publications of the Museum ofNatural History of the University of Kansas 65:1–352.
Duellman, W.E. 1982. A new species of small yel-low Hyla from Peru (Anura: Hylidae). Am-phibia-Reptilia 3: 153–160.
Duellman, W.E. 1989. New species of hylid frogsfrom the Andes of Colombia and Venezuela.Occasional Papers of the Museum of NaturalHistory, The University of Kansas 131: 1–12.
Duellman, W.E. 1999. Distribution patterns ofamphibians in South America. In W.E. Duell-man (editor), Patterns of distribution of am-phibians. A global perspective: 255–328. Bal-timore: The Johns Hopkins University Press.
Duellman, W.E. 2001. Hylid frogs of Middle
136 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
America. Ithaca, NY: Society for the Study ofAmphibians and Reptiles.
Duellman, W.E., and J.A. Campbell. 1992. Hylidfrogs of the genus Plectrohyla: systematics andphylogenetic relationships. Miscellaneous Pub-lications of the Museum of Zoology, Universityof Michigan 181: 1–32.
Duellman, W.E., and C.J. Cole. 1965. Studies ofchromosomes of some anuran amphibians (Hy-lidae and Centrolenidae). Systematic Zoology14: 139–143.
Duellman, W. E., and L.A. Coloma. 1993. Hylastaufferorum, a new species of treefrog in theHyla larinopygion group from the cloud forestof Ecuador. Occasional Papers of the Museumof Natural History, The University of Kansas161: 1–11.
Duellman, W.E., and M. Crump. 1974. Speciationin frogs of the Hyla parviceps group in the up-per Amazon basin. Occasional Papers of theMuseum of Natural History, University of Kan-sas 23: 1–40.
Duellman, W.E., and R.O. de Sa. 1988. A newgenus and species of South American hylidfrog with a highly modified tadpole. TropicalZoology 1: 117–136.
Duellman, W.E., and J.M.J. Fouquette. 1968.Middle American frogs of the Hyla microce-phala group. University of Kansas Publicationsof the Museum of Natural History 17: 517–557.
Duellman, W.E., and P. Gray. 1983. Developmen-tal biology and systematics of the egg-broodinghylid frogs, genera Flectonotus and Fritziana.Herpetologica 39: 333–359.
Duellman, W.E., and D.M. Hillis. 1990. System-atics of frogs of the Hyla larinopygion group.Occasional Papers of the Museum of NaturalHistory, University of Kansas 134: 1–23.
Duellman, W.E., and M.S. Hoogmoed. 1984. Thetaxonomy and phylogenetic relationships of thehylid frog genus Stefania. Miscellaneous Pub-lications of the Museum of Natural History ofthe University of Kansas 75: 1–39.
Duellman, W.E., and M.S. Hoogmoed. 1992.Some hylid frogs from the Guiana highlands,northeastern south America: new species, dis-tributional records, and a generic reallocation.Occasional Papers of the Museum of NaturalHistory, University of Kansas 147: 1–21.
Duellman, W.E., and J.R. Mendelson III. 1995.Amphibians and reptiles from northern Depar-tamento Loreto, Peru: taxonomy and biogeog-raphy. The University of Kansas Science Bul-letin 55: 329–376.
Duellman, W.E., and L. Trueb. 1966. Neotropicalhylid frogs, genus Smilisca. University of Kan-sas Publications Museum of Natural History17: 281–375.
Duellman, W.E., and L. Trueb. 1976. The system-atic status and relationships of the hylid frogNyctimantis rugiceps Boulenger. OccasionalPapers of the Museum of Natural History, Uni-versity of Kansas 58: 1–14.
Duellman, W.E., and L. Trueb. 1983. Frogs of theHyla columbiana group: taxonomy and phylo-genetic relationships. In A.G.J. Rhodin and K.Miyata (editors), Advances in herpetology andevolutionary biology: 33–51. Cambridge, MA:Museum of Comparative Zoology, HarvardUniversity.
Duellman, W.E., and L. Trueb. 1986. Biology ofamphibians. New York: McGraw-Hill.
Duellman, W.E., and L. Trueb. 1989. Two newtreefrogs of the Hyla parviceps group from theAmazon basin in southeastern Peru. Herpeto-logica 45: 1–10.
Duellman, W.E., and J.J. Wiens. 1992. The statusof the hylid frog genus Ololygon and the rec-ognition of Scinax Wagler, 1830. OccasionalPapers of the Museum of Natural History, Uni-versity of Kansas 151: 1–23.
Duellman, W.E., and M. Yoshpa. 1996. A newspecies of Tepuihyla (Anura: Hylidae) fromGuyana. Herpetologica 52: 275–281.
Duellman, W.E., I. De la Riva, and E.R. Wild.1997. Frogs of the Hyla armata and Hyla pul-chella groups in the Andes of South America,with definitions and analyses of phylogeneticrelationships of Andean groups of Hyla. Sci-entific Papers of the Natural History Museum,The University of Kansas 3: 1–41.
Duellman, W.E., J.E. Cadle, and D.C. Cannatella.1988a. A new species of terrestrial Phyllome-dusa (Anura: Hylidae) from southern Peru.Herpetologica 44: 91–95.
Duellman, W.E., L.R. Maxson, and C.A. Jesi-olowski. 1988b. Evolution of marsupial frogs(Hylidae: Hemiphractinae): immunological ev-idence. Copeia 1988: 527–543.
Dumeril, A.M.C., and G. Bibron. 1841. Erpeto-logie generale ou histoire naturelle completedes reptiles, tome 8. Paris: Librairie Encyclo-pedique de Roret.
Dunn, E.R. 1926. The frogs of Jamaica. Proceed-ings of the Boston Society of Natural History38: 11–130.
Dunn, E.R. 1943. An extraordinary new Hylafrom Colombia. Caldasia 1943: 309–311.
Echeverria, D.D. 1997. Microanatomy of the buc-cal apparatus and oral cavity of Hyla minutaPeters, 1872 larvae (Anura, Hylidae), with dataon feeding habits. Alytes 15: 26–36.
Eliosa Leon, H.R. 2002. Variacion geografica enHyla eximia (Anura: Hylidae). Boletin de la So-ciedad Herpetologica Mexicana 10: 59–60.
Emerson, S.B., C. Richards, R.C. Drewes, and
2005 137FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
K.M. Kjer. 2000. On the relationships amongranoid frogs: a review of the evidence. Herpe-tologica 56: 209–230.
Erspamer, V. 1994. Bioactive secretions of theamphibian integument. In H. Heatwole andG.T. Barthalmus (editors), Amphibian biology,vol. 1: 178–350. Surrey: Beatty & Sons.
Estes, R. 1970. Origin of the recent North Amer-ican lower vertebrate fauna: An inquiry into thefossil record. Forma et Functio 3: 139–163.
Estes, R., and A.M. Baez. 1985. Herpetofauna ofNorth and South America during the late Cre-taceous and Cenozoic: evidence for inter-change? In F.G. Stehli and S.D. Webb (editors),The great American biotic interchange: 139–197. New York: Plenum Press.
Estes, R., and O. Reig. 1973. The early fossil re-cord of frogs: a review of the evidence. In J.Vial (editor), Evolutionary biology of the an-urans: 11–63. Columbia, MO: University ofMissouri Press.
Eterovick, P.C., and R.A. Brandao. 2001. A de-scription of the tadpoles and advertisementcalls of members of the Hyla pseudopseudisgroup. Journal of Herpetology 35: 442–450.
Eterovick, P.C., I.S. Barros, and I. Sazima. 2002.Tadpoles of two species in the Hyla polytaeniaspecies group and comparison with other tad-poles of Hyla polytaenia and Hyla pulchellagroups (Anura, Hylidae). Journal of Herpetol-ogy 36: 512–515.
Fabrezi, M. 2001. A survey of prepollex and pre-hallux variation in anuran limbs. ZoologicalJournal of the Linnean Society 131: 227–248.
Fabrezi, M., and E.O. Lavilla. 1992. Estructuradel condrocraneo y esqueleto hiobranquial enlarvas de algunos hılidos neotropicales (Anura:Hylidae). Acta Zoologica Lilloana 41: 155–164.
Faivovich, J. 2002. A cladistic analysis of Scinax(Anura: Hylidae). Cladistics 18: 367–393.
Faivovich, J., C.A.G. Cruz, and O.L. Peixoto.2002. Identity of Hyla ehrhardti Muller, 1924(Anura, Hylidae). Journal of Herpetology 36:325–327.
Faivovich, J., P.C.A. Garcia, F. Ananias, L. Lanari,N.G. Basso, and W.C. Wheeler. 2004. A mo-lecular perspective on the phylogeny of theHyla pulchella species group (Anura, Hylidae).Molecular Phylogenetics and Evolution 32:938–950.
Farris, J.S. 1983. The logical basis of phyloge-netic analysis. In N.I. Platnick and V.A. Funk(editors), Advances in cladistics: Proceedingsof the Third Meeting of the Willi Hennig So-ciety, vol. 2: 7–36. New York: Columbia Uni-versity Press.
Farris, J.S. 1999. Likelihood and inconsistency.Cladistics 15: 199–204.
Farris, J.S. 2000. Corroboration versus ‘‘strongestevidence’’. Cladistics 16: 385–393.
Farris, J.S., V.A. Albert, M. Kallersjo, D. Lip-scomb, and A.G. Kluge. 1996. Parsimony jack-knifing outperforms neighbor-joining. Cladis-tics 12: 99–124.
Feller, A., and S.B. Hedges. 1998. Molecular ev-idence for the early history of living amphibi-ans. Molecular Phylogenetics and Evolution 9:509–516.
Firschein, I.L., and H.M. Smith. 1956. A newfringe-limbed Hyla (Amphibia: Anura) from anew faunal district of Mexico. Herpetologica12: 17–21.
Fitzinger, L.J. 1843 [1973]. Systema Reptilium.Athens, OH: SSAR, p. 128.
Ford, L.S., and D.C. Cannatella. 1993. The majorclades of frogs. Herpetological Monographs 7:94–117.
Fouquette, M.J., and A.J. Delahoussaye. 1977.Sperm morphology in the Hyla rubra group(Amphibia, Anura, Hylidae), and its bearing ongeneric status. Journal of Herpetology 11: 387–396.
Franz, R. 2003. Wet mountains and mountainfrogs. In R.W. Henderson and R. Powell (edi-tors), Islands and the sea: essays on herpetolog-ical exploration in the West Indies. Society forthe Study of the Amphibians and Reptiles. Con-tributions to Herpetology, vol. 20: 159–167.Ithaca, NY.
Frost, D.R. 1985. Amphibian species of the world:a taxonomic and geographic reference. Law-rence, KS: Association of Systematic Collec-tions and Allen Press.
Frost, D.R. 2004. Amphibian species of the world:an online reference, version 3.0 (22 August2004). Electronic database accessible at http://research.amnh.org/herpetology/amphibia/in-dex.html. American Museum of Natural His-tory.
Frost, D.R., M.T. Rodrigues, T. Grant, and T.A.Titus. 2001. Phylogenetics of the lizard genusTropidurus (Squamata: Tropiduridae: Tropidur-inae): direct optimization, descriptive efficien-cy, and sensitivity analysis of congruence be-tween molecular data and morphology. Molec-ular Phylogenetics and Evolution 21: 352–371.
Funkhouser, A. 1957. A review of the Neotropicaltree frogs of the genus Phyllomedusa. Occa-sional Papers of the Natural History Museumof Stanford University 5: 1–90.
Gallardo, J.M. 1964. Observaciones biologicassobre Hyla raddiana Fitz., en la provincia deBuenos Aires. Ciencia E Investigacion 17: 63–69.
138 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Gallardo, J.M. 1970. Estudio ecologico sobre losanfibios y reptiles del sudeste de la provinciade Buenos Aires, Argentina. Revista del MuseoArgentino de Ciencias Naturales ‘‘BernardinoRivadavia’’, Zoologia 10: 27–63.
Garcia, P.C.A. 2003. Revisao taxonomica e anal-ise filogenetica das especies do genero HylaLaurenti do complexo marginata/semiguttata(Amphibia, Anua, Hylidae). Ph.D. dissertation,Universidade Estadual Paulista, Rio Claro, SaoPaulo, Brazil.
Garcia, P.C.A., U. Caramaschi, and A. Kwet.2001a. O status taxonomico de Hyla cochranaeMertens e recaracterizacao de Aplastodiscus A.Lutz (Anura, Hylidae). Revista Brasileira deZoologia 18: 1197–1218.
Garcia, P.C.A., G. Vinciprova, and C.F.B. Haddad.2001b. Vocalizacao, girino, distribucao geo-grafica e novos comentarios sobre Hyla mar-ginata Boulenger, 1887 (Anura, Hylidae, Hyli-nae). Boletim do Museu Nacional, Nova Serie,Zoologia 460: 1–19.
Garcia, P.C.A., G. Vinciprova, and C.F.B. Haddad.2003. The taxonomic status of Hyla pulchellajoaquini B. Lutz, 1968 (Anura: Hylidae). Her-petologica 59: 350–363.
Garda, A.A., G.C. Costa, G.R. Colli, and S.N.Bao. 2004. Spermatozoa of Pseudinae (Am-phibia, Anura, Hylidae), with a test of the hy-pothesis that sperm ultrastructure correlateswith reproductive modes in anurans. Journal ofMorphology 261: 196–205.
Gaudin, A.J. 1974. An osteological analysis ofHolarctic tree frogs, family Hylidae. Journal ofHerpetology 8: 141–152.
Giribet, G. 2001. Exploring the behavior of POY,a program for direct optimization of moleculardata. Cladistics 17: S60–S70.
Goeldi, E. 1907. Description of Hyla resinifictrixGoeldi, a new Amazonian tree frog peculiar forits breeding habits. Proceedings of the Zoolog-ical Society of London 1907: 135–140.
Goin, C.J. 1961. Synopsis of the genera of hylidfrogs. Annals of the Carnegie Museum 36: 5–18.
Goin, C.J., and O.B. Goin. 1968. A new tree frogfrom Suriname. Copeia 3: 581–583.
Goin, C.J., and J.D. Woodley. 1969. A new tree-frog from Guyana. Zoological Journal of theLinnean Society 48: 135–140.
Goloboff, P.A. 1999. Analyzing large data sets inreasonable times: solutions for composite opti-ma. Cladistics 15: 415–428.
Goloboff, P.A. 2003. Parsimony, likelihood, andsimplicity. Cladistics 19: 91–103.
Goloboff, P.A., and J.S. Farris. 2001. Methods forquick consensus estimation. Cladistics 17: S26–S34.
Goloboff, P.A., J.S. Farris, and K.C. Nixon. 2000.T.N.T.: tree analysis using new technology. Pro-gram and documentation, version 1.0. Avail-able from the authors, and at www.zmuc.dk/public/phylogeny.
Gomes, M.R., and O.L. Peixoto. 1991a. Consi-deracoes sobre os girinos de Hyla senicula(Cope, 1868) e Hyla soaresi (Caramaschi eJim, 1983) (Amphibia, Anura, Hylidae). ActaBiologica Leopoldensia 13: 5–18.
Gomes, M.R., and O.L. Peixoto. 1991b. Larvasde Hyla do grupo ‘‘leucophyllata’’ com a des-cricao da de H. elegans Wied, 1824 e notassobre a variacao do padrao de colorido do ad-ulto nesta especie (Anura, Hylidae). RevistaBrasileira de Biologia 51: 257–262.
Gomes, M.R., and O.L. Peixoto. 2002. O girinode Hyla leucopygia Cruz and Peixoto, 1984(Amphibia, Anura, Hylidae). Boletim do Mu-seu de Biologia Mello Leitao, Nova Serie 13:17–25.
Gorham, S.W. 1974. Checklist of world amphib-ians up to January 1, 1970. Saint John, Canada:New Brunswick Museum.
Gorzula, S., and C. Senaris. ‘‘1998’’ [1999]. Con-tribution to the herpetofauna of the VenezuelanGuyana: I. A data base. Scientia Guianae 8: 1–270.
Grant, T., E.C. Humphrey, and C.W. Myers. 1997.The medial lingual process of frogs: a bizarrecharacter of old world ranoids discovered inSouth American dendrobatids. American Mu-seum Novitates 3212: 1–40.
Gray, J.E. 1825. A synopsis of the genera of rep-tiles and Amphibia, with a description of somenew species. Annals of Philosophy, ser. 2, 10:193–217.
Graybeal, A. 1993. The phylogenetic utility of cy-tochrome b: lessons from bufonid frogs. Mo-lecular Phylogenetics and Evolution 2: 256–269.
Graybeal, A. 1997. Phylogenetic relationships ofbufonid frogs and tests of alternate macroevo-lutionary hypotheses characterizing their radi-ation. Zoological Journal of the Linnean Soci-ety 119: 297–338.
Gruber, S.L. 2002. Estudos citogeneticos em es-pecies do genero Hyla (Anura, Hylidae) dosgrupos com 2n 5 24 e 2n 5 30 cromossomos,com tecnicas de coloracao diferencial. Master’sthesis, Universidade Estadual Paulista, Rio Cla-ro, Sao Paulo, Brazil.
Guillaume, C.P., P. Gaucher, and F.M. Catzeflis.2001. Does Hyla hadroceps belong to the genusPhrynohyas (Hylidae: Batrachia)? First ap-proach using the mitochondrial gene 12S,rRNA. 11th ordinary general meeting of Socie-tas Europaea Herpetologica (SEH), Zalec, Slov-
2005 139FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
enie, 13–17 VII 2001. Summary in Biota2(suppl.): 68–69.
Guimaraes, L.D., R. de Freitas Juliano, and R.P.Bastos. 2001. Life history notes: Hyla raniceps(NCN). Combat. Herpetological Review 32:102–103.
Haas, A. 1996. Das larvale cranium von Gastro-theca riobambae und seine metamorphose(Amphibia, Anura, Hylidae). Verhandlungendes Naturwissenschaftlichen Vereins in Ham-burg 36: 33–162.
Haas, A. 2003. Phylogeny of frogs as inferredfrom primarily larval characters (Amphibia:Anura). Cladistics 19: 23–89.
Haas, A., and S.J. Richards. 1998. Correlations ofcranial morphology, ecology, and evolution inAustralian suctorial tadpoles of the genera Li-toria and Nyctimystes (Amphibia: Anura: Hy-lidae: Pelodryadinae). Journal of Morphology238: 109–141.
Haddad, C.F.B., and R.J. Sawaya. 2000. Repro-ductive modes of Atlantic forest hylid frogs: ageneral overview and the description of a newmode. Biotropica 32: 862–871.
Haddad, C.F.B., J. Faivovich, and P.C.A. Garcia.2005. The specialized reproductive mode of thetreefrog Aplastodiscus perviridis (Anura: Hyli-dae). Amphibia-Reptilia. 26: 87–92.
Hall, T.A. 1999. BioEdit: a user-friendly biologi-cal sequence alignment editor and analysis. De-partment of Microbiology. North Carolina StateUniversity.
Hartmann, M.T., P.A. Hartmann and C.F.B. Had-dad. 2004. Visual signaling and reproductivebiology in a nocturnal treefrog genus Hyla (An-ura: Hylidae). Amphibia-Reptilia 25: 395–406.
Hass, C.A., L.R. Maxson, and S.B. Hedges. 2001.Relationships and divergence times of West In-dian amphibians and reptiles: insights from al-bumin immunology. In C.A. Woods and F.E.Sergile (editors), Biogeography of the West In-dies. Patterns and perspectives: 157–174. BocaRaton, FL: CRC Press.
Hawkins, J.A., C.E. Hughes, and R.W. Scotland.1997. Primary homology assessment, charac-ters, and character states. Cladistics 13: 275–283.
Hay, J.M., I. Ruvinsky, S.B. Hedges, and L.R.Maxson. 1995. Phylogenetic relationships ofamphibian families inferred from DNA se-quences of mitochondrial 12S and 16S ribo-somal RNA genes. Molecular Biology andEvolution 12: 928–937.
Hedges, S.B. 1986. An electrophoretic analysis ofHolarctic hylid frog evolution. Systematic Zo-ology 35: 1–21.
Hedges, S.B. 1987. Vocalization and habitat pref-erence of the Jamaican treefrog Hyla marianae
(Anura, Hylidae). Caribbean Journal of Science23: 380–384.
Hedges, S.B. 1994. Molecular evidence for theorigin of birds. Proceedings of the NationalAcademy of Sciences USA 91: 2621–2624.
Hedges, S.B. 1996. The origin of West Indian am-phibians and reptiles. In R. Powell and R.W.Henderson (editors), Contributions to West In-dian herpetology: a tribute to Albert Schwartz:95–128. Ithaca, NY: Society for the Study ofAmphibians and Reptiles.
Heyer, W.R. 1977. Taxonomic notes on frogs fromthe Madeira and the Purus rivers, Brasil. PapeisAvulsos de Zoologia 31: 141–162.
Heyer, W.R. 1980. The calls and taxonomic po-sitions of Hyla giesleri and Ololygon opalina(Amphibia: Anura: Hylidae). Proceedings ofthe Biological Society of Washington 93: 655–661.
Heyer, W.R. 1985. New species of frogs fromBoraceia, Sao Paulo, Brazil. Proceedings of theBiological Society of Washington 98: 657–671.
Heyer, W.R., A.S. Rand, C.A.G. Cruz, O.L. Pei-xoto, and C.E. Nelson. 1990. Frogs of Bora-ceia. Arquivos de Zoologia 31: 231–410.
Hillis, D.M., and M.T. Dixon. 1991. RibosomalRNA: molecular evolution and phylogenetic in-ference. Quarterly Review of Biology 66: 411–453.
Hillis, D.M., L.K. Ammerman, M.T. Dixon, andR.O. de Sa. 1993. Ribosomal DNA and thephylogeny of frogs. Herpetological Mono-graphs 7: 118–131.
Hoegg, S., M. Vences, H. Brinkmann, and A.Meyer. 2004. Phylogeny and comparative sub-stitution rates of frogs inferred from sequencesof three nuclear genes. Molecular Biology andEvolution 21: 1188–1200.
Holman, A.J. 1968. Lower Oligocene amphibiansfrom Saskatchewan. Quarterly Journal of theFlorida Academy of Sciences 31: 273–289.
Holman, A.J. 2003. Fossil frogs and toads ofNorth America. Bloomington: Indiana Univer-sity Press.
Hoogmoed, M.S. 1979. Resurrection of Hyla or-natissima Noble (Amphibia, Hylidae) and re-marks on related species of green tree frogsfrom the Guiana area. Notes on the herpetofau-na of Surinam VI. Zoologische Verhandelingen172: 1–46.
Hoogmoed, M.S. 1990. Resurrection of Hyla wav-rini Parker (Amphibia: Anura: Hylidae), a glad-iator frog from northern South America. Zool-ogische Mededeelingen 64: 71–93.
Hoogmoed, M.S., and J.E. Cadle. 1991. Naturalhistory and distribution of Agalychnis craspe-dopus (Funkhouser, 1957) (Amphibia: Anura:
140 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Hylidae). Zoologische Mededeelingen 65: 129–142.
Hutchinson, M.N., and L.R. Maxson. 1986. Im-munological evidence on relationships of someAustralian terrestrial frogs (Anura: Hylidae: Pe-lodryadinae). Australian Journal of Zoology 34:575–582.
Hutchinson, M.N., and L.R. Maxson. 1987. Phy-logenetic relationships among Australian treefrogs (Anura: Hylidae: Pelodryadinae): an im-munological approach. Australian Journal ofZoology 35: 61–74.
ICZN. 1999. International code of zoological no-menclature, 4th ed. London: International Trustfor Zoological Nomenclature.
Iturralde-Vinent, M.A., and R.D.E. MacPhee.1999. Paleogeography of the Caribbean region:implications for Cenozoic biogeography. Bul-letin of the American Museum of Natural His-tory 238: 1–95.
Izecksohn, E. 1959. Uma nova especie de ‘‘Hy-lidae’’ da baixada fluminense, Estado de Rio deJaneiro, Brasil (Amphibia, Anura). RevistaBrasileira de Biologia 19: 259–263.
Izecksohn, E. 1988. Algumas consideracoes sobreo genero Euparkerella, com a descricao de tresnovas especies (Amphibia, Anura, Leptodactyl-idae). Revista Brasileira de Biologia 48: 59–74.
Izecksohn, E. 1996. Novo genero de Hylidae bras-ileiro (Amphibia, Anura). Revista de Universi-dade Rural, Serie Ciencias da Vida 18: 47–52.
Izecksohn, E., and C.A.G. Cruz. 1976. Nova es-pecie de Phyllomedusinae do Estado do Espır-ito Santo, Brasil (Amphibia, Anura, Hylidae).Revista Brasileira de Biologia 36: 257–261.
Jared, C., M.M. Antoniazzi, E. Katchburian, R.C.Toledo, and E. Freymuller. 1999. Some aspectsof the natural history of the casque-headed treefrog Corythomantis greeningi Boulenger (Hy-lidae). Annales des Sciences Naturelles 3: 105–115.
Jim, J. 1980. Aspectos ecologicos dos anfıbios re-gistrados na regiao de Botucatu, Sao Paulo(Amphibia, Anura). Ph.D. dissertation, Univ-ersidade de Sao Paulo.
Jim, J., and U. Caramaschi. 1979. Uma nova es-pecie da regiao de Botucatu, Sao Paulo, Brasil(Amphibia, Anura). Revista Brasileira de Biol-ogia 39: 717–719.
Jungfer, K.H. 1996. Reproduction and parentalcare of the coronated treefrog, Anotheca spi-nosa (Steindachner, 1864) (Anura: Hylidae).Herpetologica 52: 25–32.
Jungfer, K.H., and W. Hodl. 2002. A new speciesof Osteocephalus from Ecuador and a rede-scription of O. leprieurii (Dumeril and Bibron,1841) (Anura: Hylidae). Amphibia-Reptilia 23:21–46.
Jungfer, K.H., and E. Lehr. 2001. A new speciesof Osteocephalus with bicoloured iris from Po-zuzo (Peru: Departamento de Pasco) (Amphib-ia: Anura: Hylidae). Zoologische Abhandlun-gen Staatliches Museum fur Tierkunde, Dres-den 51: 321–329.
Jungfer, K.H., and L.C. Schiesari. 1995. Descrip-tion of a central Amazonian and Guyanan treefrog, genus Osteocephalus (Anura, Hylidae),with oophagous tadpoles. Alytes 13: 1–13.
Jungfer, K.H., and P. Weygoldt. 1994. The repro-ductive biology of the leaf frog Phyllomedusalemur Boulenger, 1882, and a comparison withother members of the Phyllomedusinae (Anura:Hylidae). Revue Francaise de Aquariologie 21:57–64.
Jungfer, K.H., and P. Weygoldt. 1999. Biparentalcare in the tadpole-feeding Amazonian treefrogOsteocephalus oophagus. Amphibia-Reptilia20: 235–249.
Jungfer, K.H., S. Ron, R. Seipp, and A. Almen-dariz. 2000. Two new species of hylid frogs,genus Osteocephalus, from Amazonian Ecua-dor. Amphibia-Reptilia 21: 327–340.
Kaiser, H., C. Mais, F. Bolanos, C. Steinlen, W.Feichtinger, and M. Schmid. 1996. Chromo-somal investigation of three Costa Rican frogsfrom the 30-chromosome radiation of Hylawith the description of a unique geographicvariation in nucleolus organizer regions. Ge-netica 98: 95–102.
Kaplan, M. 1991. A new species of Hyla from theeastern slope of the Cordillera Oriental innorthern Colombia. Journal of Herpetology 25:313–316.
Kaplan, M. 1994. A new species of frog of thegenus Hyla from the cordillera oriental innorthern Colombia with comments on the tax-onomy of Hyla minuta. Journal of Herpetology28: 79–87.
Kaplan, M. 1997. On the status of Hyla bogertiCochran and Goin. Journal of Herpetology 31:536–541.
Kaplan, M. 1999. On the phylogenetic relation-ships of Hyla praestans and the monophyly ofthe Hyla columbiana group: morphological ob-servations on the larynx. Revista de la Acade-mia Colombiana de Ciencias Exactas, Fısicas yNaturales 23: 299–302.
Kaplan, M., and P.M. Ruız. 1997. Two new spe-cies of Hyla from the Andes of Central Colom-bia and their relationships to other small An-dean Hyla. Journal of Herpetology 31: 230–244.
Kasahara, S., A.P. Zampieri Silva, S.L. Gruber,and C.F.B. Haddad. 2003. Comparative cyto-genetic analysis on four tree frog species (An-
2005 141FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
ura, Hylidae, Hylinae) from Brazil. Cytogenet-ics and Genome Research 103: 155–162.
Kellogg, R. 1932. Mexican tailless amphibians inthe United States National Museum. Bulletin ofthe United States National Museum Bulletin160: 1–224.
Kenny, J.S. 1969. The Amphibia of Trinidad.Studies on the fauna of Curacao and other Ca-ribbean islands 108: 1–78.
King, M. 1980. A cytotaxonomic analysis of Aus-tralian hylid frogs of the genus Litoria. In C.B.Banks and A.A. Martin (editors), Proceedingsof the Melbourne Herpetological Symposium:169–175. Melbourne, Australia.
King, M., M.J. Tyler, M. Davies, and D. King.1979. Karyotypic studies on Cyclorana and as-sociated genera of Australian frogs. AustralianJournal of Zoology 27: 699–708.
Kizirian, D., L.A. Coloma, and A. Paredes-Re-calde. 2003. A new treefrog (Hylidae: Hyla)from southern Ecuador and a description of itsantipredator behavior. Herpetologica 59: 339–349.
Klappenbach, M.A. 1961. Notas herpetologicas,II. Hallazgo de Trachycephalus siemersi (Mer-tens) y Phyllomedusa iheringi Boulenger (Am-phibia, Salientia) en el Uruguay. Comunica-ciones Zoologicas del Museo de Historia Nat-ural de Montevideo 5: 1–8.
Klappenbach, M.A. 1985. Notas herpetologicas,V. Comunicaciones Zoologicas del Museo deHistoria Natural de Montevideo 11: 1–23.
Klappenbach, M.A., and J.A. Langone. 1992. Lis-ta sistematica y sinonımica de los anfibios delUruguay. Anales del Museo Nacional de His-toria Natural de Montevideo, 2a Serie, 8: 163–222.
Kluge, A.G. 1979. The gladiator frogs of MiddleAmerica and Colombia—a reevaluation of theirsystematics (Anura: Hylidae). Occasional Pa-pers of the Museum of Zoology, University ofMichigan 688: 1–24.
Kluge, A.G. 1989. A concern for evidence and aphylogenetic hypothesis of relationships amongEpicrates (Boidae, Serpentes). Systematic Zo-ology 38: 7–25.
Kocher, T.D., W.K. Thomas, A. Meyer, S.V. Ed-wards, S. Pabo, F.X. Villablanca, and A.C. Wil-son. 1989. Dynamics of mitochondrial DNAevolution in animals: amplification and se-quencing with conserved primers. Proceedingsof the National Academy of Sciences USA 86:6196–6200.
Kolaczkowski, B., and J.W. Thornton. 2004. Per-formance of maximum parsimony and likeli-hood phylogenetics when evolution is hetero-geneous. Nature 431: 980–984.
Kohler, J. 2000. Amphibian diversity in Bolivia:
a study with special reference to montane forestregions. Bonner Zoologische Monographien48: 1–243.
Kohler, J., and S. Lotters. 2001a. A new speciesof minute Hyla from the southwestern Amazonbasin (Amphibia, Anura, Hylidae). Studies onNeotropical Fauna and Environment 36: 105–112.
Kohler, J., and S. Lotters. 2001b. Description ofa small tree frog, genus Hyla (Anura: Hylidae),from humid Andean slopes of Bolivia. Sala-mandra 37: 175–184.
Kolenc, F., C. Borteiro, and M. Tedros. ‘‘2003’’[2004]. La larva de Hyla uruguaya Schmidt,1944 (Anura: Hylidae), con comentarios sobresu biologıa en Uruguay y su status taxonomico.Cuadernos de Herpetologıa 17: 87–100.
Kuramoto, M. 1980. Mating calls of treefrogs (ge-nus Hyla) in the Far East, with description of anew species from Korea. Copeia 1980: 100–108.
Kuramoto, M. 1998. Spermatozoa of several frogspecies from Japan and adjacent regions. Jap-anese Journal of Herpetology 17: 107–116.
Kuramoto, M., and A. Allison. 1991. Karyotypesof five hylid frogs from Papua New Guinea,with a discussion on their systematic implica-tions. Japanese Journal of Herpetology 14: 6–11.
Kwet, A. 2000. The genus Pseudis (Anura: Pseu-didae) in Rio Grande do Sul, southern Brazil,with description of a new species. Amphibia-Reptilia 21: 39–55.
Lacombe, C., C. Cifuentes-Diaz, I. Dunia, M. Au-ber-Thomay, P. Nicolas, and M. Amiche. 2000.Peptide secretion in the cutaneous glands ofSouth American tree frog Phyllomedusa bicol-or: an ultrastructural study. European Journalof Cell Biology 79: 634–641.
Lajmanovich, R.C., M.F. Izaguirre, M.F. Vera-Candioti, and V.H. Casco. 2000. Unique struc-tural pattern of the manicotto glandulare ofHyla nana tadpoles (Anura: Hylidae). Amphib-ia-Reptilia 21: 237–242.
La Marca, E. 1985. Systematic and ecological ob-servations on the Neotropical frogs Hyla jahniand Hyla platydactyla. Journal of Herpetology19: 227–237.
La Marca, E. 1992. Catalogo taxonomico biogeo-grafico y bibliografico de las ranas de Venezue-la. Cuadernos Geograficos 9: 1–197.
La Marca, E. 1997. Lista actualizada de los anfi-bios de Venezuela. In E. La Marca (editor),Vertebrados actuales y fosiles de Venezuela 1:103–119. Merida,Venezuela: Museo de Cienciay Tecnologıa de Merida.
La Marca, E. 1998. Biodiversidad de anfibios enlos Andes de Venezuela: analisis preliminar por
142 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
pisos de vegetacion. In G. Halffter (editor), Ladiversidad biologica de Iberoamerica III. Vol-umen Especial, Acta Zoologica Mexicana, nue-va serie: 199–210. Xalapa, Mexico: Instituto deEcologıa, A.C.
Lang, M. 1990. Annotated catalogue of the type-specimens from the herpetological collection ofthe Royal Belgian Institute of Natural Sciences,Brussels. Documents de Travail de l’InstitutRoyal des Sceinces Naturelles de Belgique 59:1–36.
Langone, J.A. 1990. Revalidacion de Hyla uru-guaya Schmidt, 1944 (Amphibia, Anura, Hyli-dae). Comunicaciones Zoologicas del Museo deHistoria Natural de Montevideo 12: 1–9.
Langone, J.A. ‘‘1994’’ [1995]. Ranas y sapos delUruguay. Montevideo: Museo Damaso AntonioLarranaga, Serie de Divulgacion n8 5, 123 pp.
Langone, J.A., and N.G. Basso. 1987. Distribu-cion geografica y sinonimia de Hyla nana Bou-lenger, 1889 y de Hyla sanborni Schmidt, 1944(Anura, Hylidae) y observaciones sobre formasafines. Comunicaciones Zoologicas del Museode Historia Natural de Montevideo 11: 1–17.
Langone, J.A., C.M. Prigioni, and L. Venturino.1985. Informe preliminar sobre el comporta-miento reproductor y otros aspectos de la biol-ogıa de Phyllomedusa iheringi, Boulenger,1885 (Anura, Hylidae). Comunicaciones Zool-ogicas del Museo de Historia Natural de Mon-tevideo 11: 1–12.
Lannoo, M.J., D.S. Townsend, and R. Wassersug.1987. Larval life in the leaves: arboreal tadpoletypes, with special attention to the morphology,ecology, and behavior of the oophagous Osteo-pilus brunneus (Hylidae) larva. Fieldiana Zo-ology, N.S. 38: 1–31.
Laurent, R.F. 1986. Sous classe de Lissamphi-biens: Lissamphibia systematique. In P.-P.Grasse and M. Delsol (editors), Traite de zool-ogie: anatomie, systematique, biologie, tomeXIV, fascicle 1B—Batraciens: 594–797. Paris:Masson.
Lavilla, E.O. 1984. Redescripcion de larvas deHyla pulchella andina (Anura: Hylidae) con unanalisis de la variabilidad interpoblacional.Neotropica 30: 19–30.
Lavilla, E.O. 1990. The tadpole of Hyla nana(Anura: Hylidae). Journal of Herpetology 24:207–209.
Lavilla, E.O. 1992. Tipos portadores de nombre ylocalidades tipo de anfibios de Argentina. ActaZoologica Lilloana 42: 61–100.
Lee, J.C. 1996. The amphibians and reptiles of theYucatan peninsula. Ithaca, NY: Cornell Univer-sity Press.
Lehr, E., and R.V. May. 2004. Rediscovery of
Hyla melanopleura Boulenger, 1912 (Amphib-ia: Anura: Hylidae). Salamandra 40: 51–58.
Lescure, J., and C. Marty. 2000. Atlas des am-phibiens de Guyane. Paris: Publications Scien-tifiques du Museum National d’Histoire Natu-relle.
Lescure, J., C. Marty, V. Marty, F. Starace, M.Auber-Thomay, and F. Letellier. 1995. Contri-bution a l’etude des amphibiens de Guyanafrancaise. X. Les Phyllomedusa (Anura, Hyli-dae). Revue Francaise de Aquariologie 22: 35–50.
Lewis, P.O. 2001. A likelihood approach to esti-mating phylogeny from discrete morphologicalcharacter data. Systematic Biology 50: 913–925.
Lichtenstein, M.H. C., and E. von Martens 1856.Nomenclator Reptilium et Amphibiorum MuseiZoologici Berolinensis. Namensverzeichnissder in der zoologischen Sammlung de Konig-lichen Universitat zu Berlin aufgestellten Artenvon Reptilien und Amphibien nach ihren Ord-nungen, Familien und Gattungen, Berlin, 48 pp.
Lips, K. 1996. New treefrog from the Cordillerade Talamanca of Central America with a dis-cussion of systematic relationships in the Hylalancasteri group. Copeia 1996: 615–626.
Luddecke, H., and O.R. Sanchez. 2002. Are trop-ical highland frog calls cold-adapted? The caseof the Andean frog Hyla labialis. Biotropica34: 281–288.
Lutz, A., and B. Lutz. 1939. I. Notes on the genusPhyllomedusa Wagler. A) Observations onsmall Phyllomedusae without vomerine teeth orconspicuous parotids found in the region of Riode Janeiro. B) Phyllomedusa bahiana Lutz. An-naes da Academia Brasileira de Sciencias 11:219–263.
Lutz, B. 1948. Anuros da Colecao Adolpho Lutzda regiao sud-este do Brasil. I. Hyla ancepsLutz, 1929. Memorias do Instituto OswaldoCruz 46: 299–313.
Lutz, B. ‘‘1948’’ [1949]. Anfibios anuros da co-lecao Adolpho Lutz II. Especies verdes do ge-nero Hyla do leste-meridional do Brasil. Me-morias do Instituto Oswaldo Cruz 46: 551–577.
Lutz, B. 1950. Anfıbios anuros da colecao Adol-pho Lutz. V. Locomocao e estrutura das ex-tremidades. V.a Phyllomedusa (P.) burmeisteridistincta A. Lutz. V.b Aplastodiscus perviridisA. Lutz. Memorias do Instituto Oswaldo Cruz48: 599–637.
Lutz, B. 1954. Anfıbios anuros do Distrito Fed-eral. Memorias do Instituto Oswaldo Cruz 52:155–238.
Lutz, B. 1963. New species of Hyla from south-eastern Brazil. Copeia 1963: 561–562.
Lutz, B. 1968. Taxonomy of Neotropical Hylidae.
2005 143FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
The Pearce-Sellards Series, Texas MemorialMuseum 11: 1–25.
Lutz, B. 1973. Brazilian Species of Hyla. Austin:University of Texas Press.
Lynch, J.D. 1971. Evolutionary relationships, os-teology, and zoogeography of leptodactyloidfrogs. Miscellaneous Publications of the Mu-seum of Natural History, University of Kansas53: 1–238.
Lynch, J.D. 1973. The transition from archaic toadvanced frogs. In J.L. Vial (editor), Evolution-ary biology of the anurans. Contemporary re-search on major problems: 131–182. Columbia:University of Missouri Press.
Lynch, J.D. 1986. Origins of the high Andean her-petological fauna. In F. Vuilleumier and M.Monasterio (editors), High altitude tropical bio-geography: 478–499. New York: Oxford Uni-versity Press.
Lynch, J.D. 1998. New species of Eleutherodac-tylus from the Cordillera Occidental of westernColombia with a synopsis of the distributionsof species in western Colombia. Revista de laAcademia Colombiana de Ciencias Exactas,Fısicas y Naturales 22: 117–148.
Lynch, J.D. 2002. A new species of the genusOsteocephalus (Hylidae: Anura) from the west-ern Amazon. Revista de la Academia Colom-biana de Ciencias Exactas, Fısicas y Naturales26: 289–292.
Lynch, J.D., and W.E. Duellman. 1997. Frogs ofthe genus Eleutherodactylus in western Ecua-dor. University of Kansas Natural History Mu-seum, Special Publication 23: 1–236.
Lynch, J.D., and A.M. Suarez-Mayorga. 2001.The distributions of the gladiator frogs (Hylaboans group) in Colombia, with comments onsize variation and sympatry. Caldasia 23: 491–507.
Lynch, J.D., P.M. Ruiz-Carranza, and C.M. Ar-dila-Robayo. 1997. Biogeographic patterns ofColombian frogs and toads. Revista de la Ac-ademia Colombiana de Ciencias Exactas, Fisi-cas y Naturales 21: 237–248.
MacCulloch, R.D., and A. Lathrop. 2002. Excep-tional diversity of Stefania (Anura: Hylidae) onMount Ayangana, Guyana: three new speciesand new distribution records. Herpetologica 58:327–346.
Manzano, A.S. 1997. Estudio comparativo de lamiologıa de la cintura pectoral de algunos Phyl-lomedusinos. Bollettino del Museo di ScienzeNaturali, Torino 15: 255–277.
Manzano, A.S., and E.O. Lavilla. 1995a. Myolog-ical peculiarities in Rhinoderma darwinii (An-ura: Rhinodermatidae). Journal of Morphology224: 125–129.
Manzano, A.S., and E.O. Lavilla. 1995b. Notas
sobre la miologıa apendicular de Phyllomedusahypocondrialis (Anura, Hylidae). Alytes 12:169–174.
Marquez, R., I. De la Riva, and J. Bosch. 1993.Advertisement calls of Bolivian species of Hyla(Amphibia, Anura, Hylidae). Biotropica 25:426–443.
Martins, M., and A.J. Cardoso. 1987. Novas es-pecies de hilıdeos do Estado do Acre (Amphib-ia: Anura). Revista Brasileira de Biologia 47:549–558.
Martins, M., and C.F.B. Haddad. 1988. Vocaliza-tions and reproductive behavior in the smithfrog, Hyla faber Wied (Amphibia: Hylidae).Amphibia-Reptilia 9: 49–60.
Martins, M., and G. Moreira. 1991. The nest andthe tadpole of Hyla wavrini, Parker (Amphibia,Anura). Memorias do Instituto Butantan 53:197–204.
Maxson, L.R. 1992. Tempo and pattern in anuranspeciation and phylogeny: an albumin perspec-tive. In K. Adler (editor), Herpetology: currentresearch on the biology of amphibians and rep-tiles: 41–57. Oxford, OH: Society for the Studyof Amphibians and Reptiles.
Maxson, L.R., D.P. Ondrula, and M.J. Tyler. 1985.An immunological perspective on evolutionaryrelationships in Australian frogs of the hylidgenus Cyclorana. Australian Journal of Zoolo-gy 33: 17–22.
Maxson, L.R., M.J. Tyler, and R.D. Maxson.1982. Phylogenetic relationships of Cycloranaand the Litoria aurea species-group (Anura:Hylidae): a molecular perspective. AustralianJournal of Zoology 30: 643–651.
McCranie, J.R., and L.D. Wilson. 1993. Taxo-nomic changes associated with the names Hylaspinipollex Schmidt and Ptychohyla meraziWilson and McCranie (Anura, Hylidae). TheSouthwestern Naturalist 38: 100–104.
McCranie, J.R., and L.D. Wilson. 2002. The am-phibians of Honduras. Ithaca, NY: Society forthe Study of Amphibians and Reptiles.
McDiarmid, R.W., and R. Altig. 1990. Descriptionof a bufonid tadpole and two hylid tadpolesfrom western Ecuador. Alytes 8: 51–60.
McDonald, K.R. and D.L. Storch. 1993. A newreproductive mode for an Australian hylid frog.Memoirs of the Queensland Museum 34: 200.
Mendelson, J.R., III, and J.A. Campbell. 1999.The taxonomic status of populations referred toHyla chaneque in southern Mexico, with thedescription of a new treefrog from Oaxaca.Journal of Herpetology 33: 80–86.
Mendelson, J.R., III, and K.R. Toal. 1995. A newspecies of Hyla (Anura: Hylidae) from cloudforest in Oaxaca, Mexico, with comments onthe status of the Hyla bistincta group. Occa-
144 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
sional Papers of the Museum of Natural His-tory, The University of Kansas 174: 1–20.
Mendelson J.R., III, and K.R. Toal III. 1996. Anew species of Hyla (Anura: Hylidae) from thesierra madre del sur of Oaxaca, Mexico, withcomments on Hyla chryses and Hyla mykter.Journal of Herpetology 30: 326–333.
Mendelson, J.R., III, H.R. da Silva, and A.M.Maglia. 2000. Phylogenetic relationshipsamong marsupial frog genera (Anura: Hylidae:Hemiphractinae) based on evidence from mor-phology and natural history. Zoological Journalof the Linnean Society 128: 125–148.
Menin, M., R.A. Silva, and A.A. Giaretta. 2004.Reproductive biology of Hyla goiana (Anura,Hylidae). Iheringia, Zoologia 94: 49–52.
Metcalf, M.M. 1923a. The origin and distributionof the Anura. American Naturalist 57: 385–411.
Metcalf, M.M. 1923b. The opalinid ciliate infu-sorians. Bulletin of the United States NationalMuseum 120: 1–484.
Metcalf, M.M. 1928. The Bell-toads and theiropalinid parasites. American Naturalist 62: 5–21.
Meyer, E., B.G.M. Jamieson, and D.M. Scheltin-ga. 1997. Sperm ultrastructure of six Australianhylid frogs from two genera (Litoria and Cy-clorana): phylogenetic implications. Journal ofSubmicroscopic Cytology and Pathology 29:443–451.
Mijares-Urrutia, A. 1992a. Sobre el renacuajo deHyla alemani Rivero (Anura: Hylidae). ActaBiologica Venezuelica 13: 35–39.
Mijares-Urrutia, A. 1992b. El renacuajo de Hylalascinia, con aportes al conocimiento de losrenacuajos de Hyla jahni e Hyla platydactyla(Hylidae) de los andes venezolanos. Alytes 10:91–98.
Mijares-Urrutia, A. 1997. Renacuajos de tres es-pecies de ranas (Amphibia: Bufonidae, Hyli-dae) de Colombia y Venezuela. Acta CientıficaVenezolana 48: 139–144.
Mijares-Urrutia, A. 1998. Una nueva especie derana arborıcola (Amphibia: Hylidae) de unbosque nublado del oeste de Venezuela. RevistaBrasileira de Biologia 58: 659–663.
Mijares-Urrutia, A., and R.A. Rivero. 2000. Anew treefrog from the Sierra de Aroa, northernVenezuela. Journal of Herpetology 34: 80–84.
Mijares-Urrutia, A., J. Manzanilla-Puppo, and E.La Marca. 1999. Una nueva especie de Tepui-hyla (Anura: Hylidae) del noroeste de Vene-zuela, con comentarios sobre su biogeografıa.Revista de Biologia Tropical 47: 1099–1110.
Miranda-Ribeiro, A. 1926. Notas para servirem aoestudo do gymnobatrachios (Anura) brasileiros.
Archivos do Museu Nacional do Rio de Janeiro27: 1–227.
Miranda-Ribeiro, A. 1937. Alguns batrachios no-vos das colecoes do Museu Nacional. O Campo1937(Maio): 66–69.
Misuraca, G., G. Prota, J.T. Bagnara, and S.K.Frost. 1977. Identification of the leaf–frog me-lanophore pigment, rhodomelanochrome, aspterorhodin. Comparative Biochemistry andPhysiology 57B: 41–43.
Morand, M., and A. Hernando. 1996. Cariotipo yregion organizadora del nucleolo en Argenteo-hyla siemersi pederseni (Anura: Hylidae). FA-CENA 12: 141–144.
Morescalchi, A. 1973. Amphibia. In A.B. Chi-arelli and E. Capanna (editors), Cytotaxonomyand vertebrate evolution: 233–348. London:Academic Press.
Moriarty, E.C., and D.C. Cannatella. 2004. Phy-logenetic relationships of the north americanchorus frogs (Pseudacris: Hylidae). MolecularPhylogenetics and Evolution 30: 409–420.
Moritz, C., C.J. Schneider, and D.B. Wake. 1992.Evolutionary relationships within the Ensatinaeschscholtzii complex confirm the ring speciesinterpretation. Systematic Biology 41: 273–291.
Muller, L., and W. Hellmich. 1936. Amphibienund Reptilien I. Teil: Amphibia, Chelonia, Lor-icata. In H. Krieg (editor), WissenschaftlicheErgebnisse der Deutschen Gran Chaco-Expe-dition: 120 pp. Stuttgart: Stecker und Schroder.
Myers, C.W., and M.A. Donnelly. 1997. A tepuiherpetofauna on a granitic mountain (Tamacu-ari) in the borderland between Venezuela andBrazil: report from the Phipps Tapirapeco Ex-pedition. American Museum Novitates 3213:1–71.
Myers, C.W., and R. Stothers. MS. The myth ofHylas revisited: the frog name Hyla, and othercommentary on the Specimen Medicum of J. N.Laurenti, the ‘‘father of herpetology’’.
Myers, C.W., A.O. Paolillo, and J.W. Daly. 1991.Discovery of a defensively malodorous andnocturnal frog in the family Dendrobatidae:phylogenetic significance of a new genus andspecies from the Venezuelan Andes. AmericanMuseum Novitates 3002: 1–33.
Napoli, M.F. 2000. Taxonomia, variacao morfo-logica e distribuicao geografica das especies dogrupo de Hyla circumdata (Cope, 1870) (Am-phibia, Anura, Hylidae). Ph.D. dissertation,Universidade Federal do Rio de Janeiro.
Napoli, M.F., and U. Caramaschi. 1998. Duas no-vas especies de Hyla Laurenti, 1768 do Brasilcentral afins de H. tritaeniata Bokermann, 1965(Amphibia, Anura, Hylidae). Boletim do Mu-seu Nacional, Nova Serie, Zoologia 391: 1–12.
2005 145FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Napoli, M.F., and U. Caramaschi. 1999. The tax-onomic status of Hyla elongata Lutz, 1925.Journal of Herpetology 33: 484–487.
Napoli, M.F., and U. Caramaschi. 2004. Two newspecies of the Hyla circumdata group from Se-rra do Mar and Serra da Mantiqueira, south-eastern Brazil, with description of the adver-tisement call of Hyla ibitipoca (Anura, Hyli-dae). Copeia 2004: 534–545.
Napoli, M.F., and B.V.S. Pimenta. 2003. Nova es-pecie do grupo de Hyla circumdata (Cope,1870) do sul da Bahia, Brasil (Amphibia, An-ura, Hylidae). Arquivos do Museu Nacional 61:189–194.
Nicholls, G.C. 1916. The structure of the vertebralcolumn in the Anura Phaneroglossa and its im-portance as a basis of classification. Proceed-ings of the Linnean Society of London 128:80–92.
Nieden, F. 1923. Anura I. Subordo Aglossa undPhanerglossa sectio 1 Arcifera. Das Tierreich46: 1–584.
Nixon, K.C. 1999a. The parsimony ratchet, a newmethod for rapid parsimony analysis. Cladistics15: 407–414.
Nixon, K.C. 1999b. WinClada. Computer soft-ware and documentation, available at: http://www.Cladistics.com.
Nixon, K.C., and J.M. Carpenter. 1996. On si-multaneous analysis. Cladistics 12: 221–241.
Noble, G.K. 1922. The phylogeny of the Salientia.I. The osteology and the thigh musculature;their bearing on classification and phylogeny.Bulletin of the American Museum of NaturalHistory 46: 1–87.
Noble, G.K. 1927. The value of life history datain the study of evolution of the Amphibia. An-nals of the New York Academy of Sciences 30:31–128.
Noble, G.K. 1931. The biology of the Amphibia.New York: McGraw-Hill.
Page, R.D.M., and C. Lydeard. 1994. Towards acladistic biogeography of the Caribbean. Cla-distics 10: 21–41.
Page, T.J. 1859. La Plata, the Argentine Confed-eration, and Paraguay. Being a narrative of theexploration of the tributaries of the river LaPlata and adjacent countries during the years1853, ’54, ’55, and ’56, under the orders of theUnited States Government. New York: Harperand Brothers.
Palumbi, S.R., A. Martin, W.O. McMillan, L.Stice, and G. Grabowski. 1991. The simplefool’s guide to PCR, version 2.0. Privately pub-lished.
Paolillo, A.O., and J. Cerda. 1981. Nuevos hal-lazgos de Aparasphenodon venezolanus (Mer-tens) (Salientia, Hylidae) en el Territorio Fe-
deral Amazonas, Venezuela, con anotacionessobre su biologıa. Memoria de la Sociedad deCiencias Naturales La Salle 41: 77–95.
Parker, H.W. 1940. The Australian frogs of thefamily Leptodactylidae. Novitates Zoologicae42: 1–106.
Parmalee, J.R. 1999. Trophic ecology of a tropicalanuran assemblage. Scientific Papers of theNatural History Museum, The University ofKansas 11: 1–59.
Peixoto, O.L. 1981. Nova especie de Hyla da Se-rra dos Orgaos, Estado de Rio de Janeiro, Brasil(Amphibia, Anura, Hylidae). Revista Brasileirade Biologia 41: 515–520.
Peixoto, O.L. 1987. Caracterizacao do grupo‘‘perpusilla’’ e revalidacao da posicao taxon-omica de Ololygon perpusilla perpusilla eOlolygon perpusilla v-signata (Amphibia, An-ura, Hylidae). Arquivos de Universidade Fed-eral Rural do Rio de Janeiro 10: 37–49.
Peixoto, O.L., and C.A.G. Cruz. 1983. Girinos deespecies de Hyla do grupo ‘‘albomarginata’’ doSudeste brasileiro (Amphibia, Anura, Hylidae).Arquivos de Universidade Federal Rural do Riode Janeiro 6: 155–163.
Peixoto, O.L., and C.A.G. Cruz. 1988. Descricaode duas especies novas do genero PhyllodytesWagler (Amphibia, Anura, Hylidae). RevistaBrasileira de Biologia 48: 265–272.
Peixoto, O.L., and M.R. Gomes. 1999. The tad-pole of Hyla nahdereri Lutz and Bokermann,1963. Journal of Herpetology 33: 477–479.
Peixoto, O.L., U. Caramaschi, and E.M.X. Freire.2003. Two new species of Phyllodytes (Anura:Hylidae) from the state of Alagoas, northeast-ern Brazil. Herpetologica 59: 235–246.
Peters, W.C.H. ‘‘1872’’ [1873]. Uber eine, zweineue Gattungen enthaltende, Sammlung vonBatrachiern des Hrn. Dr. O. Wucherer aus Ba-hia, so wie uber einige neue oder weniger be-kannte Saurier. Monatsberichte der koniglichAkademie der Wissenschaften zu Berlin 1872:768–776.
Peters, W.C.H. 1873. Uber eine neue Schildkro-tenart, Cinosternon effeldtii und einige andereneue oder weniger bekannte Amphibien. Mon-atsberichte der koniglich Akademie der Wis-senschaften zu Berlin 1873: 603–618.
Pol, D., and M.E. Siddall. 2001. Biases in maxi-mum likelihood and parsimony: a simulationapproach to a 10-taxon case. Cladistics 17:266–281.
Pombal, J.P., Jr. 1993. A new species of Apara-sphenodon (Anura: Hylidae) from southeasternBrazil. Copeia 1993: 1088–1091.
Pombal, J.P., Jr., and R.P. Bastos. 1998. Nova es-pecie de Hyla Laurenti, 1768 do centro-oestebrasileiro e a posicao taxonomica de H. micro-
146 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
cephala werneri Cochran, 1952 e H. microce-phala meridiana B. Lutz, 1952 (Anura, Hyli-dae). Boletim do Museu Nacional, Nova Serie,Zoologia 390: 1–14.
Pombal, J.P., Jr., and U. Caramaschi. 1995. Posi-cao taxonomica de Hyla pseudopseudis Miran-da-Ribeiro, 1937 e Hyla saxicola Bokermann,1964 (Anura, Hylidae). Boletim do Museu Na-cional, Nova Serie, Zoologia 363: 1–8.
Pombal, J.P., Jr., and M. Gordo. 2004. Anfıbiosanuros da Jureia. In O.A.V. Marques and W.Duleba (editors), Estacao ecologica Jureia-Ita-tins: ambiente fısico, flora e fauna: 243–256.Ribeirao Preto, Sao Paulo, Brasil: Holos Edi-tora.
Pombal, J.P., Jr., and C.F.B. Haddad. 1992. Espe-cies de Phyllomedusa do grupo burmeisteri doBrasil oriental, com descricao de uma especienova (Amphibia: Hylidae). Revista Brasileirade Biologia 52: 217–229.
Pombal, J.P., Jr., and C.F.B. Haddad. 1993. Hylaluctuosa, a new treefrog from southeastern Bra-zil (Amphibia: Hylidae). Herpetologica 49: 16–21.
Pombal, J.P., Jr., C.F.B. Haddad, and C.A.G. Cruz.2003. New species of Phrynohyas from Atlan-tic rain forest of southeastern Brazil (Anura,Hylidae). Copeia 2003: 379–383.
Pope, C.H. 1931. Notes on the amphibians fromFukien, Hainan, and other parts of China. Bul-letin of the American Museum of Natural His-tory 61: 397–611.
Porras, L., and L.D. Wilson. 1987. A new speciesof Hyla from the highlands of Honduras and ElSalvador. Copeia 1987: 478–482.
Powell, R., and R.W. Henderson. 2003a. Somehistorical perspectives. In R.W. Henderson andR. Powell (editors), Islands and the sea: essayson herpetological exploration in the West In-dies. Society for the Study of Amphibians andReptiles, Contributions to Herpetology, 20: 9–20. Ithaca, NY.
Powell, R., and R.W. Henderson. 2003b. A secondset of addenda to the checklist of West Indianamphibians and Reptiles. Herpetological Re-view 34: 341–345.
Prado, G., J.H. Borgo, P.A. Abrunhosa, and H.Wogel. 2003. Comportamento reprodutivo, vo-calizacao e redescricao do girino de Phrynoh-yas mesophaea (Hensel, 1867) do Sudeste doBrasil (Amphibia, Anura, Hylidae). Boletim doMuseu Nacional, Nova Serie, Zoologia 510: 1–11.
Prasad, G.V.R., and J.C. Rage. 1995. Amphibiansand squamates from the Maastrichtian of Nas-kal, India. Cretaceous Research 16: 95–107.
Prendini, L. 2001. Species or supraspecific taxa asterminals in cladistic analysis? Groundplans
versus exemplars revisited. Systematic Biology50: 290–300.
Pugliese, A., A.C.R. Alves, and S.P. Carvalho eSilva. 2000. The tadpoles of Hyla oliveirai andHyla decipiens with notes on the Hyla micro-cephala group (Anura, Hylidae). Alytes 18: 73–80.
Pugliese, A., A.C.R. Alves, and J.P. Pombal, Jr.2001. The tadpole of Hyla rubicundula. Journalof Herpetology 35: 686–688.
Pyburn, W.F. 1977. A new hylid frog (Amphibia,Anura, Hylidae) from the Vaupes River of Co-lombia with comments on related species. Jour-nal of Herpetology 11: 405–410.
Pyburn, W.F. 1978. The voice and relationships ofthe treefrog Hyla hobbsi (Anura: Hylidae). Pro-ceedings of the Biological Society of Washing-ton 91: 123–131.
Pyburn, W.F. 1984. A new stream-inhabiting tree-frog (Anura: Hylidae) from southeastern Co-lombia. Herpetologica 40: 366–372.
Pyburn, W.F. 1993. A new species of dimorphictree frog, genus Hyla (Amphibia: Anura: Hy-lidae) from the Vaupes River of Colombia. Pro-ceedings of the Biological Society of Washing-ton 106: 46–50.
Rabello, M.N. 1970. Chromosomal studies in Bra-zilian anurans. Caryologia 23: 45–59.
Ralin, D.B. 1970. Genetic compatibility and aphylogeny of the temperate North Americanhylid fauna. Ph.D. dissertation, University ofTexas, Austin. [Not seen, cited from Hedges,1986].
Ramos, A.D., and J.L. Gasparini. 2004. Anfibiosdo Gioapaba-Acu, Fundao, Estado do EspıritoSanto. Vitoria: Grafica Santo Antonio.
Read, K., J.S. Keogh, I.A.W. Scott, J.D. Roberts,and P. Doughty. 2001. Molecular phylogeny ofthe australian frog genera Crinia, Geocrinia,and allied taxa (Anura, Myobatrachidae). Mo-lecular Phylogenetics and Evolution 21: 294–308.
Reynolds, R.P., and M.S. Foster. 1992. Four newspecies of frogs and one new species of snakefrom the Chapare region of Bolivia, with noteson other species. Herpetological Monographs 6:83–104.
Richards, S.J. 1993. Functional significance ofnest construction by an australian rainforestfrog: a preliminary analysis. Memoirs of theQueensland Museum 34: 89–93.
Richards, S.J., and G.R. Johnston. 1993. A newspecies of Nyctimystes (Anura: Hylidae) fromthe Star mountains, Papua New Guinea. Mem-oirs of the Queensland Museum 33: 73–76.
Rivero, J.A. 1961. Salientia of Venezuela. Bulle-tin of the Museum of Comparative Zoology126: 1–207.
2005 147FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Rivero, J.A. 1964. Salientios (Amphibia) en la co-leccion de la Sociedad de Ciencias Naturales laSalle de Venezuela. Caribbean Journal of Sci-ence 4: 297–305.
Rivero, J.A. 1968. Los Centrolenidos de Vene-zuela (Amphibia: Salientia). Memoria de la So-ciedad de Ciencias Naturales La Salle 28: 301–334.
Rivero, J.A. 1969. A new species of Hyla (Am-phibia, Salientia) from the region of Paramo deTama, Venezuela. Caribbean Journal of Science9: 145–150.
Rivero, J.A. 1970. On the origin, endemism anddistribution of the genus Stefania Rivero (Am-phibia, Salientia) with a description of a newspecies from southeastern Venezuela. Boletinde la Sociedad Venezolana de Ciencias Natur-ales 28: 456–481.
Rivero, J.A. ‘‘1971’’ [1972]. Notas sobre los an-fibios de Venezuela. I. Sobre los hilidos de laGuayana venezolana. Caribbean Journal of Sci-ences 11: 181–188.
Rivero, J.A. 1985. Nuevos centrolenidos de Co-lombia y Venezuela. Brenesia 23: 335–373.
Rodrıguez, L.O., and W.E. Duellman. 1994.Guide to the frogs of the Iquitos region, Ama-zonian Peru. The University of Kansas NaturalHistory Museum Special Publication 22: i–ii,1–80.
Romero de Perez, G., and P.M. Ruiz-Carranza.1996. Histologia, histoquımica y estructura finade la glandula mentoniana de dos especies deHyla (grupo bogotensis) y del antebrazo dePhrynopus adenobatrachius. Revista de la Ac-ademia Colombiana de Ciencias Exactas, Fısi-cas y Naturales 20: 575–584.
Ron, S., and J.B. Pramuk. 1999. A new speciesof Osteocephalus (Anura, Hylidae) from Am-azonian Ecuador and Peru. Herpetologica 55:433–446.
Ruiz-Carranza, P.M., J.I. Hernandez-Camacho,and J.V. Rueda-Almonacid. 1988. Una nuevaespecie de Phyllomedusa Wagler 1830 (Am-phibia: Anura: Hylidae) del noroeste de Colom-bia. Trianea 2: 373–382.
Ruiz-Carranza, P.M., and J.D. Lynch. 1991. RanasCentrolenidae de Colombia I. Lozania 57: 1–32.
Ruvinsky, I., and L.R. Maxson. 1996. Phyloge-netic relationships among bufonoid frogs (An-ura: Neobatrachia) inferred from mitochondrialDNA sequences. Molecular Phylogenetics andEvolution 5: 533–547.
Sachsse, R., E. Izecksohn, and S.P. Carvalho eSilva. 1999. The systematic status of Hyla al-bolineata Lutz and Lutz, 1939 (Amphibia: An-ura: Hylidae). Herpetologica 55: 401–406.
Salducci, M.D., C. Marty, R. Chappaz, and A.
Gilles. 2002. Molecular phylogeny of FrenchGuiana Hylinae: implications for the systematicand biodiversity of the Neotropical frogs.Compte Rendu des Seances de la Societe dePhysique et d’Histoire Naturelle, Biologie 325:141–153.
Sanchiz, B. 1998a. Salientia. In P. Wellnhofer (ed-itor), Encyclopedia of paleoherpetology 4: 1–276. Munchen: Verlag Dr. Friedrich Pfeil.
Sanchiz, B. 1998b. Vertebrates from the earlyMiocene lignite deposits of the opencast mineOberdorf (Western Styrian Basin, Austria): 2.Amphibia. Annalen des Naturhistorischen Mu-seums in Wien 99A: 13–29.
Sanderson, M.J., and J. Kim. 2000. Parametricphylogenetics? Systematic Biology 49: 817–829.
Sankoff, D. 1975. Minimal mutation trees of se-quence. SIAM Journal of Applied Mathematics21: 35–42.
Sankoff, D., and R.J. Cedergren. 1983. Simulta-neous comparisons of three or more sequencesrelated by a tree. In D. Sankoff and J. Kruskal(editors), Time warps, string edits, and macro-molecules. The theory and practice of sequencecomparisons: 253–264. Reading, MA: Addi-son-Wesley.
Sanmartın, I., and F. Ronquist. 2004. Southernhemisphere biogeography inferred by event-based models: plant versus animal patterns.Systematic Biology 53: 216–243.
Santos, C.S., A.C.R. Alves, and S.P. Carvalho eSilva. 1998. Description of the tadpoles of Hylagiesleri and Hyla microps from southern Brazil.Journal of Herpetology 32: 61–66.
Santos, J.C., L.A. Coloma, and D.C. Cannatella.2003. Multiple, recurring origins of aposema-tism and diet specialization in poison frogs.Proceedings of the National Academy of Sci-ences USA 100: 12792–12797.
Savage, J.M. 1966. The origins and history of theCentral American herpetofauna. Copeia 1966:719–766.
Savage, J.M. 1973. The geographic distribution offrogs: patterns and predictions. In J.L. Vial (ed-itor), Evolutionary biology of the Anurans:contemporary research on major problems:351–445. Columbia: University of MissouriPress.
Savage, J.M. 1982. The enigma of Central Amer-ican herpetofauna: dispersals or vicariance?Annals of the Missouri Botanical Garden 69:464–547.
Savage, J.M. 2002a. The amphibians and reptilesof Costa Rica. A herpetofauna between twocontinents, between two seas. Chicago: Univer-sity of Chicago Press.
148 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Savage, J.M. 2002b. The hylid frogs of MiddleAmerica [book review]. Copeia 2002: 545–552.
Savage, J.M., and A.L.D. Carvalho. 1953. Thefamily position of neotropical frogs currentlyreferred to the genus Pseudis. Zoologica, Sci-entific Contributions of the New York Zoolog-ical Society 38: 193–200.
Savage, J.M., and W.R. Heyer. ‘‘1968’’ [1969].The tree-frogs (family Hylidae) of Costa Rica:diagnosis and distribution. Revista de BiologiaTropical 16: 1–127.
Sazima, I., and W.C.A. Bokermann. 1977. Anfı-bios da Serra do Cipo, Minas Gerais, Brasil. 3:Observacoes sobre a biologia de Hyla alvar-engai Bok. (Anura, Hylidae). Revista Brasileirade Biologia 37: 413–417.
Schiesari, L.C., B. Grillitsch, and C. Vogl. 1996.Comparative morphology of phytotelmonousand pond-dwelling larvae of four neotropicaltreefrog species (Anura, Hylidae, Osteocephal-us oophagus, Osteocephalus taurinus, Phry-nohyas resinifictrix, Phrynohyas venulosa).Alytes 13: 109–139.
Schmidt, K.P. 1953. A checklist of North Ameri-can amphibians and reptiles, 6th ed. Chicago:American Society of Ichthyologists and Her-petologists.
Senaris, J.C., and J. Ayarzaguena. 2002. A newHyla (Anura: Hylidae) from the highlands ofVenezuelan Guyana. Journal of Herpetology36: 634–640.
Senaris, J.C., J. Ayarzaguena, and S. Gorzula.‘‘1996’’ [1997]. Revision taxonomica del ge-nero Stefania (Anura; Hylidae) en Venezuelacon la descripcion de cinco nuevas especies.Publicaciones de la Asociacion de Amigos deDonana 7: 1–57.
Sheil, C.A., J.R. Mendelson, and H.R. da Silva.2001. Phylogenetic relationships of the speciesof Neotropical horned frogs, genus Hemiphrac-tus (Anura: Hylidae: Hemiphractinae), based onevidence from morphology. Herpetologica 57:203–214.
Siddall, M.E. 1998. Success of parsimony in thefour-taxon case: long-branch repulsion by like-lihood in the Farris zone. Cladistics 14: 209–220.
Siddall, M.E., and A.G. Kluge. 1999. Letter to theeditor. Cladistics 15: 439–440.
Simmons, M.P., K.M. Pickett, and M. Miya. 2004.How meaningful are Bayesian support values?Molecular Biology and Evolution 21: 188–199.
Skuk, G., and J.A. Langone. 1991. Los cromo-somas de cuatro especies del genero Hyla (An-ura, Hylidae) con un numero diploide de 2n 530. Acta Zoologica Lilloana 41: 165–171.
Smith, E.N., and B.P. Noonan. 2001. A new spe-cies of Osteocephalus (Anura: Hylidae) from
Guyana. Revista de Biologia Tropical 49: 347–357.
Smith, H.M., and E.H. Taylor. 1948. An annotatedchecklist and key to the Amphibia of Mexico.Bulletin of the United States National Museum194: 1–118.
Smith, T.F., M.S. Waterman, and W.M. Fitch.1981. Comparative biosequence metrics. Jour-nal of Molecular Evolution 18: 38–46.
Snyder, D.H. 1972. Hyla juanitae, a new treefrogfrom southern Mexico, and its relationship toH. pinorum. Journal of Herpetology 6: 5–15.
Spirandeli Cruz, E.F. 1991. Estudo comparativoda morfologia oral interna de larvas de anfıbiosanuros que ocorrem na regiao de Botucatu, SaoPaulo (Amphibia, Anura). Instituto de Biocien-cias. Ph.D. dissertation, Universidade de SaoPaulo, Sao Paulo
Starrett, P.H. 1968. The phylogenetic significanceof the jaw musculature in anuran amphibians.Ph.D. dissertation University of Michigan, AnnArbor.
Steel, M. 2002. Some statistical aspects of themaximum parsimony method. In R. De Salle,G. Giribet, and W. Wheeler (editors), Molecularsystematics and evolution: theory and practice:125–139. Basel, Switzerland: Birkhauser Ver-lag.
Steel, M., and D. Penny. 2000. Parsimony, like-lihood, and the role of models in molecularphylogenetics. Molecular Biology and Evolu-tion 17: 839–850.
Steel, M.A., M.D. Hendy, and D. Penny. 1993.Parsimony can be consistent! Systematic Biol-ogy 42: 581–587.
Steel, M., L. Szekely, and M. Hendy. 1994. Re-constructing trees from sequences whose sitesevolve at variable rates. Journal of ComparativeBiology 1: 153–163.
Stejneger, L. 1907. Herpetology of Japan and ad-jacent territory. Bulletin of the United StatesNational Museum 58: 1–577.
Stewart, M.M. 2003. Recollections of Jamaica. InR.W. Henderson and R. Powell (editors), Is-lands and the sea: essays on herpetological ex-ploration in the West Indies: 121–132. Contri-butions to Herpetology, no. 20. Ithaca, NY: So-ciety for the Study of Amphibians and Reptiles.
Strong, E.E., and D.L. Lipscomb. 1999. Charactercoding and inapplicable data. Cladistics 15:363–371.
Suarez-Mayorga, A.M., and J.D. Lynch. 2001a.Redescription of the tadpole of Hyla vigilans(Anura: Hylidae) and notes about possible tax-onomic relationships. Caribbean Journal of Sci-ences 37: 116–119.
Suarez-Mayorga, A.M., and J.D. Lynch. 2001b.Los renacuajos colombianos de Sphaenorhyn-
2005 149FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
chus (Hylidae): descripciones, anotaciones sis-tematicas y ecologicas. Revista de la AcademiaColombiana de Ciencias Exactas, Fısicas y Na-turales 25: 411–420.
Taboga, S.R., and H. Dolder. 1993. Ultrastructuralanalysis of the Hyla ranki spermatozoan tail(Amphibia, Anura, Hylidae). Cytobios 75: 85–95.
Taylor, E.H. 1948. Two new hylid frogs from Cos-ta Rica. Copeia 1948: 233–238.
Taylor, E.H. 1952. The frogs and toads of CostaRica. University of Kansas Science Bulletin 35:577–942.
Taylor, E.H. 1954. Frog-egg eating tadpoles ofAnotheca coronata (Stejneger) (Salientia, Hy-lidae). University of Kansas Science Bulletin36: 589–595.
Teixeira, R.L., J.A.P. Schneider, and G.I. Almeida.2002. The occurrence of amphibians in bro-meliads from a southeastern Brazilian restingahabitat, with special reference to Aparasphen-odon brunoi (Anura, Hylidae). Brazilian Jour-nal of Biology 62: 263–268.
Thomas, E.O., L. Tsang, and P. Licht. 1993. Com-parative histochemistry of the sexually dimor-phic skin glands of anuran amphibians. Copeia1993: 133–143.
Titus, T.A., and A. Larson. 1996. Molecular phy-logenetics of desmognathine salamanders (Cau-data: Plethodontidae): a reevaluation of evolu-tion in ecology, life history, and morphology.Systematic Biology 45: 229–238.
Toal, K.R., III. 1994. A new species of Hyla (An-ura: Hylidae) from the Sierra de Juarez, Oaxa-ca, Mexico. Herpetologica 50: 187–193.
Toledo, R.C., and C. Jared. 1995. Cutaneous gran-ular glands and amphibian venoms. Compara-tive Biochemistry and Physiology 111A: 1–29.
Trewavas, E. 1933. The hyoid and larynx of theAnura. Philosophical Transactions of the RoyalSociety of London 222: 401–527.
Trueb, L. 1969. Pternohyla, P. dentata, P. fodiens.Catalogue of American Amphibians and Rep-tiles 77: 1–4.
Trueb, L. 1970a. Evolutionary relationships ofcasque-headed tree frogs with co-ossified skulls(Family Hylidae). University of Kansas Publi-cations, Museum of Natural History 18: 547–716.
Trueb, L. 1970b. The generic status of Hyla sie-mersi Mertens. Herpetologica 26: 254–267.
Trueb, L. 1974. Systematic relationships of Neo-tropical horned frogs, genus Hemiphractus(Anura: Hylidae). Occasional Papers of theMuseum of Natural History, University of Kan-sas 29: 1–60.
Trueb, L., and W.E. Duellman. 1971. A synopsisof Neotropical hylid frogs, genus Osteocephal-
us. Occasional Papers of the Museum of Nat-ural History, The University of Kansas 1: 1–47.
Trueb, L., and M.J. Tyler. 1974. Systematics andevolution of the greater Antillean hylid frogs.Occasional Papers of the Museum of NaturalHistory, University of Kansas 24: 1–60.
Tschudi, J.J. von. 1838 (1967). Classification derBatrachier, mit berucksichtigung der fossilenthiere dieser abtheilung der reptilien. Athens,OH: Society for the Study of Amphibians andReptiles.
Tuffley, C., and M. Steel. 1997. Links betweenmaximum likelihood and maximum parsimonyunder a simple model of site substitution. Bul-letin of Mathematical Biology 59: 581–607.
Tyler, M.J. 1963. A taxonomic study of amphib-ians and reptiles of the Central Highlands ofNew Guinea, with notes on their ecology andbiology 2. Anura: Ranidae and Hylidae. Trans-actions of the Royal Society of South Australia86: 105–130.
Tyler, M.J. 1968. Papuan hylid frogs of the genusHyla. Zoologische Verhandelingen: 1–203.
Tyler, M.J. 1971. The phylogenetic significance ofvocal sac structure in hylid frogs. OccasionalPapers of the Museum of Natural History, TheUniversity of Kansas 19: 319–360.
Tyler, M.J. 1972. Superficial mandibular muscu-lature, vocal sacs and the phylogeny of Austra-lo-Papuan leptodactylid frogs. Records of theSouth Australian Museum 16: 1–20.
Tyler, M.J. 1978. Amphibians of South Australia.Government Printer, South Australia.
Tyler, M.J. 1979. Herpetofaunal relationships ofSouth America with Australia. In W.E. Duell-man (editor), The South American herpetofau-na: its origins, evolution, and dispersal: 73–106. University of Kansas, Museum of NaturalHistory Monographs, 7.
Tyler, M.J. 1982. The hylid frog genus LitoriaTschudi: an overview. In D.G. Newman (edi-tor), New Zealand herpetology. Proceedings ofa symposium held at Victoria University ofWellington 29–31 January 1980: 103–112.Wellington, N.Z.: New Zealand Wildlife Ser-vice, Occasional Publication No. 2.
Tyler, M.J. 1991. A large new species of Litoria(Anura: Hylidae) from the Tertiary of Queens-land. Transactions of the Royal Society ofSouth Australia 115: 103–105.
Tyler, M.J., and M. Davies. 1978a. Phylogeneticrelationships of Australian hyline and Neotrop-ical phyllomedusine frogs of the family Hyli-dae. Herpetologica 34: 219–224.
Tyler, M.J., and M. Davies. 1978b. Species-groups within the Australopapuan hylid frog
150 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
genus Litoria Tschudi. Australian Journal ofZoology Supplement 63: 1–47.
Tyler, M.J., and M. Davies. 1979. Redefinitionand evolutionary origin of the Australopapuanhylid frog genus Nyctimystes Stejneger. Austra-lian Journal of Zoology 27: 755–772.
Tyler, M.J., and M. Davies. 1993. Family Hylidae.In C.J. Glasby, G.J.B. Ross, and P.L. Beesley(editors), Fauna of Australia, vol. 2A. Amphib-ia and Reptilia: 58–63. Canberra: AustralianGovernment Publishing Service.
Tyler, M.J., G.F. Watson, and A.A. Martin. 1981.The Amphibia: diversity and distribution. In A.Keast (editor), Ecological biogeography ofAustralia: 1277–1301. Hague: Dr. W. Junk.
Ustach, P.C., J.R. Mendelson III, R.W. Mc-Diarmid, and J.A. Campbell. 2000. A new spe-cies of Hyla (Anura: Hylidae) from the SierraMixes, Oaxaca, Mexico, with comments on on-togenetic variation in tadpoles. Herpetologica56: 239–250.
Vellard, J. 1948. Batracios del Chaco Argentino.Acta Zoologica Lilloana 5: 137–174.
Vences, M., and F. Glaw. 2001. Systematic reviewand molecular phylogenetic relationships of thedirect developing Malagasy anurans of theMantidactylus asper group. Alytes 19: 107–139.
Vences, M., J. Kosuch, F. Glaw, W. Bohme, andM. Veith. 2003a. Molecular phylogeny of hy-peroliid treefrogs: biogeographic origin of Mal-agasy and Seychelean taxa and re-analysis offamilial paraphyly. Journal of Zoological Sys-tematics and Evolutionary Research 41: 205–215.
Vences, M., J. Kosuch, R. Boistel, C.F.B. Haddad,E. La Marca, S. Lotters, and M. Veith. 2003b.Convergent evolution of aposematic colorationin Neotropical poison frogs: a molecular phy-logenetic perspective. Organisms, Diversity,and Evolution 3: 215–226.
Vences, M., D.R. Vieites, F. Glaw, H. Brinkmann,J. Kosuch, M. Veith, and A. Meyer. 2003c.Multiple overseas dispersal in amphibians. Pro-ceedings of the Royal Society of London B270: 2435–2442.
Vences, M., J. Kosuch, S. Lotters, A. Widmer,K.H. Jungfer, J. Kohler, and M. Veith. 2000.Phylogeny and classification of poison frogs,(Amphibia: Dendrobatidae), based on mito-chondrial 16S and 12S ribosomal RNA genesequences. Molecular Phylogenetics and Evo-lution 15: 34–40.
Vera Candioti, M.F. 2004. Morphology of pre-metamorphic larvae of Lysapsus limellus (An-ura: Pseudinae). Amphibia-Reptilia 25: 41–54.
Vera Candioti, M.F., E.O. Lavilla, and D.D. Eche-verria. 2004. Feeding mechanism in two tree-
frogs, Hyla nana and Scinax nasicus (Anura:Hylidae). Journal of Morphology 261: 206–224.
Vigle, G.O., and D.C.I. Goberdhan-Vigle. 1990.A new species of small colorful Hyla from thelowland rainforest of Amazonian Ecuador. Her-petologica 46: 467–473.
Wagler, J. 1830. Naturliches System der Amphi-bien: mit vorangehender Classification der Sau-gethiere und Vogel: ein Beitrag zur verglei-chenden Zoologie. Cotta’schen, Munchen.
Wassersug, R. 1980. Internal oral features of lar-vae from eight anuran families: functional, sys-tematic, evolutionary and ecological consider-ations. Miscellaneous Publications of the Mu-seum of Natural History, University of Kansas68: 1–146.
Wassersug, R.J., and W.E. Duellman. 1984. Oralstructures and their development in egg-brood-ing hylid frog embryos and larvae: evolution-ary and ecological implications. Journal ofMorphology 182: 1–37.
Werner, F. 1903. Neue reptilien und batrachier ausdem naturhistorischen museum in Brussel.Nebst bemerkungen uber einige andere arten.Zoologische Anzeiger 26: 246–253.
Weygoldt, P. 1984. Durch nachzucht erhalten: dermakifrosch Phyllomedusa exilis. Aquarien-Ma-gazin 1984: 147–150.
Weygoldt, P. 1991. Zur biologie und zum verhal-ten von Phyllomedusa marginata Izecksohnand Da Cruz, 1976 im terrarium. Salamandra27: 83–96.
Weygoldt, P., and S.P. Carvalho e Silva. 1991. Ob-servations on mating, oviposition, egg sac for-mation and development in the egg-broodingfrog, Fritziana goeldii. Amphibia-Reptilia 12:67–80.
Weygoldt, P., and O.L. Peixoto. 1987. Hyla rus-chii n. sp., a new frog from the Atlantic forestdomain in the State of Espırito Santo, Brazil(Amphibia, Hylidae). Studies on NeotropicalFauna and Environment 22: 237–247.
Wheeler, W.C. 1996. Optimization alignment: theend of multiple sequence alignment in phylo-genetics? Cladistics 12: 1–9.
Wheeler, W.C. 1998. Alignment characters, dy-namic programming and heuristic solutions. InR. DeSalle and B. Schierwater (editors), Mo-lecular approaches to ecology and evolution:243–251. Basel: Birkhauser Verlag.
Wheeler, W.C. 2002. Optimization alignment:down, up, error, and improvements. In R. DeSalle, G. Giribet, and W.C. Wheeler (editors),Techniques in molecular systematics and evo-lution: 55–69. Bassel: Birkhauser Verlag.
Wheeler, W.C. 2003a. Iterative pass optimizationof sequence data. Cladistics 19: 254–260.
2005 151FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Wheeler, W.C. 2003b. Implied alignment: a syn-apomorphy-based multiple-sequence alignmentmethod and its use in cladogram search. Cla-distics 19: 261–268.
Wheeler, W.C., D. Gladstein, and J. De Laet.2002. POY, version 3.0. Program available atftp.amnh.org/pub.molecular/poy.
Wild, E.R. 1992. The tadpoles of Hyla fasciataand H. allenorum, with a key to the tadpoles ofthe Hyla parviceps group (Anura: Hylidae).Herpetologica 48: 439–447.
Wiley, J.E. 1982. Chromosome banding patternsof treefrogs (Hylidae) of the eastern UnitedStates. Herpetologica 38: 507–520.
Wilkinson, J.A., M. Matsui, and T. Terachi. 1996.Geographic variation in a Japanese tree frog(Rhacophorus arboreus) revealed by PCR-aid-ed restriction site analysis of mtDNA. Journalof Herpetology 30: 418–423.
Williams, J.D., and A. Bosso. 1994. Estado sis-tematico y distribucion geografica de Argenteo-hyla siemersi (Mertens, 1937) en la RepublicaArgentina (Anura: Hylidae). Cuadernos de Her-petologia 8: 57–62.
Wilson, L.D., and J.R. McCranie. 1989. A newspecies of Ptychohyla of the euthysanota groupfrom Honduras, with comments on the status ofthe genus Ptychohyla Taylor (Anura: Hylidae).Herpetologica 45: 10–17.
Wilson, L.D., J.R. McCranie, and K.L. Williams.1985. Two new species of fringe-limbed hylidfrogs from nuclear Central America. Herpeto-logica 41: 141–150.
Wilson, L.D., J.R. McCranie, and G.A. Cruz.1994a. A new species of Plectrohyla (Anura:Hylidae) from a premontane rainforest in north-ern Honduras. Proceedings of the BiologicalSociety of Washington 107: 67–78.
Wilson, L.D., J.R. McCranie, and K.L. Williams.1985. Two new species of fringe-limbed hylidfrogs from nuclear Central America. Herpeto-logica 41: 141–150.
Wilson, L.D., J.R. McCranie, and K.L. Williams.1994b. Commentary on the reproductive biol-ogy and systematic relationships of Hyla inso-lita McCranie, Wilson, and Williams. Caribbe-an Journal of Science 30: 214–221.
Wogel, H., P.A. Abrunhosa, and J.P. Pombal, Jr.2000. Girinos de cinco especies do Sudeste doBrasil (Amphibia: Hylidae, Leptodactylidae,Microhylidae). Boletim do Museu Nacional,Nova Serie, Zoologia: 1–16.
Yang, Z. 1997. How often do frog models producebetter phylogenies? Molecular Biology andEvolution 414: 105–108.
Zweifel, R.G. 1983. Two new hylid frogs fromthe Papua New Guinea and a discussion of theNyctimystes papua species group. AmericanMuseum Novitates 2759: 1–21.
152 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
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Hyl
aal
bovi
ttat
aL
icht
enst
ein
and
Mar
tens
,18
56U
nass
igne
dH
ypsi
boas
pulc
hell
usH
ypsi
boas
pulc
hell
usgr
oup
Hyl
aal
eman
iR
iver
o,19
64H
.gr
anos
agr
oup
Hyp
sibo
asal
eman
iH
ypsi
boas
punc
tatu
sgr
oup
2005 153FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
aal
leno
rum
Due
llm
anan
dT
rueb
,19
89H
.pa
rvic
eps
grou
pD
endr
opso
phus
alle
noru
mD
endr
opso
phus
parv
icep
sgr
oup
Hyl
aal
tipo
tens
Due
llm
an,
1968
H.
taen
iopu
sgr
oup
Cha
radr
ahyl
aal
tipo
tens
—H
yla
alva
reng
aiB
oker
man
n,19
56U
nass
igne
dB
oker
man
nohy
laal
vare
ngai
Bok
erm
anno
hyla
pseu
dops
eudi
sgr
oup
Hyl
aal
ytol
ylax
Due
llm
an,
1972
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
alyt
olyl
axH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
amei
both
alam
eC
anse
co-M
rque
z,M
ende
lson
,an
dG
utie
rrez
-May
en,
2002
H.
bist
inct
agr
oup
Ple
ctro
hyla
amei
both
alam
eP
lect
rohy
labi
stin
cta
grou
p
Hyl
aam
eric
ana
(Dum
eril
and
Bib
ron,
1841
)U
nass
igne
dN
omen
dubi
um—
Hyl
aam
icor
umM
ijar
es-U
rrut
ia,
1998
Una
ssig
ned
Den
drop
soph
usam
icor
umU
nass
igne
dH
yla
anat
alia
sias
iB
oker
man
n,19
72H
.ru
bicu
ndul
agr
oup
Den
drop
soph
usan
atal
iasi
asi
Den
drop
soph
usm
icro
ceph
alus
grou
p,D
.ru
bicu
ndul
uscl
ade
Hyl
aan
ceps
A.
Lut
z,19
29U
nass
igne
dD
endr
opso
phus
ance
psD
endr
opso
phus
leuc
ophy
llat
usgr
oup
Hyl
aan
ders
onii
Bai
rd,
1854
H.
vers
icol
orgr
oup
Hyl
aan
ders
onii
Hyl
aex
imia
grou
pH
yla
andi
naM
ulle
r,19
24H
.pu
lche
lla
grou
pH
ypsi
boas
andi
nus
Hyp
sibo
aspu
lche
llus
grou
pH
yla
angu
stil
inea
taT
aylo
r,19
52H
.ps
eudo
pum
agr
oup
Isth
moh
yla
angu
stil
inea
taIs
thm
ohyl
aps
eudo
pum
agr
oup
Hyl
aan
nect
ans
(Jer
don,
1870
)H
.ar
bore
agr
oup
Hyl
aan
nect
ans
Hyl
aar
bore
agr
oup
Hyl
aap
erom
eaD
uell
man
,19
82H
.m
inim
agr
oup
Den
drop
soph
usap
erom
eus
Den
drop
soph
usm
inim
usgr
oup
Hyl
aar
agua
yaN
apol
ian
dC
aram
asch
i,19
98H
.ru
bicu
ndul
agr
oup
Den
drop
soph
usar
agua
yaD
endr
opso
phus
mic
roce
phal
usgr
oup,
D.
rubi
cund
ulus
clad
eH
yla
arbo
rico
laT
aylo
r,19
41H
.ex
imia
grou
pH
yla
arbo
rico
laH
.ex
imia
grou
pH
yla
arbo
rea
(Lin
naeu
s,17
58)
H.
arbo
rea
grou
pH
yla
arbo
rea
Hyl
aar
bore
agr
oup
Hyl
aar
bore
scan
dens
Tay
lor,
1939
‘‘19
38’’
H.
mio
tym
panu
mgr
oup
Ple
ctro
hyla
arbo
resc
ande
nsP
lect
rohy
labi
stin
cta
grou
pH
yla
aren
icol
orC
ope,
1866
H.
exim
iagr
oup
Hyl
aar
enic
olor
Hyl
aex
imia
grou
pH
yla
aril
dae
Cru
zan
dP
eixo
to,
1987
‘‘19
85’’
H.
albo
fren
ata
com
plex
,H
.al
bom
argi
nata
grou
pA
plas
todi
scus
aril
dae
Apl
asto
disc
usal
bofr
enat
usgr
oup
Hyl
aar
mat
aB
oule
nger
,19
02H
.ar
mat
agr
oup
Hyl
osci
rtus
arm
atus
Hyl
osci
rtus
arm
atus
grou
pH
yla
arom
atic
aA
yarz
agae
Úna
and
Sen
aris
,19
94‘‘
1993
’’H
.ar
omat
ica
grou
pM
yers
iohy
laar
omat
ica
—
Hyl
aas
tart
eaB
oker
man
n,19
67H
.ci
rcum
data
grou
pB
oker
man
nohy
laas
tart
eaB
oker
man
nohy
laci
rcum
data
grou
pH
yla
atla
ntic
aC
aram
asch
ian
dV
elos
a,19
96H
.pu
ncta
tagr
oup
Hyp
sibo
asat
lant
icus
Hyp
sibo
aspu
ncta
tus
grou
pH
yla
aura
ria
Pet
ers,
1873
Una
ssig
ned
Nom
endu
bium
—H
yla
aviv
oca
Vio
sca,
1928
H.
vers
icol
orgr
oup
Hyl
aav
ivoc
aH
yla
vers
icol
orgr
oup
Hyl
aba
lzan
iB
oule
nger
,18
98H
.pu
lche
lla
grou
pH
ypsi
boas
balz
ani
Hyp
sibo
aspu
lche
llus
grou
pH
yla
batt
ersb
yiR
iver
o,19
61U
nass
igne
dD
endr
opso
phus
batt
ersb
yiD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
abe
cker
iC
aram
asch
ian
dC
ruz,
2004
H.
pulc
hell
agr
oup,
H.
poly
taen
iacl
ade
Hyp
sibo
asbe
cker
iH
ypsi
boas
pulc
hell
usgr
oup,
H.
poly
taen
ius
clad
eH
yla
beni
tezi
Riv
ero,
1961
Una
ssig
ned
Hyp
sibo
asbe
nite
ziH
ypsi
boas
beni
tezi
grou
p
154 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
abe
rtha
lutz
aeB
oker
man
n,19
62H
yla
deci
pien
sgr
oup
Den
drop
soph
usbe
rtha
lutz
aeD
endr
opso
phus
mic
roce
phal
usgr
oup,
D.
deci
pien
scl
ade
Hyl
abi
furc
aA
nder
sson
,19
45H
.le
ucop
hyll
ata
grou
pD
endr
opso
phus
bifu
rcus
Den
drop
soph
usle
ucop
hyll
atus
grou
pH
yla
bipu
ncta
taS
pix,
1824
H.
mic
roce
phal
agr
oup
Den
drop
soph
usbi
punc
tatu
sD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
abi
scho
ffiB
oule
nger
,18
87H
.pu
lche
lla
grou
pH
ypsi
boas
bisc
hoffi
Hyp
sibo
aspu
lche
llus
grou
pH
yla
bist
inct
aC
ope,
1878
‘‘18
77’’
H.
bist
inct
agr
oup
Ple
ctro
hyla
bist
inct
aP
lect
rohy
labi
stin
cta
grou
pH
yla
bivo
cata
Due
llm
anan
dH
oyt,
1961
H.
mio
tym
panu
mgr
oup
Exe
rodo
nta
bivo
cata
Una
ssig
ned
Hyl
abo
ans
(Lin
naeu
s,17
58)
H.
boan
sgr
oup
Hyp
sibo
asbo
ans
Hyp
sibo
asse
mil
inea
tus
grou
pH
yla
boco
urti
Moc
quar
d,18
99H
.ex
imia
grou
pH
yla
boco
urti
Hyl
aex
imia
grou
pH
yla
boge
rti
Coc
hran
and
Goi
n,19
70H
.co
lum
bian
agr
oup
Den
drop
soph
usbo
gert
iD
endr
opso
phus
colu
mbi
anus
grou
pH
yla
bogo
tens
is(P
eter
s,18
82)
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
bogo
tens
isH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
boke
rman
niG
oin,
1960
H.
parv
icep
sgr
oup
Den
drop
soph
usbo
kerm
anni
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
bran
neri
Coc
hran
,19
48H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
bran
neri
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
brev
ifro
nsD
uell
man
and
Cru
mp,
1974
H.
parv
icep
sgr
oup
Den
drop
soph
usbr
evif
rons
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
brom
elia
cia
Sch
mid
t,19
33H
.br
omel
iaci
agr
oup
Bro
mel
iohy
labr
omel
iaci
a—
Hyl
abu
riti
Car
amas
chi
and
Cru
z,19
99H
.pu
lche
lla
grou
p,H
.po
lyta
enia
clad
eH
ypsi
boas
buri
tiH
ypsi
boas
pulc
hell
usgr
oup,
H.
poly
taen
ius
clad
eH
yla
cach
imbo
Nap
oli
and
Car
amas
chi,
1999
H.
rubi
cund
ula
grou
pD
endr
opso
phus
cach
imbo
Den
drop
soph
usm
icro
ceph
alus
grou
p,D
.ru
bicu
ndul
uscl
ade
Hyl
aca
ingu
aC
arri
zo,
1991
‘‘19
90’’
H.
pulc
hell
agr
oup
Hyp
sibo
asca
ingu
aH
ypsi
boas
pulc
hell
usgr
oup
Hyl
aca
lcar
ata
Tro
sche
l,18
48H
.ge
ogra
phic
agr
oup
Hyp
sibo
asca
lcar
atus
Hyp
sibo
asal
bopu
ncta
tus
grou
pH
yla
call
ipez
aD
uell
man
,19
89H
.bo
gote
nsis
grou
pH
ylos
cirt
usca
llip
eza
Hyl
osci
rtus
bogo
tens
isgr
oup
Hyl
aca
llip
leur
aB
oule
nger
,19
02H
.pu
lche
lla
grou
pH
ypsi
boas
call
iple
ura
Hyp
sibo
aspu
lche
llus
grou
pH
yla
call
ipyg
iaC
ruz
and
Pei
xoto
,19
8519
84H
.al
bosi
gnat
aco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Apl
asto
disc
usca
llip
ygiu
sA
plas
todi
scus
albo
sign
atus
grou
p
Hyl
aca
lthu
laU
stac
h,M
ende
lson
,M
cDia
rmid
,an
dC
ampb
ell,
2000
H.
bist
inct
agr
oup
Ple
ctro
hyla
calt
hula
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
aca
lvic
olli
naT
oal,
1994
H.
bist
inct
agr
oup
Ple
ctro
hyla
calv
icol
lina
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
aca
lyps
aL
ips,
1996
H.
pict
ipes
grou
pIs
thm
ohyl
aca
lyps
aIs
thm
ohyl
api
ctip
esgr
oup
Hyl
aca
ram
asch
iiN
apol
i,20
05H
.ci
rcum
data
grou
pB
oker
man
nohy
laca
ram
asch
iiB
oker
man
nohy
laci
rcum
data
grou
pH
yla
carn
ifex
Due
llm
an,
1969
H.
colu
mbi
ana
grou
pD
endr
opso
phus
carn
ifex
Den
drop
soph
usco
lum
bian
usgr
oup
Hyl
aca
rval
hoi
Pei
xoto
,19
81H
.ci
rcum
data
grou
pB
oker
man
nohy
laca
rval
hoi
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
aca
trac
haP
orra
san
dW
ilso
n,19
87H
.m
ioty
mpa
num
grou
pE
xero
dont
aca
trac
haU
nass
igne
dH
yla
cauc
ana
Ard
ila-
Rob
ayo,
Rui
z-C
arra
nza,
and
Roa
-Tru
jill
o,19
93H
.la
rino
pygi
ongr
oup
Hyl
osci
rtus
cauc
anus
Hyl
osci
rtus
lari
nopy
gion
grou
p
2005 155FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
aca
vico
laC
ruz
and
Pei
xoto
,19
85‘‘
1984
’’H
.al
bosi
gnat
aco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Apl
asto
disc
usca
vico
laA
plas
todi
scus
albo
sign
atus
grou
p
Hyl
ace
lata
Toa
lan
dM
ende
lson
,19
95H
.bi
stin
cta
grou
pP
lect
rohy
lace
lata
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
ace
mbr
aC
aldw
ell,
1974
H.
bist
inct
agr
oup
Ple
ctro
hyla
cem
bra
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
ace
rrad
ensi
sN
apol
ian
dC
aram
asch
i,19
98H
.ru
bicu
ndul
agr
oup
Den
drop
soph
usce
rrad
ensi
sD
endr
opso
phus
mic
roce
phal
usgr
oup,
D.
rubi
cund
ulus
clad
eH
yla
chan
eque
Due
llm
an,
1961
H.
taen
iopu
sgr
oup
Cha
radr
ahyl
ach
aneq
ue—
Hyl
ach
arad
rico
laD
uell
man
,19
64H
.bi
stin
cta
grou
pP
lect
rohy
lach
arad
rico
laP
lect
rohy
labi
stin
cta
grou
pH
yla
char
azan
iV
ella
rd,
1970
H.
arm
ata
grou
pH
ylos
cirt
usch
araz
ani
Hyl
osci
rtus
arm
atus
grou
pH
yla
chim
alap
aM
ende
lson
and
Cam
pbel
l,19
94H
.su
mic
hras
tigr
oup
Exe
rodo
nta
chim
alap
aE
xero
dont
asu
mic
hras
tigr
oup
Hyl
ach
inen
sis
Gun
ther
,18
58H
.ar
bore
agr
oup
Hyl
ach
inen
sis
Hyl
aar
bore
agr
oup
Hyl
ach
loro
stea
Rey
nold
san
dF
oste
r,19
92U
nass
igne
dIn
cert
ase
dis
—H
yla
chry
ses
Adl
er,
1965
H.
bist
inct
agr
oup
Ple
ctro
hyla
chry
ses
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
ach
ryso
scel
isC
ope,
1880
H.
vers
icol
orgr
oup
Hyl
ach
ryso
scel
isH
yla
vers
icol
orgr
oup
Hyl
aci
nere
a(S
chne
ider
,17
99)
H.
cine
rea
grou
pH
yla
cine
rea
Hyl
aci
nere
agr
oup
Hyl
aci
poen
sis
B.
Lut
z,19
68H
.pu
lche
lla
grou
p,H
.po
lyta
enia
clad
eH
ypsi
boas
cipo
ensi
sH
ypsi
boas
pulc
hell
usgr
oup,
H.
poly
taen
ius
clad
eH
yla
circ
umda
ta(C
ope,
1871
)H
.ci
rcum
data
grou
pB
oker
man
nohy
laci
rcum
data
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
acl
ares
igna
taA
.L
utz
and
B.
Lut
z,19
39H
.cl
ares
igna
tagr
oup
Bok
erm
anno
hyla
clar
esig
nata
Bok
erm
anno
hyla
clar
esig
nata
grou
pH
yla
clep
sydr
aA
.L
utz,
1925
H.
clar
esig
nata
grou
pB
oker
man
nohy
lacl
epsy
dra
Bok
erm
anno
hyla
clar
esig
nata
grou
pH
yla
colu
mbi
ana
Boe
ttge
r,18
92H
.co
lum
bian
agr
oup
Den
drop
soph
usco
lum
bian
usD
endr
opso
phus
colu
mbi
anus
grou
pH
yla
coly
mba
Dun
n,19
31H
.bo
gote
nsis
grou
pH
ylos
cirt
usco
lym
baH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
cord
obae
Bar
rio,
1965
H.
pulc
hell
agr
oup
Hyp
sibo
asco
rdob
aeH
ypsi
boas
pulc
hell
usgr
oup
Hyl
acr
assa
(Bro
cchi
,18
77)
H.
bist
inct
agr
oup
Ple
ctro
hyla
cras
saP
lect
rohy
labi
stin
cta
grou
pH
yla
crep
itan
sW
ied-
Neu
wie
d,18
24H
.bo
ans
grou
pH
ypsi
boas
crep
itan
sH
ypsi
boas
fabe
rgr
oup
Hyl
acr
uzi
Pom
bal
and
Bas
tos,
1998
H.
mic
roce
phal
agr
oup
Den
drop
soph
uscr
uzi
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
cyan
omm
aC
aldw
ell,
1974
H.
bist
inct
agr
oup
Ple
ctro
hyla
cyan
omm
aP
lect
rohy
labi
stin
cta
grou
pH
yla
cycl
ada
Cam
pbel
lan
dD
uell
man
,20
00H
.m
ioty
mpa
num
grou
pP
lect
rohy
lacy
clad
aP
lect
rohy
labi
stin
cta
grou
pH
yla
cym
balu
mB
oker
man
n,19
63H
.pu
lche
lla
grou
pH
ypsi
boas
cym
balu
mH
ypsi
boas
pulc
hell
usgr
oup
Hyl
ade
bili
sT
aylo
r,19
52H
.pi
ctip
esgr
oup
Isth
moh
yla
debi
lis
Isth
moh
yla
pict
ipes
grou
pH
yla
deci
pien
sA
.L
utz,
1925
H.
deci
pien
sgr
oup
Den
drop
soph
usde
cipi
ens
Den
drop
soph
usm
icro
ceph
alus
grou
p,D
.de
cipi
ens
clad
eH
yla
dela
riva
iK
hler
and
Ltt
ers,
2001
H.
min
uta
grou
pD
endr
opso
phus
dela
riva
iD
endr
opso
phus
min
utus
grou
pH
yla
dend
roph
asm
aC
ampb
ell,
Sm
ith,
and
Ace
vedo
,20
00H
.m
ilia
ria
grou
pP
tych
ohyl
ade
ndro
phas
ma
—
Hyl
ade
ndro
scar
taT
aylo
r,19
40H
.br
omel
iaci
agr
oup
Bro
mel
iohy
lade
ndro
scar
ta—
156 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
ade
ntei
Bok
erm
ann,
1967
H.
geog
raph
ica
grou
pH
ypsi
boas
dent
eiH
ypsi
boas
albo
punc
tatu
sgr
oup
Hyl
ade
ntic
ulen
taD
uell
man
,19
72H
.bo
gote
nsis
grou
pH
ylos
cirt
usde
ntic
ulen
tus
Hyl
osci
rtus
bogo
tens
isgr
oup
Hyl
ado
lloi
Wer
ner,
1903
Una
ssig
ned
Scin
axdo
lloi
Scin
axru
ber
clad
eH
yla
dutr
aiG
omes
and
Pei
xoto
,19
96H
.m
arm
orat
agr
oup
Den
drop
soph
usdu
trai
Den
drop
soph
usm
arm
orat
usgr
oup
Hyl
aeb
racc
ata
Cop
e,18
74H
.le
ucop
hyll
ata
grou
pD
endr
opso
phus
ebra
ccat
usD
endr
opso
phus
leuc
ophy
llat
usgr
oup
Hyl
aec
hina
taD
uell
man
,19
62H
.tu
berc
ulos
agr
oup
Ecn
omio
hyla
echi
nata
—H
yla
ehrh
ardt
iM
ulle
r,19
24H
.al
bofr
enat
aco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Apl
asto
disc
useh
rhar
dti
Apl
asto
disc
usal
bofr
enat
usgr
oup
Hyl
ael
egan
sW
ied-
Neu
wie
d,18
24H
.le
ucop
hyll
ata
grou
pD
endr
opso
phus
eleg
ans
Den
drop
soph
usle
ucop
hyll
atus
grou
pH
yla
elia
neae
Nap
oli
and
Car
amas
chi,
2000
H.
rubi
cund
ula
grou
pD
endr
opso
phus
elia
neae
Den
drop
soph
usm
icro
ceph
alus
grou
p,D
.ru
bicu
ndul
uscl
ade
Hyl
aer
icae
Car
amas
chi
and
Cru
z,20
00H
.pu
lche
lla
grou
pH
ypsi
boas
eric
aeH
ypsi
boas
pulc
hell
usgr
oup
Hyl
aeu
phor
biac
eaG
unth
er,
1858
H.
exim
iagr
oup
Hyl
aeu
phor
biac
eaH
yla
exim
iagr
oup
Hyl
aex
asti
sC
aram
asch
ian
dR
odri
gues
,20
03H
.bo
ans
grou
pH
ypsi
boas
exas
tis
Hyp
sibo
asfa
ber
grou
pH
yla
exim
iaB
aird
,18
54H
.ex
imia
grou
pH
yla
exim
iaH
yla
exim
iagr
oup
Hyl
afa
ber
Wie
d-N
euw
ied,
1821
H.
boan
sgr
oup
Hyp
sibo
asfa
ber
Hyp
sibo
asfa
ber
grou
pH
yla
fasc
iata
Gun
ther
,18
58H
.ge
ogra
phic
agr
oup
Hyp
sibo
asfa
scia
tus
Hyp
sibo
asal
bopu
ncta
tus
grou
pH
yla
feio
iN
apol
ian
dC
aram
asch
i,20
04H
.ci
rcum
data
grou
pB
oker
man
nohy
lafe
ioi
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
afe
mor
alis
Bos
c,18
00H
.ci
nere
agr
oup
Hyl
afe
mor
alis
Una
ssig
ned
Hyl
afim
brim
embr
aT
aylo
r,19
48H
.m
ilia
ria
grou
pE
cnom
iohy
lafim
brim
embr
a—
Hyl
aflu
min
eaC
ruz
and
Pei
xoto
,19
85‘‘
1984
’’H
.al
bofr
enat
aco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Apl
asto
disc
usflu
min
eus
Apl
asto
disc
usal
bofr
enat
usgr
oup
Hyl
afr
eica
neca
eC
arna
val
and
Pei
xoto
,20
04H
.pu
lche
lla
grou
pH
ypsi
boas
frei
cane
cae
Hyp
sibo
aspu
lche
llus
grou
pH
yla
fuen
tei
C.
Goi
nan
dO
.G
oin,
1968
Una
ssig
ned
Hyp
sibo
asfu
ente
iU
nass
igne
dH
yla
fusc
aL
aure
nti,
1768
Una
ssig
ned
Nom
endu
bium
—H
yla
gara
goen
sis
Kap
lan,
1991
H.
gara
goen
sis
grou
pD
endr
opso
phus
gara
goen
sis
Den
drop
soph
usga
rago
ensi
sgr
oup
Hyl
aga
uche
riL
escu
rean
dM
arty
,20
00H
.pa
rvic
eps
grou
pD
endr
opso
phus
gauc
heri
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
geog
raph
ica
Spi
x,18
24H
.ge
ogra
phic
agr
oup
Hyp
sibo
asge
ogra
phic
usH
ypsi
boas
sem
ilin
eatu
sgr
oup
Hyl
agi
esle
riM
erte
ns,
1950
H.
parv
icep
sgr
oup
Den
drop
soph
usgi
esle
riD
endr
opso
phus
parv
icep
sgr
oup
Hyl
ago
dman
iG
unth
er,
1901
H.
godm
ani
grou
pT
lalo
cohy
lago
dman
i—
Hyl
ago
iana
B.
Lut
z,19
68H
.pu
lche
lla
grou
p,H
.po
lyta
enia
clad
eH
ypsi
boas
goia
nus
Hyp
sibo
aspu
lche
llus
grou
p
Hyl
ago
uvea
iP
eixo
toan
dC
ruz,
1992
H.
circ
umda
tagr
oup
Bok
erm
anno
hyla
gouv
eai
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
agr
acea
eM
yers
and
Due
llm
an,
1982
H.
pseu
dopu
ma
grou
pIs
thm
ohyl
agr
acea
eIs
thm
ohyl
aps
eudo
pum
agr
oup
Hyl
agr
andi
sona
eG
oin,
1966
H.
mic
roce
phal
agr
oup
Den
drop
soph
usgr
andi
sona
eD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
agr
anos
aB
oule
nger
,18
82H
.gr
anos
agr
oup
Hyp
sibo
asgr
anos
usH
ypsi
boas
punc
tatu
sgr
oup
2005 157FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
agr
atio
saL
eCon
te,
1857
‘‘18
56’’
H.
cine
rea
grou
pH
yla
grat
iosa
Hyl
aci
nere
agr
oup
Hyl
agr
ylla
taD
uell
man
,19
73H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
gryl
latu
sD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
agu
enth
eri
Bou
leng
er,
1886
H.
pulc
hell
agr
oup
Hyp
sibo
asgu
enth
eri
Hyp
sibo
aspu
lche
llus
grou
pH
yla
hadd
adi
Bas
tos
and
Pom
bal,
1996
H.
deci
pien
sgr
oup
Den
drop
soph
usha
ddad
iD
endr
opso
phus
mic
roce
phal
usgr
oup,
D.
deci
pien
scl
ade
Hyl
aha
llow
elli
iT
hom
pson
,19
12H
.ar
bore
agr
oup
Hyl
aha
llow
elli
iH
yla
arbo
rea
grou
pH
yla
hara
ldsc
hult
ziB
oker
man
n,19
62U
nass
igne
dD
endr
opso
phus
hara
ldsc
hult
ziU
nass
igne
dH
yla
haze
lae
Tay
lor,
1940
H.
mio
tym
panu
mgr
oup
Ple
ctro
hyla
haze
lae
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
ahe
ilpr
ini
Nob
le,
1923
Una
ssig
ned
Hyp
sibo
ashe
ilpr
ini
Hyp
sibo
asal
bopu
ncta
tus
grou
pH
yla
hele
nae
Rut
hven
,19
19U
nass
igne
dIn
cert
ase
dis
—H
yla
hobb
siC
ochr
anan
dG
oin,
1970
H.
punc
tata
grou
pH
ypsi
boas
hobb
siH
ypsi
boas
punc
tatu
sgr
oup
Hyl
ahu
tchi
nsi
Pyb
urn
and
Hal
l,19
84H
.ge
ogra
phic
agr
oup
Hyp
sibo
ashu
tchi
nsi
Hyp
sibo
asbe
nite
zigr
oup
Hyl
ahy
lax
Hey
er,
1985
H.
circ
umda
tagr
oup
Bok
erm
anno
hyla
hyla
xB
oker
man
nohy
laci
rcum
data
grou
pH
yla
hyps
elop
s(C
ope,
1871
)U
nass
igne
dN
omen
dubi
um—
Hyl
aib
irap
itan
gaC
ruz,
Pim
enta
and
Sil
vano
,20
03H
.al
bosi
gnat
aco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Apl
asto
disc
usib
irap
itan
gaA
plas
todi
scus
albo
sign
atus
grou
p
Hyl
aib
itig
uara
Car
doso
,19
83H
.ps
eudo
pseu
dis
grou
pB
oker
man
nohy
laib
itig
uara
Bok
erm
anno
hyla
pseu
dops
eudi
sgr
oup
Hyl
aib
itip
oca
Car
amas
chi
and
Fei
o,19
90H
.ci
rcum
data
grou
pB
oker
man
nohy
laib
itip
oca
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
aim
itat
or(B
arbo
uran
dD
unn,
1921
)U
nass
igne
dIn
cert
ase
dis
—H
yla
imm
acul
ata
Boe
ttge
r,18
88H
.ar
bore
agr
oup
Hyl
aim
mac
ulat
aH
yla
arbo
rea
grou
pH
yla
infr
amac
ulat
aB
oule
nger
,18
82U
nass
igne
dIn
cert
ase
dis
—H
yla
infu
cata
Due
llm
an,
1968
H.
pseu
dopu
ma
grou
pIs
thm
ohyl
ain
fuca
taIs
thm
ohyl
aps
eudo
pum
agr
oup
Hyl
ain
parq
uesi
Aya
rzag
uena
and
Sen
aris
,19
94‘‘
1993
’’H
.ar
omat
ica
grou
pM
yers
iohy
lain
parq
uesi
—
Hyl
ain
soli
taM
cCra
nie,
Wil
son,
and
Wil
liam
s,19
93H
.pi
ctip
esgr
oup
Isth
moh
yla
inso
lita
Isth
moh
yla
pict
ipes
grou
p
Hyl
ain
term
edia
Bou
leng
er,
1882
H.
arbo
rea
grou
pH
yla
inte
rmed
iaH
yla
arbo
rea
grou
pH
yla
izec
ksoh
niJi
man
dC
aram
asch
i,19
79H
.ci
rcum
data
grou
pB
oker
man
nohy
laiz
ecks
ohni
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
aja
hni
Riv
ero,
1961
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
jahn
iH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
japo
nica
Gun
ther
,18
59‘‘
1858
’’H
.ar
bore
agr
oup
Hyl
aja
poni
caH
yla
exim
iagr
oup
Hyl
aji
mi
Nap
oli
and
Car
amas
chi,
1999
H.
rubi
cund
ula
grou
pD
endr
opso
phus
jim
iD
endr
opso
phus
mic
roce
phal
usgr
oup,
D.
rubi
cund
ulus
clad
eH
yla
joan
nae
Koh
ler
and
Lot
ters
,20
01H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
joan
nae
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
joaq
uini
B.
Lut
z,19
68H
.pu
lche
lla
grou
pH
ypsi
boas
joaq
uini
Hyp
sibo
aspu
lche
llus
grou
pH
yla
juan
itae
Sny
der,
1972
H.
mio
tym
panu
mgr
oup
Exe
rodo
nta
juan
itae
—H
yla
kana
ima
Goi
nan
dW
oodl
ey,
1969
H.
geog
raph
ica
grou
pM
yers
iohy
laka
naim
a—
158 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
aka
rena
nnea
eP
ybur
n,19
93U
nass
igne
dSc
inax
kare
nann
aeSc
inax
rube
rcl
ade
Hyl
ako
echl
ini
Due
llm
anan
dT
rueb
,19
89H
.pa
rvic
eps
grou
pD
endr
opso
phus
koec
hlin
iD
endr
opso
phus
parv
icep
sgr
oup
Hyl
ala
beda
ctyl
aM
ende
lson
and
Toa
l,19
96H
.bi
stin
cta
grou
pP
lect
rohy
lala
beda
ctyl
aP
lect
rohy
labi
stin
cta
grou
pH
yla
labi
alis
Pet
ers,
1863
H.
labi
alis
grou
pD
endr
opso
phus
labi
alis
Den
drop
soph
usla
bial
isgr
oup
Hyl
ala
ncas
teri
Bar
bour
,19
28H
.pi
ctip
esgr
oup
Isth
moh
yla
lanc
aste
riIs
thm
ohyl
api
ctip
esgr
oup
Hyl
ala
ncif
orm
is(C
ope,
1871
)H
.al
bopu
ncta
tagr
oup
Hyp
sibo
asla
ncif
orm
isH
ypsi
boas
albo
punc
tatu
sgr
oup
Hyl
ala
ngei
Bok
erm
ann,
1965
H.
mar
tins
igr
oup
Bok
erm
anno
hyla
lang
eiB
oker
man
nohy
lam
arti
nsi
grou
pH
yla
lari
nopy
gion
Due
llm
an,
1973
H.
lari
nopy
gion
grou
pH
ylos
cirt
usla
rino
pygi
onH
ylos
cirt
usla
rino
pygi
ongr
oup
Hyl
ala
scin
iaR
iver
o,19
69H
.bo
gote
nsis
grou
pH
ylos
cirt
usla
scin
ius
Hyl
osci
rtus
bogo
tens
isgr
oup
Hyl
ala
tist
riat
aC
aram
asch
ian
dC
ruz,
2004
H.
pulc
hell
agr
oup,
H.
poly
taen
iacl
ade
Hyp
sibo
asla
tist
riat
usH
ypsi
boas
pulc
hell
usgr
oup,
H.
poly
taen
ius
clad
eH
yla
leal
iB
oker
man
n,19
64H
.m
inim
agr
oup
Den
drop
soph
usle
ali
Den
drop
soph
usm
inim
usgr
oup
Hyl
ale
mai
Riv
ero,
1971
Una
ssig
ned
Hyp
sibo
asle
mai
Hyp
sibo
asbe
nite
zigr
oup
Hyl
ale
ptol
inea
taP.
Bra
unan
dC
.B
raun
,19
77H
.pu
lche
lla
grou
p,H
.po
lyta
enia
clad
eH
ypsi
boas
lept
olin
eatu
sH
ypsi
boas
pulc
hell
usgr
oup,
H.
poly
taen
ius
clad
eH
yla
leuc
oche
ila
Car
amas
chi
and
Nie
mey
er,
2003
H.
albo
punc
tata
grou
pH
ypsi
boas
leuc
oche
ilus
Hyp
sibo
asal
bopu
ncta
tus
grou
pH
yla
leuc
ophy
llat
a(B
eire
is,
1783
)H
.le
ucop
hyll
ata
grou
pD
endr
opso
phus
leuc
ophy
llat
usD
endr
opso
phus
leuc
ophy
llat
usgr
oup
Hyl
ale
ucop
ygia
Cru
zan
dP
eixo
to,
1985
1984
.H
.al
bosi
gnat
aco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Apl
asto
disc
usle
ucop
ygiu
sA
plas
todi
scus
albo
sign
atus
grou
p
Hyl
ali
mai
Bok
erm
ann,
1962
Una
ssig
ned
Den
drop
soph
usli
mai
Den
drop
soph
usm
inut
usgr
oup
Hyl
ali
ndae
Due
llm
anan
dA
ltig
,19
78H
.la
rino
pygi
ongr
oup
Hyl
osci
rtus
lind
aeH
ylos
cirt
usla
rino
pygi
ongr
oup
Hyl
alo
quax
Gai
gean
dS
tuar
t,19
34H
.go
dman
igr
oup
Tla
loco
hyla
loqu
ax—
Hyl
alo
veri
dgei
Riv
ero,
1961
Una
ssig
ned
Mye
rsio
hyla
love
ridg
ei—
Hyl
alu
cian
aeN
apol
ian
dP
imen
ta,
2003
H.
circ
umda
tagr
oup
Bok
erm
anno
hyla
luci
anae
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
alu
ctuo
saP
omba
lan
dH
adda
d,19
93H
.ci
rcum
data
grou
pB
oker
man
nohy
lalu
ctuo
saB
oker
man
nohy
laci
rcum
data
grou
pH
yla
lund
iiB
urm
eist
er,
1856
H.
boan
sgr
oup
Hyp
sibo
aslu
ndii
Hyp
sibo
asfa
ber
grou
pH
yla
lute
ooce
llat
aR
oux,
1927
H.
parv
icep
sgr
oup
Den
drop
soph
uslu
teoo
cell
atus
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
lync
hiR
uiz-
Car
ranz
aan
dA
rdil
a-R
obay
o,19
91H
.bo
gote
nsis
grou
pH
ylos
cirt
usly
nchi
Hyl
osci
rtus
bogo
tens
isgr
oup
Hyl
am
argi
nata
Bou
leng
er,
1887
H.
pulc
hell
agr
oup
Hyp
sibo
asm
argi
natu
sH
ypsi
boas
pulc
hell
usgr
oup
Hyl
am
aria
nita
eC
arri
zo,
1992
H.
pulc
hell
agr
oup
Hyp
sibo
asm
aria
nita
eH
ypsi
boas
pulc
hell
usgr
oup
Hyl
am
arm
orat
a(L
aure
nti,
1768
)H
.m
arm
orat
agr
oup
Den
drop
soph
usm
arm
orat
usD
endr
opso
phus
mar
mor
atus
grou
pH
yla
mar
tins
iB
oker
man
n,19
64H
.m
arti
nsi
grou
pB
oker
man
nohy
lam
arti
nsi
Bok
erm
anno
hyla
mar
tins
igr
oup
Hyl
am
athi
asso
niC
ochr
anan
dG
oin,
1970
H.
mic
roce
phal
agr
oup
Den
drop
soph
usm
athi
asso
niD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
am
elan
argy
rea
Cop
e,18
87H
.m
arm
orat
agr
oup
Den
drop
soph
usm
elan
argy
reus
Den
drop
soph
usm
arm
orat
usgr
oup
Hyl
am
elan
omm
aT
aylo
r,19
40H
.m
ioty
mpa
num
grou
pE
xero
dont
am
elan
omm
aU
nass
igne
d
2005 159FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
am
elan
ople
ura
Bou
leng
er,
1912
H.
pulc
hell
agr
oup
Hyp
sibo
asm
elan
ople
urus
Hyp
sibo
aspu
lche
llus
grou
pH
yla
mel
anor
abdo
ta(S
chne
ider
,17
99)
Una
ssig
ned
Nom
endu
bium
—H
yla
mer
iden
sis
Riv
ero,
1961
H.
labi
alis
grou
pD
endr
opso
phus
mer
iden
sis
Den
drop
soph
usla
bial
isgr
oup
Hyl
am
erid
iana
B.
Lut
z,19
54H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
mer
idia
nus
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
mer
idio
nali
sB
oett
ger,
1874
H.
arbo
rea
grou
pH
yla
mer
idio
nali
sH
yla
arbo
rea
grou
pH
yla
mic
roce
phal
aC
ope,
1886
H.
mic
roce
phal
agr
oup
Den
drop
soph
usm
icro
ceph
alus
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
mic
rode
rma
Pyb
urn,
1977
H.
geog
raph
ica
grou
pH
ypsi
boas
mic
rode
rma
Hyp
sibo
asbe
nite
zigr
oup
Hyl
am
icro
psP
eter
s,18
72H
.pa
rvic
eps
grou
pD
endr
opso
phus
mic
rops
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
mil
iari
a(C
ope,
1886
)H
.tu
berc
ulos
agr
oup
Ecn
omio
hyla
mil
iari
a—
Hyl
am
iner
aW
ilso
n,M
cCra
nie,
and
Wil
liam
s,19
85H
.tu
berc
ulos
agr
oup
Ecn
omio
hyla
min
era
—
Hyl
am
inim
aA
hl,
1933
H.
min
ima
grou
pD
endr
opso
phus
min
imus
Den
drop
soph
usm
inim
usgr
oup
Hyl
am
inus
cula
Riv
ero,
1971
H.
mic
roce
phal
agr
oup
Den
drop
soph
usm
inus
culu
sD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
am
inut
aP
eter
s,18
72H
.m
inut
agr
oup
Den
drop
soph
usm
inut
usD
endr
opso
phus
min
utus
grou
pH
yla
mio
tym
panu
mC
ope,
1863
H.
mio
tym
panu
mgr
oup
Ecn
omio
hyla
mio
tym
panu
m—
Hyl
am
ixe
Due
llm
an,
1965
H.
mix
omac
ulat
agr
oup
Meg
asto
mat
ohyl
am
ixe
—H
yla
mix
omac
ulat
aT
aylo
r,19
50H
.m
ixom
acul
ata
grou
pM
egas
tom
atoh
yla
mix
omac
ulat
a—
Hyl
am
iyat
aiV
igle
and
Gob
erdh
an-V
igle
,19
90H
.m
inim
agr
oup
Den
drop
soph
usm
iyat
aiD
endr
opso
phus
min
imus
grou
pH
yla
mol
itor
Sch
mid
t,18
57U
nass
igne
dN
omen
dubi
um—
Hyl
am
ulti
fasc
iata
Gun
ther
,18
59‘‘
1858
’’H
.al
bopu
ncta
tagr
oup
Hyp
sibo
asm
ulti
fasc
iatu
sH
ypsi
boas
albo
punc
tatu
sgr
oup
Hyl
am
usic
aB
.L
utz,
1948
H.
albo
fren
ata
com
plex
,H
.al
bom
argi
nata
grou
pA
plas
todi
scus
mus
icus
Apl
asto
disc
usal
bofr
enat
usgr
oup
Hyl
am
ykte
rA
dler
and
Den
nis,
1972
H.
bist
inct
agr
oup
Ple
ctro
hyla
myk
ter
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
ana
hder
eri
B.
Lut
zan
dB
oker
man
n,19
63H
.m
arm
orat
agr
oup
Den
drop
soph
usna
hder
eri
Den
drop
soph
usm
arm
orat
usgr
oup
Hyl
ana
naB
oule
nger
,18
89H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
nanu
sD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
ana
nuza
eB
oker
man
nan
dS
azim
a,19
73H
.ci
rcum
data
grou
pB
oker
man
nohy
lana
nuza
eB
oker
man
nohy
laci
rcum
data
grou
pH
yla
neph
ila
Men
dels
onan
dC
ampb
ell,
1999
H.
taen
iopu
sgr
oup
Cha
radr
ahyl
ane
phil
a—
Hyl
ano
vais
iB
oker
man
n,19
68H
.m
arm
orat
agr
oup
Den
drop
soph
usno
vais
iD
endr
opso
phus
mar
mor
atus
grou
pH
yla
nubi
cola
Due
llm
an,
1964
‘‘19
63’’
H.
mix
omac
ulat
agr
oup
Meg
asto
mat
ohyl
anu
bico
la—
Hyl
aol
ivei
rai
Bok
erm
ann,
1963
H.
deci
pien
sgr
oup
Den
drop
soph
usol
ivei
rai
Den
drop
soph
usm
icro
ceph
alus
grou
p,D
.de
cipi
ens
clad
eH
yla
orna
tiss
ima
Nob
le,
1923
H.
gran
osa
grou
pH
ypsi
boas
orna
tiss
imus
Hyp
sibo
aspu
ncta
tus
grou
pH
yla
pach
aD
uell
man
and
Hil
lis,
1990
H.
lari
nopy
gion
grou
pH
ylos
cirt
uspa
cha
Hyl
osci
rtus
lari
nopy
gion
grou
pH
yla
pach
yder
ma
Tay
lor,
1942
H.
bist
inct
agr
oup
Ple
ctro
hyla
pach
yder
ma
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
apa
drel
una
Kap
lan
and
Rui
z,19
97H
.ga
rago
ensi
sgr
oup
Den
drop
soph
uspa
drel
una
Den
drop
soph
usga
rago
ensi
sgr
oup
160 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
apa
laes
tes
Due
llm
an,
De
La
Riv
a,an
dW
ild,
1997
H.
pulc
hell
agr
oup
Hyp
sibo
aspa
laes
tes
Hyp
sibo
aspu
lche
llus
grou
p
Hyl
apa
llia
taC
ope,
1863
Una
ssig
ned
Nom
endu
bium
—H
yla
palm
eri
Bou
leng
er,
1908
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
palm
eri
Hyl
osci
rtus
bogo
tens
isgr
oup
Hyl
apa
ntos
tict
aD
uell
man
and
Ber
ger,
1982
H.
lari
nopy
gion
grou
pH
ylos
cirt
uspa
ntos
tict
usH
ylos
cirt
usla
rino
pygi
ongr
oup
Hyl
apa
rdal
isS
pix,
1824
H.
boan
sgr
oup
Hyp
sibo
aspa
rdal
isH
ypsi
boas
fabe
rgr
oup
Hyl
apa
rvic
eps
Bou
leng
er,
1882
H.
parv
icep
sgr
oup
Den
drop
soph
uspa
rvic
eps
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
paui
nien
sis
Hey
er,
1977
H.
mic
roce
phal
agr
oup
Den
drop
soph
uspa
uini
ensi
sD
endr
opso
phus
parv
icep
sgr
oup
Hyl
ape
lidn
aD
uell
man
,19
89H
.la
bial
isgr
oup
Den
drop
soph
uspe
lidn
aD
endr
opso
phus
labi
alis
grou
pH
yla
pell
ita
Due
llm
an,
1968
H.
mix
omac
ulat
agr
oup
Meg
asto
mat
ohyl
ape
llit
a—
Hyl
ape
lluc
ens
Wer
ner,
1901
H.
albo
mar
gina
taco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Hyp
sibo
aspe
lluc
ens
Hyp
sibo
aspe
lluc
ens
grou
p
Hyl
ape
nthe
ter
Adl
er,
1965
H.
bist
inct
agr
oup
Ple
ctro
hyla
pent
hete
rP
lect
rohy
labi
stin
cta
grou
pH
yla
perk
insi
Cam
pbel
lan
dB
rodi
e,19
92H
.m
ioty
mpa
num
grou
pE
xero
dont
ape
rkin
siU
nass
igne
dH
yla
phae
ople
ura
Car
amas
chi
and
Cru
z,20
00H
.pu
lche
lla
grou
p,H
.po
lyta
enia
clad
eH
ypsi
boas
phae
ople
ura
Hyp
sibo
aspu
lche
llus
grou
p,H
.po
lyta
eniu
scl
ade
Hyl
aph
anta
smag
oria
Dun
n,19
43H
.tu
berc
ulos
agr
oup
Ecn
omio
hyla
phan
tasm
agor
ia—
Hyl
aph
lebo
des
Ste
jneg
er,
1906
H.
mic
roce
phal
agr
oup
Den
drop
soph
usph
lebo
des
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
phyl
logn
atha
Mel
in,
1941
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
phyl
logn
athu
sH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
pica
doi
Dun
n,19
37H
.pi
ctip
esgr
oup
Isth
moh
yla
pica
doi
Isth
moh
yla
pict
ipes
grou
pH
yla
pice
igul
aris
Rui
z-C
arra
nza
and
Lyn
ch,
1982
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
pice
igul
aris
Hyl
osci
rtus
bogo
tens
isgr
oup
Hyl
api
cta
(Gun
ther
,19
01)
H.
godm
ani
grou
pT
lalo
cohy
lapi
cta
—H
yla
pict
ipes
Cop
e,18
75‘‘
1876
’’H
.pi
ctip
esgr
oup
Isth
moh
yla
pict
ipes
Isth
moh
yla
pict
ipes
grou
pH
yla
pict
urat
aB
oule
nger
,18
99H
.ge
ogra
phic
agr
oup
Hyp
sibo
aspi
ctur
atus
Hyp
sibo
aspu
ncta
tus
grou
pH
yla
pini
ma
Bok
erm
ann
and
Saz
ima,
1973
H.
urug
uaya
grou
pSc
inax
pini
ma
Scin
axru
ber
clad
e,S.
urug
uayu
sgr
oup
Hyl
api
noru
mT
aylo
r,19
37H
.m
ioty
mpa
num
grou
pE
xero
dont
api
noru
mU
nass
igne
dH
yla
plat
ydac
tyla
Bou
leng
er,
1905
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
plat
ydac
tylu
sH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
plic
ata
Bro
cchi
,18
77H
.ex
imia
grou
pH
yla
plic
ata
Hyl
aex
imia
grou
pH
yla
poly
taen
iaC
ope,
1870
‘‘18
69’’
H.
pulc
hell
agr
oup,
H.
poly
taen
iacl
ade
Hyp
sibo
aspo
lyta
eniu
sH
ypsi
boas
pulc
hell
usgr
oup,
H.
poly
taen
ius
clad
eH
yla
pom
bali
Car
amas
chi,
Pim
enta
,an
dF
eio,
2004
H.
geog
raph
ica
grou
pH
ypsi
boas
pom
bali
Hyp
sibo
asse
mil
inea
tus
grou
p
Hyl
apr
aest
ans
Due
llm
anan
dT
rueb
,19
83U
nass
igne
dD
endr
opso
phus
prae
stan
sD
endr
opso
phus
gara
goen
sis
grou
pH
yla
pras
ina
Bur
mei
ster
,18
56H
.pu
lche
lla
grou
pH
ypsi
boas
pras
inus
Hyp
sibo
aspu
lche
llus
grou
pH
yla
psar
olai
ma
Due
llm
anan
dH
illi
s,19
90H
.la
rino
pygi
ongr
oup
Hyl
osci
rtus
psar
olai
mus
Hyl
osci
rtus
lari
nopy
gion
grou
pH
yla
psar
osem
aC
ampb
ell
and
Due
llm
an,
2000
H.
bist
inct
agr
oup
Ple
ctro
hyla
psar
osem
aP
lect
rohy
labi
stin
cta
grou
p
2005 161FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
aps
eudo
mer
idia
naC
ruz,
Car
amas
chi,
and
Dia
s,20
00H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
pseu
dom
erid
ianu
sD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
aps
eudo
pseu
dis
Mir
anda
-Rib
eiro
,19
37H
.ps
eudo
pseu
dis
grou
pB
oker
man
nohy
laps
eudo
pseu
dis
Bok
erm
anno
hyla
pseu
dops
eudi
sgr
oup
Hyl
aps
eudo
pum
aG
unth
er,
1901
H.
pseu
dopu
ma
grou
pIs
thm
ohyl
aps
eudo
pum
aIs
thm
ohyl
aps
eudo
pum
agr
oup
Hyl
apt
ycho
dact
yla
Due
llm
anan
dH
illi
s,19
90H
.la
rino
pygi
ongr
oup
Hyl
osci
rtus
ptyc
hoda
ctyl
usH
ylos
cirt
usla
rino
pygi
ongr
oup
Hyl
apu
gnax
Sch
mid
t,18
57H
.bo
ans
grou
pH
ypsi
boas
pugn
axH
ypsi
boas
fabe
rgr
oup
Hyl
apu
lche
lla
Dum
eril
and
Bib
ron,
1841
H.
pulc
hell
agr
oup
Hyp
sibo
aspu
lche
llus
Hyp
sibo
aspu
lche
llus
grou
pH
yla
puli
doi
(Riv
ero,
1961
)U
nass
igne
dH
ypsi
boas
puli
doi
Hyp
sibo
asbe
nite
zigr
oup
Hyl
apu
ncta
ta(S
chne
ider
,17
99)
H.
punc
tata
grou
pH
ypsi
boas
punc
tatu
sH
ypsi
boas
punc
tatu
sgr
oup
Hyl
aqu
adri
line
ata
(Sch
neid
er,
1799
)U
nass
igne
dN
omen
dubi
um—
Hyl
ara
nice
ps(C
ope,
1862
)H
.al
bopu
ncta
tagr
oup
Hyp
sibo
asra
nice
psH
ypsi
boas
albo
punc
tatu
sgr
oup
Hyl
ara
vida
Car
amas
chi,
Nap
oli,
and
Ber
nard
es,
2001
H.
circ
umda
tagr
oup
Bok
erm
anno
hyla
ravi
daB
oker
man
nohy
laci
rcum
data
grou
p
Hyl
arh
eaN
apol
ian
dC
aram
asch
i,19
99H
.ru
bicu
ndul
agr
oup
Den
drop
soph
usrh
eaD
endr
opso
phus
mic
roce
phal
usgr
oup,
D.
rubi
cund
ulus
clad
eH
yla
rhod
opep
laG
unth
er,
1858
H.
mic
roce
phal
agr
oup
Den
drop
soph
usrh
odop
eplu
sD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
arh
ythm
icus
Sen
aris
,an
dA
yarz
ague
na.
2002
Una
ssig
ned
Hyp
sibo
asrh
ythm
icus
Hyp
sibo
asbe
nite
zigr
oup
Hyl
ari
ojan
aK
oslo
wsk
y,18
95H
.pu
lche
lla
grou
pH
ypsi
boas
rioj
anus
Hyp
sibi
oas
pulc
hell
usgr
oup
Hyl
ari
vero
iC
ochr
anan
dG
oin,
1970
H.
min
ima
grou
pD
endr
opso
phus
rive
roi
Den
drop
soph
usm
inim
usgr
oup
Hyl
ari
vula
ris
Tay
lor,
1952
H.
pict
ipes
grou
pIs
thm
ohyl
ari
vula
ris
Isth
moh
yla
pict
ipes
grou
pH
yla
robe
rtm
erte
nsi
Tay
lor,
1937
H.
mic
roce
phal
agr
oup
Den
drop
soph
usro
bert
mer
tens
iD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
aro
bert
soru
mT
aylo
r,19
40H
.bi
stin
cta
grou
pP
lect
rohy
laro
bert
soru
mP
lect
rohy
labi
stin
cta
grou
pH
yla
roes
chm
anni
DeG
rys,
1938
Una
ssig
ned
Nom
endu
bium
—H
yla
rora
ima
Due
llm
anan
dH
oogm
oed,
1992
H.
geog
raph
ica
grou
pH
ypsi
boas
rora
ima
Hyp
sibo
asbe
nite
zigr
oup
Hyl
aro
senb
ergi
Bou
leng
er,
1898
H.
boan
sgr
oup
Hyp
sibo
asro
senb
ergi
Hyp
sibo
asfa
ber
grou
pH
yla
ross
alle
niG
oin,
1959
H.
leuc
ophy
llat
agr
oup
Den
drop
soph
usro
ssal
leni
Den
drop
soph
usle
ucop
hyll
atus
grou
pH
yla
rubi
cund
ula
Rei
nhar
dtan
dL
utke
n,18
62‘‘
1861
’’H
.ru
bicu
ndul
agr
oup
Den
drop
soph
usru
bicu
ndul
usD
endr
opso
phus
mic
roce
phal
usgr
oup,
D.
rubi
cund
ulus
clad
eH
yla
rubr
acyl
aC
ochr
anan
dG
oin,
1970
H.
albo
mar
gina
taco
mpl
ex,
H.
albo
mar
gina
tagr
oup
Hyp
sibo
asru
brac
ylus
Hyp
sibo
aspe
lluc
ens
grou
p
Hyl
aru
fitel
aF
ouqu
ette
,19
61H
.al
bom
argi
nata
com
plex
,H
.al
bom
argi
nata
grou
pH
ypsi
boas
rufit
elus
Hyp
sibo
aspe
lluc
ens
grou
p
Hyl
aru
schi
iW
eygo
ldt
and
Pei
xoto
,19
87H
.pa
rvic
eps
grou
pD
endr
opso
phus
rusc
hii
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
sabr
ina
Cal
dwel
l,19
74H
.bi
stin
cta
grou
pP
lect
rohy
lasa
brin
aP
lect
rohy
labi
stin
cta
grou
pH
yla
salv
aje
Wil
son,
McC
rani
e,an
dW
illi
ams,
1985
H.
tube
rcul
osa
grou
pE
cnom
iohy
lasa
lvaj
e—
162 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
asa
nbor
niS
chm
idt,
1944
H.
mic
roce
phal
agr
oup
Den
drop
soph
ussa
nbor
niD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
asa
nchi
ange
nsis
Pop
e,19
29H
.ar
bore
agr
oup
Hyl
asa
nchi
ange
nsis
Hyl
aar
bore
agr
oup
Hyl
asa
ram
pion
aR
uiz-
Car
ranz
aan
dL
ynch
,19
82H
.la
rino
pygi
ongr
oup
Hyl
osci
rtus
sara
mpi
ona
Hyl
osci
rtus
lari
nopy
gion
grou
pH
yla
sara
yacu
ensi
sS
hrev
e,19
35H
.le
ucop
hyll
ata
grou
pD
endr
opso
phus
sara
yacu
ensi
sD
endr
opso
phus
leuc
ophy
llat
usgr
oup
Hyl
asa
rda
(De
Bet
ta,
1853
)H
.ar
bore
agr
oup
Hyl
asa
rda
Hyl
aar
bore
agr
oup
Hyl
asa
rtor
iS
mit
h,19
51H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
sart
ori
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
savi
gnyi
Aud
ouin
,18
27H
.ar
bore
agr
oup
Hyl
asa
vign
yiH
yla
arbo
rea
grou
pH
yla
saxi
cola
Bok
erm
ann,
1964
H.
pseu
dops
eudi
sgr
oup
Bok
erm
anno
hyla
saxi
cola
Bok
erm
anno
hyla
pseu
dops
eudi
sgr
oup
Hyl
asa
zim
aiC
ardo
soan
dA
ndra
de,
1983
‘‘19
82’’
H.
circ
umda
tagr
oup
Bok
erm
anno
hyla
sazi
mai
Bok
erm
anno
hyla
circ
umda
tagr
oup
Hyl
asc
huba
rti
Bok
erm
ann,
1963
H.
min
ima
grou
pD
endr
opso
phus
schu
bart
iD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
ase
cede
nsB
.L
utz,
1963
H.
pulc
hell
agr
oup
Hyp
sibo
asse
cede
nsH
ypsi
boas
pulc
hell
usgr
oup
Hyl
ase
mig
utta
taA
.L
utz,
1925
H.
pulc
hell
agr
oup
Hyp
sibo
asse
mig
utta
tus
Hyp
sibo
aspu
lche
llus
grou
pH
yla
sem
ilin
eata
Spi
x,18
24H
.ge
ogra
phic
agr
oup
Hyp
sibo
asse
mil
inea
tus
Hyp
sibo
asse
mil
inea
tus
grou
pH
yla
seni
cula
Cop
e,18
68H
.m
arm
orat
agr
oup
Den
drop
soph
usse
nicu
lus
Den
drop
soph
usm
arm
orat
usgr
oup
Hyl
asi
bila
taC
ruz,
Pim
enta
,an
dS
ilva
no,
2003
H.
albo
sign
ata
com
plex
,H
.al
bom
argi
nata
grou
pA
plas
todi
scus
sibi
latu
sA
plas
todi
scus
albo
sign
atus
grou
p
Hyl
asi
bles
ziR
iver
o,19
71H
.gr
anos
agr
oup
Hyp
sibo
assi
bles
ziH
ypsi
boas
punc
tatu
sgr
oup
Hyl
asi
mm
onsi
Due
llm
an,
1989
H.
bogo
tens
isgr
oup
Hyl
osci
rtus
sim
mon
siH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
sim
plex
Boe
ttge
r,19
01H
.ar
bore
agr
oup
Hyl
asi
mpl
exH
yla
arbo
rea
grou
pH
yla
siop
ela
Due
llm
an,
1968
H.
bist
inct
agr
oup
Ple
ctro
hyla
siop
ela
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
asm
arag
dina
Tay
lor,
1940
H.
sum
ichr
asti
grou
pE
xero
dont
asm
arag
dina
Exe
rodo
nta
sum
ichr
asti
grou
pH
yla
smit
hii
Bou
leng
er,
1901
H.
godm
ani
grou
pT
lalo
cohy
lasm
ithi
i—
Hyl
aso
ares
iC
aram
asch
ian
dJi
m,
1983
H.
mar
mor
ata
grou
pD
endr
opso
phus
soar
esi
Den
drop
soph
usm
arm
orat
usgr
oup
Hyl
asq
uire
lla
Bos
c,18
00H
.ci
nere
agr
oup
Hyl
asq
uire
lla
Hyl
aci
nere
agr
oup
Hyl
ast
auff
eror
umD
uell
man
and
Col
oma,
1993
H.
lari
nopy
gion
grou
pH
ylos
cirt
usst
auff
eror
umH
ylos
cirt
usla
rino
pygi
ongr
oup
Hyl
ast
enoc
epha
laC
aram
asch
ian
dC
ruz,
1999
H.
pulc
hell
agr
oup,
H.
poly
taen
iacl
ade
Hyp
sibo
asst
enoc
epha
lus
Hyp
sibo
aspu
lche
llus
grou
p,H
.po
lyta
eniu
scl
ade
Hyl
ast
ingi
Kap
lan,
1994
Una
ssig
ned
Den
drop
soph
usst
ingi
Una
ssig
ned
Hyl
ast
uder
aeC
arva
lho
eS
ilva
,C
arva
lho
eS
ilva
,an
dIz
ecks
ohn,
2003
H.
mic
roce
phal
agr
oup
Den
drop
soph
usst
uder
aeD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
asu
bocu
lari
sD
unn,
1934
H.
parv
icep
sgr
oup
Den
drop
soph
ussu
bocu
lari
sD
endr
opso
phus
parv
icep
sgr
oup
Hyl
asu
mic
hras
ti(B
rocc
hi,
1879
)H
.su
mic
hras
tigr
oup
Exe
rodo
nta
sum
ichr
asti
Exe
rodo
nta
sum
ichr
asti
grou
pH
yla
suri
nam
ensi
sD
audi
n,18
02U
nass
igne
dN
omen
dubi
um—
Hyl
asu
weo
nens
isK
uram
oto,
1980
H.
arbo
rea
grou
pH
yla
suw
eone
nsis
Hyl
aex
imia
grou
pH
yla
taen
iopu
sG
unth
er,
1901
H.
taen
iopu
sgr
oup
Cha
radr
ahyl
ata
enio
pus
—
2005 163FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Hyl
ata
pich
alac
aK
izir
ian,
Col
oma,
and
Par
edes
-R
ecal
de,
2003
H.
lari
nopy
gion
grou
pH
ylos
cirt
usta
pich
alac
aH
ylos
cirt
usla
rino
pygi
ongr
oup
Hyl
ath
orec
tes
Adl
er,
1965
H.
pict
ipes
grou
pP
lect
rohy
lath
orec
tes
Ple
ctro
hyla
bist
inct
agr
oup
Hyl
ath
ysan
ota
Due
llm
an,
1966
H.
tube
rcul
osa
grou
pE
cnom
iohy
lath
ysan
ota
—H
yla
tica
Sta
rret
t,19
66H
.pi
ctip
esgr
oup
Isth
moh
yla
tica
Isth
moh
yla
pict
ipes
grou
pH
yla
tim
beba
Mar
tins
and
Car
doso
,19
87H
.pa
rvic
eps
grou
pD
endr
opso
phus
tim
beba
Den
drop
soph
uspa
rvic
eps
grou
pH
yla
tint
inna
bulu
mM
elin
,19
41U
nass
igne
dD
endr
opso
phus
tint
inna
bulu
mU
nass
igne
dH
yla
torr
enti
cola
Due
llm
anan
dA
ltig
,19
78H
.bo
gote
nsis
grou
pH
ylos
cirt
usto
rren
tico
laH
ylos
cirt
usbo
gote
nsis
grou
pH
yla
tria
ngul
umG
unth
er,
1869
‘‘18
68’’
H.
leuc
ophy
llat
agr
oup
Den
drop
soph
ustr
iang
ulum
Den
drop
soph
usle
ucop
hyll
atus
grou
pH
yla
trit
aeni
ata
Bok
erm
ann,
1965
H.
rubi
cund
ula
grou
pD
endr
opso
phus
trit
aeni
atus
Den
drop
soph
usm
icro
ceph
alus
grou
p,D
.ru
bicu
ndul
uscl
ade
Hyl
atr
uxA
dler
and
Den
nis,
1972
H.
taen
iopu
sgr
oup
Cha
radr
ahyl
atr
ux—
Hyl
ats
inli
ngen
sis
Liu
and
Hu,
1966
H.
arbo
rea
grou
pH
yla
tsin
ling
ensi
sH
yla
arbo
rea
grou
pH
yla
tube
rcul
osa
Bou
leng
er,
1882
H.
tube
rcul
osa
grou
pE
cnom
iohy
latu
berc
ulos
a—
Hyl
aur
ugua
yaS
chm
idt,
1944
H.
urug
uaya
grou
pSc
inax
urug
uayu
sSc
inax
rube
rcl
ade,
S.ur
ugua
yus
grou
pH
yla
ussu
rien
sis
Nik
olsk
y,19
18H
.ar
bore
agr
oup
Hyl
aus
suri
ensi
sH
yla
arbo
rea
grou
pH
yla
vala
ncif
erF
irsc
hein
and
Sm
ith,
1956
H.
tube
rcul
osa
grou
pE
cnom
iohy
lava
lanc
ifer
—H
yla
vare
lae
Car
rizo
,19
92U
nass
igne
dH
ypsi
boas
vare
lae
Una
ssig
ned
Hyl
ave
rsic
olor
LeC
onte
,18
25H
.ve
rsic
olor
grou
pH
yla
vers
icol
orH
yla
vers
icol
orgr
oup
Hyl
avi
gila
nsS
olan
o,19
71U
nass
igne
dIn
cert
ase
dis
—H
yla
viro
line
nsis
Kap
lan
and
Rui
z,19
97H
.ga
rago
ensi
sgr
oup
Den
drop
soph
usvi
roli
nens
isD
endr
opso
phus
gara
goen
sis
grou
pH
yla
wal
ford
iB
oker
man
n,19
62H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
wal
ford
iD
endr
opso
phus
mic
roce
phal
usgr
oup
Hyl
aw
alke
riS
tuar
t,19
54H
.ex
imia
grou
pH
yla
wal
keri
Hyl
aex
imia
grou
pH
yla
war
reni
Due
llm
anan
dH
oogm
oed,
1992
Una
ssig
ned
Ince
rtae
sedi
s—
Hyl
aw
avri
niP
arke
r,19
36H
.bo
ans
grou
pH
ypsi
boas
wav
rini
Hyp
sibo
asse
mil
inea
tus
grou
pH
yla
wer
neri
Coc
hran
,19
52H
.m
icro
ceph
ala
grou
pD
endr
opso
phus
wer
neri
Den
drop
soph
usm
icro
ceph
alus
grou
pH
yla
wey
gold
tiC
ruz
and
Pei
xoto
,19
87‘‘
1985
’’.
H.
albo
fren
ata
com
plex
,H
.al
bom
argi
nata
grou
pA
plas
todi
scus
wey
gold
tiA
plas
todi
scus
albo
fren
atus
grou
p
Hyl
aw
righ
toru
mT
aylo
r,19
39‘‘
1938
’’H
.ex
imia
grou
pH
yla
wri
ghto
rum
Hyl
aex
imia
grou
pH
yla
xant
host
icta
Due
llm
an,
1968
H.
pict
ipes
grou
pIs
thm
ohyl
axa
ntho
stic
taIs
thm
ohyl
api
ctip
esgr
oup
Hyl
axa
puri
ensi
sM
arti
nsan
dC
ardo
so,
1987
H.
min
uta
grou
pD
endr
opso
phus
xapu
rien
sis
Den
drop
soph
usm
inut
usgr
oup
Hyl
axe
raM
ende
lson
and
Cam
pbel
l,19
94H
.su
mic
hras
tigr
oup
Exe
rodo
nta
xera
Exe
rodo
nta
sum
ichr
asti
grou
pH
yla
yara
cuya
naM
ijar
es-U
rrut
iaan
dR
iver
o,20
00U
nass
igne
dD
endr
opso
phus
yara
cuya
nus
Una
ssig
ned
Hyl
aze
teki
Gai
ge,
1929
H.
pict
ipes
grou
pIs
thm
ohyl
aze
teki
Isth
moh
yla
pict
ipes
grou
pH
yla
zhao
ping
ensi
sT
ang
and
Zha
ng,
1984
H.
arbo
rea
grou
pH
yla
zhao
ping
ensi
sH
yla
arbo
rea
grou
pL
ysap
sus
cara
yaG
alla
rdo,
1964
—L
ysap
sus
cara
ya—
164 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Lys
apsu
sla
evis
Par
ker,
1935
—L
ysap
sus
laev
is—
Lys
apsu
sli
mel
lum
Cop
e,18
62—
Lys
apsu
sli
mel
lum
—N
ycti
man
tis
rugi
ceps
Bou
leng
er,
1882
—N
ycti
man
tis
rugi
ceps
—O
steo
ceph
alus
buck
leyi
(Bou
leng
er,
1882
)—
Ost
eoce
phal
usbu
ckle
yi—
Ost
eoce
phal
usca
brer
ai(C
ochr
anan
dG
oin,
1970
)—
Ost
eoce
phal
usca
brer
ai—
Ost
eoce
phal
usde
ride
nsJu
ngfe
r,R
on,
Sei
pp,
and
Alm
enda
riz,
2000
—O
steo
ceph
alus
deri
dens
—
Ost
eoce
phal
usel
keju
ngin
gera
e(H
enle
,19
81)
—O
steo
ceph
alus
elke
jung
inge
rae
—O
steo
ceph
alus
exop
htha
lmus
Sm
ith
and
Noo
nan,
2001
—O
steo
ceph
alus
exop
htha
lmus
—
Ost
eoce
phal
usfu
scif
acie
sJu
ngfe
r,R
on,
Sei
pp,
and
Alm
enda
riz,
2000
—O
steo
ceph
alus
fusc
ifac
ies
—
Ost
eoce
phal
ushe
yeri
Lyn
ch,
2002
—O
steo
ceph
alus
heye
ri—
Ost
eoce
phal
usla
ngsd
orffi
i(D
umer
ilan
dB
ibro
n,18
41)
—It
apot
ihyl
ala
ngsd
orffi
i—
Ost
eoce
phal
usle
onia
eJu
ngfe
ran
dL
ehr,
2001
—O
steo
ceph
alus
leon
iae
—O
steo
ceph
alus
lepr
ieur
ii(D
umla
ril
and
Bib
ron,
1841
)—
Ost
eoce
phal
usle
prie
urii
—
Ost
eoce
phal
usm
utab
orJu
ngfe
ran
dH
odl,
2002
—O
steo
ceph
alus
mut
abor
—O
steo
ceph
alus
ooph
agus
Jung
fer
and
Sch
iesa
ri,
1995
—O
steo
ceph
alus
ooph
agus
—
Ost
eoce
phal
uspe
arso
ni(G
aige
,19
29)
—O
steo
ceph
alus
pear
soni
—O
steo
ceph
alus
plan
icep
sC
ope,
1874
—O
steo
ceph
alus
plan
icep
s—
Ost
eoce
phal
ussu
btil
isM
arti
nsan
dC
ardo
so,
1987
—O
steo
ceph
alus
subt
ilis
—O
steo
ceph
alus
taur
inus
Ste
inda
chne
r,18
62—
Ost
eoce
phal
usta
urin
us—
Ost
eoce
phal
usve
rruc
iger
(Wer
ner,
1901
)—
Ost
eoce
phal
usve
rruc
iger
—O
steo
ceph
alus
yasu
niR
onan
dP
ram
uk,
1999
—O
steo
ceph
alus
yasu
ni—
Ost
eopi
lus
brun
neus
(Gos
se,
1851
)—
Ost
eopi
lus
brun
neus
—O
steo
pilu
scr
ucia
lis
(Har
lan,
1826
)—
Ost
eopi
lus
cruc
iali
s—
Ost
eopi
lus
dom
inic
ensi
s(T
schu
di,
1838
)—
Ost
eopi
lus
dom
inic
ensi
s—
Ost
eopi
lus
mar
iana
e(D
unn,
1926
)—
Ost
eopi
lus
mar
iana
e—
Ost
eopi
lus
pulc
hril
inea
tus
Cop
e,18
70‘‘
1869
’’—
Ost
eopi
lus
pulc
hril
inea
tus
—O
steo
pilu
sse
pten
trio
nali
s(D
umer
ilan
dB
ibro
n,18
41)
—O
steo
pilu
sse
pten
trio
nali
s—
Ost
eopi
lus
vast
us(C
ope,
1871
)—
Ost
eopi
lus
vast
us—
Ost
eopi
lus
wil
deri
(Dun
n,19
25)
—O
steo
pilu
sw
ilde
ri—
2005 165FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Phr
ynoh
yas
cori
acea
(Pet
ers,
1867
)—
Tra
chyc
epha
lus
cori
aceu
s—
Phr
ynoh
yas
hadr
ocep
s(D
uell
man
and
Hoo
gmoe
d,19
92)
—T
rach
ycep
halu
sha
droc
eps
—
Phr
ynoh
yas
imit
atri
x(M
iran
da-R
ibei
ro,
1926
)—
Tra
chyc
epha
lus
imit
atri
x—
Phr
ynoh
yas
mes
opha
ea(H
ense
l,18
67)
—T
rach
ycep
halu
sm
esop
haeu
s—
Phr
ynoh
yas
resi
nific
trix
(Goe
ldi,
1907
)—
Tra
chyc
epha
lus
resi
nific
trix
—P
hryn
ohya
sve
nulo
sa(L
aure
nti,
1768
)—
Tra
chyc
epha
lus
venu
losu
s—
Phy
llod
ytes
acum
inat
usB
oker
man
n,19
66P
hyll
odyt
eslu
teol
usgr
oup
Phy
llod
ytes
acum
inat
usP
hyll
odyt
eslu
teol
usgr
oup
Phy
llod
ytes
aura
tus
(Bou
leng
er,
1917
)P
.au
ratu
sgr
oup
Phy
llod
ytes
aura
tus
P.
aura
tus
grou
pP
hyll
odyt
esbr
evir
ostr
isP
eixo
toan
dC
ruz,
1988
P.
lute
olus
grou
pP
hyll
odyt
esbr
evir
ostr
isP
.lu
teol
usgr
oup
Phy
llod
ytes
edel
moi
Pei
xoto
,C
aram
asch
i,an
dF
reir
e,20
03P
.lu
teol
usgr
oup
Phy
llod
ytes
edel
moi
P.
lute
olus
grou
p
Phy
llod
ytes
gyri
naet
hes
Pei
xoto
,C
aram
asch
ian
dF
reir
e,20
03P
.gy
rina
ethe
sgr
oup
Phy
llod
ytes
gyri
naet
hes
P.
gyri
naet
hes
grou
p
Phy
llod
ytes
kaut
skyi
Pei
xoto
and
Cru
z,19
88P
.lu
teol
usgr
oup
Phy
llod
ytes
kaut
skyi
P.
lute
olus
grou
pP
hyll
odyt
eslu
teol
us(W
ied-
Neu
wie
d,18
24)
P.
lute
olus
grou
pP
hyll
odyt
eslu
teol
usP
.lu
teol
usgr
oup
Phy
llod
ytes
mel
anom
ysta
xC
aram
asch
i,da
Sil
va,
and
Bri
tto-
Per
eira
,19
92P
.lu
teol
usgr
oup
Phy
llod
ytes
mel
anom
ysta
xP
.lu
teol
usgr
oup
Phy
llod
ytes
punc
tatu
sC
aram
asch
ian
dP
eixo
to,
2004
P.
tube
rcul
osus
grou
pP
hyll
odyt
espu
ncta
tus
P.
tube
rcul
osus
grou
p
Phy
llod
ytes
tube
rcul
osus
Bok
erm
ann,
1966
P.
tube
rcul
osus
grou
pP
hyll
odyt
estu
berc
ulos
usP
.tu
berc
ulos
usgr
oup
Phy
llod
ytes
wuc
here
ri(P
eter
s,18
73)
P.
aura
tus
grou
pP
hyll
odyt
esw
uche
reri
P.
aura
tus
grou
pP
lect
rohy
laac
anth
odes
Due
llm
anan
dC
ampb
ell,
1992
—P
lect
rohy
laac
anth
odes
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
avia
Stu
art,
1952
—P
lect
rohy
laav
iaP
lect
rohy
lagu
atem
alen
sis
grou
pP
lect
rohy
lach
ryso
pleu
raW
ilso
n,M
cCra
nie,
and
Cru
z,19
94—
Ple
ctro
hyla
chry
sopl
eura
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
dasy
pus
McC
rani
ean
dW
ilso
n,19
81—
Ple
ctro
hyla
dasy
pus
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
exqu
isit
aM
cCra
nie
and
Wil
son,
1998
—P
lect
rohy
laex
quis
ita
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
glan
dulo
sa(B
oule
nger
,18
83)
—P
lect
rohy
lagl
andu
losa
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
guat
emal
ensi
sB
rocc
hi,
1877
—P
lect
rohy
lagu
atem
alen
sis
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
hart
weg
iD
uell
man
,19
68—
Ple
ctro
hyla
hart
weg
iP
lect
rohy
lagu
atem
alen
sis
grou
pP
lect
rohy
laix
ilS
tuar
t,19
42—
Ple
ctro
hyla
ixil
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
lace
rtos
aB
umza
hem
and
Sm
ith,
1954
—P
lect
rohy
lala
cert
osa
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
mat
udai
Har
tweg
,19
41—
Ple
ctro
hyla
mat
udai
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
166 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Ple
ctro
hyla
poko
mch
iD
uell
man
and
Cam
pbel
l,19
84—
Ple
ctro
hyla
poko
mch
iP
lect
rohy
lagu
atem
alen
sis
grou
p
Ple
ctro
hyla
psil
oder
ma
McC
rani
ean
dW
ilso
n,19
99—
Ple
ctro
hyla
psil
oder
ma
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
pycn
ochi
laR
abb,
1959
—P
lect
rohy
lapy
cnoc
hila
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
quec
chi
Stu
art,
1942
—P
lect
rohy
laqu
ecch
iP
lect
rohy
lagu
atem
alen
sis
grou
pP
lect
rohy
lasa
goru
mH
artw
eg,
1941
—P
lect
rohy
lasa
goru
mP
lect
rohy
lagu
atem
alen
sis
grou
pP
lect
rohy
late
cunu
man
iD
uell
man
and
Cam
pbel
l,19
84—
Ple
ctro
hyla
tecu
num
ani
Ple
ctro
hyla
guat
emal
ensi
sgr
oup
Ple
ctro
hyla
teuc
hest
esD
uell
man
and
Cam
pbel
l,19
92—
Ple
ctro
hyla
teuc
hest
esP
lect
rohy
lagu
atem
alen
sis
grou
p
Pse
udac
ris
brac
hyph
ona
(Cop
e,18
89)
Pse
udac
ris
nigr
ita
clad
eP
seud
acri
sbr
achy
phon
aP
seud
acri
sni
grit
acl
ade
Pse
udac
ris
brim
leyi
Bra
ndt
and
Wal
ker,
1933
P.
nigr
ita
clad
eP
seud
acri
sbr
imle
yiP
.ni
grit
acl
ade
Pse
udac
ris
cada
veri
na(C
ope,
1866
)P
.re
gill
acl
ade
Pse
udac
ris
cada
veri
naP
.re
gill
acl
ade
Pse
udac
ris
clar
kii
(Bai
rd,
1854
)P
.ni
grit
acl
ade
Pse
udac
ris
clar
kii
P.
nigr
ita
clad
eP
seud
acri
scr
ucif
er(W
ied-
Neu
wie
d,18
38)
P.
cruc
ifer
clad
eP
seud
acri
scr
ucif
erP
.cr
ucif
ercl
ade
Pse
udac
ris
feri
arum
(Bai
rd,
1854
)P
.ni
grit
acl
ade
Pse
udac
ris
feri
arum
P.
nigr
ita
clad
eP
seud
acri
sil
lino
ensi
sS
mit
h,19
51P
.or
nata
clad
eP
seud
acri
sil
lino
ensi
sP
.or
nata
clad
eP
seud
acri
sm
acul
ata
(Aga
ssiz
,18
50)
P.
nigr
ita
clad
eP
seud
acri
sm
acul
ata
P.
nigr
ita
clad
eP
seud
acri
sni
grit
a(L
eCon
te,
1825
)P
.ni
grit
acl
ade
Pse
udac
ris
nigr
ita
P.
nigr
ita
clad
eP
seud
acri
soc
ular
is(B
osc
and
Dau
din,
1801
)P
.cr
ucif
ercl
ade
Pse
udac
ris
ocul
aris
P.
cruc
ifer
clad
eP
seud
acri
sor
nata
(Hol
broo
k,18
36)
P.
orna
tacl
ade
Pse
udac
ris
orna
taP
.or
nata
clad
eP
seud
acri
sre
gill
a(B
aird
and
Gir
ard,
1852
)P
.re
gill
acl
ade
Pse
udac
ris
regi
lla
P.
regi
lla
clad
eP
seud
acri
sst
reck
eri
A.
A.
Wri
ght
and
A.
H.
Wri
ght,
1933
P.
orna
tacl
ade
Pse
udac
ris
stre
cker
iP
.or
nata
clad
e
Pse
udac
ris
tris
eria
ta(W
ied-
Neu
wie
d,18
38)
P.
nigr
ita
clad
eP
seud
acri
str
iser
iata
P.
nigr
ita
clad
eP
seud
isbo
lbod
acty
laA
.L
utz,
1925
—P
seud
isbo
lbod
acty
la—
Pse
udis
card
osoi
Kw
et,
2000
—P
seud
isca
rdos
oi—
Pse
udis
fusc
aG
arm
an,
1883
—P
seud
isfu
sca
—P
seud
ism
inut
aG
unth
er,
1858
—P
seud
ism
inut
a—
Pse
udis
para
doxa
(Lin
naeu
s,17
58)
—P
seud
ispa
rado
xa—
Pse
udis
toca
ntin
sC
aram
asch
ian
dC
ruz,
1998
—P
seud
isto
cant
ins
—P
tern
ohyl
ade
ntat
aS
mit
h,19
57—
Smil
isca
dent
ata
—P
tern
ohyl
afo
dien
sB
oule
nger
,18
82—
Smil
isca
fodi
ens
—P
tych
ohyl
aac
roch
orda
Cam
pbel
lan
dD
uell
man
,20
00—
Pty
choh
yla
acro
chor
da—
2005 167FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Pty
choh
yla
eryt
hrom
ma
(Tay
lor,
1937
)—
Pty
choh
yla
eryt
hrom
ma
—P
tych
ohyl
aeu
thys
anot
a(K
ello
gg,
1928
)—
Pty
choh
yla
euth
ysan
ota
—P
tych
ohyl
ahy
pom
ykte
rM
cCra
nie
and
Wil
son,
1993
—P
tych
ohyl
ahy
pom
ykte
r—
Pty
choh
yla
legl
eri
(Tay
lor,
1958
)—
Pty
choh
yla
legl
eri
—P
tych
ohyl
ale
onha
rdsc
hult
zei
(Ahl
,19
34)
—P
tych
ohyl
ale
onha
rdsc
hult
zei
—P
tych
ohyl
am
acro
tym
panu
m(T
anne
r,19
57)
—P
tych
ohyl
am
acro
tym
panu
m—
Pty
choh
yla
panc
hoi
Due
llm
anan
dC
ampb
ell,
1982
—P
tych
ohyl
apa
ncho
i—
Pty
choh
yla
salv
ador
ensi
s(M
erte
ns,
1952
)—
Pty
choh
yla
salv
ador
ensi
s—
Pty
choh
yla
sanc
taec
ruci
sC
ampb
ell
and
Sm
ith,
1992
—P
tych
ohyl
asa
ncta
ecru
cis
—
Pty
choh
yla
spin
ipol
lex
(Sch
mid
t,19
36)
—P
tych
ohyl
asp
inip
olle
x—
Pty
choh
yla
zoph
odes
Cam
pbel
lan
dD
uell
man
,20
00—
Pty
choh
yla
zoph
odes
—
Scar
thyl
ago
inor
um(B
oker
man
n,19
64)
—Sc
arth
yla
goin
orum
—Sc
inax
acum
inat
us(C
ope,
1862
)S.
rube
rcl
ade
Scin
axac
umin
atus
S.ru
ber
clad
eSc
inax
agil
is(C
ruz
&P
eixo
to,
1983
)S.
cath
arin
aecl
ade
Scin
axag
ilis
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup.
Scin
axal
bica
ns(B
oker
man
n,19
67)
S.ca
thar
inae
clad
eSc
inax
albi
cans
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axal
catr
az(B
.L
utz,
1973
)S.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axal
catr
azS.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axal
tae
(Dun
n,19
33)
S.ru
ber
clad
eSc
inax
alta
eS.
rube
rcl
ade
Scin
axal
ter
(B.
Lut
z,19
73)
S.ru
ber
clad
eSc
inax
alte
rS.
rube
rcl
ade
Scin
axan
gren
sis
(B.
Lut
z,19
73)
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axan
gren
sis
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axar
duou
s(P
eixo
to,
2001
)S.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axar
duou
sS.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axar
gyre
orna
tus
(Mir
anda
-Rib
eiro
,19
26)
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axar
gyre
orna
tus
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axar
iadn
e(B
oker
man
n,19
67)
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axar
iadn
eS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
arom
othy
ella
Fai
vovi
ch,
2005
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axar
omot
hyel
laS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
atra
tus
(Pei
xoto
,‘‘
1988
’’19
89)
S.ca
thar
inae
clad
e,S.
perp
usil
lus
grou
pSc
inax
atra
tus
S.ca
thar
inae
clad
e,S.
perp
usil
lus
grou
p
168 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Scin
axau
ratu
s(W
ied,
1821
)S.
rube
rcl
ade
Scin
axau
ratu
sS.
rube
rcl
ade
Scin
axba
umga
rdne
ri(R
iver
o,19
61)
S.ru
ber
clad
eSc
inax
baum
gard
neri
S.ru
ber
clad
eSc
inax
bert
hae
(Bar
rio,
1962
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
bert
hae
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axbl
airi
(Fou
quet
tean
dP
ybur
n,19
72)
S.ru
ber
clad
eSc
inax
blai
riS.
rube
rcl
ade
Scin
axbo
esem
ani
(Goi
n,19
66)
S.ru
ber
clad
eSc
inax
boes
eman
iS.
rube
rcl
ade
Scin
axbo
ulen
geri
(Cop
e,18
87)
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axbo
ulen
geri
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axbr
ieni
(De
Wit
te,
1930
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
brie
niS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
cald
arum
(B.
Lut
z,19
68)
S.ru
ber
clad
eSc
inax
cald
arum
S.ru
ber
clad
eSc
inax
cana
stre
nsis
(Car
doso
and
Had
dad,
1982
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
cana
stre
nsis
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axca
rdos
oi(C
arva
lho
eS
ilva
and
Pei
xoto
,19
91)
S.ru
ber
clad
eSc
inax
card
osoi
S.ru
ber
clad
e
Scin
axca
rnev
alli
i(C
aram
asch
ian
dK
iste
umac
her,
1989
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
carn
eval
liS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
cast
rovi
ejoi
De
La
Riv
a,19
93S.
rube
rcl
ade
Scin
axca
stro
viej
oiS.
rube
rcl
ade
Scin
axca
thar
inae
(Bou
leng
er,
1888
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
cath
arin
aeS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
cent
rali
sP
omba
lan
dB
asto
s,19
96S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
cent
rali
sS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
chiq
uita
nus
(De
La
Riv
a,19
90)
S.ru
ber
clad
eSc
inax
chiq
uita
nus
S.ru
ber
clad
eSc
inax
cros
pedo
spil
us(A
.L
utz,
1925
)S.
rube
rcl
ade
Scin
axcr
ospe
dosp
ilus
S.ru
ber
clad
eSc
inax
crue
ntom
mus
(Due
llm
an,
1972
)S.
rube
rcl
ade
Scin
axcr
uent
omm
usS.
rube
rcl
ade
Scin
axcu
rici
caP
ugli
ese,
Pom
bal,
and
Saz
ima,
2004
S.ru
ber
clad
eSc
inax
curi
cica
S.ru
ber
clad
e
Scin
axcu
spid
atus
(A.
Lut
z,19
25)
S.ru
ber
clad
eSc
inax
cusp
idat
usS.
rube
rcl
ade
Scin
axda
nae
(Due
llm
an,
1986
)S.
rube
rcl
ade
Scin
axda
nae
S.ru
ber
clad
eSc
inax
duar
tei
(B.
Lut
z,19
51)
S.ru
ber
clad
eSc
inax
duar
tei
S.ru
ber
clad
eSc
inax
elae
ochr
ous
(Cop
e,18
75)
S.ru
ber
clad
eSc
inax
elae
ochr
ous
S.ru
ber
clad
eSc
inax
eury
dice
(Bok
erm
ann,
1968
)S.
rube
rcl
ade
Scin
axeu
rydi
ceS.
rube
rcl
ade
Scin
axex
iguu
s(D
uell
man
,19
86)
S.ru
ber
clad
eSc
inax
exig
uus
S.ru
ber
clad
eSc
inax
flavi
dus
La
Mar
ca,
2004
S.ru
ber
clad
eSc
inax
flavi
dus
S.ru
ber
clad
eSc
inax
flavo
gutt
atus
(A.
Lut
zan
dB
.L
utz,
1939
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
flavo
gutt
atus
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
2005 169FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Scin
axfu
nere
us(C
ope,
1874
)S.
rube
rcl
ade
Scin
axfu
nere
usS.
rube
rcl
ade
Scin
axfu
scom
argi
natu
s(A
.L
utz,
1925
)S.
rube
rcl
ade
Scin
axfu
scom
argi
natu
sS.
rube
rcl
ade
Scin
axfu
scov
ariu
s(A
.L
utz,
1925
)S.
rube
rcl
ade
Scin
axfu
scov
ariu
sS.
rube
rcl
ade
Scin
axga
rbei
(Mir
anda
-Rib
eiro
,19
26)
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axga
rbei
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axgr
anul
atus
(Pet
ers,
1871
)S.
rube
rcl
ade
Scin
axgr
anul
atus
S.ru
ber
clad
eSc
inax
hayi
i(B
arbo
ur,
1909
)S.
rube
rcl
ade
Scin
axha
yii
S.ru
ber
clad
eSc
inax
heye
ri(P
eixo
toan
dW
eygo
ldt,
1986
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
heye
riS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
hiem
alis
(Had
dad
and
Pom
bal,
1987
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
hiem
alis
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axhu
mil
is(B
.L
utz,
1954
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
hum
ilis
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axic
teri
cus
Due
llm
anan
dW
iens
,19
93S.
rube
rcl
ade
Scin
axic
teri
cus
S.ru
ber
clad
eSc
inax
joly
iL
escu
rean
dM
arty
,20
00S.
rube
rcl
ade,
S.ro
stra
tus
grou
pSc
inax
joly
iS.
rube
rcl
ade,
S.ro
stra
tus
grou
p
Scin
axju
reia
(Pom
bal
and
Gor
do,
1991
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
jure
iaS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
kaut
skyi
(Car
valh
oe
Sil
vaan
dP
eixo
to,
1991
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
kaut
skyi
S.ca
thar
inae
Scin
axke
nned
yi(P
ybur
n19
73)
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axke
nned
yiS.
rube
rcl
ade,
S.ro
stra
tus
grou
p
Scin
axli
ndsa
yiP
ybur
n,19
92S.
rube
rcl
ade
Scin
axli
ndsa
yiS.
rube
rcl
ade
Scin
axli
ttor
alis
(Pom
bal
and
Gor
do,
1991
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
litt
oral
isS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
litt
oreu
s(P
exot
o,19
88)
S.ca
thar
inae
clad
e,S.
perp
usil
lus
grou
pSc
inax
litt
oreu
sS.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axlo
ngil
ineu
s(B
.L
utz,
1968
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
long
ilin
eus
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axlu
izot
avio
i(C
aram
asch
ian
dK
iste
umac
her,
1989
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
luiz
otav
ioi
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axm
acha
doi
(Bok
erm
ann
and
Saz
ima,
1973
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
mac
hado
iS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
man
riqu
eiB
arri
o-A
mor
os,
Ore
llan
a,an
dC
haco
n,20
04S.
rube
rcl
ade
Scin
axm
anri
quei
S.ru
ber
clad
e
Scin
axm
arac
aya
(Car
doso
and
Saz
ima,
1980
)S.
rube
rcl
ade
Scin
axm
arac
aya
S.ru
ber
clad
e
170 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Scin
axm
egap
odiu
s(M
iran
da-R
ibei
ro,
1926
)S.
rube
rcl
ade
Scin
axfu
scov
ariu
sS.
rube
rcl
ade
Scin
axm
ello
i(P
eixo
to,
1989
)S.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axm
ello
iS.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axna
sicu
s(C
ope,
1862
)S.
rube
rcl
ade
Scin
axna
sicu
sS.
rube
rcl
ade
Scin
axne
bulo
sus
(Spi
x,18
24)
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axne
bulo
sus
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axob
tria
ngul
atus
(B.
Lut
z,19
73)
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axob
tria
ngul
atus
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axor
eite
sD
uell
man
and
Wie
ns,
1993
S.ru
ber
clad
eSc
inax
orei
tes
S.ru
ber
clad
eSc
inax
pach
ycru
s(M
iran
da-R
ibei
ro,
1937
)S.
rube
rcl
ade
Scin
axpa
chyc
rus
S.ru
ber
clad
eSc
inax
park
eri
(Gai
ge,
1926
)S.
rube
rcl
ade
Scin
axpa
rker
iS.
rube
rcl
ade
Scin
axpe
drom
edin
ae(H
enle
,19
91)
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axpe
drom
edin
aeS.
rube
rcl
ade,
S.ro
stra
tus
grou
p
Scin
axpe
rere
caP
omba
l,H
adda
d,an
dK
asah
ara,
1995
S.ru
ber
clad
eSc
inax
pere
reca
S.ru
ber
clad
e
Scin
axpe
rpus
illu
s(A
.L
utz
and
B.
Lut
z,19
39)
S.ca
thar
inae
clad
e,S.
perp
usil
lus
grou
pSc
inax
perp
usil
lus
S.ca
thar
inae
clad
e,S.
perp
usil
lus
grou
pSc
inax
prob
osci
deus
(Bro
nger
sma,
1933
)S.
rube
rcl
ade,
S.ro
stra
tus
grou
pSc
inax
prob
osci
deus
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axqu
inqu
efas
ciat
us(F
owle
r,19
13)
S.ru
ber
clad
eSc
inax
quin
quef
asci
atus
S.ru
ber
clad
eSc
inax
rank
i(A
ndra
dean
dC
ardo
so,
1987
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
rank
iS.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
rizi
bili
s(B
oker
man
n,19
64)
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axri
zibi
lis
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
Scin
axro
stra
tus
(Pet
ers,
1863
)S.
rube
rcl
ade,
S.ro
stra
tus
grou
pSc
inax
rost
ratu
sS.
rube
rcl
ade,
S.ro
stra
tus
grou
p
Scin
axru
ber
(Lau
rent
i,17
68)
S.ru
ber
clad
eSc
inax
rube
rS.
rube
rcl
ade
Scin
axsi
mil
is(C
ochr
an,
1952
)S.
rube
rcl
ade
Scin
axsi
mil
isS.
rube
rcl
ade
Scin
axsq
uali
rost
ris
(A.
Lut
z,19
26)
S.ru
ber
clad
eSc
inax
squa
liro
stri
sS.
rube
rcl
ade
Scin
axst
auff
eri
(Cop
e,18
65)
S.ru
ber
clad
eSc
inax
stau
ffer
iS.
rube
rcl
ade
Scin
axst
rigi
latu
s(S
pix,
1824
)S.
cath
arin
aecl
ade
Scin
axst
rigi
latu
sS.
cath
arin
aecl
ade
Scin
axsu
gill
atus
(D
uell
man
,19
73)
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axsu
gill
atus
S.ru
ber
clad
e,S.
rost
ratu
sgr
oup
Scin
axtr
achy
thor
ax(M
ulle
ran
dH
ellm
ich,
1936
)S.
rube
rcl
ade
Scin
axfu
scov
ariu
sS.
rube
rcl
ade
Scin
axtr
apic
heir
oi(B
.L
utz,
1954
)S.
cath
arin
aecl
ade,
S.ca
thar
inae
grou
pSc
inax
trap
iche
iroi
S.ca
thar
inae
clad
e,S.
cath
arin
aegr
oup
2005 171FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Scin
axtr
ilin
eatu
s(H
oogm
oed
and
Gor
zula
,19
77)
S.ru
ber
clad
eSc
inax
tril
inea
tus
S.ru
ber
clad
eSc
inax
v-si
gnat
us(B
.L
utz,
1968
)S.
cath
arin
aecl
ade,
S.pe
rpus
illu
sgr
oup
Scin
axv-
sign
atus
S.ca
thar
inae
clad
e,S.
perp
usil
lus
grou
pSc
inax
wan
dae
(Pyb
urn
and
Fou
quet
te,
1971
)S.
rube
rcl
ade
Scin
axw
anda
eS.
rube
rcl
ade
Scin
axx-
sign
atus
(Spi
x,18
24)
S.ru
ber
clad
eSc
inax
x-si
gnat
usS.
rube
rcl
ade
Smil
isca
baud
inii
(Dum
eril
and
Bib
ron,
1841
)—
Smil
isca
baud
inii
—Sm
ilis
cacy
anos
tict
a(S
mit
h,19
53)
—Sm
ilis
cacy
anos
tict
a—
Smil
isca
phae
ota
(Cop
e,18
62)
—Sm
ilis
caph
aeot
a—
Smil
isca
pum
a(C
ope,
1885
)—
Smil
isca
pum
a—
Smil
isca
sila
Due
llm
anan
dT
rueb
,19
66—
Smil
isca
sila
—Sm
ilis
caso
rdid
a(P
eter
s,18
63)
—Sm
ilis
caso
rdid
a—
Spha
enor
hync
hus
brom
elic
ola
Bok
erm
ann,
1966
—Sp
haen
orhy
nchu
sbr
omel
icol
a—
Spha
enor
hync
hus
carn
eus
(Cop
e,18
68)
—Sp
haen
orhy
nchu
sca
rneu
s—
Spha
enor
hync
hus
dori
sae
(Goi
n,19
57)
—Sp
haen
orhy
nchu
sdo
risa
e—
Spha
enor
hync
hus
lact
eus
(Dau
din,
1801
)—
Spha
enor
hync
hus
lact
eus
—Sp
haen
orhy
nchu
sor
ophi
lus
(A.
Lut
zan
dB
.L
utz,
1938
)—
Spha
enor
hync
hus
orop
hilu
s—
Spha
enor
hync
hus
palu
stri
sB
oker
man
n,19
66—
Spha
enor
hync
hus
palu
stri
s—
Spha
enor
hync
hus
paul
oalv
ini
Bok
erm
ann,
1973
—Sp
haen
orhy
nchu
spa
uloa
lvin
i—
Spha
enor
hync
hus
plan
icol
a(A
.L
utz
and
B.
Lut
z,19
38)
—Sp
haen
orhy
nchu
spl
anic
ola
—
Spha
enor
hync
hus
plat
ycep
halu
s(W
erne
r,18
94)
—Sp
haen
orhy
nchu
spl
atyc
epha
lus
—Sp
haen
orhy
nchu
spr
asin
usB
oker
man
n,19
73—
Spha
enor
hync
hus
pras
inus
—Sp
haen
orhy
nchu
ssu
rdus
(Coc
hran
,19
53)
—Sp
haen
orhy
nchu
ssu
rdus
—T
epui
hyla
aeci
i(A
yarz
ague
na,
Sen
aris
,an
dG
orzu
la,
1993
)—
Tep
uihy
laae
cii
—
Tep
uihy
lace
lsae
Mij
ares
-Urr
utia
,M
anza
nill
a-P
uppo
,an
dL
aM
arca
,19
99—
Tep
uihy
lace
lsae
—
Tep
uihy
laed
elca
e(A
yarz
ague
na,
Sen
aris
,an
dG
orzu
la,
1993
)—
Tep
uihy
laed
elca
e—
Tep
uihy
laga
lani
(Aya
rzag
uena
,S
enar
is,
and
Gor
zula
,19
93)
—T
epui
hyla
gala
ni—
Tep
uihy
lalu
teol
abri
s(A
yarz
ague
na,
Sen
aris
,an
dG
orzu
la,
1993
)—
Tep
uihy
lalu
teol
abri
s—
Tep
uihy
lari
mar
um(A
yarz
ague
na,
Sen
aris
,an
dG
orzu
la,
1993
)—
Tep
uihy
lari
mar
um—
172 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Tep
uihy
laro
drig
uezi
(Riv
ero,
1968
)—
Tep
uihy
laro
drig
uezi
—T
epui
hyla
talb
erga
eD
uell
man
and
Yos
hpa,
1996
—T
epui
hyla
talb
erga
e—
Tra
chyc
epha
lus
atla
sB
oker
man
n,19
66—
Tra
chyc
epha
lus
atla
s—
Tra
chyc
epha
lus
jord
ani
(Ste
jneg
eran
dT
est,
1891
)—
Tra
chyc
epha
lus
jord
ani
—T
rach
ycep
halu
sni
grom
acul
atus
Tsc
hudi
,18
38—
Tra
chyc
epha
lus
nigr
omac
ulat
us—
Tri
prio
npe
tasa
tus
(Cop
e,18
65)
—T
ripr
ion
peta
satu
s—
Tri
prio
nsp
atul
atus
Gun
ther
,18
82—
Tri
prio
nsp
atul
atus
—X
enoh
yla
euge
nioi
Car
amas
chi,
1998
—X
enoh
yla
euge
nioi
—X
enoh
yla
trun
cata
(Ize
ckso
hn,
1959
)—
Xen
ohyl
atr
unca
ta—
Phy
llom
edus
inae
Aga
lych
nis
anna
e(D
uell
man
,19
63)
—A
galy
chni
san
nae
—A
galy
chni
sca
lcar
ifer
Bou
leng
er,
1902
—C
ruzi
ohyl
aca
lcar
ifer
—A
galy
chni
sca
llid
ryas
(Cop
e,18
62)
—A
galy
chni
sca
llid
ryas
—A
galy
chni
scr
aspe
dopu
s(F
unkh
ouse
r,19
57)
—C
ruzi
ohyl
acr
aspe
dopu
s—
Aga
lych
nis
lito
drya
s(D
uell
man
and
Tru
eb,
1967
)—
Aga
lych
nis
lito
drya
s—
Aga
lych
nis
mor
elet
ii(D
umer
il,
1853
)—
Aga
lych
nis
mor
elet
ii—
Aga
lych
nis
salt
ator
Tay
lor,
1955
—A
galy
chni
ssa
ltat
or—
Aga
lych
nis
spur
rell
iB
oule
nger
,19
14‘‘
1913
’’—
Aga
lych
nis
spur
rell
i—
Hyl
oman
tis
aspe
raP
eter
s,18
73‘‘
1872
’’—
Hyl
oman
tis
aspe
raH
ylom
anti
sas
pera
grou
pH
ylom
anti
sgr
anul
osa
(Cru
z,19
89)
—H
ylom
anti
sgr
anul
osa
Hyl
oman
tis
aspe
ragr
oup
Pac
hym
edus
ada
cnic
olor
(Cop
e,18
64)
—P
achy
med
usa
dacn
icol
or—
Pha
smah
yla
coch
rana
e(B
oker
man
n,19
66)
—P
hasm
ahyl
aco
chra
nae
—P
hasm
ahyl
aex
ilis
(Cru
z,19
80)
—P
hasm
ahyl
aex
ilis
—P
hasm
ahyl
agu
ttat
a(A
.L
utz,
1925
)—
Pha
smah
yla
gutt
ata
—P
hasm
ahyl
aja
ndai
a(B
oker
man
nan
dS
azim
a,19
78)
—P
hasm
ahyl
aja
ndai
a—
Phr
ynom
edus
aap
pend
icul
ata
(A.
Lut
z,19
25)
—P
hryn
omed
usa
appe
ndic
ulat
a—
Phr
ynom
edus
abo
kerm
anni
Cru
z,19
91—
Phr
ynom
edus
abo
kerm
anni
—P
hryn
omed
usa
fimbr
iata
Mir
anda
-Rib
eiro
,19
23—
Phr
ynom
edus
afim
bria
ta—
Phr
ynom
edus
am
argi
nata
(Ize
ckso
hnan
dC
ruz,
1976
)—
Phr
ynom
edus
am
argi
nata
—
Phr
ynom
edus
ava
nzol
inii
Cru
z,19
91—
Phr
ynom
edus
ava
nzol
inii
—P
hyll
omed
usa
atel
opoi
des
Due
llm
an,
Cad
le,
and
Can
nate
lla,
1988
Una
ssig
ned
Phy
llom
edus
aat
elop
oide
sU
nass
igne
d
Phy
llom
edus
aay
eaye
(B.
Lut
z,19
66)
P.
hypo
chon
dria
lis
grou
pP
hyll
omed
usa
ayea
yeP
.hy
poch
ondr
iali
sgr
oup
2005 173FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
1(C
onti
nued
)
Cur
rent
taxo
nom
yS
peci
esgr
oup
incu
rren
tta
xono
my
New
taxo
nom
yS
peci
esgr
oup
inne
wta
xono
my
Phy
llom
edus
aba
ltea
Due
llm
anan
dT
oft,
1979
P.
peri
neso
sgr
oup
Phy
llom
edus
aba
ltea
P.
peri
neso
sgr
oup
Phy
llom
edus
abi
colo
r(B
odda
ert,
1772
)U
nass
igne
dP
hyll
omed
usa
bico
lor
Una
ssig
ned
Phy
llom
edus
abo
livi
ana
Bou
leng
er,
1902
P.
tars
ius
grou
pP
hyll
omed
usa
boli
vian
aP
.ta
rsiu
sgr
oup
Phy
llom
edus
abu
ckle
yiB
oule
nger
,18
82P
.bu
ckle
yigr
oup
Hyl
oman
tis
buck
leyi
Hyl
oman
tis
buck
leyi
grou
pP
hyll
omed
usa
burm
eist
eri
Bou
leng
er,
1882
P.
burm
eist
eri
grou
pP
hyll
omed
usa
burm
eist
eri
P.
burm
eist
eri
grou
pP
hyll
omed
usa
cam
baD
eL
aR
iva,
2000
P.
tars
ius
grou
pP
hyll
omed
usa
cam
baP
.ta
rsiu
sgr
oup
Phy
llom
edus
ace
ntra
lis
Bok
erm
ann,
1965
P.
hypo
chon
dria
lis
grou
pP
hyll
omed
usa
cent
rali
sP
.hy
poch
ondr
iali
sgr
oup
Phy
llom
edus
aco
eles
tis
(Cop
e,18
74)
Una
ssig
ned
Phy
llom
edus
aco
eles
tis
Una
ssig
ned
Phy
llom
edus
ada
niel
iR
uiz-
Car
ranz
a,H
erna
ndez
-C
amac
ho,
and
Rue
da-A
lmon
acid
,19
88P
.bu
ckle
yigr
oup
Hyl
oman
tis
dani
eli
Hyl
oman
tis
buck
leyi
grou
p
Phy
llom
edus
adi
stin
cta
B.
Lut
z,19
50P
.bu
rmei
ster
igr
oup
Phy
llom
edus
adi
stin
cta
P.
burm
eist
eri
grou
pP
hyll
omed
usa
duel
lman
iC
anna
tell
a,19
82P
.pe
rine
sos
grou
pP
hyll
omed
usa
duel
lman
iP
.pe
rine
sos
grou
pP
hyll
omed
usa
ecua
tori
ana
Can
nate
lla,
1982
P.
peri
neso
sgr
oup
Phy
llom
edus
aec
uato
rian
aP
.pe
rine
sos
grou
pP
hyll
omed
usa
hull
iD
uell
man
and
Men
dels
on,
1995
P.
buck
leyi
grou
pH
ylom
anti
shu
lli
Hyl
oman
tis
buck
leyi
grou
p
Phy
llom
edus
ahy
poch
ondr
iali
s(D
audi
n,18
00)
P.
hypo
chon
dria
lis
grou
pP
hyll
omed
usa
hypo
chon
dria
lis
P.
hypo
chon
dria
lis
grou
pP
hyll
omed
usa
iher
ingi
iB
oule
nger
,18
85P
.bu
rmei
ster
igr
oup
Phy
llom
edus
aih
erin
gii
P.
burm
eist
eri
grou
pP
hyll
omed
usa
lem
urB
oule
nger
,18
82P
.bu
ckle
yigr
oup
Hyl
oman
tis
lem
urH
ylom
anti
sbu
ckle
yigr
oup
Phy
llom
edus
am
egac
epha
la(M
iran
da-R
ibei
ro,
1926
)P
.hy
poch
ondr
iali
sgr
oup
Phy
llom
edus
am
egac
epha
laP
.hy
poch
ondr
iali
sgr
oup
Phy
llom
edus
am
edin
aiF
unkh
ouse
r,19
62P
.bu
ckle
yigr
oup
Hyl
oman
tis
med
inai
Hyl
oman
tis
buck
leyi
grou
pP
hyll
omed
usa
orea
des
Bra
ndao
,20
02P
.hy
poch
ondr
iali
sgr
oup
Phy
llom
edus
aor
eade
sP
.hy
poch
ondr
iali
sgr
oup
Phy
llom
edus
apa
llia
taP
eter
s,18
73‘‘
1872
’’U
nass
igne
dP
hyll
omed
usa
pall
iata
Una
ssig
ned
Phy
llom
edus
ape
rine
sos
Due
llm
an,
1973
P.
peri
neso
sgr
oup
Phy
llom
edus
ape
rine
sos
P.
peri
neso
sgr
oup
Phy
llom
edus
aps
ilop
ygio
nC
anna
tell
a,19
80P
.bu
ckle
yigr
oup
Hyl
oman
tis
psil
opyg
ion
Hyl
oman
tis
buck
leyi
grou
pP
hyll
omed
usa
rohd
eiM
erte
ns,
1926
P.
hypo
chon
dria
lis
grou
pP
hyll
omed
usa
rohd
eiP
.hy
poch
ondr
iali
sgr
oup
Phy
llom
edus
asa
uvag
iiB
oule
nger
,18
82P
.ta
rsiu
sgr
oup
Phy
llom
edus
asa
uvag
iiP
.ta
rsiu
sgr
oup
Phy
llom
edus
ata
rsiu
s(C
ope,
1868
)P
.ta
rsiu
sgr
oup
Phy
llom
edus
ata
rsiu
sP
.ta
rsiu
sgr
oup
Phy
llom
edus
ate
trap
loid
eaP
omba
lan
dH
adda
d,19
92P
.bu
rmei
ster
igr
oup
Phy
llom
edus
ate
trap
loid
eaP
.bu
rmei
ster
igr
oup
Phy
llom
edus
ato
mop
tern
a(C
ope,
1868
)U
nass
igne
dP
hyll
omed
usa
tom
opte
rna
Una
ssig
ned
Phy
llom
edus
atr
init
atis
Mer
tens
,19
26U
nass
igne
dP
hyll
omed
usa
trin
itat
isU
nass
igne
dP
hyll
omed
usa
vail
lant
iiB
oule
nger
,18
82U
nass
igne
dP
hyll
omed
usa
vail
lant
iiU
nass
igne
dP
hyll
omed
usa
venu
sta
Due
llm
anan
dT
rueb
,19
67U
nass
igne
dP
hyll
omed
usa
venu
sta
Una
ssig
ned
174 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 2
LOCALITY DATA AND GENBANK ACCESSIONS
The table on the following pages lists all spec-imens, collection numbers, localities, and Gen-Bank accessions of the sequences included in thisanalysis. The current taxonomy of hylids isused here, as these names were used for theGenBank submission; see appendix 1 for thenew taxonomy. All species followed by an aster-isk (*) correspond to sequences retrieved fromGenBank. In a few cases, the tissues have separatenumbers of official tissue collections. These aregiven as footnotes. The list includes collection ab-breviations for (1) vouchers and or tissues em-ployed in this project, (2) preserved specimens re-ferred to in this paper, and (3) vouchers from se-quences retrieved from GenBank, followed by anasterisk (*), with their locality data taken fromDarst and Cannatella (2004).
Collection abbreviations are as follows: AF,Laboratorio de Citogenetica de Vertebrados, In-stituto de Biociencias, Universidade de Sao Paulo(to be accesioned in MZUSP); AM, AustralianMuseum, Sidney, Australia; AM-CC, AmbroseMonell Cryo Collection; AMNH, American Mu-seum of Natural History, New York; BMNH,British Museum (Natural History), London;CFBH, Collection Celio F.B. Haddad, Universi-dade Estadual Paulista, Rio Claro, Sao Paulo, Bra-zil; CWM, Field number of Charles W. Myers (tobe accessioned in AMNH); DFCH-USFQ, Univ-ersidad San Francisco de Quito, Quito, Ecuador;DLR, field numbers of Ignacio De la Riva (to beaccessioned in the Museo Nacional de CienciasNaturales, Madrid, Spain); IRSNB, Institut Royaldes Sciences Naturelles de Belgique, Bruxelles;ITH, field numbers used in the project Levanta-mento da Fauna de Vertebrados Terrestres do areasob influencia da Linha de Transmissao (LT) Ita-bera-Tijuco Preto III (to be accessioned inMZUSP); IWK, field numbers used by MaureenA. Donnelly (to be accessioned in the Herpeto-logical Collection of the Florida InternationalUniversity); JF, field numbers of Julian Faivovich(to be accessioned in MACN); JPC, field num-bers of Janalee P. Caldwell; KRL, field numbersof Karen Lips; KU, University of Kansas, Mu-seum of Natural History, Lawrence, KS; LSUMZH, tissue collection, Louisiana State UniversityMuseum of Zoology, Baton Rouge, LA; MACN,Museo Argentino de Ciencias Naturales ‘‘Bernar-dino Rivadavia’’, Buenos Aires, Argentina;MAD, field numbers of Maureen Donnelly;
MCN, Museo de Ciencias Naturales de la Univ-ersidad Nacional de Salta, Salta, Argentina; MCP,Museu de Ciencias e Tecnologia da PontificiaUniversidade Catolica de Rio Grande do Sul, Bra-zil; MCZ, Museum of Comparative Zoology,Harvard University, Cambridge, MA; MJH, fieldnumbers of Martin J. Henzl; MLP-A, Museo deLa Plata, La Plata, Argentina; MLP-DB, Collec-tion Diego Baldo, at MLP; MNCN ADN, collec-tion of DNA samples of the Museo Nacional deCiencias Naturales, Madrid, Spain; MNHN. Mu-seum National d’Histoire Naturelle, Paris; MNK,Museo ‘‘Noel Kempf Mercado’’, Santa Cruz, Bo-livia; MRT, field numbers of Miguel Trefaut Ro-drigues (to be accesioned in MZUSP); MVZ, Mu-seum of Vertebrate Zoology, University of Cali-fornia, Berkeley, CA; MVZFC, MVZ frozen tis-sue collection; MZFC, Museo de Zoologıa de laFacultad de Ciencias, Universidad Nacional Au-tonoma de Mexico; MZUSP, Museu de Zoologia,Universidade de Sao Paulo, Sao Paulo, Brazil;NHMG, Naturhistoriska Museet, Goteborg, Swe-den; NMP6V, National Museum, Prague, CzechRepublic; QCAZ, Museo de Zoologıa de la Pon-tificia Universidad Catolica del Ecuador; QULC,Queen’s University Laboratory Collection, King-ston, Canada; RdS, field numbers of Rafael de Sa;RNF, field numbers of Robert N. Fisher; ROM,Royal Ontario Museum, Toronto, Canada; RWM,field numbers of Roy W. McDiarmid (specimensto be accessioned in USNM); SAMA, South Aus-tralia Museum, Adelaide, South Australia; SIUC-H, Department of Zoology and Center for Sys-tematic Biology, Southern Illinois University atCarbondale, IL; TMSA, Transvaal Museum, Pre-toria, South Africa; TNHC, Texas Memorial Mu-seum, Texas Natural History Collection, Austin,TX; UMMZ, University of Michigan, Museum ofZoology, Ann Arbor, MI; USNM, National Mu-seum of Natural History, Smithsonian Institution,Washington DC; UTA, University of Texas at Ar-lington, TX; ZFMK-H, tissue collection of theZoologisches Forschungsinstitut und Museum Al-exander Koenig, Bonn, Germany; ZMB, Zoolo-gisches Museum–Universitat Humboldt, Berlin;ZSM, Zoologisches Staatssammlung, Munich,Germany; ZUEC, Museu de Historia Natural,Universidade de Campinas, Campinas, Sao Paulo,Brazil; ZUFRJ, Departamento de Zoologia, Insti-tuto de Biologia, Universidade Federal do Rio deJaneiro, Rio de Janeiro, Brazil.
2005 175FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
LIS
TO
FS
PE
CIM
EN
S,
CO
LL
EC
TIO
NN
UM
BE
RS,
LO
CA
LIT
IES,
AN
DG
EN
BA
NK
AC
CE
SS
ION
SO
FS
EQ
UE
NC
ES
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
LS
UM
ZH
-216
4A
cris
crep
itan
sA
Y84
3559
AY
8437
82A
Y84
4533
AY
8443
58A
Y84
4019
AY
8447
62A
Y84
4194
US
A:
Ala
bam
a:D
eK
alb
Co.
:po
wer
line
acce
ss,
0.1
mi
Wof
Loo
kout
Mt.
Boy
sC
amp
Rd.
AM
NH
A-1
6842
2A
cris
gryl
lus
AY
8435
60A
Y84
3783
AY
8445
34A
Y84
4359
AY
8440
20A
Y84
4763
—U
SA
:F
lori
da:
Oka
loos
aC
o.:
Egl
inA
FB
,J.
R.
Wal
ton
Pon
d.30
841.
599N
,86
828.
369W
EN
S10
039
Ano
thec
asp
inos
aA
Y84
3566
AY
8437
88A
Y84
4540
AY
8443
63A
Y84
4022
AY
8447
68A
Y84
4198
Mex
ico:
Oax
aca:
Ixtl
ande
Juar
ez:
San
tiag
oC
omal
tepe
c,V
ista
Her
mos
aC
FB
H27
15A
para
sphe
nodo
nbr
unoi
AY
8435
67A
Y84
3789
AY
8445
41A
Y84
4364
AY
8440
23A
Y84
4769
AY
8441
99B
razi
l:E
spır
ito
San
to:
Ara
cruz
CF
BH
3001
Apl
asto
disc
usco
chra
nae
AY
8435
68A
Y84
3790
AY
8445
42A
Y84
4365
AY
8440
24A
Y84
4770
AY
8442
00B
razi
l:S
anta
Cat
arin
a:R
anch
oQ
ueim
ado
MA
CN
3779
1A
plas
todi
scus
perv
irid
isA
Y84
3569
AY
8437
91A
Y84
4543
AY
8443
66A
Y84
4025
AY
8447
71A
Y84
4201
Arg
enti
na:
Mis
ione
s:G
uara
ni:
San
Vic
ente
:C
ampo
Ane
xoIN
TA
‘‘C
uart
elR
ioV
icto
ria’
’M
AC
N38
644
Arg
ente
ohyl
asi
emer
siA
Y84
3570
AY
8437
92A
Y84
4544
AY
8443
67A
Y84
4026
AY
8447
72A
Y84
4202
Arg
enti
na:
Cor
rien
tes:
Bel
laV
ista
:In
ters
ecci
onR
.N.
12y
Rio
San
taL
ucia
CF
BH
2968
Cor
ytho
man
tis
gree
ning
iA
Y84
3578
AY
8438
00A
Y84
4551
AY
8443
74A
Y84
4030
AY
8447
79A
Y84
4209
Bra
zil:
Ala
goas
:R
epre
sade
Xin
go,
Pir
anha
sM
VZ
2071
93a
Due
llm
anoh
yla
rufio
culi
sA
Y54
9315
AY
5493
68A
Y84
4556
AY
8443
77A
Y84
4033
AY
8447
82A
Y84
4212
Cos
taR
ica:
Gua
naca
ste:
Vol
can
Cac
aoU
TA
A-5
0812
Due
llm
anoh
yla
sora
lia
AY
8435
84A
Y84
3806
AY
8445
57A
Y84
4378
AY
8440
34A
Y84
4783
—G
uate
mal
a:Iz
abal
:M
oral
es,
Sie
rra
deC
aral
,F
inca
Que
brad
as–C
erro
Poz
ode
Agu
aC
FB
H59
15H
yla
sp.
1(a
f.H
.eh
rhar
dti)
AY
8436
69A
Y84
3913
AY
8446
60A
Y84
4456
—A
Y84
4875
—B
razi
l:S
aoP
aulo
:U
batu
ba(P
icin
guab
a)
176 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
US
NM
3030
22H
yla
aril
dae
AY
8436
04A
Y84
3825
AY
8445
78A
Y84
4392
AY
8440
49A
Y84
4803
AY
8442
23B
razi
l:S
aoP
aulo
:N
ear
Sal
esop
olis
,E
stac
oB
iolo
gica
deB
orac
eia
AF
0068
Hyl
aw
eygo
ldti
AY
8436
85A
Y84
3931
AY
8446
78A
Y84
4467
—A
Y84
4887
—B
razi
l:E
spır
ito
San
to:
Dom
ingo
sM
arti
ns:
Sao
Pau
lodo
Ara
caU
SN
M28
4519
Hyl
aal
bom
argi
nata
AY
5493
16A
Y54
9369
AY
8445
68A
Y84
4384
—A
Y84
4794
AY
8442
18B
razi
l:P
erna
mbu
co:
Nea
rC
arau
rucu
,on
way
toS
erra
dos
Cav
alos
KU
2027
34H
yla
pell
ucen
s*A
Y32
6058
——
——
——
Ecu
ador
:P
ichi
ncha
:1.
8km
SS
ES
anJu
an,
3420
mK
RL
798
Hyl
aru
fitel
aA
Y84
3662
AY
8439
05A
Y84
4652
—A
Y84
4105
AY
8448
67A
Y84
4282
Pan
ama:
Coc
le:
El
Cop
e:P
arqu
eN
acio
nal
‘‘O
mar
Tor
rijo
s’’
CF
BH
3184
Hyl
aal
bosi
gnat
aA
Y84
3596
AY
8438
17A
Y84
4570
AY
8443
85A
Y84
4042
AY
8447
96A
Y84
4219
Bra
zil:
San
taC
atar
ina:
Sao
Ben
todo
Sul
(Rio
Ver
mel
ho)
CF
BH
3909
Hyl
aca
llip
ygia
AY
8436
14A
Y84
3840
AY
8445
92A
Y84
4402
AY
8440
58A
Y84
4813
AY
8442
36B
razi
l:M
inas
Ger
ais:
Mon
teV
erde
AF
0070
Hyl
aca
vico
laA
Y84
3617
AY
8438
43A
Y84
4594
AY
8444
05—
AY
8448
14—
Bra
zil:
Esp
ırit
oS
anto
:D
omin
gos
Mar
tins
US
NM
3030
38H
yla
leuc
opyg
iaA
Y84
3638
AY
8438
73A
Y84
4622
AY
8444
25A
Y84
4084
AY
8448
40A
Y84
4261
Bra
zil:
Sao
Pau
lo:
Nea
rS
ales
opol
is,
Est
acao
Bio
logi
cade
Bor
acei
aZ
UE
C12
053
Hyl
aal
bopu
ncta
taA
Y54
9317
AY
5493
70A
Y84
4569
—A
Y84
4041
AY
8447
95—
Bra
zil:
Sao
Pau
lo:
Cam
pina
sM
JH56
4H
yla
lanc
ifor
mis
AY
8436
36A
Y84
3870
AY
8446
19—
AY
8440
81A
Y84
4837
AY
8442
58P
eru:
Lor
eto:
Alp
ahua
yoA
MN
HA
-141
040b
Hyl
am
ulti
fasc
iata
AY
8436
48A
Y84
3887
AY
8446
33A
Y84
4436
AY
8440
93A
Y84
4851
AY
8442
70G
uyan
a:D
emer
ara:
Cei
baS
tati
on,
Mad
ewin
iR
iver
,ca
.3
mi
(lin
ear)
ET
imeh
riA
irpo
rt
2005 177FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MA
CN
3779
5H
yla
rani
ceps
AY
8436
57A
Y84
3900
AY
8446
46—
AY
8441
03A
Y84
4863
—A
rgen
tina
:S
anta
Fe:
Ver
a:E
a.‘‘
Las
Gam
as’’
RO
M40
304
Hyl
aan
nect
ans
AY
8436
00A
Y84
3821
AY
8445
74A
Y84
4388
AY
8440
45A
Y84
4800
—V
ietn
am:
Lao
Cai
:S
aP
aN
/AH
yla
arbo
rea
AY
8436
01A
Y84
3822
AY
8445
75A
Y84
4389
AY
8440
46—
AY
8442
21P
ettr
ade
LS
UM
ZH
-230
Hyl
aja
poni
caA
Y84
3633
AY
8438
66A
Y84
4615
AY
8444
20A
Y84
4078
AY
8448
33A
Y84
4255
Japa
n:H
iros
him
aP
refe
ctur
e:H
iros
him
aci
tyY
asuf
utui
chi-
cho,
Ait
aN
/Ac
Hyl
asa
vign
yiA
Y84
3665
AY
8439
07A
Y84
4654
—A
Y84
4107
—A
Y84
4284
Yem
en:
Yar
im(2
813
m)
AM
NH
A-1
6516
3H
yla
arm
ata
AY
5493
21A
Y54
9374
AY
8445
79A
Y84
4393
AY
8440
50A
Y84
4804
AY
8442
24B
oliv
ia:
San
taC
ruz:
Cab
alle
ro:
Can
ton
San
Juan
:A
mbo
roN
atio
nal
Par
kA
MN
H-A
1651
32d
Hyl
ach
araz
ani
AY
8436
18A
Y84
3844
AY
8445
95A
Y84
4406
AY
8440
61—
AY
8442
39B
oliv
ia:
La
Paz
:B
auti
sta
Saa
vedr
a:C
anto
nC
hara
zani
,st
ream
2R
WM
1768
8H
yla
inpa
rque
siA
Y84
3672
—A
Y84
4663
—A
Y84
4114
AY
8448
76A
Y84
4291
Ven
ezue
la:
Am
azon
as:
Cer
rode
laN
ebli
naJA
C22
650
Hyl
abi
stin
cta
AY
8436
09A
Y84
3834
AY
8445
87A
Y84
4399
AY
8440
54—
AY
8442
30M
exic
o:O
axac
a:M
po.
Cui
catl
an,
Tut
epet
ongo
,16
19m
JAC
2116
7H
yla
calt
hula
AY
8436
15A
Y84
3841
AY
8445
93A
Y84
4403
AY
8440
59—
AY
8442
37M
exic
o:O
axac
a:C
arre
tera
Coc
onal
es-
Zac
atep
ec,
1360
mR
WM
1774
6H
yla
boan
sA
Y84
3610
AY
8438
35A
Y84
4588
—A
Y84
4055
AY
8448
09A
Y84
4231
Ven
ezue
la:
Am
azon
as:
Can
oA
gua
Bla
nca:
3.5
kmS
EN
ebli
naba
seca
mp
onR
ioB
aria
CF
BH
2966
Hyl
acr
epit
ans
AY
8436
21A
Y84
3850
AY
8446
01A
Y84
4412
AY
8440
67—
—B
razi
l:A
lago
as:
Mun
icıp
iode
Pir
anha
s:R
epre
sade
Xin
go
178 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MA
CN
3700
0H
yla
fabe
rA
Y54
9334
AY
5493
87A
Y84
4607
——
AY
8448
25—
Arg
enti
na:
Mis
ione
s:G
uara
ni:
San
Vic
ente
:C
ampo
Ane
xoIN
TA
‘‘C
uart
elR
ioV
icto
ria’
’C
FB
H40
00H
yla
lund
iiA
Y84
3639
AY
8438
74A
Y84
4623
—A
Y84
4085
AY
8448
41A
Y84
4262
Bra
zil:
Sao
Pau
lo:
Rio
Cla
ro(I
tape
)U
SN
M30
3046
Hyl
apa
rdal
isA
Y84
3651
AY
8438
91A
Y84
4637
—A
Y84
4096
AY
8448
55—
Bra
zil:
Sao
Pau
lo:
near
Sal
esop
olis
,E
stac
aoB
iolo
gica
deB
orac
eia
SIU
CH
-707
9H
yla
coly
mba
AY
8436
20A
Y84
3848
AY
8445
99A
Y84
4410
AY
8440
65A
Y84
4818
AY
8442
43P
anam
a:C
ocle
:P
arqu
eN
acio
nal
El
Cop
eS
IUC
H-6
924
Hyl
apa
lmer
iA
Y84
3650
AY
8438
90A
Y84
4636
AY
8444
39A
Y84
4095
AY
8448
54A
Y84
4273
Pan
ama:
El
Cop
e:P
arqu
eN
acio
nal
‘‘O
mar
Tor
rijo
s’’
UT
AA
-507
71H
yla
brom
elia
cia
AY
8436
12A
Y84
3837
AY
8445
90A
Y84
4401
AY
8440
56A
Y84
4811
AY
8442
33G
uate
mal
a:H
uehu
eten
ango
:S
ierr
ade
los
Cuc
hum
atan
es:
Fin
caC
hibl
ac(n
owA
ldea
Bue
nos
Air
es)
N/A
(AM
CC
1256
27)
Hyl
aan
ders
onii
AY
8435
98A
Y84
3819
AY
8445
72—
AY
8440
44A
Y84
4798
—U
SA
:F
lori
da:
San
taR
osa
Co.
:C
o.R
d.19
1N
ofM
unso
nM
VZ
1453
85e
Hyl
aci
nere
aA
Y54
9327
AY
5493
80A
Y84
4597
AY
8444
08A
Y84
4063
AY
8448
16A
Y84
4241
US
A:
Tex
as:
Tra
vis
Co.
Aus
tin,
Mun
icip
alG
olf
Cou
rse
AM
NH
A-1
6840
4H
yla
grat
iosa
AY
8436
30A
Y84
3862
AY
8446
11A
Y84
4418
AY
8440
76A
Y84
4829
AY
8442
52U
SA
:F
lori
da:
San
taR
osa
Co.
:C
o.R
d.19
1N
ofM
unso
n.30
853.
289N
,86
853.
269W
AM
NH
A-1
6842
7H
yla
squi
rell
aA
Y84
3678
AY
8439
23A
Y84
4670
AY
8444
62A
Y84
4119
AY
8448
82A
Y84
4295
US
A:
Flo
rida
:A
lach
uaC
o.:
Co.
Rd.
346
1.2
mi
121,
2983
0.11
9N,
8282
5.62
9W
2005 179FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
US
NM
3030
32H
yla
asta
rtea
AY
5493
22A
Y54
9375
AY
8445
80—
——
AY
8442
25B
razi
l:S
aoP
aulo
:ne
arS
ales
opol
is,
Est
acao
Bio
logi
cade
Bor
acei
aC
FB
H36
21H
yla
circ
umda
taA
Y54
9328
AY
5493
81A
Y84
4598
AY
8444
09A
Y84
4064
AY
8448
17A
Y84
4242
Bra
zil:
San
taC
atar
ina:
Sao
Ben
todo
Sul
US
NM
3030
36H
yla
hyla
xA
Y54
9338
AY
5493
91A
Y84
4614
AY
8444
19A
Y84
4077
AY
8448
32A
Y84
4254
Bra
zil:
Sao
Pau
lo:
near
Sal
esop
olis
,E
stac
aoB
iolo
gica
deB
orac
eia
CF
BH
5766
Hyl
asp
.3
AY
8436
73A
Y84
3916
AY
8446
64—
AY
8441
15—
—B
razi
l:R
iode
Jane
iro:
Ang
rado
sR
eis
CF
BH
5917
Hyl
asp
.4
AY
8436
74A
Y84
3917
AY
8446
65A
Y84
4458
AY
8441
16A
Y84
4877
—B
razi
l:S
aoP
aulo
:U
batu
ba:
Pic
ingu
aba
DF
CH
-US
FQ
899
Hyl
aca
rnif
exA
Y84
3616
AY
8438
42—
AY
8444
04A
Y84
4060
—A
Y84
4238
Ecu
ador
:P
ichi
ncha
:T
anda
yapa
CF
BH
5418
Hyl
abe
rtha
lutz
aeA
Y84
3607
AY
8438
31A
Y84
4584
AY
8443
97A
Y84
4052
AY
8448
07A
Y84
4228
Bra
zil:
Rio
deJa
neir
o:D
uque
deC
axia
sU
MM
Z77
55H
yla
aren
icol
orA
Y84
3603
AY
8438
24A
Y84
4577
AY
8443
91A
Y84
4048
AY
8448
02—
US
A:
Ari
zona
:G
ila
Co.
:H
oust
oncr
eek
just
NH
wy
260,
appr
ox.
4m
iE
ofP
ayso
nU
TA
A-5
4763
Hyl
aeu
phor
biac
eaA
Y84
3625
AY
8438
55A
Y84
4606
—A
Y84
4072
AY
8448
23A
Y84
4248
Mex
ico:
Oax
aca:
Car
rete
raM
itla
-Ayu
tla,
1950
mM
ZF
C48
14H
yla
exim
iaA
Y84
3626
AY
8438
56—
—A
Y84
4073
AY
8448
24A
Y84
4249
Mex
ico:
Pue
bla:
Chi
gnah
uapa
n,20
kmS
AM
NH
A-1
6840
6H
yla
wal
keri
AY
8436
84A
Y84
3930
AY
8446
77A
Y84
4466
AY
8441
25—
—P
ettr
ade
NM
P6V
7125
0H
yla
calc
arat
aA
Y84
3613
AY
8438
39—
——
—A
Y84
4235
Per
u:A
ngui
lla,
50km
Wof
Iqui
tos
MA
D44
0H
yla
fasc
iata
AY
5493
35A
Y54
9388
AY
8446
08—
——
—G
uyan
a:Iw
okra
ma:
Cow
fly
cam
p,80
mA
MN
H-A
1410
54f
Hyl
age
ogra
phic
aA
Y84
3628
——
——
——
Guy
ana:
War
niab
oC
reek
,4
mi
(by
rd)
SW
Dub
ulay
Ran
chho
use
RO
M39
582
Hyl
aka
naim
aA
Y84
3634
AY
8438
68A
Y84
4617
AY
8444
22A
Y84
4079
AY
8448
35—
Guy
ana:
Mou
ntA
yang
anna
180 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
NM
P6V
7125
8/1
Hyl
am
icro
derm
aA
Y84
3644
AY
8438
81—
——
—A
Y84
4267
Per
u:A
ngui
lla,
50km
Wof
Iqui
tos
KU
2027
37H
yla
pict
urat
a*A
Y32
6055
——
——
——
Ecu
ador
:P
ichi
ncha
:T
inal
andi
a:15
.5km
SE
San
toD
omin
gode
los
Col
orad
os,
700
mR
OM
3962
4H
yla
rora
ima
AY
8436
60A
Y84
3903
AY
8446
50A
Y84
4448
AY
8441
04A
Y84
4866
AY
8442
80G
uyan
a:M
ount
Aya
ngan
naC
FB
H54
24H
yla
sem
ilin
eata
AY
8437
78A
Y84
3779
AY
8439
09A
Y84
4656
AY
8444
53A
Y84
4108
AY
8448
71A
Y84
4286
Bra
zil:
Rio
deJa
neir
o:D
uque
deC
axia
sR
dS60
6H
yla
pict
aA
Y84
3654
AY
8438
94A
Y84
4640
AY
8444
42A
Y84
4099
AY
8448
58A
Y84
4276
Bel
ize:
Sta
nnC
reek
Dis
tric
t:C
oksc
omb
Bas
inW
ildl
ife
San
tuar
yU
TA
A-5
4773
Hyl
asm
ithi
iA
Y84
3668
AY
8439
12A
Y84
4659
—A
Y84
4111
AY
8448
74—
Mex
ico:
Oax
aca:
Sie
rra
Mad
rede
lS
ur;
Car
rete
raP
ochu
tla,
288
mM
AD
085
Hyl
agr
anos
aA
Y54
9336
AY
8438
61A
Y84
4610
——
AY
8448
28—
Guy
ana:
Iwok
ram
a:M
uri
Scr
ubca
mp,
80m
RO
M39
570
Hyl
ale
mai
AY
8436
37A
Y84
3871
AY
8446
20A
Y84
4423
AY
8440
82A
Y84
4838
AY
8442
59G
uyan
a:M
ount
Aya
ngan
naR
OM
3956
1H
yla
sibl
eszi
AY
8436
67A
Y84
3911
AY
8446
58A
Y84
4455
AY
8441
10A
Y84
4873
AY
8442
88G
uyan
a:M
ount
Aya
ngan
naQ
UL
C97
005
Hyl
ala
bial
isA
Y84
3635
AY
8438
69A
Y84
4618
—A
Y84
4080
AY
8448
36A
Y84
4257
Col
ombi
a:P
arqu
eN
atur
alN
acio
nal
Chi
ngaz
aK
U20
2760
Hyl
apa
cha*
AY
3260
57—
——
——
—E
cuad
or:
Azu
ay:
2km
SS
EP
alm
as,
2340
mK
U20
2732
Hyl
apa
ntos
tict
a*A
Y32
6052
——
——
——
Ecu
ador
:N
apo:
18km
ES
anta
Bar
bara
QC
AZ
1670
4H
yla
tapi
chal
aca
AY
5636
25A
Y84
3925
AY
8446
72—
AY
8441
21—
AY
8442
97E
cuad
or:
Zam
ora-
Chi
nchi
pe:
Res
erva
Tap
icha
laca
:ro
adbe
twee
nY
anga
naan
dV
alla
doli
d
2005 181FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
RdS
790
Hyl
aeb
racc
ata
AY
8436
24A
Y84
3853
AY
8446
04A
Y84
4415
AY
8440
70A
Y84
4822
AY
8442
47B
eliz
e:S
tann
Cre
ekD
istr
ict:
Cok
scom
bB
asin
Wil
dlif
eS
antu
ary
(cap
tive
-ra
ised
tadp
oles
from
adul
tsco
llec
ted
inth
islo
cali
ty)
MJH
7143
Hyl
asa
raya
cuen
sis
AY
8436
64—
—A
Y84
4451
—A
Y84
4869
—P
eru:
Hua
nuco
:R
ioL
lull
apic
his:
Pan
guan
aM
JH38
44H
yla
tria
ngul
umA
Y84
3680
AY
8439
26A
Y84
4673
AY
8444
64A
Y84
4122
—A
Y84
4298
Bra
zil:
Acr
e:L
ago
Cat
alao
:Il
haX
ibor
ena
AF
414
Hyl
am
arti
nsi
AY
8436
41A
Y84
3878
AY
8446
26—
AY
8440
86A
Y84
4844
AY
8442
64B
razi
l:M
inas
Ger
ais:
San
taB
arba
raM
JH71
16H
yla
mar
mor
ata
AY
8436
40A
Y84
3877
—A
Y84
4428
——
—P
eru:
Hua
nuco
:R
ioL
lull
apic
his:
Pan
guan
aC
FB
H57
61H
yla
seni
cula
AY
8436
66A
Y84
3910
AY
8446
57A
Y84
4454
AY
8441
09A
Y84
4872
AY
8442
87B
razi
l:R
iode
Jane
iro:
Ang
rado
sR
eis
MR
T59
46H
yla
bipu
ncta
taA
Y84
3608
AY
8438
32A
Y84
4585
—A
Y84
4053
AY
8448
08A
Y84
4229
Bra
zil:
Bah
ia:
Juss
ari:
Ser
rado
Tei
mos
oU
TA
A-5
0632
Hyl
am
icro
ceph
ala
AY
8436
43A
Y84
3880
AY
8446
28A
Y84
4430
—A
Y84
4846
AY
8442
66H
ondu
ras:
Atl
anti
da:
Cor
dill
era
Nom
bre
deD
ios,
Ald
eaR
ioV
iejo
MA
CN
3778
5H
yla
nana
AY
5493
46A
Y84
3888
AY
8446
34A
Y84
4437
—A
Y84
4852
AY
8442
71A
rgen
tina
:E
ntre
Rio
s:D
epto
.Is
las
del
Ibic
uyM
HZ
462
Hyl
arh
odop
epla
AY
8436
58—
AY
8446
47A
Y84
4446
—A
Y84
4864
—P
eru:
Lor
eto:
Jena
roH
erre
raM
AC
N38
638
Hyl
asa
nbor
niA
Y84
3663
AY
8439
06A
Y84
4653
AY
8444
50A
Y84
4106
AY
8448
68A
Y84
4283
Arg
enti
na:
Ent
reR
ios:
Dep
to.
Isla
sde
lIb
icuy
:R
uta
12vi
eja,
entr
eB
razo
Lar
goy
Arr
oyo
Luc
iano
MJH
129
Hyl
aw
alfo
rdi
AY
8436
83A
Y84
3929
AY
8446
76—
—A
Y84
4886
—B
razi
l:U
nkno
wn
JPC
1077
2gH
yla
miy
atai
AY
8436
47A
Y84
3886
AY
8446
32A
Y84
4435
AY
8440
92A
Y84
4850
—E
cuad
or:
Suc
umbi
os
182 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MA
CN
3379
9H
yla
min
uta
AY
5493
45A
Y54
9398
—A
Y84
4432
AY
8440
89—
—A
rgen
tina
:M
isio
nes:
Gua
rani
:S
anV
icen
te:
Cam
poA
nexo
INT
A‘‘
Cua
rtel
Rio
Vic
tori
a’’
UT
AA
-562
83H
yla
arbo
resc
ande
nsA
Y84
3602
AY
8438
23A
Y84
4576
AY
8443
90A
Y84
4047
AY
8448
01A
Y84
4222
Mex
ico:
Pue
bla:
Sie
rra
Neg
ra,
1833
mU
TA
A-5
4762
Hyl
acy
clad
aA
Y84
3622
AY
8438
51A
Y84
4602
AY
8444
13A
Y84
4068
AY
8448
20A
Y84
4245
Mex
ico:
Oax
aca:
Sie
rra
Mix
e,2.
0m
iW
Tot
onte
pec,
2045
mU
TA
A-5
4766
Hyl
am
elon
omm
aA
Y84
3642
AY
8438
79A
Y84
4627
AY
8444
29A
Y84
4087
AY
8448
45A
Y84
4265
Mex
ico:
Oax
aca:
Sie
rra
Mad
rede
lS
ur;
Car
rete
raP
ochu
tla,
681m
JAC
2243
8H
yla
mio
tym
panu
mA
Y84
3645
AY
8438
84A
Y84
4630
AY
8444
33A
Y84
4090
AY
8448
48—
Mex
ico:
Pue
bla:
Sie
rra
Nor
te,
Cue
tzal
an,
Hot
elV
illa
sC
uetz
alan
,12
50m
UT
AA
-547
21H
yla
perk
insi
AY
8436
53A
Y84
3893
AY
8446
39A
Y84
4441
AY
8440
98A
Y84
4857
AY
8442
75G
uate
mal
a:H
uehu
eten
ango
:B
aril
las,
Ald
eaN
ucaq
uexi
sJA
C21
583
Hyl
am
ixe
AY
8436
46A
Y84
3885
AY
8446
31A
Y84
4434
AY
8440
91A
Y84
4849
AY
8442
69M
exic
o:O
axac
a:S
ierr
aM
ixe;
Mun
.S
anti
ago
Zac
atep
ec:
carr
eter
aZ
acat
epec
-Jes
usC
arra
nza,
28m
ide
entr
onqu
eco
nca
rret
era
aT
oton
tepe
c,80
3m
MJH
7101
Hyl
abr
evif
rons
AY
8436
11A
Y84
3836
AY
8445
89A
Y84
4400
—A
Y84
4810
AY
8442
32P
eru:
Hua
nuco
:R
ioL
lull
apic
his:
angu
ana
CF
BH
S/N
Hyl
agi
esle
riA
Y84
3629
AY
8438
60—
AY
8444
17A
Y84
4075
AY
8448
27A
Y84
4251
Bra
zil:
Sao
Pau
lo:
Uba
tuba
:P
icin
guab
a
2005 183FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
AM
NH
A-1
3931
5H
yla
parv
icep
sA
Y84
3652
AY
8438
92A
Y84
4638
AY
8444
40A
Y84
4097
AY
8448
56A
Y84
4274
Bra
zil:
Acr
e:C
entr
oE
xper
imen
tal
daU
nive
rsid
ade
doA
cre
atkm
23on
Rio
Bra
nco-
Por
toV
elho
Roa
dM
VZ
1497
50h
Hyl
ari
vula
ris
AY
8436
59A
Y84
3902
AY
8446
49—
AY
8441
17—
—C
osta
Ric
a:H
ered
ia:
‘‘C
hom
pipe
’’vi
cini
tyof
Vol
can
Ban
baJA
C22
24H
yla
sp.
5(a
ff.
H.
thor
ecte
s)A
Y84
3675
AY
8439
18A
Y84
4666
AY
8444
59A
Y84
4118
AY
8448
78—
Mex
ico:
Gue
rrer
o:C
arre
tera
Nue
vaD
elhi
-L
aG
uita
rra,
1900
mC
FB
H56
42H
yla
sp.
6(a
ff.
H.
pseu
dops
eudi
s)A
Y84
3676
AY
8439
19A
Y84
4667
AY
8444
60A
Y84
4101
AY
8448
79A
Y84
4292
Bra
zil:
Bah
ia:
Mun
icıp
iode
Len
cois
:R
ioG
risa
nte,
Ser
rado
Sin
cora
,C
hapa
daD
iam
anti
naM
VZ
1497
64H
yla
pseu
dopu
ma
AY
8436
56A
Y84
3897
AY
8446
43A
Y84
4444
AY
8440
75A
Y84
4861
AY
8442
77C
osta
Ric
a:P
unta
Are
nas:
Mon
teve
rde
ML
PA21
38H
yla
andi
naA
Y54
9319
AY
5493
72A
Y84
4573
AY
8443
87—
AY
8447
99—
Arg
enti
na:
Tuc
uman
:T
afide
lV
alle
DL
R41
19H
yla
balz
ani
AY
5493
23A
Y54
9376
AY
8445
82A
Y84
4395
—A
Y84
4806
AY
8442
26B
oliv
ia:
Dep
to.
La
Paz
:P
rov.
Nor
yung
as:
Ser
rani
aB
ella
vist
aC
FB
H33
56H
yla
bisc
hoffi
AY
5493
24A
Y54
9377
AY
8445
86A
Y84
4398
——
—B
razi
l:S
anta
Cat
arin
a:R
anch
oQ
ueim
ado
ML
P-D
B10
84H
yla
cain
gua
AY
5493
26A
Y54
9379
AY
8445
91—
AY
8440
57A
Y84
4812
AY
8442
34A
rgen
tina
:M
isio
nes:
Pos
adas
MA
CN
3769
2H
yla
cord
obae
AY
5493
30A
Y54
9383
AY
8446
00A
Y84
4411
AY
8440
66A
Y84
4819
AY
8442
44A
rgen
tina
:S
anL
uis:
Dep
to.
Cha
cabu
co:
Vil
laE
lena
:A
rroy
o‘‘
La
cale
ra’’
MA
CN
3779
4H
yla
sp.
7(a
ff.
H.
sem
igut
tata
)A
Y54
9359
AY
5494
12—
——
AY
8448
80—
Arg
enti
na:
Mis
ione
s:S
anV
icen
te:
Cam
poA
nexo
INT
A‘‘
Cua
rtel
Rio
Vic
tori
a’’
184 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MZ
US
P11
1556
Hyl
ala
tist
riat
aA
Y54
9360
AY
5494
13A
Y84
4668
——
—A
Y84
4293
Bra
zil:
Min
asG
erai
s:M
unic
ıpio
Itam
onte
sC
FB
H35
99H
yla
eric
aeA
Y54
9332
AY
5493
85A
Y84
4605
AY
8444
16A
Y84
4071
——
Bra
zil:
Goi
as:
Alt
oP
arai
sode
Goi
asC
FB
H33
86H
yla
guen
ther
iA
Y84
3631
AY
5493
90A
Y84
4612
——
AY
8448
30A
Y84
4253
Bra
zil:
Rio
Gra
nde
doS
ul:
Ter
rade
Are
iaC
FB
H36
25H
yla
joaq
uini
AY
5493
39A
Y54
9392
AY
8446
16A
Y84
4421
—A
Y84
4834
AY
8442
56B
razi
l:S
anta
Cat
arin
a:U
rubi
ciC
FB
H38
48H
yla
lept
olin
eata
AY
5493
41A
Y54
9394
AY
8446
21A
Y84
4424
AY
8440
83A
Y84
4839
AY
8442
60B
razi
l:S
anta
Cat
arin
a:M
unic
ipio
deS
aoD
omin
gos
CF
BH
3098
Hyl
am
argi
nata
AY
5493
42A
Y54
9395
AY
8446
24A
Y84
4426
—A
Y84
4842
AY
8442
63B
razi
l:R
ioG
rand
edo
Sul
:S
aoF
ranc
iso
deP
aula
MV
0249
Hyl
am
aria
nita
eA
Y54
9344
AY
5493
97A
Y84
4625
AY
8444
27—
AY
8448
43—
Arg
enti
na:
Sal
ta:
Bar
itu
CF
BH
5752
Hyl
apo
lyta
enia
AY
8436
55A
Y84
3895
AY
8446
41A
Y84
4443
—A
Y84
4859
—B
razi
l:R
iode
Jane
iro:
Itat
iaia
:M
arin
gaC
FB
H33
88H
yla
pras
ina
AY
5493
47A
Y54
9400
AY
8446
42—
AY
8441
00A
Y84
4860
—B
razi
l:S
anta
Cat
arin
a:R
ioV
erm
elho
MA
CN
3778
8H
yla
pulc
hell
aA
Y54
9352
AY
5494
05A
Y84
4644
AY
8444
45A
Y84
4102
AY
8448
62A
Y84
4278
Arg
enti
na:
Bue
nos
Air
es:
Car
ilo
MA
CN
3750
9H
yla
rioj
ana
AY
5493
55A
Y54
9408
AY
8446
48A
Y84
4447
—A
Y84
4865
AY
8442
79A
rgen
tina
:L
aR
ioja
:S
anog
asta
:E
lH
uaco
deA
rrib
aC
FB
H35
79H
yla
sem
igut
tata
AY
5493
57A
Y54
9410
AY
8446
55A
Y84
4452
—A
Y84
4870
AY
8442
85B
razi
l:P
aran
a:P
iraq
uara
MA
CN
3779
2H
yla
punc
tata
AY
5493
53A
Y54
9406
AY
8446
45—
——
—A
rgen
tina
:C
haco
:R
esis
tenc
ia:
Cam
ino
aIs
lade
lC
erri
to,
altu
raP
eaje
Gra
l.B
elgr
ano
IT-H
0653
Hyl
aru
bicu
ndul
aA
Y84
3661
AY
8439
04A
Y84
4651
AY
8444
49—
—A
Y84
4281
Bra
zil:
Sao
Pau
lo:
Bur
iJA
C21
736
Hyl
ach
imal
apa
AY
8436
19A
Y84
3845
AY
8445
96A
Y84
4407
AY
8440
62A
Y84
4815
AY
8442
40M
exic
o:C
hiap
as:
Col
onia
Rod
ulfo
Fig
uero
a,E
lC
arri
zal,
1475
m
2005 185FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
JAC
2237
1H
yla
xera
AY
8436
86A
Y84
3932
AY
8446
79A
Y84
4468
AY
8441
26A
Y84
4888
AY
8443
00M
exic
o:P
uebl
a:C
arre
tera
Tex
cala
-Z
apot
itla
nS
alin
as,
5227
ftJA
C21
531
Hyl
ane
phil
aA
Y84
3649
AY
8438
89A
Y84
4635
AY
8444
38A
Y84
4094
AY
8448
53A
Y84
4272
Mex
ico:
Oax
aca:
Col
onia
Rod
ulfo
Fig
uero
a,E
lC
arri
zal,
1475
mJA
C24
443
Hyl
ata
enio
pus
AY
8436
79A
Y84
3924
AY
8446
71A
Y84
4463
AY
8441
20A
Y84
4883
AY
8442
96M
exic
o:P
uebl
a:S
ierr
aN
orte
,C
uetz
alan
,H
otel
Vil
las
Cue
tzal
an,
1250
mU
TA
A-5
1838
Hyl
ade
ndro
phas
ma
AY
8436
23A
Y84
3852
AY
8446
03A
Y84
4414
AY
8440
69A
Y84
4821
AY
8442
46G
uate
mal
a:H
uehu
eten
ango
:F
inca
San
Fra
ncis
co,
near
Ald
eaY
alam
bojo
chS
IUC
H-0
6998
Hyl
am
ilia
ria
AY
8437
76A
Y84
3777
AY
8438
82A
Y84
4629
AY
8444
31A
Y84
4088
AY
8448
47A
Y84
4268
Pan
ama:
Coc
le:
El
Cop
e:P
arqu
eN
acio
nal
Om
arT
orri
jos
CF
BH
5788
Hyl
aur
ugua
yaA
Y84
3681
AY
8439
27A
Y84
4674
—A
Y84
4123
AY
8448
84A
Y84
4299
Bra
zil:
Rio
Gra
nde
doS
ul:
Cam
bara
doS
ulA
MN
HA
-168
423
Hyl
aav
ivoc
aA
Y84
3605
AY
8438
28A
Y84
4581
AY
8443
94A
Y84
4051
AY
8448
05—
US
A:
Flo
rida
:A
lach
uaC
o.:
Loc
hloo
saW
ildl
ife
Mgm
tar
eane
arR
iver
Sty
x.29
831.
259N
,82
813.
049
AM
NH
A-1
6842
5H
yla
fem
oral
isA
Y84
3627
AY
8438
59A
Y84
4609
—A
Y84
4074
AY
8448
26A
Y84
4250
US
A:
Flo
rida
:O
kalo
osa
Co.
:U
SH
wy
90at
Yel
low
Riv
erfl
oodp
lain
UM
FS
5545
Hyl
ave
rsic
olor
AY
8436
82A
Y84
3928
AY
8446
75A
Y84
4465
AY
8441
24A
Y84
4885
—P
ettr
ade
CW
M19
512
Hyl
asp
.8
AY
8436
71A
Y84
3915
AY
8446
62—
AY
8441
13—
AY
8442
90V
enez
uela
:B
oliv
ar:
Cer
roG
uana
y
186 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
CF
BH
5652
Hyl
asp
.9
(aff
.H
.al
vare
ngai
)A
Y84
3677
AY
8439
22A
Y84
4669
AY
8444
61—
AY
8448
81A
Y84
4294
Bra
zil:
Bah
ia:
Mun
icıp
iode
Len
cois
:R
ioG
risa
nte,
Ser
rado
Sin
cora
,C
hapa
daD
iam
anti
naC
FB
H57
97H
yla
ance
psA
Y84
3597
AY
8438
18A
Y84
4571
AY
8443
86A
Y84
4043
AY
8447
97A
Y84
4220
Bra
zil:
Esp
ırit
oS
anto
:L
inha
res
(Pov
oaca
o)U
SN
M30
2435
Hyl
abe
nite
ziA
Y84
3606
AY
8438
30A
Y84
4583
AY
8443
96—
—A
Y84
4227
Bra
zil:
Ror
aim
a:V
illa
Pac
arai
ma,
bord
erm
arke
rB
V(B
razi
l–V
enez
uela
)8
AM
NH
A-1
6840
5H
yla
heil
prin
iA
Y84
3632
AY
8438
64A
Y84
4613
——
AY
8448
31—
Pet
trad
eN
MP
6V71
202/
2H
yla
sp.
2A
Y84
3670
AY
8439
14A
Y84
4661
AY
8444
57A
Y84
4112
—A
Y84
4289
Per
u:50
kmW
ofIq
uito
sN
/Ai
Lys
apsu
sla
evis
AY
8436
96A
Y84
3941
AY
8446
89A
Y84
4476
AY
8441
33A
Y84
4896
AY
8443
05G
uyan
a:S
outh
ern
Rup
unun
iS
avan
ah:
Ais
halt
on(o
nK
uban
awan
Cre
ek)
MA
CN
3864
5L
ysap
sus
lim
ellu
mA
Y84
3697
AY
8439
42A
Y84
4690
AY
8444
77—
AY
8448
97—
Arg
enti
na:
Cor
rien
tes:
Bel
laV
ista
:In
ters
ecci
onR
.N.
12y
Rio
San
taL
ucia
N/A
Nyc
tim
anti
sru
gice
psA
Y84
3780
AY
8437
81A
Y84
3945
——
——
—E
cuad
or:
Nap
o:Ja
tun
Sac
ha(e
ggs
laid
inca
ptiv
ity
from
adul
tsco
llec
ted
ther
e)JP
C13
178j
Ost
eoce
phal
usca
brer
aiA
Y84
3705
AY
8439
50A
Y84
4696
AY
8444
81A
Y84
4136
AY
8449
02A
Y84
4310
Bra
zil:
Acr
e:5
kmN
Por
toW
alte
r,in
land
from
Rio
Juru
aM
AC
N38
643
Ost
eoce
phal
usla
ngsd
orffi
iA
Y84
3706
AY
8439
51A
Y84
4697
AY
8444
82A
Y84
4137
AY
8449
03A
Y84
4311
Arg
enti
na:
Mis
ione
s:G
ener
alB
elgr
ano:
10km
NB
erna
rdo
deIr
igoy
en:
Sal
toA
ndre
sito
2005 187FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
AM
NH
A-1
3125
4O
steo
ceph
alus
lepr
ieur
iiA
Y54
9361
AY
8439
52A
Y84
4698
AY
8444
83A
Y84
4138
AY
8449
04A
Y84
4312
Ven
ezue
la:
Am
azon
as:
Neb
lina
Bas
eC
amp
onR
ioM
awar
inum
a(5
Rio
Bar
ia),
140
mM
NH
N20
01.0
828
Ost
eoce
phal
usoo
phag
usA
Y84
3708
AY
8439
53A
Y84
4699
AY
8444
84A
Y84
4139
——
Fre
nch
Guy
ana:
Kaw
road
:04
842N
/528
18W
AM
NH
-A13
1245
Ost
eoce
phal
usta
urin
usA
Y84
3709
AY
8439
54A
Y84
4700
AY
8444
85A
Y84
4140
AY
8449
05A
Y84
4313
Ven
ezue
la:
Am
azon
as:
Neb
lina
Bas
eC
amp
onR
ioM
awar
inum
a(5
Rio
Bar
ia),
140
MN
/AO
steo
pilu
scr
ucia
lis
AY
8437
10A
Y84
3955
——
——
AY
8443
14Ja
mai
ca:
Man
ches
ter
Par
ish:
Man
devi
lle
(egg
sla
idin
capt
ivit
yfr
omad
ults
coll
ecte
dth
ere)
AM
NH
A-1
6841
0O
steo
pilu
sdo
min
icen
sis
AY
8437
11A
Y84
3956
AY
8447
01A
Y84
4486
AY
8441
41—
AY
8443
15P
ettr
ade
US
NM
3178
30O
steo
pilu
sse
pten
trio
nali
sA
Y84
3712
AY
8439
57—
AY
8444
87A
Y84
4142
AY
8449
06A
Y84
4316
Cub
a:G
uant
anam
o:G
uant
anam
oB
ayA
MN
HA
-168
415
Ost
eopi
lus
vast
usA
Y84
3713
AY
8439
58—
—A
Y84
4143
AY
8449
07A
Y84
4317
Pet
trad
eM
NH
N20
01.0
814
Phr
ynoh
yas
hadr
ocep
sA
Y84
3717
AY
8439
62A
Y84
4704
AY
8444
90A
Y84
4146
—A
Y84
4319
Fre
nch
Guy
ana:
Kaw
road
:04
8429
N,
5281
89W
CF
BH
5780
Phr
ynoh
yas
mes
opha
eaA
Y84
3718
AY
8439
63A
Y84
4705
AY
8444
91A
Y84
4147
AY
8449
10A
Y84
4320
Bra
zil:
Rio
deJa
neir
o:P
arat
iA
MN
H-A
1312
01k
Phr
ynoh
yas
resi
nifr
icti
xA
Y84
3719
AY
8439
64A
Y84
4706
AY
8444
92A
Y84
4148
AY
8449
11A
Y84
4321
Ven
ezue
la:
Am
azon
as:
Neb
lina
base
cam
pon
Rio
Maw
arin
uma,
140
mA
MN
H-A
1411
42P
hryn
ohya
sve
nulo
saA
Y54
9362
AY
5494
15A
Y84
4707
AY
8444
93A
Y84
4149
AY
8449
12A
Y84
4322
Guy
ana:
Dub
ulay
Ran
chon
the
Ber
bice
Riv
er,
200
ftN
/Al
Phy
llod
ytes
lute
olus
AY
8437
21A
Y84
3965
AY
8447
08A
Y84
4494
AY
8441
50A
Y84
4913
AY
8443
24B
razi
l:E
spır
ito
San
to:
Set
iba,
Gua
rapa
riM
RT
6144
Phy
llod
ytes
sp.
AY
8437
22A
Y84
3966
AY
8447
09—
AY
8441
51A
Y84
4914
AY
8443
25B
razi
l:B
ahia
:U
ruci
-Una
188 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MV
Z14
9403
Ple
ctro
hyla
glan
dulo
saA
Y84
3730
AY
8439
67A
Y84
4718
AY
8445
00A
Y84
4159
AY
8449
23A
Y84
4331
Gua
tem
ala:
San
Mar
cos:
ridg
eab
ove
El
Rin
con
UT
AA
-551
40P
lect
rohy
lagu
atem
alen
sis
AY
8437
31A
Y84
3976
AY
8447
19A
Y84
4501
AY
8441
60A
Y84
4924
AY
8443
32G
uate
mal
a:G
uate
mal
a:D
onJu
sto:
San
taR
osal
ia:
km12
.5ca
rret
era
aE
lS
alva
dor
JAC
2170
7P
lect
rohy
lam
atud
aiA
Y84
3732
AY
8439
77A
Y84
4720
AY
8445
02A
Y84
4161
AY
8449
25A
Y84
4333
Mex
ico:
Chi
apas
:ca
min
oC
olon
iaR
odul
foF
igue
roa,
Dia
zO
rdaz
–1.
4m
ide
Rod
ulfo
Fig
uero
aR
NF
2424
Pse
udac
ris
cada
veri
naA
Y84
3734
AY
8439
78A
Y84
4722
—A
Y84
4162
—A
Y84
4334
US
A:
Cal
ifor
nia:
San
Die
goC
o:A
nza
Bor
rego
N/A
Pse
udac
ris
cruc
ifer
AY
8437
35A
Y84
3980
AY
8447
23—
AY
8441
63A
Y84
4927
—P
ettr
ade
AM
NH
A-1
6847
2P
seud
acri
soc
ular
isA
Y84
3736
AY
8439
81A
Y84
4724
—A
Y84
4164
——
US
A:
Flo
rida
:C
olum
bia
Co.
,S
R25
02
mi
WI-
10R
NF
3255
Pse
udac
ris
regi
lla
AY
8437
37A
Y84
3982
AY
8447
25A
Y84
4504
AY
8441
65—
—U
SA
:C
alif
orni
a:S
anD
iego
Co.
:B
owde
nC
anyo
nA
MN
HA
-168
468
Pse
udac
ris
tris
eria
taA
Y84
3738
AY
8439
83A
Y84
4726
—A
Y84
4166
AY
8449
28A
Y84
4335
US
A:
Uta
h:C
ache
:A
mal
gaM
AC
N37
786
Pse
udis
min
uta
AY
8437
39A
Y84
3984
—A
Y84
4505
—A
Y84
4929
AY
8443
36A
rgen
tina
:E
ntre
Rio
s:D
epto
.Is
las
del
Ibic
uy:
Rut
a12
viej
aM
AC
N38
642
Pse
udis
para
doxa
AY
8437
40A
Y84
3985
AY
8447
27A
Y84
4506
AY
8441
67—
AY
8443
37A
rgen
tina
:C
orri
ente
s:D
epto
.B
ella
vist
a:C
amin
oS
anR
oque
-B
ella
vist
aM
VZ
1329
95P
tern
ohyl
afo
dien
sA
Y84
3743
AY
8439
86A
Y84
4730
AY
8445
08A
Y84
4169
AY
8449
32A
Y84
4339
Mex
ico:
Son
ora:
Vic
init
yA
lam
osU
TA
A-5
4786
Pty
choh
yla
euth
ysan
ota
AY
8437
44A
Y84
3989
AY
8447
31A
Y84
4509
AY
8441
70A
Y84
4933
AY
8443
40M
exic
o:C
hiap
as:
Cer
roE
lB
aul,
Col
onia
Rod
ulfo
Fig
uero
a,13
07m
2005 189FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
EN
S84
86P
tych
ohyl
ahy
pom
ykte
rA
Y84
3745
AY
8439
90A
Y84
4732
——
——
Gua
tem
ala:
Izab
al
UT
AA
-547
82P
tych
ohyl
ale
onha
rdsc
hult
zei
AY
8437
46A
Y84
3991
AY
8447
33A
Y84
4510
AY
8441
71A
Y84
4934
AY
8443
41M
exic
o:O
axac
a:S
ierr
aM
adre
del
Sur
;C
arre
tera
Poc
hutl
a,68
1m
US
NM
5143
81P
tych
ohyl
asp
inip
olle
xA
Y84
3748
AY
8439
92A
Y84
4735
AY
8445
12A
Y84
4173
AY
8449
36A
Y84
4343
Hon
dura
s:A
tlan
tida
:P
arqu
eN
acio
nal
Pic
oB
onit
o,Q
uebr
ada
deO
ro(t
ribu
tary
ofR
ioV
iejo
)U
TA
A-5
4784
Pty
choh
yla
zoph
odes
AY
8437
49A
Y84
3994
AY
8447
36A
Y84
4513
AY
8441
74A
Y84
4937
AY
8443
44M
exic
o:O
axac
a:S
ierr
aM
ixe;
Mun
.S
anti
ago
Zac
atep
ec,
carr
eter
aZ
acat
epec
–Jes
usC
arra
nza,
1182
mJA
C21
606
Pty
choh
yla
sp.
AY
8437
47A
Y84
3993
AY
8447
34A
Y84
4511
AY
8441
72A
Y84
4935
AY
8443
42M
exic
o:O
axac
a:S
ierr
aM
ixe
QU
LC
2340
Scar
thyl
ago
inor
umA
Y84
3752
AY
8439
97A
Y84
4738
AY
8445
14—
AY
8449
38—
Bra
zil:
Am
azon
as:
Igar
ape
Nov
aE
mpr
esa
MA
CN
3864
9Sc
inax
acum
inat
usA
Y84
3753
AY
8439
98A
Y84
4739
AY
8445
15A
Y84
4176
AY
8449
39—
Arg
enti
na:
Cor
rien
tes:
Pas
ode
laP
atri
aM
LPA
2137
Scin
axbe
rtha
eA
Y84
3754
AY
8439
99A
Y84
4740
——
AY
8449
40A
Y84
4345
Arg
enti
na:
Bue
nos
Air
es:
Ata
laya
MV
Z20
7215
mSc
inax
boul
enge
riA
Y84
3755
AY
8440
00A
Y84
4741
AY
8445
16A
Y84
4177
——
Cos
taR
ica:
Gua
naca
ste:
ca.
0.2
kmW
Cos
taR
ica
Hw
y1
onfi
rst
pave
dro
ad10
kmN
entr
ance
San
taR
osa
Nat
iona
lP
ark
alon
gH
wy
1M
CP
3734
Scin
axca
thar
inae
AY
8437
56A
Y84
4001
AY
8447
42A
Y84
4517
—A
Y84
4941
AY
8443
46B
razi
l:R
ioG
rand
eD
oS
ul:
Pro
-Mat
a
190 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MV
Z20
3919
nSc
inax
elae
ochr
ous
AY
8437
57A
Y84
4002
AY
8447
43A
Y84
4518
AY
8441
78A
Y84
4942
—C
osta
Ric
a:H
ered
ia:
Sta
rkey
9sW
oods
,1.
5–3.
0km
ER
ioF
rio
Rd.
,at
1km
NW
entr
ance
Est
acio
nB
iolo
gica
La
Sel
vaJF
1973
Scin
axfu
scov
ariu
sA
Y84
3758
AY
8440
03A
Y84
4744
AY
8445
19A
Y84
4179
AY
8449
43A
Y84
4347
Arg
enti
na:
Mis
ione
s:G
uara
ni:
San
Vic
ente
:C
ampo
Ane
xoIN
TA
‘‘C
uart
elR
ioV
icto
ria’
’M
AC
N38
650
Scin
axna
sicu
sA
Y84
3759
AY
8440
04A
Y84
4745
AY
8445
20A
Y84
4180
—A
Y84
4348
Arg
enti
na:
Bue
nos
Air
es:
Bar
ader
o:E
stan
cia
‘‘E
lR
eton
o’’
IWK
109
Scin
axru
ber
AY
5493
65A
Y54
9418
AY
8447
46A
Y84
4521
AY
8441
81A
Y84
4944
—G
uyan
a:Iw
okra
ma:
Mur
iS
crub
cam
pM
AC
N38
241
Scin
axsq
uali
rost
ris
AY
8437
60—
AY
8447
47A
Y84
4522
AY
8441
82A
Y84
4945
AY
8443
49A
rgen
tina
:E
ntre
Rio
s:D
epto
.Is
las
del
Ibic
uy:
Rut
a12
viej
a,en
tre
Bra
zoL
argo
yA
rroy
oL
ucia
noU
TA
A-5
0749
Scin
axst
auff
eri
AY
8437
61A
Y84
4006
AY
8447
48A
Y84
4523
AY
8441
83—
—G
uate
mal
a:Z
acap
a:2.
9km
ST
ecul
utan
,on
road
toH
uit
MV
Z13
3014
oSm
ilis
caba
udin
iiA
Y54
9366
AY
8440
07A
Y84
4749
——
AY
8449
46—
Mex
ico:
Son
ora:
10.6
mi
W(b
yro
ad)
Ala
mos
JAC
2262
4Sm
ilis
cacy
anos
tict
aA
Y84
3763
AY
8440
08A
Y84
4750
AY
8445
24A
Y84
4184
AY
8449
47A
Y84
4350
Mex
ico:
Ver
acru
z:M
po.
San
And
res
Tux
tla,
Vol
can
San
Mar
tin,
Mar
tire
sde
Chi
cago
RdS
786
Smil
isca
phae
ota
AY
8437
64A
Y84
4009
AY
8447
51—
AY
8441
85A
Y84
4948
AY
8443
51C
apti
vera
ised
from
bree
ding
adul
tsat
Bal
tim
ore
Nat
iona
lA
quar
ium
2005 191FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
VC
R17
9Sm
ilis
capu
ma
AY
8437
65A
Y84
4010
AY
8447
52A
Y84
4525
AY
8441
86A
Y84
4949
—C
osta
Ric
a:P
rovi
ncia
Her
edia
:C
hila
mat
e(1
5km
WP
uert
oV
iejo
):R
eser
va‘‘
El
Vej
uco’
’M
JH46
Spha
enor
hync
hus
dori
sae
AY
8437
66A
Y84
4011
AY
8447
53A
Y84
4526
AY
8441
87—
—B
razi
l:A
maz
onas
:M
anau
s:L
ago
Jana
uri
US
NM
1521
36Sp
haen
orhy
nchu
sla
cteu
sA
Y54
9367
AY
8440
12A
Y84
4754
AY
8445
27A
Y84
4188
—A
Y84
4352
Per
u:M
adre
deD
ios:
30km
(air
line
)S
SW
Pue
rto
Mal
dona
do,
Tam
bopa
taR
eser
veM
NH
NP
1998
-311
Tep
uihy
laed
elca
eA
Y84
3770
——
AY
8445
30—
——
Ven
ezue
la:
Est
ado
Bol
ivar
:A
uyan
tepu
i,20
15m
UM
MZ
2189
14T
rach
ycep
halu
sjo
rdan
iA
Y84
3771
AY
8440
15A
Y84
4758
AY
8445
31A
Y84
4190
AY
8449
53A
Y84
4356
No
data
N/A
lT
rach
ycep
halu
sni
grom
acul
atus
AY
8437
72A
Y84
4016
AY
8447
59—
AY
8441
91—
—B
razi
l:E
spır
ito
San
to:
Set
iba,
Gua
rapa
riR
dS74
9T
ripr
ion
peta
satu
sA
Y84
3774
AY
8440
17A
Y84
4761
AY
8445
32A
Y84
4193
AY
8449
55A
Y84
4357
Bel
ize:
Hum
min
gbir
dH
wy,
9.5
kmfr
omW
este
rnH
ighw
aytu
rnpo
int
(cap
tive
-ra
ised
tadp
oles
,fr
omad
ults
coll
ecte
din
this
loca
lity
)N
/Al
Xen
ohyl
atr
unca
taA
Y84
3775
AY
8440
18—
——
——
Bra
zil:
Rio
deJa
neir
o:R
esti
nga
deM
aric
aH
ylid
ae,
Hem
iphr
acti
nae
Cry
ptob
atra
chus
sp.*
AY
3260
50—
——
——
—C
olom
bia:
San
tand
er:
Mun
icip
ioS
anG
il:
7km
byro
adS
WS
anG
ilC
FB
H57
20F
lect
onot
ussp
.A
Y84
3589
AY
8438
09A
Y84
4562
AY
8443
79A
Y84
4038
AY
8447
88A
Y84
4215
Bra
zil:
San
taC
atar
ina:
San
toA
mar
oda
Impe
ratr
iz
192 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
KR
L79
9G
astr
othe
caco
rnut
aA
Y84
3591
AY
8438
11—
—A
Y84
4040
——
Pan
ama:
Coc
le:
El
Cop
e:P
arqu
eN
acio
nal
‘‘O
mar
Tor
rijo
s’’
JLG
90G
astr
othe
cafis
sipe
sA
Y84
3592
—A
Y84
4564
AY
8443
81—
AY
8447
90—
Bra
zil:
Esp
ırit
oS
anto
:S
etib
a,G
uara
pari
MN
K52
86G
astr
othe
cacf
.m
arsu
piat
aA
Y84
3590
AY
8438
10A
Y84
4563
AY
8443
80A
Y84
4039
AY
8447
89—
Bol
ivia
:S
anta
Cru
z:C
abal
lero
:C
anto
nS
anJu
an:
Am
boro
Nar
iona
lP
ark
TN
HC
6249
2G
astr
othe
caps
eust
es*
AY
3260
51—
——
——
—E
cuad
or:
Chi
mbo
razo
:3.
3km
ST
ixan
,29
90m
MJH
3689
Hem
iphr
actu
she
lioi
AY
8435
94A
Y84
3813
AY
8445
66A
Y84
4382
—A
Y84
4792
—P
eru:
Uca
yali
:3
kmS
km65
onC
arre
tera
Fed
eric
oB
asad
reat
Ivit
aA
MN
HA
-164
211
Stef
ania
evan
siA
Y84
3767
—A
Y84
4755
—A
Y84
4189
AY
8449
50A
Y84
4353
Guy
ana:
Iwok
ram
a:P
akat
auC
reek
(85
m,
4845
9N,
5980
19W
)M
NH
N20
02.6
92St
efan
iasc
hube
rti
AY
8437
68A
Y84
4013
AY
8447
56A
Y84
4528
—A
Y84
4951
AY
8443
54V
enez
uela
:E
stad
oB
oliv
ar:
Auy
ante
pui,
2325
mH
ylid
ae,
Phy
llom
edus
inae
KR
L80
0A
galy
chni
sca
lcar
ifer
AY
8435
62A
Y84
3785
AY
8445
36—
——
AY
8441
96P
anam
a:C
ocle
:E
lC
ope:
Par
que
Nac
iona
l‘‘
Om
arT
orri
jos’
’R
dS53
7A
galy
chni
sca
llid
ryas
AY
8435
63—
AY
8445
37—
—A
Y84
4765
—B
eliz
e:S
tann
Cre
ekD
istr
ict:
Cok
scom
bB
asin
Wil
dlif
eS
antu
ary
QC
AZ
1321
7A
galy
chni
sli
todr
yas*
AY
3260
43—
——
——
—E
cuad
or:
Unk
now
n
2005 193FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MV
Z20
3768
Aga
lych
nis
salt
ator
*A
Y32
6044
——
——
——
Cos
taR
ica:
Her
edia
:S
tark
ey9s
Woo
ds:
1.5–
3.0
kmE
Rio
Fri
oR
dat
1km
NW
entr
ance
toE
stac
ion
Bio
logi
caL
aS
elva
ZU
FR
J79
26H
ylom
anti
sgr
anul
osa
AY
8436
87A
Y84
3933
AY
8446
80A
Y84
4469
AY
8441
27A
Y84
4889
AY
8443
01B
razi
l:P
erna
mbu
co:
Jaqu
eira
JAC
2200
9P
achy
med
usa
dacn
icol
orA
Y84
3714
AY
8439
59A
Y84
4702
AY
8444
88A
Y84
4144
AY
8449
08A
Y84
4318
Mex
ico:
Gue
rrer
o:C
arre
tera
Tie
rra
Col
orad
a-A
yutl
a,18
7m
CF
BH
7307
Pha
smah
yla
coch
rana
eA
Y84
3715
AY
8439
60—
——
——
Bra
zil:
Min
asG
erai
s:P
ocos
deC
alda
sC
FB
H57
56P
hasm
ahyl
agu
ttat
aA
Y84
3716
AY
8439
61A
Y84
4703
AY
8444
89A
Y84
4145
AY
8449
09—
Bra
zil:
Sao
Pau
lo:
Uba
tuba
:P
icin
guab
aA
MN
HA
1684
59P
hyll
omed
usa
bico
lor
AY
8437
23A
Y84
3968
AY
8447
10A
Y84
4495
AY
8441
52A
Y84
4915
—P
ettr
ade
AM
NH
-A14
1109
pP
hyll
omed
usa
hypo
chon
dria
lis
AY
8437
24A
Y84
3969
AY
8447
11A
Y84
4496
AY
8441
53A
Y84
4916
—G
uyan
a:D
ubul
ayR
anch
onth
eB
erbi
ceR
iver
KR
L95
5P
hyll
omed
usa
lem
urA
Y84
3687
AY
8439
70A
Y84
4712
—A
Y84
4154
AY
8449
17—
Pan
ama:
Coc
le:
El
Cop
e:P
arqu
eN
acio
nal
‘‘O
mar
Tor
rijo
s’’
KU
2054
20P
hyll
omed
usa
pall
iata
*A
Y32
6046
——
——
——
Per
u:M
adre
deD
ios:
Cuz
coA
maz
onic
oM
JH67
Phy
llom
edus
ata
rsiu
sA
Y84
3726
AY
8439
71A
Y84
4713
—A
Y84
4155
AY
8449
18A
Y84
4326
Bra
zil:
Am
azon
as:
Man
aus:
Res
erva
Duk
eM
AC
N37
796
Phy
llom
edus
ate
trap
loid
eaA
Y84
3727
AY
8439
72A
Y84
4714
—A
Y84
4156
AY
8449
19A
Y84
4327
Arg
enti
na:
Mis
ione
s:G
uara
ni:
San
Vic
ente
:C
ampo
Ane
xoIN
TA
‘‘C
uart
elR
ioV
icto
ria’
’M
JH70
76P
hyll
omed
usa
tom
opte
rna
AY
8437
28A
Y84
3973
AY
8447
15A
Y84
4497
AY
8441
57A
Y84
4920
AY
8443
28P
eru:
Hua
nuco
:R
ioL
lull
apic
his:
Pan
guan
a
194 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
AM
NH
-A16
6288
qP
hyll
omed
usa
vail
lant
iA
Y54
9363
AY
5494
16A
Y84
4716
AY
8444
98A
Y84
4158
AY
8449
21A
Y84
4329
Guy
ana:
Ber
bice
Riv
erca
mp
atca
.18
mi
(lin
ear)
SW
Kw
akw
ani
(ca.
2m
ido
wnr
iver
from
Kur
undi
Riv
er)
confl
uenc
e),
200
ftH
ylid
ae,
Pel
odry
adin
aeS
AM
1690
6C
yclo
rana
aust
rali
sA
Y84
3580
AY
8438
02A
Y84
4553
AY
8443
76—
——
Aus
tral
ia:
No
data
TN
HC
5193
6L
itor
iaar
faki
ana*
AY
3260
39—
——
——
—P
apua
New
Gui
nea:
Mad
ang:
ca.
10km
NW
Sim
bai,
Kai
ronk
Vil
lage
,20
00m
AM
5274
4L
itor
iaau
rea
AY
8436
91A
Y84
3937
AY
8446
84—
AY
8441
30A
Y84
4892
—N
ewC
aled
onia
:P
rovi
nce
Nor
d:V
alle
Pha
aye,
Nom
acR
iver
,8
kmE
Pou
mA
MN
HA
-168
409
Lit
oria
caer
ulea
AY
8436
92A
Y84
3938
AY
8446
85—
AY
8441
31A
Y84
4893
—P
ettr
ade
SA
MA
1226
0L
itor
iafr
eicy
neti
AY
8436
93A
Y84
3939
AY
8446
86A
Y84
4473
—A
Y84
4894
—A
ustr
alia
:N
ewS
outh
Wal
es:
16km
ER
etre
atN
/AL
itor
iain
fraf
rena
taA
Y84
3694
AY
8439
40A
Y84
4687
AY
8444
74—
—A
Y84
4304
Pet
trad
eS
AM
A17
215
Lit
oria
mei
rian
aA
Y84
3695
—A
Y84
4688
AY
8444
75A
Y84
4132
AY
8448
95—
Aus
tral
ia:
Wes
tern
Aus
tral
ia:
Bla
ckR
ock,
near
Kun
unur
raS
AM
A45
336
Nyc
tim
iste
spu
lche
rA
Y84
3701
AY
8439
46A
Y84
4692
—A
Y84
4134
——
Pap
uaN
ewG
uine
a:M
agid
obo,
SH
PA
MN
HA
-828
22r
Nyc
tim
yste
sku
bori
AY
8437
02A
Y84
3947
AY
8446
93A
Y84
4479
——
—P
apua
New
Gui
nea:
Mor
oba:
Wau
AM
NH
A-8
2845
sN
ycti
mys
tes
nari
nosu
sA
Y84
3703
AY
8439
48A
Y84
4694
—A
Y84
4135
—A
Y84
4308
Pap
uaN
ewG
uine
a:T
ambu
lA
llop
hryn
idae
MA
D15
12A
llop
hryn
eru
thve
niA
Y84
3564
AY
8437
86A
Y84
4538
AY
8443
61—
AY
8447
66—
Guy
ana:
Kab
ocal
ica
mp
(101
m,
4817
.109
N,
5883
0.56
9W)
2005 195FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
DP
L39
32t
Ast
ylos
tern
idae
Tri
chob
atra
chus
robu
stus
AY
8437
73—
AY
8447
60—
AY
8441
92A
Y84
4954
—C
amer
oon:
Sou
thw
est
Pro
vinc
e:nn
hill
sout
hof
Man
yem
enB
ufon
idae
MV
Z22
3279
Ate
lopu
sva
rius
*A
Y32
5996
——
——
——
Cos
taR
ica:
Sou
thof
Las
Alt
uras
MA
CN
3863
9B
ufo
aren
arum
AY
8435
73A
Y84
3795
AY
8445
47A
Y84
4370
—A
Y84
4775
AY
8442
05A
rgen
tina
:S
anL
uis,
ruta
20en
tre
Bar
das
Bla
ncas
ykm
330
MJH
7095
Den
drop
hryn
iscu
sm
inut
usA
Y84
3582
AY
8438
04A
Y84
4555
——
——
Per
u:H
uanu
co:
Rio
Llu
llap
ichi
s:P
angu
ana
N/A
Dyd
inam
ipus
sjoe
sted
ti*
AY
3259
91—
——
——
—C
amer
oon:
Unk
now
n
MA
CN
3853
1M
elan
ophr
ynis
cus
klap
penb
achi
AY
8436
99A
Y84
3944
—A
Y84
4478
—A
Y84
4899
AY
8443
06A
rgen
tina
:C
haco
:pr
oxim
idad
esde
Res
iste
ncia
QC
AZ
4580
Oso
rnop
hryn
egu
acam
ayo*
AY
3260
36—
——
——
—E
cuad
or:
Nap
o:L
ago
Sum
aco,
Vol
can
Sum
aco
N/A
Ped
osti
bes
hoss
i*A
Y32
5993
——
——
——
Mal
aysi
a:P
ahan
g:K
rau
Wil
dlif
eR
eser
ve,
Peh
ang
mai
nre
sear
chfi
eld
stat
ion,
ca.
13km
NW
Kua
laK
rau
atco
nflue
nce
Kra
uan
dL
ompa
tR
iver
sT
NH
C62
001
Schi
smad
erm
aca
rens
*A
Y32
5997
——
——
——
Tan
zani
a:D
odom
an
Bra
chyc
epha
lida
eN
/AB
rach
ycep
halu
sep
hipp
ium
*A
Y32
6008
——
——
——
Bra
zil:
noda
ta
Cen
trol
enid
aeS
IUC
H-7
053
Cen
trol
ene
pros
oble
pon
AY
8435
74A
Y84
3796
AY
8445
48A
Y84
4371
—A
Y84
4776
AY
8442
06P
anam
a:C
ocle
:E
lC
ope:
Par
que
Nac
iona
l‘‘
Om
arT
orri
jos’
’
196 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
MN
K52
42C
ochr
anel
labe
jara
noi
AY
8435
76A
Y84
3798
AY
8445
49A
Y84
4372
AY
8440
29A
Y84
4777
AY
8442
08B
oliv
ia:
San
taC
ruz:
Cab
alle
ro:
Can
ton
San
Juan
:A
mbo
roN
atio
nal
Par
kC
FB
H57
29H
yali
noba
trac
hium
eury
gnat
hum
AY
8435
95A
Y84
3814
AY
8445
67A
Y84
4383
—A
Y84
4793
AY
8442
17B
razi
l:M
inas
Ger
ais:
Itam
onte
sD
endr
obat
idae
US
NM
-FS
5205
5C
olos
teth
usta
lam
anca
eA
Y84
3577
AY
8437
99A
Y84
4550
AY
8443
73—
AY
8447
78—
Pan
ama:
Boc
asde
lT
oro
US
NM
3131
8D
endr
obat
esau
ratu
sA
Y84
3581
AY
8438
03A
Y84
4554
—A
Y84
4032
AY
8447
81A
Y84
4211
Pan
ama:
Boc
asde
lT
oro
TN
HC
6248
8P
hyll
obat
esbi
colo
r*A
Y32
6031
——
——
——
No
data
Hel
eoph
ryni
dae
TM
SA
8415
7H
eleo
phry
nepu
rcel
liA
Y84
3593
AY
8438
12A
Y84
4565
——
AY
8447
91A
Y84
4216
Sou
thA
fric
a:W
este
rnC
ape
Pro
vinc
e:C
edar
berg
Mou
ntai
nR
ange
:he
adof
Kro
mR
iver
Hem
isot
idae
TN
HC
6248
9H
emis
usm
arm
orat
um*
AY
3260
70—
AY
3643
97—
——
—S
eeD
arst
and
Can
nate
lla
(200
4)an
dB
iju
and
Bos
suyt
(200
3)L
epto
dact
ylid
ae,
Cer
atop
hryi
nae
JF92
9C
erat
ophr
yscr
anw
elli
AY
8435
75A
Y84
3797
——
——
AY
8442
07A
rgen
tina
:S
anta
Fe:
Ver
a:E
a.‘‘
Las
Gam
as’’
JF18
91O
dont
ophr
ynus
amer
ican
usA
Y84
3704
AY
8439
49A
Y84
4695
AY
8444
80—
AY
8449
01A
Y84
4309
Arg
enti
na:
Bue
nos
Air
es:
Esc
obar
:L
oma
Ver
de:
Ea.
‘‘L
osC
ipre
ses’
’L
epto
dact
ylid
ae,
Cyc
lora
mph
inae
ML
PA14
14C
ross
odac
tylu
ssc
hmid
tiA
Y84
3579
AY
8438
01A
Y84
4552
AY
8443
75A
Y84
4031
AY
8447
80A
Y84
4210
Arg
enti
na:
Mis
ione
s:A
rist
obul
ode
lV
alle
:B
alne
ario
Cun
apir
u
2005 197FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
2(C
onti
nued
)
Vou
cher
Spe
cies
12S
-tR
NA
vali
ne-1
6SC
ytoc
hrom
eb
Rho
dops
inex
on1
RA
G-1
Tyr
osin
ase
Sev
enth
inab
sent
ia28
SL
ocal
ity
AM
NH
-A16
5195
Lep
toda
ctyl
idae
,E
leut
hero
dact
ylin
aeE
leut
hero
dact
ylus
pluv
ican
orus
AY
8435
86—
AY
8445
59—
AY
8440
35A
Y84
4785
AY
8442
13B
oliv
ia:
San
taC
ruz:
Cab
alle
ro:
Can
ton
San
Juan
:A
mbo
roN
atio
nal
Par
kK
U20
2519
Ele
uthe
roda
ctyl
usth
ymel
ensi
s*A
Y32
6009
——
——
——
Ecu
ador
:C
arch
i:12
kmW
Tufi
no,
3520
mA
MN
H-A
1651
08P
hryn
opus
sp.
AY
8437
20—
——
——
AY
8443
23B
oliv
ia:
La
Paz
:B
auti
sta
Saa
vedr
a:C
anto
nC
hara
zani
Lep
toda
ctyl
idae
,L
epto
dact
ylin
aeA
MN
H-A
1663
12A
deno
mer
asp
.A
Y84
3561
AY
8437
84A
Y84
4535
AY
8443
60A
Y84
4021
—A
Y84
4195
Guy
ana:
Ber
ebic
eR
iver
cam
pat
ca.
18m
i(l
inea
r)S
WK
wak
wan
i(c
a.2
mi
dow
nriv
erfr
omK
urun
diR
iver
confl
uenc
e)M
JH70
82E
dalo
rhin
ape
rezi
AY
8435
85A
Y84
3807
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198 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORYA
PP
EN
DIX
2(C
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nued
)
Vou
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2005 199FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAEA
PP
EN
DIX
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)
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200 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 3
MORPHOLOGICAL CHARACTERS
Based on Burton’s (2004) collection of obser-vations on hylid foot musculature, we built a dataset. Burton (2004) included two appendices, Aand B, with the taxa he examined for the differentcharacters. In the text, however, it is not imme-diately clear which of the appendices he refers toin some cases. Burton (personal commun.) gra-ciously provided us with a precise list of the ap-pendix to which each character refers. This infor-mation is reproduced below in a character list thatincludes a brief discussion when relevant. Burton(2004) numbered his characters as a continuationof Duellman’s (2001) character list. We are notincorporating Duellman’s (2001) characters be-cause he only summarized character states for thehylid subfamilies without any reference to the in-dividual taxa, therefore assuming monophyly.
Character List
0 (Burton’s char. 25): Insertions of the m. flexordigitorum brevis superficialis. (0) Three inser-tions. (1) Two insertions.
1 (Burton’s char. 26): Structure of the tendonof the m. flexor digitorum brevis superficialis. (0)Undivided. (1) Divided along its length into a me-dial tendon, from which arises tendo superficialisIV, and a lateral tendon from which arise tendinessuperficiales III and V, with cross tendons be-tween the divisions. (2) Divided along its lengthinto a medial tendon, from which arise tendo su-perficialis IV and m. lumbricalis longus digiti V,and a lateral tendon from which arise tendo su-perficialis V and m. lumbricalis longus digiti IV.
2 (Burton’s char. 27): Structure of the terminiof tendines superficiales. (0) Tendon neither ex-panded nor bifurcated. (1) Tendon bifid.
3 (Burton’s char. 28): Origin of the tendo su-perficialis hallucis. (0) Tendo superficialis hallucisarises from the mediodistal corner of the aponeu-rosis plantaris. (1) (Burton’s char. 28.2). The ten-do superficialis hallucis tapers from an expandedcorner of the aponeurosis plantaris; fibers of them. transversus plantae distalis originating on dis-tal tarsal 2–3 insert on the lateral side of the ten-don. (2) (Burton’s char. 28.3). The tendo superfi-cialis hallucis arises from the aponeurosis, but theorigin is proximal to that of the m. lumbricalisbrevis hallucis, so that the tendo passes at an an-gle across that muscle as it passes along the toe,neither along its marginor straight along the mus-cle. (3) (Burton’s char. 28.4). The tendo superfi-cialis hallucis comes from a massive muscle thatarises from distal tarsal 2–3. COMMENT: Burton(2004: 218) included within this character a state
(1) pertaining to the morphology of the tendo su-perficialis hallucis, not to its origin. For this rea-son, we consider it to be a different character andthat is why we excluded it here.
4 (Burton’s char. A): Structure of m. contrahen-tis hallucis. (0) Present and conspicuous. (1) Ab-sent or reduced.
5 (Burton’s char. B): Origin of m. contrahentishallucis (if present). (0) Origin on distal tarsal 1.(1) Origin on distal tarsal 2–3.
6 (Burton’s char. 29): Presence or absence ofm. flexor teres hallucis. (0) Absent. (1) Present.
7 (Burton’s char. 30): Presence or absence ofm. abductor brevis plantae hallucis. (0) Present.(1) Absent. COMMENT: Burton (2004: 219) includ-ed states pertaining to presence or absence, aswell as structure, in the same character. FollowingHawkins et al. (1997) and Strong and Lipscomb(1999), we consider them as two different char-acters.
8 (Burton’s char. 30): Structure of the m. ab-ductor brevis plantae hallucis. (0) Narrow. (1)Broad.
9 (Burton’s char. 31): Origin of tendo superfi-cialis pro digiti II. (0) Tendon arising from thedistal edge of the aponeurosis plantaris. (1) Ten-don arising from a deep, triangular muscle, whichoriginates on the distal tarsal 2–3. (2) Tendonbroad at the base, acting as a point of insertion ofa portion of the m. transversus plantae distalis, sothat the tendo superficialis pro digiti II appears tobe the tendon of insertion of the m. transversusplantae distalis.
10 (Burton’s char. 32): Origin of tendo super-ficialis pro digiti III. (0) Tendo superficialis prodigiti III arising from the m. flexor digitorumbrevis superficialis, but with the distal margin ofthe aponeurosis wrapped around the tendon. (1)Tendo superficialis pro digiti III arising from the.flexor digitorum brevis superficialis, with no con-tribution from the aponeurosis plantaris. (2) Tendosuperficialis pro digiti III arising in part from m.flexor digitorum brevis superficialis or the tendosuperficialis pro digiti IV, and in part by a super-ficial tendon (from which also cutaneous tendonsarise) that emerges from centrally on the plantarsurface of the aponeurosis plantaris. (3) Tendo su-perficialis pro digiti III arising entirely from themargin of the aponeurosis plantaris. (4) Tendo su-perficialis pro digiti III arising entirely from thesuperficial tendon.
11 (Burton’s char. 33): Origin of m. flexor ossismetatarsi II. (0) Tendon arising from the condyleof the fibulare alone, in common with the tendon
2005 201FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
of origin of the m. flexor ossis metatarsi III. (1)Tendon arising from distal tarsal 2–3 only. (2)Muscle with two origins—a long distal sectionarising from a tendon on the distal condyle of thefibulare, and a short proximal section arising fromdistal tarsal 2–3.
12 (Burton’s char. D): Accessory tendon of or-igin of the m. lumbricalis longus digiti III. (0)Absent. (1) Present.
13 (Burton’s char. 34): Number of tendons ofinsertion of m. lumbricalis longissimus digiti IV.(0) Two tendons. (1) One tendon.
14 (Burton’s char. 35, refers to taxa listed onhis appendix B): Relationship between the originof m. flexor ossis metatarsi IV and the joint ten-don of origin of m. flexores ossum metatarsorumII and III varies. (0) The tendons are adjacent attheir origins. (1) The tendons cross each other.COMMENT: Burton (2004: 222) included a thirdstate in this character, whose taxonomic distribu-tion does not match any of the taxa included inthis study.
15 (Burton’s char. F): Structure of m. flexor os-sis metatarsi IV. (0) Inserting on metatarsus IVonly. (1) Inserting on both metatarsi IV and V.
16 (Burton’s char. 36): Length variation of them. flexor ossis metatarsi IV. (0) Very short, in-serting on the proximal two thirds of metatarsalIV or less only. (1) Extending the entire length ofMetatarsal IV.
17 (Burton’s char. 37, refers to taxa listed in hisappendix A): Tendons of insertion of m. lumbri-calis longus digiti V. (0) One tendon. (1) Two ten-dons arising equally from both sides of an undi-vided muscle. (2) Two tendons arising from twoequal muscle slips. (3) (Burton’s char. 37.4). Twotendons, the medial of which arises from a small,distal slip. COMMENT: The state 3 of this characterdescribed by Burton is present only in Cochra-nella siren, a taxon not included in this analysis.
18 (Burton’s char. 38, refers to taxa listed in hisappendix A): Number of slips of the medial m.lumbricalis brevis digiti V. (0) Two slips. (1) Oneslip. COMMENT: Burton included a third state thatis present in Gastrophryne, a taxon not includedin this analysis.
19 (Burton’s char. 39, refers to taxa listed inboth appendices): Presence or absence of a tendonfrom the m. flexor digitorum brevis superficialisto the medial slip of the medial m. lumbricalisbrevis digiti V. (0) Absent. (1) Present.
20 (Burton’s char. 40): Insertion of the lateralslip of the medial m. lumbricalis brevis digiti V.(0) Insertion by tendon onto the basal phalanx. (1)Insertion pennate.
21 (Burton’s char. 41, refers to taxa listed in hisappendix A): Width and orientation of the m.transversus metatarsus II. (0) Narrow, occupying
less than 80% of the length of metatarsus II. (1)Broad, occupying the entire length of metatarsusII. (2) Oblique, with a narrow, proximal connec-tion onto metatarsus III, and a broad, distal con-nection to metatarsus II. COMMENT: Burton (inlitt., 19 July 2004) warned us that the numbers ofthe metatarsals in the description of state 2 ofchars. 41 and 42 were accidentally reversed in hispaper; this reversal is corrected here.
22 (Burton’s char. 42): Position of m. transver-sus metatarsus III. (0) Relatively distal, not oc-cupying the proximal 15% of either metatarsus.(1) Proximal. (2) Oblique, with a narrow, proxi-mal connection onto metatarsus IV, and a broad,distal connection to metatarsus III.
23 (Burton’s char. 43, refers to taxa listed in hisappendix A): Breadth of m. transversus metatar-sus III. (0) Narrow, extending less than 70% ofthe length of metatarsus III. (1) Broad, occupyingmore than 75% of the length of metatarsus III.
24 (Burton’s char. 44): Presence or absenceof m. transversus metatarsus IV. (0) Present. (1)Absent. COMMENTS: Burton included presence,absence, and several modifications within a sin-gle character. We are unsure as to the definitionsand limits of the states describing the differentorigins and insertions of the muscle when pre-sent, and thus we only score its presence or ab-sence.
25 (Burton’s char. 45): Relationship of the m.flexor ossis metatarsi IV with m. transversusmetatarsus IV, if present. (0) M. flexor ossis meta-tarsi IV dorsal to m. transversus metatarsus IV.(1) M. flexor ossis metatarsi IV ventral to m.transversus metatarsus IV.
26 (Burton’s char. 46, refers to taxa listed in hisappendix A): Nature of origin of m. extensor dig-itorum comunis longus. (0) Fibrous origin fromthe medial surface of the m. tarsalis anticus. (1)A long, straplike tendon arising on the lateral sideof the distal end of the tibiofibula, close to theorigin of the m. tarsalis anticus.
27 (Burton’s char. 47, refers to taxa listed in hisappendix A): Nature of insertion of m. extensordigitorum comunis longus. (0) Flat tendon ontothe fascia of one or more dorsal superficial mus-cles. (1) Strong tendon(s) directly to the dorsa ofone or more metatarsi.
28 (Burton’s char. 48): Insertion of m. extensordigitorum comunis longus on metatarsal II. (0)Present. (1) Absent. COMMENTS: Burton definedthe states of character 48 in a complex way, com-bining the different points of insertions into singlestates. We consider it more informative and lessredundant to consider the different points of in-sertion as different characters. The insertion onmetatarsal III is uninformative in the present con-
202 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
text, since it is reported absent only in Stefaniascalae, a taxon not included in the analysis.
29 (Burton’s char. 48): Insertion of m. extensordigitorum comunis longus on metatarsal IV. (0)Present. (1) Absent.
30 (Burton’s char. 49): Origins of the m. exten-sor brevis medius hallucis and m. extensor brevismedius digiti II. (0) Separate origins. (1) Adjacentorigins.
31 (Burton’s char. G): Insertion of the m. ex-tensor brevis medius hallucis. (0) Insertion in thebasal phalanx of digit I only. (1) Insertion by aflat tendon onto the base of the basal phalanx ofdigit I, plus a strong, narrow tendon that passesalong the medial margin of digit II, inserting onthe basal phalanx.
32 (Burton’s char. 50, refers to taxa listed in hisappendix A): Position and nature of insertions ofm. abductor brevis dorsalis digiti V. (0) Insertionpennate, along the proximal half of the lateralmargin of metatarsal V, displacing the origins ofmm. extensores breves profundi digiti V, so thatthe lateral m. extensor brevis profundus originateson the mediodorsal surface of the metatarsal V.(1) Insertion by a short tendon to the dorsum ofthe metatarsal V. COMMENT: Burton described athird state that is present in Hyperolius, a taxonnot included in this analysis.
33 (Burton’s char. 51, refers to taxa listed onhis appendix A): Nature of the origin of the m.extensor brevis superficialis digiti III on the distalend of the fibulare. (0) Fibrous origin. (1) Originby a flat tendon.
34 (Burton’s char. 52, refers to taxa listed in hisappendix A): Number of insertions of m. extensorbrevis superficialis digiti III. (0) Single insertiononto the dorsum of the m. extensor brevis mediusdigiti III. (1) Single insertion via a long tendonproximally on the dorsum of basal phalanx III. (2)Two insertions, a flat tendon onto basal phalanxIII, as state 1, and a pennate insertion on meta-tarsus III.
35 (Burton’s char. 53, refers to taxa listed in hisappendix A): Presence or absence of m. extensorbrevis medius digiti III. (0) Present. (1) Absent.
36 (Burton’s char. 54): Number of slips in them. extensor brevis superficialis digiti IV. (0) Twoorigins, two separate muscles. (1) One origin, bel-ly undivided.
37 (Burton’s char. 55, refers to taxa listed in hisappendix A): Nature of the origin of the mm. ex-tensores breves superficiales digiti IV. (0) Bothorigins pennate. (1) (Burton’s char. 55.2): Bothorigins from long, flat tendons. COMMENT: Burtondescribed a third state present in Rhacophorusmaculatus, a taxon not included here.
Characters C and E were not included becausethe taxonomic distribution of their states is irrel-evant for hylids. Character 56 was excluded be-cause it actually includes several characters (pres-ence or absence of mm. extensores breves distalesin each digit), and there is no information as towhich of the slips (medial or lateral) is present.
Morphological Synapomorphies Common toAll Trees
(node numbers as in figs. 2–5)
Allophryne ruthveni, 10: 2 → 4. Anotheca spi-nosa, 3: 0 → 1. Cyclorana australis, 17: 3 → 0.Dendropsophus sarayacuensis, 6: 1 → 0. Hyla eu-phorbiacea, 28: 1 → 0. Hypsiboas calcaratus, 3:1 → 0. Hypsiboas granosus, 9: 2 → 0. Hypsiboaspellucens, 3: 1 → 0, 9: 2 → 0. Hypsiboas pictur-atus, 6: 0 → 1. Hypsiboas punctatus, 3: 1 → 0.Hypsiboas raniceps, 5: 0 → 1. Hypsiboas rufite-lus, 6: 0 → 1. Litoria aurea, 21: 0 → 1. Litoriafreycineti, 18: 0 → 1, 28: 1 → 0. Osteopilus sep-tentrionalis, 6: 0 → 1, 29: 1 → 0. Osteopilus vas-tus, 31: 1 → 0. Plectrohyla guatemalensis, 12: 0→ 1. Scarthyla goinorum, 25: 0 → 1. Node 289,1: 0 → 1, 10: 2 → 1. Node 337, 21: 0 → 2. Node338, 1: 1 → 0. Node 352, 11: 0 → 1. Node 372,3: 0 → 1. Node 375, 1: 1 → 0. Node 390, 28: 1→ 0. Node 422, 10: 1 → 0, 14: 1 → 0, 16: 1 →0, 19: 1 → 0. 22: 1 → 2. Node 462, 28: 1 → 0.Node 465, 33: 1 → 0. Node 498, 21: 0 → 1. Node507, 7: 0 → 1.
APPENDIX 4
ADDITIONAL COMMENTS ON SOME SPECIES
Hyla albovittata Lichtenstein and Martens,1856: Lichtenstein and Martens (1856) describedthis species from ‘‘Brazil’’, without further data.Subsequently, it was not included in Nieden’s(1923) catalog; Duellman (1977) stated that theholotype was unknown; however, the holotype ishoused at the Zoologisches Museum–Universitat
Humboldt, Berlin. The specimen (ZMB 3140), anadult male, is in a remarkably good state of pres-ervation, retaining details of pattern and colora-tion. On study, it is evident that it is a junior syn-onym of Hyla pulchella Dumeril and Bibron,1841.
Hyla auraria Peters, 1873: This species was de-
2005 203FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
scribed in detail based on a single specimen re-ported as coming from ‘‘angeblich aus Sudamer-ika’’ (Peters, 1873). Subsequently, Boulenger(1882) presented a brief description without fur-ther comment. Duellman (1977) reported that theholotype was ‘‘formerly at ZSM, now lost.’’Duellman (in Frost 1985) stated that the name hasnever been associated with a population of an-urans. However, the holotype is extant. The ho-lotype of H. auraria (ZSM 1175/0), presumablya female (it lacks vocal slits), is in relatively goodstate of preservation, but it is very faded. Wecould not associate it with any known species ofHylinae nor could we associate it with any of thegenera recognized in this work.
Hyla palliata Cope, 1863: Cope (1863) de-scribed this species, stating that it was a specimenfrom the Page collection, with provenance ‘‘Par-aguay’’.32 He also provided the Smithsonian Mu-seum number 6225. The species was not includedin the list of type specimens of the USNM (Coch-ran, 1961). According to Heyer (personal com-mun. to J.F., 21 Jan. 2004), the USNM catalogentry indicates that there were originally twospecimens cataloged as 6225, and in a card fileinitiated by Cochran of missing type specimens,there is a card indicating that the specimens couldnot be found in 1957. Because the specimens hadnot been found since, Heyer noted, ‘‘the most rea-sonable conclusion is that the specimens are in-deed lost.’’ The description provided by Cope in-dicates a character combination that could be in-dicative of a species of Hypsiboas (no species ofBokermannohyla are known for the area surveyedby the Page expedition): ‘‘All the digits of pos-terior extremity palmate to penultimate phalanx;of the anterior the three external are one thirdwebbed. Metacarpus of inner digit with a largetubercle.’’ Nevertheless, in the absence of otherevidence, we prefer to consider this species as anomen dubium.
Hyloscirtus estevesi (Rivero, 1968): This spe-cies was originally described as a member of theCentrolenidae by Rivero (1968), and was later(Rivero, 1985) included in the Centrolenella pul-idoi species group. Savage (in Frost, 1985) con-sidered this species to be a member of the Hyli-dae, and it was not included in the taxonomic re-arrangement of Centrolenidae by Ruiz-Carranzaand Lynch (1991). Myers and Donnelly (1997)and Frost (2002) considered it as incerta sedis.However, La Marca (1992) listed it as a centro-
32 Although the provenance is ‘‘Paraguay’’, thisshould be taken in a very wide sense, since the Pageexpedition travelled from Buenos Aires, through the Pa-rana river up to Corumba, State of Matto Grosso, Brazil(Page, 1859).
lenid, and later (La Marca, 1997; 1998) employedthe combination Hyalinobatrachium estevesi, mis-takenly attributing it to Ruiz-Carranza and Lynch(1991). The study of its holotype (MCZ 72498)indicates that it is a juvenile of an unidentifiedspecies that we associate with the Hyloscirtus bo-gotensis species group. While it could well be ajuvenile of either H. jahni or H. platydactylus, andtherefore a potential junior synonym of either ofthese species, we tentatively recognize it as a val-id species pending additional work.
Hypsiboas pulidoi (Rivero, 1968): This specieswas described from ‘‘Monte Duida, 2000 pies,Territorio Amazonas, Venezuela’’ as a Centrolen-idae by Rivero (1968). Later Rivero (1985) in-cluded in its own species group, the Centrolenellapulidoi group. Savage (in Frost, 1985) consideredthis species to be a hylid. Furthermore, Rivero(1985: 361) stated that ‘‘the resemblance of C.pulidoi with some specimens with the same col-oration of H. benitezi (with which it is syntopic)is so remarkable that for some time it was thoughtto consider them synonyms, but C. pulidoi hasfused tarsal bones, a different coloration, and theeyes are red in living specimens’’ (freely trans-lated from the Spanish). This species was not in-cluded in the taxonomic rearrangement of Centro-lenidae by Ruiz-Carranza and Lynch (1991), butAyarzaguena (1992) and La Marca (1992) stilllisted it as a centrolenid, and Gorzula and Senaris(1999) referred to it as ‘‘Centrolenella’’ pulidoi.Duellman (1999) employed the combination Hylapulidoi without further comment; Myers and Don-nelly (1997) and Frost (2002) considered it as in-certae sedis. The study of the holotype (MCZ72499), a female according to Rivero (1968), in-dicates that it is a species close to Hypsiboas ben-itezi. The specimen has a slightly enlarged pre-pollex, similar to the situation seen in females ofH. benitezi. Considering its small size (20.3 mm),we are unsure as to whether it is an adult femaleor a juvenile. We tentatively consider this speciesas valid, pending a careful comparison with ju-veniles of Hypsiboas benitezi. The only apparentdifference between Hypsiboas benitezi and H.pulidoi is that the latter has a red iris (Rivero,1968), whereas the iris of the former was de-scribed by Myers and Donnelly (1997) as ‘‘lightbronzy brown with fine balck venation.’’ How-ever, this issue should be addressed cautiously,because as noted by Myers and Donnelly (1997),at least two species are probably confounded un-der the name H. benitezi, and these authors de-scribed specimens from Tamacuari, that they con-sidered morphologically different from the topo-types, and iris coloration in topotypic material re-mains undescribed.
Scinax dolloi (Werner, 1903): Werner (1903)
204 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
described H. dolloi, based on two specimens,whose provenance was stated as Brazil, adding‘‘unfortunately it is unknown with more preci-sion’’ (from the German). The species was in-cluded in Nieden (1923) with a brief description,and without additional comments by Bokermann(1966b), Gorham (1974), and Duellman (1977).B. Lutz (1973) included it as a ‘‘doubtful spe-cies’’. Duellman (in Frost, 1985, 2002) stated thatthe name had not been associated with any knownpopulation.
Lang (1990) included it on the list of type spec-imens housed at the Institut Royal des SciencesNaturelles in Brussels, Belgium, commenting that‘‘although the original type-description lists spe-cifically that no specific locality is know the reg-ister has ‘Haut Maringa, Bresil’ as locality data’’.The two syntypes (IRSNB 6481) are in a fairlygood state of preservation, and their study showsthat they clearly belong to the genus Scinax, and,within it, they seem to be related to the ruberclade (Faivovich, 2002). Scinax dolloi is morpho-logically close to Scinax hayii and S. perereca.While its exact status remains to be elucidated, inthe meantime we consider it a valid species ofScinax.
Scinax karenanneae Pyburn, 1993: Pyburn(1993) described this species from ‘‘near Timbo,Department of Vaupes, Colombia’’. Although theoriginal description and accompanying figuressuggest that the species is superficially very sim-ilar to many species of Scinax, Pyburn (1993)ruled out the possibility that this species belongedto Scinax due to the sperm with a single tail fil-ament. Problems with the interpretation of spermmorphology data have been discussed earlier, andin this particular case the poorly known taxonom-ic distribution of this character state in Hylinaeprecludes any interpretation of polarity.
The paratypes of Hyla karenanneae (UTA A-3768–79) show the synapomorphies of Scinaxlisted by Faivovich (2002) that can be seen with-out deep dissections. Therefore, we propose itsinclusion in Scinax. This species has a large sub-gular vocal sac, which in the phylogenetic anal-ysis of Scinax of Faivovich (2002) optimized asa synapomorphy of a clade composed by S. stauf-feri, S. cruentommus, S. fuscomarginatus, and anundescribed species from northern Brazil.
Scinax megapodius: This species was figuredby Miranda-Ribeiro (1926), but formally de-scribed only later (Miranda-Ribeiro, 1937). Orig-inally described from the localities of Sao Luiz deCaceres and Porto Esperidiao, Matto Grosso (Mi-
randa-Ribeiro, 1937), this species was considereda synonym of Hyla fuscovaria by Lutz (1973).However, Fouquette and Delahoussaye (1977)consider it a valid species, based on differencesin sperm morphology from H. fuscovaria. On thatbasis, they included it in their Ololygon cathari-nae group. Almeida and Cardoso (1985) present-ed an analysis of intraspecific variation amongspermatozoa of O. fuscovaria, and suggested thatthe differences observed among O. fuscovaria andO. megapodia were the same as those seen amongdifferent specimens of the same population of O.fuscovarius. Without comment, Duellman andWiens (1992) considered Hyla megapodia to be asynonym of Scinax fuscovarius, as did Lavilla(1992). Based on the lack of comments by theseauthors, Frost (2002) continued recognition ofScinax megapodius as a valid species. Based onthe results of Almeida and Cardoso (1985), andconsidering that there is no published evidencethat Scinax megapodius is different from S. fus-covarius, following Lutz (1973), we treat it as ajunior synonym of the latter.
Scinax trachythorax: This species was describedby Muller and Hellmich (1936) from the localityof ‘‘Apa-Bergland (San Luis)’’, Paraguay, also re-ferred as ‘‘Estancia San Luis de la Sierra, Apa-Bergland’’ (Muller and Hellmich, 1936: 114). Lutz(1973) considered this species a junior synonym ofHyla fuscovaria. Fouquette and Delahoussaye(1977) considered it a valid species, based on dif-ferences in sperm morphology from H. fuscovaria,and recognized it as Ololygon trachthorax. Almei-da and Cardoso (1985) presented an analysis ofintraspecific variation in spermatozoa of O. fusco-varia, and suggested that the differences observedamong O. fuscovaria and O. trachythorax were thesame as those seen among different specimenswithin the same population of O. fuscovaria. How-ever, O. trachythorax was recognized as a validspecies by Duellman and Wiens (1992), who em-ployed the combination Scinax trachythorax. Lav-illa (1992) considered it a synonym of S. fusco-varius, as did De la Riva et al. (2000), who ex-plicitly followed Lutz (1973) and Cei (1980). (Thispublication did not incorporate changes introducedby Fouquette and Delahoussaye [1977]; Cei [1987]later recognized S. trachythorax as a valid species.)Considering the comments of Almeida and Car-doso (1985), and in light of the absence of anypublished diagnostic character state between S. fus-covarius and S. trachythorax, following Lutz(1973), we tentatively consider the latter a juniorsynonym of the former.
2005 205FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5SYNAPOMORPHIES INVOLVING DNA SEQUENCES, FOR ALL TAXONOMIC GROUPS OF HYLIDAE WITH THE EXCEPTION OF SPECIES
GROUPS WHOSE MONOPHYLY WAS NOT TESTED IN THIS ANALYSIS
Hylidae
Cytochrome bPos. 14: T → A12SPos. 646: A → TPos. 1270: G → APos. 1423: T → AtRNA valinePos. 6: A → GPos. 105: T → C16SPos. 162: T → CPos. 172: T → APos. 369: C → GapPos. 399: A → TPos. 483: C → APos. 1458: Gap → APos. 1649: Gap → TPos. 1842: CT → APos. 1883: G → CPos. 1948: A → TPos. 1951: T → APos. 2381: Gap → TPos. 2844: C → APos. 2866: Gap → APos. 3032: C → TPos. 3230: C → TPos. 3234: T → A28SPos. 249: Gap → CPos. 505: Gap → GPos. 783: Gap → CPos. 1049: C → GRAG-1Pos. 6 A → TPos. 65 T → CRhodopsinPos. 281: C → TPos. 296: C → TPos. 316: T → ASIAPos. 331: G → ATyrosinasePos. 160: C → TPos. 470: T → APos. 483: A → T
Hylinae
12SPos. 733: T → APos. 1296: T → CPos. 1409: A → GPos. 1426: T → C16SPos. 392: C → APos. 665: T → APos. 1237: A → GapPos. 1705: T → CPos. 1718: A → CPos. 2058: C → APos. 2875: A → TPos. 3173: A → GPos. 3239: T → APos. 3274: A → GPos. 3301: T → C28SPos. 863: Gap → GRAG-1Pos. 2: A → CPos. 53: G → A
Pos. 341: T → CPos. 410: C → ARhodopsinPos. 194: T → CSIAPos. 169: A → TTyrosinasePos. 73: T → APos. 150: T → CPos. 270: A → GPos. 300: C → APos. 324: A → GPos. 348: T → CPos. 376: G → TPos. 511: A → G
Cophomantini
Cytochrome bPos. 13: A → TPos. 153: C → TPos. 238: T → A12SPos. 159: C → APos. 297: T → CPos. 379: A → GPos. 436: T → CPos. 571: A → CTPos. 622: C → TPos. 636: G → APos. 716: A → CTPos. 729: T → CPos. 838: A → TPos. 1002: C → TPos. 1066: C → TPos. 1203: Gap → TPos. 1264: G → APos. 1266: A → TPos. 1332: G → APos. 1343: CT → GapPos. 1374: C → TPos. 1412: C → TPos. 1509: A → G16SPos. 294: A → CPos. 498: T → CPos. 876: A → GPos. 877: G → APos. 1049: T → CPos. 1228: A → CPos. 1402: A → GapPos. 1530: A → CPos. 1616: Gap → CPos. 1795: T → APos. 1874: T → CPos. 1925: A → CPos. 1998: A → CPos. 2605: G → APos. 2711: C → GapPos. 2736: A → GapPos. 2802: T → CPos. 2866: A → CPos. 3018: T → APos. 3027: T → GPos. 3328: A → T28SPos. 150: A → CPos. 152: C → TPos. 153: T → GPos. 178: G → APos. 211: Gap → C
Pos. 212: Gap → TPos. 230: Gap → APos. 302: G → APos. 343: Gap → APos. 344: Gap → GPos. 654: Gap → TPos. 1032: C → GRAG-1Pos. 357 C → TRhodopdsinPos. 93: C → TPos. 163: C → TSIAPos. 112: C → TPos. 304: T → CTyrosinasePos. 174: G → APos. 258: A → TPos. 508: T → GPos. 532: T → CAplastodiscus
Cytochrome bPos. 31: A → GPos. 79: A → CPos. 137: C → APos. 220: T → CPos. 310: C → A12SPos. 117: A → TPos. 152: A → GapPos. 174: G → APos. 206: A → TPos. 392: T → CPos. 414: C → TPos. 558: C → APos. 571: CT → APos. 655: Gap → APos. 823: A → TPos. 880: A → CPos. 1116: A → TPos. 1187: T → CPos. 1330: A → GPos. 1397: A → GapPos. 1408: A → GPos. 1427: T → CPos. 1431: T → CPos. 1539: C → TPos. 1573: A → GPos. 1590: Gap → AtRNA valinePos. 6: G → APos. 105: C → T16SPos. 376: A → TPos. 399: T → CPos. 664: A → TPos. 800: C → TPos. 803: C → TPos. 974: A → CPos. 984: T → APos. 1298: C → TPos. 1403: Gap → CPos. 1710: T → GPos. 1788: Gap → TPos. 1842: A → GapPos. 2019: A → TPos. 2030: A → GPos. 2105: A → CPos. 2130: C → TPos. 2240: A → G
Pos. 2280: A → TPos. 2289: A → GPos. 2301: G → APos. 2313: T → CPos. 2392: A → GapPos. 2398: A → TPos. 2551: A → GPos. 2563: T → CPos. 2820: A → CPos. 3032: T → CPos. 3148: G → APos. 3287: T → CPos. 3322: C → TPos. 3330: A → CPos. 3486: A → G28SPos. 390: C → TRAG-1Pos. 125 A → GPos. 284 A → GPos. 419 C → TSIAPos. 391: C → TTyrosinasePos. 6: G → APos. 130: C → TPos. 157: C → GPos. 233: C → TPos. 315: C → APos. 330: T → CPos. 513: G → A
Aplastodiscus albofrenatus group
Cytochrome bPos. 10: A → TPos. 23: T → CPos. 101: C → TPos. 109: A → GPos. 111: C → GPos. 132: A → GPos. 135: G → CPos. 239: C → TPos. 250: T → CPos. 259: C → TPos. 268: T → CPos. 290: C → TPos. 322: C → APos. 334: A → G12SPos. 37: T → CPos. 94: Gap → TPos. 218: Gap → TPos. 251: G → APos. 264: C → TPos. 333: C → TPos. 460: T → APos. 462: T → CPos. 484: A → TPos. 550: T → CPos. 587: T → CPos. 625: G → APos. 641: A → GPos. 654: T → CPos. 692: T → CPos. 696: A → GPos. 788: A → GPos. 802: T → CPos. 830: A → GPos. 835: T → CPos. 943: A → G
206 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 1036: C → GapPos. 1094: A → GPos. 1105: A → CPos. 1128: T → APos. 1141: T → CPos. 1211: T → CPos. 1217: T → CPos. 1402: A → CPos. 1413: C → APos. 1417: G → TPos. 1441: C → APos. 1496: T → CPos. 1674: A → TPos. 1688: A → GPos. 1762: A → GPos. 1763: T → CPos. 1775: T → CtRNA valinePos. 55: A → CPos. 56: T → C16SPos. 74: Gap → APos. 234: T → APos. 355: C → TPos. 483: A → TPos. 517: A → TPos. 566: A → TPos. 702: A → GPos. 763: G → APos. 808: T → CPos. 839: C → TPos. 864: C → TPos. 908: T → APos. 1021: T → APos. 1092: A → TPos. 1203: A → GapPos. 1211: A → GapPos. 1360: A → CPos. 1574: A → TPos. 1643: A → GPos. 1654: T → GapPos. 1688: T → CPos. 1799: C → APos. 1823: A → TPos. 1888: C → TPos. 1900: C → TPos. 1945: G → APos. 1962: A → GPos. 1981: Gap → TPos. 2059: A → CPos. 2285: A → TPos. 2308: C → TPos. 2406: T → APos. 2457: G → APos. 2515: A → TPos. 2545: G → APos. 2663: A → GapPos. 2664: A → GapPos. 2672: T → GapPos. 2685: A → GapPos. 2704: G → GapPos. 2742: T → GapPos. 2780: A → GapPos. 2844: A → GPos. 2875: A → GapPos. 2901: T → GapPos. 2906: T → GapPos. 2929: C → GapPos. 2946: G → TPos. 3010: A → TPos. 3054: CT → G
Pos. 3056: T → GPos. 3071: A → TPos. 3088: A → TPos. 3118: T → CPos. 3145: C → APos. 3310: A → TPos. 3516: T → GRAG-1Pos. 428 A → TSIAPos. 280: C → TPos. 349: T → A
Aplastodiscus albosignatus group
Cytochrome bPos. 49: A → CPos. 339: A → CPos. 376: T → APos. 405: CG → T12SPos. 227: T → CPos. 452: T → APos. 521: A → TPos. 611: A → GPos. 875: C → APos. 908: A → TPos. 958: A → GPos. 1048: T → APos. 1579: C → TPos. 1589: C → APos. 1645: C → TPos. 1674: A → C16SPos. 94: C → TPos. 308: A → GapPos. 378: Gap → CPos. 419: Gap → APos. 648: G → TPos. 667: A → CPos. 668: CT → APos. 780: A → GPos. 887: C → TPos. 985: G → APos. 1049: C → TPos. 1174: T → APos. 1278: C → GPos. 1503: C → APos. 1742: Gap → CPos. 1780: T → CPos. 1795: A → CPos. 2029: T → APos. 2115: T → APos. 2195: T → CPos. 2400: Gap → GPos. 2663: A → TPos. 3299: C → TPos. 3321: A → TTyrosinasePos. 25: T → APos. 166: G → AAplastodiscus perviridis group
Cytochrome bPos. 46: C → TPos. 64: T → APos. 161: A → GPos. 183: A → TPos. 189: C → TPos. 201: T → CPos. 229: C → TPos. 362: T → APos. 399: C → T
12SPos. 103: Gap → GPos. 132: Gap → CPos. 164: T → GapPos. 169: A → GPos. 310: T → CPos. 368: A → GPos. 615: T → APos. 646: T → APos. 650: T → CPos. 880: C → TPos. 1036: C → TPos. 1326: G → APos. 1392: C → TPos. 1626: Gap → GtRNA valinePos. 58: T → C16SPos. 136: A → GapPos. 336: C → TPos. 589: T → CPos. 618: A → GPos. 636: T → CPos. 656: T → CPos. 749: C → TPos. 876: G → APos. 968: G → APos. 979: C → TPos. 1277: G → APos. 1286: A → GPos. 1314: A → GPos. 1649: C → TPos. 1718: C → TPos. 1788: T → CPos. 1799: C → TPos. 1819: A → TPos. 1953: T → CPos. 2364: A → GapPos. 2381: T → CPos. 2552: T → CPos. 2616: A → TPos. 2802: C → GapPos. 2821: Gap → TPos. 3004: A → GapPos. 3077: T → CPos. 3181: G → APos. 3266: C → TPos. 3281: A → GPos. 3297: A → CPos. 3516: T → CRAG-1Pos. 311 C → TTyrosinasePos. 438: C → T
Bokermannohyla
Cytochrome bPos. 116: C → TPos. 361: T → GapPos. 370: Gap → APos. 376: T → C12SPos. 140: C → GapPos. 171: G → GapPos. 181: Gap → TPos. 322: C → TPos. 729: C → TPos. 890: A → CPos. 943: A → GPos. 1084: T → CPos. 1145: T → A
Pos. 1243: A → CTPos. 1338: Gap → CPos. 1342: A → GPos. 1366: C → GapPos. 1369: T → GPos. 1568: A → TPos. 1599: A → GPos. 1630: A → CtRNA valinePos. 78: T → APos. 109: T → C16SPos. 0: G → APos. 166: Gap → APos. 443: C → ATPos. 731: C → GapPos. 749: C → TPos. 829: C → TPos. 860: G → APos. 1041: Gap → CPos. 1103: T → GapPos. 1129: G → APos. 1330: C → APos. 1436: A → GapPos. 1757: T → CPos. 1783: C → TPos. 1865: T → APos. 1985: A → CPos. 1998: C → GPos. 2041: A → TPos. 2059: A → TPos. 2570: A → GPos. 2618: T → GapPos. 2694: T → CPos. 2697: T → CPos. 2966: A → TPos. 3027: G → APos. 3173: G → GapPos. 3324: T → CPos. 3330: A → GapPos. 3425: C → T28SPos. 443: G → GapRAG-1Pos. 251 G → APos. 380 T → CRhodopsinPos. 78: G → CPos. 93: T → APos. 220: G → ATyrosinasePos. 31: A → GPos. 300: A → GPos. 315: C → GPos. 357: T → CPos. 387: T → CPos. 403: G → APos. 506: C → GBokermannohyla circumdata group
Cytochrome bPos. 49: A → CPos. 109: A → GPos. 120: A → GPos. 201: A → CPos. 330: C → TPos. 334: A → TPos. 356: T → CPos. 367: C → TPos. 399: C → T12SPos. 33: G → A
2005 207FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 69: C → TPos. 131: T → CPos. 227: T → CPos. 370: A → TPos. 512: T → CPos. 547: A → TPos. 729: T → APos. 823: A → GPos. 869: C → TPos. 1024: C → TPos. 1144: T → GPos. 1272: G → APos. 1348: T → CPos. 1408: A → GPos. 1427: T → CPos. 1645: A → C16SPos. 11: T → CPos. 411: C → TPos. 498: C → TPos. 630: T → APos. 692: A → GPos. 726: A → GPos. 751: C → APos. 887: T → APos. 1049: C → TPos. 1085: T → CPos. 1492: T → APos. 1649: T → APos. 1654: T → APos. 1688: T → CPos. 1880: A → CPos. 2110: C → TPos. 2176: C → TPos. 2209: Gap → CPos. 2229: C → TPos. 2611: Gap → APos. 2825: T → APos. 2866: C → TPos. 3222: T → CPos. 3516: T → C28SPos. 240: Gap → CPos. 498: Gap → G
Bokermannohyla pseudopseudisgroup
Cytochrome bPos. 76: A → GPos. 111: C → TPos. 135: G → APos. 140: A → TPos. 253: C → TPos. 334: A → CPos. 347: A → TPos. 374: C → TPos. 381: C → T12SPos. 83: A → GPos. 159: A → TPos. 164: T → CPos. 297: C → TPos. 370: A → GPos. 452: T → CPos. 615: T → CPos. 1048: T → APos. 1181: Gap → APos. 1317: C → APos. 1326: G → A16SPos. 285: Gap → C
Pos. 286: Gap → CPos. 691: G → APos. 702: A → GPos. 774: T → APos. 960: G → APos. 974: A → TPos. 1140: A → TPos. 1607: A → GPos. 1710: T → GPos. 1953: T → APos. 2025: C → TPos. 2229: C → APos. 2250: C → TPos. 2262: G → APos. 2330: C → TPos. 2473: T → APos. 2518: A → GPos. 2608: Gap → TPos. 2646: G → APos. 2856: T → APos. 3097: C → APos. 3380: T → CPos. 3381: T → APos. 3403: C → TRAG-1Pos. 365: T → CPos. 422: T → CPos. 428: A → C
Hyloscirtus
Cytochrome bPos. 14: A → CTPos. 101: C → AGPos. 134: C → TPos. 302: A → CPos. 322: C → A12SPos. 460: T → CPos. 462: T → APos. 490: Gap → CPos. 565: A → GapPos. 636: A → TPos. 817: G → APos. 842: Gap → CPos. 1174: C → APos. 1211: T → APos. 1440: C → APos. 1532: Gap → CTtRNA valinePos. 82: C → TPos. 89: A → CPos. 99: G → A16SPos. 254: T → GapPos. 272: T → ACPos. 325: Gap → TPos. 427: T → APos. 576: T → CPos. 602: A → CPos. 854: A → TPos. 889: A → GPos. 914: C → APos. 949: T → CPos. 1062: A → GapPos. 1192: Gap → CPos. 1577: A → TPos. 1587: T → APos. 1616: C → APos. 1631: T → GPos. 1673: AC → GapPos. 1705: C → A
Pos. 1718: C → GapPos. 1985: A → TPos. 2258: A → TPos. 2544: G → APos. 2756: Gap → TPos. 3140: C → APos. 3288: A → TPos. 3326: T → C28SPos. 303: Gap → TPos. 346: Gap → CPos. 391: Gap → TPos. 744: Gap → TRAG-1Pos. 257 C → TPos. 338 C → TPos. 425 G → ARhodopdsinPos. 256: C → TTyrosinasePos. 3: C → TPos. 72: C → TPos. 257: C → T
Hyloscirtus armatus group
Cytochrome bPos. 3: C → TPos. 23: T → CPos. 67: C → TPos. 127: C → TPos. 135: G → APos. 167: C → TPos. 178: C → TPos. 189: C → TPos. 248: C → APos. 290: C → APos. 369: T → C12SPos. 22: G → APos. 26: A → GPos. 176: A → TPos. 194: T → CPos. 227: T → CPos. 411: T → CPos. 414: C → TPos. 455: AC → TPos. 547: A → TPos. 558: C → TPos. 793: T → CPos. 835: T → CPos. 869: C → TPos. 1043: C → TPos. 1084: T → CPos. 1095: C → TPos. 1100: G → APos. 1101: C → TPos. 1141: C → TPos. 1153: T → GPos. 1187: T → CPos. 1203: T → APos. 1303: A → TPos. 1446: A → CPos. 1515: A → TPos. 1574: Gap → APos. 1575: Gap → CPos. 1630: A → TPos. 1636: C → APos. 1649: A → CtRNA valinePos. 95: C → T16S
Pos. 139: Gap → APos. 175: Gap → CPos. 220: A → TPos. 244: A → TPos. 498: C → TPos. 543: T → CPos. 726: A → TPos. 731: C → TPos. 767: A → CPos. 822: A → TPos. 862: C → TPos. 996: A → TPos. 1104: Gap → APos. 1140: AC → TPos. 1175: Gap → APos. 1491: A → TPos. 1503: C → APos. 1534: A → CPos. 1572: T → CPos. 1688: C → APos. 1780: T → CPos. 1865: T → APos. 1894: T → APos. 1897: T → CPos. 1928: C → APos. 1949: T → APos. 1961: A → GPos. 2122: G → APos. 2308: T → APos. 2432: A → GPos. 2461: C → TPos. 2476: C → TPos. 2552: T → CPos. 3059: A → TPos. 3140: A → TPos. 3280: A → CPos. 3281: A → TPos. 3288: T → CPos. 3375: T → APos. 3431: C → TPos. 3442: C → T28SPos. 302: A → GPos. 1000: G → GapRhodopdsinPos. 63: G → APos. 135: C → TPos. 270: A → TPos. 279: C → ATyrosinasePos. 47: A → GPos. 52: C → TPos. 56: A → CPos. 109: C → TPos. 167: G → TPos. 170: A → CPos. 188: G → APos. 258: T → APos. 273: C → GPos. 279: C → TPos. 297: C → TPos. 343: G → APos. 344: T → CPos. 529: C → T
Hyloscirtus bogotensis group
Cytochrome bPos. 126: G → APos. 153: T → CPos. 168: A → TPos. 220: T → C
208 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 232: C → APos. 253: C → TPos. 338: T → CPos. 376: T → A12SPos. 28: A → GPos. 45: T → CPos. 68: A → GPos. 110: A → GapPos. 152: A → GPos. 164: T → CPos. 295: A → CPos. 447: Gap → CPos. 550: T → CPos. 619: G → TPos. 622: T → APos. 650: T → CPos. 767: C → TPos. 991: Gap → APos. 1094: A → GPos. 1118: C → TPos. 1270: A → GPos. 1307: T → CPos. 1509: G → APos. 1521: G → APos. 1568: A → TPos. 1657: G → AtRNA valinePos. 55: A → GPos. 80: T → GapPos. 85: G → GapPos. 90: A → GapPos. 98: A → G16SPos. 12: C → TPos. 107: A → GapPos. 392: A → CPos. 411: C → GapPos. 452: T → GapPos. 665: A → GPos. 691: G → APos. 876: G → APos. 891: G → APos. 904: A → TPos. 947: C → TPos. 952: A → TPos. 969: A → GPos. 1049: C → GapPos. 1480: T → CPos. 1534: A → TPos. 1780: T → APos. 1819: A → GapPos. 1948: T → CPos. 1998: C → APos. 2041: A → TPos. 2125: T → CPos. 2457: G → APos. 2533: T → CPos. 2663: A → GapPos. 2664: A → GapPos. 2672: C → GapPos. 2719: CT → APos. 2820: A → GapPos. 2856: T → GPos. 2966: A → CPos. 2990: A → CPos. 3039: A → TPos. 3048: A → GapPos. 3063: G → APos. 3106: T → CPos. 3385: T → C
Pos. 3458: T → CPos. 3524: C → TPos. 3538: G → ARAG-1Pos. 4 C → APos. 5 A → GPos. 143 G → APos. 168 G → APos. 169 T → CPos. 170 G → APos. 305 C → TPos. 311 C → TPos. 323 T → CRhodopdsinPos. 133: C → TPos. 135: C → APos. 291: G → APos. 308: T → ATyrosinasePos. 183: A → GPos. 214: C → TPos. 220: A → GPos. 294: C → TPos. 299: A → CPos. 315: C → TPos. 342: C → T
Hyloscirtus larinopygion group
12SPos. 65: C → TPos. 454: T → CPos. 490: C → TPos. 601: T → CPos. 908: A → TPos. 1113: T → CPos. 1116: A → TPos. 1174: A → TPos. 1216: C → APos. 1321: T → CPos. 1407: G → APos. 1409: G → APos. 1426: C → TPos. 1429: C → TPos. 1678: A → GtRNA valinePos. 12: C → T16SPos. 347: C → TPos. 742: T → APos. 819: T → 4Pos. 911: A → GPos. 920: G → APos. 1092: A → TPos. 1228: C → 4Pos. 1265: A → GPos. 1325: A → CPos. 1510: A → GPos. 1561: C → TPos. 2103: A → TPos. 2395: A → CPos. 2450: T → CPos. 2768: CT → APos. 3287: T → C
Hypsiboas
Cytochrome bPos. 376: T → A12SPos. 45: T → CPos. 476: C → APos. 650: T → APos. 958: A → G
Pos. 1048: T → APos. 1107: C → TPos. 1407: G → APos. 1429: C → T16SPos. 163: Gap → TPos. 188: A → TPos. 272: T → APos. 876: G → APos. 996: A → GPos. 1169: Gap → TPos. 1692: Gap → APos. 1894: T → CPos. 1978: Gap → APos. 2182: Gap → APos. 2217: T → GapPos. 2812: T → CPos. 3086: A → TPos. 3166: C → APos. 3239: A → CPos. 3249: A → T28SPos. 355: Gap → CPos. 358: Gap → CPos. 57: C → TPos. 373: C → TTyrosinasePos. 257: C → TPos. 288: C → TPos. 407: T → A
Hypsiboas albopunctatus group
Cytochrome bPos. 63: T → CPos. 76: A → TPos. 109: A → GPos. 126: C → APos. 128: T → CPos. 153: T → CPos. 180: C → APos. 253: C → TPos. 271: T → CPos. 327: A → TPos. 415: C → T12SPos. 131: T → CPos. 227: T → CPos. 650: A → CPos. 1045: T → APos. 1158: Gap → CPos. 1159: Gap → CPos. 1217: T → CPos. 1376: T → CPos. 1573: A → TPos. 1670: C → T16SPos. 107: A → GapPos. 173: Gap → APos. 308: A → GapPos. 998: C → TPos. 1092: A → TPos. 1125: A → GPos. 1129: G → APos. 1183: A → GapPos. 1300: A → GPos. 1419: T → APos. 1737: C → APos. 1757: C → TPos. 1810: T → APos. 1840: Gap → TPos. 2187: A → T
Pos. 2203: A → TPos. 2694: T → CPos. 2846: Gap → CPos. 3054: T → CPos. 3189: G → APos. 3222: T → GapPos. 3516: T → A
Hypsiboas faber group
Cytochrome bPos. 173: A → TPos.186: A → G12SPos. 131: T → C16SPos. 11: T → CPos. 206: A → TPos. 347: C → APos. 367: C → APos. 573: A → TPos. 663: A → CPos. 775: C → TPos. 780: C → APos. 821: T → CPos. 904: T → APos. 1421: Gap → APos. 1458: A → TPos. 1510: A → GapPos. 1673: A → TPos. 1874: T → CPos. 2381: T → APos. 2602: G → TPos. 2663: A → TPos. 2820: A → CPos. 2977: A → TPos. 2995: A → CPos. 3081: A → TPos. 3351: Gap → ARAG-1Pos. 1 C → ASIAPos. 283: G → A
Hypsiboas pellucens group
12SPos. 42: T → CPos. 183: A → TPos. 297: C → TPos. 319: A → TPos. 320: A → TPos. 342: T → CPos. 392: T → CPos. 455: C → TPos. 541: T → CPos. 550: T → CPos. 641: A → TPos. 646: T → APos. 661: G → APos. 673: C → TPos. 716: C → GapPos. 731: Gap → APos. 767: T → CPos. 841: A → TPos. 1023: T → CPos. 1122: A → TPos. 1187: T → CPos. 1203: T → APos. 1214: Gap → APos. 1233: G → APos. 1316: A → GPos. 1321: T → C
2005 209FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 1337: T → CPos. 1369: T → GPos. 1392: C → TPos. 1441: A → CPos. 1445: C → TPos. 1549: A → TPos. 1579: C → TPos. 1605: A → TPos. 1630: A → CtRNA valinePos. 101: A → G16SPos. 12: C → TPos. 136: A → TPos. 206: A → GapPos. 334: Gap → APos. 498: C → APos. 573: A → TPos. 610: A → TPos. 691: G → APos. 736: C → APos. 761: T → CPos. 790: G → APos. 826: A → GPos. 836: A → CPos. 839: C → TPos. 892: G → APos. 904: T → APos. 911: A → GPos. 928: T → GPos. 946: C → TPos. 980: T → CPos. 1013: T → APos. 1062: C → TPos. 1173: Gap → CPos. 1235: C → APos. 1245: Gap → GPos. 1261: T → CPos. 1345: T → APos. 1389: A → TPos. 1458: A → GapPos. 1468: A → TPos. 1480: A → TPos. 1510: A → CPos. 1532: T → CPos. 1607: A → GapPos. 1628: Gap → TPos. 1641: Gap → APos. 1727: A → TPos. 1813: A → CPos. 1847: A → CPos. 1932: C → APos. 1953: A → TPos. 2029: T → CPos. 2030: G → APos. 2035: C → APos. 2077: A → TPos. 2086: C → TPos. 2091: A → CPos. 2094: A → CPos. 2109: Gap → APos. 2117: A → GapPos. 2267: T → APos. 2308: C → TPos. 2450: T → CPos. 2510: T → CPos. 2517: G → APos. 2518: A → GPos. 2525: A → TPos. 2538: C → TPos. 2569: T → C
Pos. 2583: A → GPos. 2613: T → APos. 2618: T → APos. 2656: A → GPos. 2719: C → TPos. 2906: T → CPos. 2953: A → GPos. 2956: A → TPos. 3041: G → APos. 3077: T → APos. 3093: T → CPos. 3111: A → GPos. 3166: A → CPos. 3234: T → APos. 3249: T → CPos. 3285: G → APos. 3286: C → TPos. 3426: A → TPos. 3454: T → GPos. 3518: Gap → T
Hypsiboas pulchellus group
Cytochrome bPos. 26: T → APos. 126: C → TPos. 145: C → TPos. 214: C → TPos. 330: C → TPos. 347: A → C12SPos. 180: C → TPos. 601: T → APos. 650: A → TPos. 1317: A → TPos. 1423: A → TPos. 1645: C → TtRNA valinePos. 4: A → TPos. 101: A → G16SPos. 71: A → CPos. 130: T → CPos. 254: T → APos. 566: A → TPos. 602: A → TPos. 667: A → CPos. 702: A → TPos. 732: A → GPos. 788: C → TPos. 1169: T → APos. 1235: C → APos. 1315: T → APos. 1436: A → TPos. 1468: A → CPos. 1572: T → APos. 1579: Gap → CPos. 1607: A → TPos. 1705: T → GPos. 1810: T → CPos. 1813: A → CPos. 1878: C → TPos. 1887: C → TPos. 1952: T → APos. 2025: C → TPos. 2089: A → TPos. 2262: G → APos. 2588: C → TPos. 2754: Gap → CPos. 2825: T → APos. 2856: T → APos. 2866: C → A
Pos. 2985: T → APos. 3239: C → GapRhodopsinPos. 10: G → ATyrosinasePos. 91: C → TPos. 154: C → TPos. 177: C → TPos. 178: C → GPos. 182: G → TPos. 211: C → GPos. 444: G → A
Hypsiboas punctatus group
Cytochrome bPos. 49: A → CPos. 109: A → CPos. 192: T → CPos. 253: C → T12SPos. 268: T → CPos. 1045: T → GapPos. 1317: C → T16SPos. 56: C → GapPos. 272: A → GPos. 294: C → TPos. 543: T → CPos. 920: G → APos. 1103: T → GapPos. 1259: CT → APos. 1654: T → CPos. 1878: C → TPos. 1887: C → TPos. 2308: C → TPos. 2719: C → GapPos. 2742: T → GapPos. 2753: A → GapPos. 2768: C → GapPos. 2906: T → APos. 2975: C → GapPos. 2982: C → GapPos. 2995: A → GapPos. 3054: T → APos. 3140: C → TPos. 3222: T → GapPos. 3516: T → A
Hypsiboas semilineatus group
Cytochrome bPos. 1: A → TPos. 46: C → TPos. 63: T → CPos. 76: A → TPos. 153: T → CPos. 167: C → TPos. 180: C → TPos. 181: T → APos. 195: A → CPos. 232: C → TPos. 280: C → TPos. 290: C → APos. 347: A → CPos. 353: T → CPos. 374: C → TPos. 381: C → T12SPos. 33: G → APos. 271: T → APos. 332: C → APos. 348: T → APos. 370: A → C
Pos. 414: C → TPos. 528: A → CPos. 602: A → TPos. 619: G → APos. 650: A → TPos. 673: C → TPos. 677: A → TPos. 765: A → GPos. 774: A → GPos. 797: G → APos. 815: T → CPos. 836: T → CPos. 875: C → TPos. 1001: A → CPos. 1215: A → TPos. 1330: A → TPos. 1336: C → TPos. 1372: G → APos. 1378: A → TPos. 1392: C → TPos. 1413: C → TPos. 1439: C → APos. 1670: C → AtRNA valinePos. 94: C → APos. 95: C → T16SPos. 134: Gap → APos. 211: Gap → TPos. 294: C → APos. 323: C → APos. 472: A → GapPos. 579: A → TPos. 580: A → TPos. 606: C → APos. 611: A → GPos. 654: T → APos. 679: C → TPos. 732: A → GPos. 819: T → GapPos. 911: A → GPos. 914: C → TPos. 1004: T → GPos. 1021: T → APos. 1160: C → APos. 1242: T → APos. 1246: T → APos. 1325: A → CPos. 1436: A → TPos. 1510: A → TPos. 1530: C → TPos. 1596: A → TPos. 1616: C → APos. 1692: A → CPos. 1727: A → CPos. 1799: C → APos. 1847: A → TPos. 1895: A → TPos. 1922: C → APos. 1950: T → APos. 1970: A → TPos. 2035: C → APos. 2086: C → TPos. 2089: A → CPos. 2125: T → APos. 2156: C → APos. 2182: A → TPos. 2229: C → APos. 2310: A → CPos. 2313: T → APos. 2406: T → Gap
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APPENDIX 5(Continued)
Pos. 2441: C → APos. 2454: A → TPos. 2457: G → APos. 2551: A → CPos. 2672: T → GapPos. 2856: T → APos. 2929: C → GapPos. 2953: A → GapPos. 3014: A → GPos. 3125: A → TPos. 3127: C → TPos. 3199: C → TPos. 3249: T → GapPos. 3291: A → GPos. 3330: A → T28SPos. 355: C → GapPos. 431: G → GapRhodopsinPos. 155: T → CPos. 270: A → TPos. 271: G → CPos. 308: T → CSIAPos. 151: C → TPos. 241: G → CPos. 253: T → CPos. 268: A → GPos. 313: G → APos. 322: C → TTyrosinasePos. 70: T → GPos. 96: G → APos. 156: C → TPos. 174: A → GPos. 260: C → APos. 262: T → CPos. 299: A → CPos. 417: C → GPos. 429: C → TPos. 475: C → TPos. 514: G → A
Myersiohyla
12SPos. 39: A → GPos. 271: T → CPos. 342: T → CPos. 356: T → CPos. 392: T → CPos. 452: T → CPos. 547: A → CPos. 615: T → APos. 672: C → APos. 835: T → CPos. 1043: C → TPos. 1645: A → GapPos. 1674: A → CtRNA valinePos. 56: T → C16SPos. 94: C → APos. 272: T → GPos. 392: A → TPos. 434: A → TPos. 517: A → CPos. 606: C → TPos. 777: A → CPos. 848: A → TPos. 1043: Gap → CPos. 1062: A → C
Pos. 1160: T → APos. 1278: T → CPos. 1614: A → CPos. 1865: T → CPos. 1951: A → TPos. 2127: G → APos. 2203: A → TPos. 2392: A → CPos. 2398: A → CPos. 2449: T → APos. 2509: C → TPos. 2566: T → CPos. 2618: T → APos. 2780: A → GapPos. 2812: T → CPos. 2856: T → CPos. 2906: T → CPos. 2997: C → GapPos. 3041: G → GapPos. 3056: T → CPos. 3106: T → CPos. 3222: T → CPos. 3287: T → CPos. 3385: T → C
Dendropsophini 1 Hylini 1
Lophiohylini
Cytochrome bPos. 11: A → CPos. 376: T → A12SPos. 1174: C → TPos. 1440: C → APos. 1670: T → C16SPos. 107: A → TPos. 399: T → CPos. 853: T → APos. 854: A → TPos. 914: C → APos. 1062: A → TPos. 1436: A → TPos. 1468: T → APos. 1847: A → GPos. 2267: T → APos. 2844: A → TPos. 2966: A → T28SPos. 233: G → GapRAG-1Pos. 5 A → GPos. 125 A → GRhodopsinPos. 10: A → GPos. 105: G → APos. 316: A → GTyrosinasePos. 210: G → APos. 407: T → APos. 408: T → APos. 506: C → G
Dendropsophini
Cytochrome bPos. 137: T → CPos. 201: A → TPos. 220: T → C12SPos. 451: T → CPos. 547: A → CPos. 641: A → GapPos. 841: A → C
Pos. 869: C → T16SPos. 162: C → TPos. 259: Gap → APos. 1799: A → CPos. 2025: C → TPos. 2680: A → TPos. 2820: A → GapPos. 3173: G → GapPos. 3230: T → APos. 3326: T → C28SPos. 400: T → CPos. 419: C → GPos. 1014: G → GapRhodopsinPos. 256: C → TTyrosinasePos. 145: T → CPos. 303: A → G
Dendropsophus
Cytochrome bPos. 3: C → APos. 183: A → TPos. 253: T → CPos. 369: T → ACPos. 414: T → C12SPos. 636: G → APos. 869: T → CPos. 1008: T → CPos. 1267: A → TPos. 1402: A → GPos. 1407: G → APos. 1429: C → T16SPos. 299: Gap → TPos. 392: A → CPos. 551: T → GapPos. 573: A → TPos. 579: A → TPos. 686: G → APos. 761: T → CPos. 777: A → CPos. 876: A → GPos. 877: G → APos. 979: C → TPos. 1469: Gap → TPos. 1949: A → CPos. 2080: A → GPos. 2156: A → CPos. 2217: T → CPos. 2244: T → CPos. 2583: A → GPos. 2802: T → CPos. 3048: T → CPos. 3230: A → C
Dendropsophus leucophyllatus group
Cytochrome bPos. 36: T → CPos. 71: C → APos. 116: C → TPos. 125: C → TPos. 134: C → TPos. 153: C → TPos. 173: A → TPos. 253: C → TPos. 405: A → C12SPos. 37: T → C
Pos. 159: C → TPos. 678: Gap → CPos. 716: A → CPos. 729: C → TPos. 838: T → APos. 897: A → CPos. 943: A → GPos. 965: T → APos. 1011: A → TPos. 1094: T → CPos. 1270: A → GPos. 1273: A → GPos. 1402: G → APos. 1581: Gap → APos. 1675: A → TPos. 1678: T → C16SPos. 1308: T → CPos. 1480: T → CPos. 1491: T → CPos. 1878: C → TPos. 1922: C → APos. 2086: C → TPos. 2091: A → TPos. 2105: A → TPos. 2107: C → TPos. 2378: Gap → APos. 2618: T → APos. 2652: T → CPos. 2780: T → CPos. 2859: Gap → GPos. 2866: A → TPos. 2982: A → CPos. 3059: A → TPos. 3166: C → TPos. 3181: G → APos. 3268: C → TPos. 3287: T → CPos. 3326: C → APos. 3452: A → G28SPos. 295: Gap → GPos. 330: Gap → CPos. 413: Gap → GPos. 544: T → GPos. 569: A → CPos. 763: Gap → CPos. 764: Gap → CPos. 974: Gap → CPos. 975: Gap → CPos. 1025: Gap → GPos. 1032: C → GRAG-1Pos. 77 T → CPos. 215 G → TPos. 245 C → TRhodopsinPos. 115: T → CPos. 262: G → APos. 281: C → TSIAPos. 112: C → TPos. 113: C → TPos. 175: C → TTyrosinasePos. 73: G → APos. 120: C → TPos. 121: C → TPos. 170: A → CPos. 174: A → GPos. 382: A → C
2005 211FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 507: T → C
Dendropsophus marmoratus group
Cytochrome bPos. 11: C → APos. 97: A → TPos. 103: C → TPos. 109: G → APos. 204: C → APos. 220: C → TPos. 311: A → C12SPos. 10: A → GPos. 28: A → GPos. 44: A → GPos. 145: C → TPos. 173: G → APos. 319: A → TPos. 347: T → CPos. 370: A → GPos. 528: A → CPos. 651: C → TPos. 672: C → APos. 769: G → APos. 821: C → TPos. 1233: G → APos. 1509: A → GPos. 1539: T → CPos. 1561: A → TtRNA valinePos. 95: C → T16SPos. 188: A → GapPos. 206: C → APos. 259: A → GapPos. 347: A → CPos. 443: C → TPos. 580: A → GPos. 606: C → APos. 625: T → CPos. 660: Gap → GPos. 667: A → CPos. 668: T → CPos. 767: A → CPos. 818: C → GapPos. 996: A → GPos. 1015: T → APos. 1826: C → APos. 1839: T → APos. 1894: T → CPos. 1895: A → GPos. 2161: A → CPos. 2229: T → APos. 2250: C → TPos. 2268: C → APos. 2313: T → CPos. 2356: Gap → APos. 2381: T → CPos. 2551: A → GPos. 2652: T → CPos. 2663: T → CPos. 2704: G → APos. 2812: T → APos. 2875: C → APos. 2889: A → GPos. 2966: T → CPos. 3046: C → TPos. 3054: T → APos. 3056: T → GPos. 3118: C → APos. 3130: A → T
Pos. 3326: C → TPos. 3425: C → TPos. 3455: G → APos. 3485: T → CPos. 3491: C → TPos. 3549: A → GRAG-1Pos. 17 A → GPos. 77 T → CPos. 230 C → T
Dendropsophus microcephalus group
Cytochrome bPos. 140: A → CPos. 241: A → TPos. 277: T → CPos. 330: T → CPos. 334: A → GPos. 381: C → T12SPos. 140: C → TPos. 338: T → CPos. 427: T → CPos. 515: C → GapPos. 553: Gap → CPos. 654: T → CPos. 875: C → T16SPos. 192: C → APos. 254: T → GapPos. 299: T → APos. 392: C → APos. 443: C → APos. 464: Gap → CPos. 573: T → CPos. 580: A → CPos. 914: A → CPos. 1183: A → GapPos. 1402: A → TPos. 1727: T → APos. 1737: T → CPos. 1839: CT → APos. 1880: CT → APos. 1950: T → APos. 1951: A → TPos. 2080: G → APos. 2112: T → CPos. 2151: T → CPos. 2250: C → TPos. 2663: T → APos. 2713: T → GapPos. 2719: T → CPos. 2736: A → CPos. 2966: T → APos. 3032: T → APos. 3051: Gap → APos. 3054: T → A
Dendropsophus parviceps group
Cytochrome bPos. 76: A → CPos. 105: A → CPos. 244: A → TPos. 331: C → TPos. 420: A → C12SPos. 28: A → GPos. 142: C → TPos. 174: G → APos. 528: A → CPos. 1630: A → CPos. 1640: T → A
16SPos. 220: C → GPos. 427: T → APos. 876: G → APos. 980: C → TPos. 1176: Gap → CPos. 1281: T → CPos. 1423: Gap → APos. 1491: T → GapPos. 1519: T → CPos. 1532: A → CPos. 2151: T → APos. 2376: C → TPos. 2618: T → APos. 3313: T → CTyrosinasePos. 438: C → TPos. 471: C → T
Lysapsus
Cytochrome bPos. 111: C → APos. 149: A → CPos. 180: C → APos. 238: CT → APos. 265: C → APos. 274: C → TPos. 334: A → GPos. 420: AC → T12SPos. 10: G → APos. 115: A → TPos. 148: C → APos. 615: T → CPos. 716: A → GPos. 733: A → TPos. 890: A → TPos. 1561: T → C16SPos. 254: T → GapPos. 370: Gap → CPos. 580: A → GPos. 848: A → TPos. 1077: C → APos. 1468: C → TPos. 1705: C → GapPos. 1744: Gap → APos. 1824: C → TPos. 1826: C → APos. 2192: Gap → CPos. 2229: T → CPos. 2395: T → CPos. 2831: Gap → APos. 2913: T → APos. 2966: T → APos. 2982: T → APos. 3081: A → GPos. 3266: C → TPos. 3280: A → CPos. 3297: T → ARAG-1Pos. 34 C → TPos. 149 A → CPos. 164 G → APos. 221 T → CPos. 326 G → APos. 392 T → GRhodopsinPos. 87: A → GPos. 185: C → ASIA
Pos. 289: C → TPos. 331: A → G
Pseudis
Cytochrome bPos. 119: A → CPos. 192: T → CPos. 290: C → APos. 369: T → APos. 399: C → T12SPos. 36: T → GPos. 227: T → CPos. 235: AT → CPos. 452: T → C16SPos. 162: T → CPos. 236: Gap → APos. 434: T → CPos. 517: A → GPos. 854: T → CPos. 1300: A → GPos. 1928: CT → APos. 1953: T → APos. 1955: A → CPos. 2091: T → CPos. 2127: G → APos. 2179: C → TPos. 2240: A → GPos. 2697: T → APos. 2768: C → TRAG-1Pos. 5 G → APos. 20 G → APos. 169 T → APos. 251 G → A
Scarthyla goinorum
Cytochrome bPos. 6: C → TPos. 10: A → TPos. 39: T → CPos. 49: C → TPos. 52: A → CPos. 57: C → APos. 67: C → TPos. 68: C → TPos. 91: T → CPos. 108: T → GPos. 145: C → TPos. 158: A → TPos. 164: C → TPos. 189: C → TPos. 201: T → APos. 202: G → APos. 204: C → APos. 211: T → CPos. 262: A → GPos. 271: C → TPos. 292: T → CPos. 311: A → CPos. 319: C → TPos. 331: C → TPos. 376: A → CPos. 415: C → T12SPos. 13: G → APos. 22: G → APos. 24: A → CPos. 28: A → GPos. 33: A → GPos. 61: C → T
212 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 98: T → CPos. 117: A → CPos. 159: C → APos. 169: A → TPos. 183: A → GPos. 231: C → TPos. 295: A → CPos. 320: A → GPos. 452: T → GPos. 454: T → CPos. 550: T → CPos. 571: A → TPos. 600: A → TPos. 646: T → APos. 650: T → CPos. 675: A → TPos. 716: A → CPos. 761: T → APos. 814: C → TPos. 838: A → TPos. 900: T → GPos. 1010: T → CPos. 1026: T → CPos. 1084: T → CPos. 1122: T → CPos. 1145: T → CPos. 1211: CT → APos. 1266: A → GPos. 1270: A → GPos. 1332: G → APos. 1342: A → GapPos. 1366: C → APos. 1374: C → TPos. 1392: T → CPos. 1405: A → GPos. 1431: T → CPos. 1439: C → TPos. 1517: A → TPos. 1521: G → APos. 1531: C → TPos. 1579: A → TPos. 1589: T → APos. 1615: Gap → TPos. 1657: G → GapPos. 1659: C → TPos. 1762: A → GtRNA valinePos. 22: Gap → TPos. 39: C → TPos. 82: C → TPos. 99: G → A16SPos. 3: A → CPos. 56: C → GapPos. 254: T → APos. 301: Gap → CPos. 355: C → TPos. 383: T → APos. 427: T → CPos. 452: T → CPos. 483: A → CPos. 634: A → TPos. 636: T → CPos. 667: A → GPos. 668: T → CPos. 686: G → APos. 689: A → TPos. 809: T → CPos. 814: T → APos. 844: A → CPos. 848: A → C
Pos. 853: A → TPos. 854: T → APos. 892: G → APos. 946: C → TPos. 952: A → CPos. 996: A → GPos. 1057: T → GapPos. 1062: T → GPos. 1085: T → CPos. 1092: T → CPos. 1103: T → CPos. 1113: A → CPos. 1118: A → CPos. 1125: T → CPos. 1188: Gap → TPos. 1189: Gap → TPos. 1200: A → TPos. 1211: A → TPos. 1235: C → APos. 1247: Gap → APos. 1248: Gap → GPos. 1249: Gap → APos. 1278: T → CPos. 1329: Gap → CPos. 1332: A → CPos. 1337: T → GapPos. 1424: Gap → CPos. 1436: T → GPos. 1451: T → GPos. 1480: T → GapPos. 1506: Gap → APos. 1532: C → APos. 1585: A → GPos. 1587: T → CPos. 1607: A → GPos. 1614: A → CPos. 1649: A → GapPos. 1654: A → TPos. 1673: A → TPos. 1727: A → TPos. 1757: T → GapPos. 1795: T → GPos. 1799: C → APos. 1823: T → CPos. 1865: T → APos. 1874: T → APos. 1887: T → CPos. 1894: T → CPos. 1971: C → TPos. 1997: G → CPos. 1999: Gap → TPos. 2019: A → TPos. 2057: A → CPos. 2058: A → TPos. 2059: A → TPos. 2069: A → TPos. 2070: A → CPos. 2086: C → TPos. 2089: C → GPos. 2112: T → APos. 2250: C → TPos. 2288: A → GPos. 2308: T → CPos. 2310: A → CPos. 2315: A → GPos. 2364: A → GapPos. 2407: Gap → CPos. 2510: T → APos. 2518: A → TPos. 2544: G → APos. 2548: T → C
Pos. 2566: T → CPos. 2575: G → APos. 2588: C → TPos. 2604: G → APos. 2605: G → APos. 2618: T → CPos. 2680: T → APos. 2768: C → APos. 2789: C → APos. 2866: A → GapPos. 2889: A → GapPos. 2929: C → GapPos. 2946: G → GapPos. 2956: C → GapPos. 2966: T → GapPos. 3036: T → CPos. 3054: T → CPos. 3071: A → TPos. 3125: A → TPos. 3127: T → APos. 3138: A → TPos. 3202: Gap → TPos. 3260: C → TPos. 3285: G → TPos. 3287: T → CPos. 3290: A → GPos. 3292: C → APos. 3318: T → CPos. 3357: A → TPos. 3375: T → APos. 3380: T → APos. 3429: T → CPos. 3451: A → GPos. 3487: T → CPos. 3491: C → ARAG-1Pos. 5 G → TPos. 29 G → APos. 137 T → CPos. 173 C → TPos. 182 A → GPos. 200 C → APos. 260 C → TPos. 297 T → CPos. 368 C → TPos. 395 T → CRhodopsinPos. 54: A → TPos. 135: C → GPos. 181: A → GPos. 244: G → TPos. 299: T → APos. 305: C → TSIAPos. 21: A → GPos. 169: T → APos. 385: T → C
Scinax
Cytochrome bPos. 97: A → CPos. 192: T → C12SPos. 333: C → APos. 636: G → APos. 745: Gap → CPos. 1113: C → TPos. 1147: A → GPos. 1245: T → APos. 1267: A → TPos. 1670: C → A
tRNA valinePos. 5: T → CPos. 87: G → TPos. 107: A → G16SPos. 234: T → CPos. 349: Gap → APos. 376: A → CPos. 606: C → APos. 611: A → GPos. 914: A → TPos. 1088: Gap → CPos. 1118: A → TPos. 1183: A → GapPos. 1219: A → TPos. 1324: A → CPos. 1332: A → GPos. 1389: T → APos. 1472: Gap → TPos. 1510: A → TPos. 1587: T → APos. 1807: A → CPos. 1826: C → TPos. 1847: G → TPos. 1861: T → CPos. 1987: Gap → TPos. 2156: A → TPos. 2406: T → APos. 2443: Gap → APos. 2660: C → TPos. 2829: Gap → APos. 2856: T → CPos. 2889: A → GPos. 2906: T → CPos. 2975: A → CPos. 2982: T → CPos. 3035: C → TPos. 3036: T → GPos. 3056: T → CPos. 3090: G → APos. 3175: Gap → APos. 3199: A → CPos. 3268: C → TPos. 3283: A → GPos. 3294: T → CPos. 3361: Gap → CT28SPos. 219: Gap → APos. 238: G → CPos. 322: Gap → APos. 327: T → APos. 394: Gap → TPos. 397: Gap → TPos. 403: Gap → GPos. 542: Gap → APos. 558: G → TPos. 569: A → CPos. 1144: C → APos. 1145: G → ARAG-1Pos. 4 C → GPos. 11 A → GPos. 116 C → GPos. 165 C → APos. 169 T → CPos. 170 G → APos. 185 T → CPos. 194 G → APos. 410 A → CPos. 425 G → ARhodopsin
2005 213FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 10: G → APos. 123: T → CSIAPos. 211: C → TPos. 218: C → TPos. 241: G → CPos. 250: C → TPos. 280: C → T
Scinax catharinae clade
Cytochrome bPos. 3: C → APos. 10: A → GPos. 53: G → APos. 109: G → CPos. 152: T → CPos. 199: G → APos. 239: C → TPos. 244: A → GPos. 288: A → TPos. 299: T → CPos. 327: C → TPos. 353: C → TPos. 359: A → TPos. 376: A → C12SPos. 227: T → GapPos. 241: T → APos. 297: T → CPos. 414: C → TPos. 460: T → APos. 659: Gap → CPos. 965: T → CPos. 1122: T → CPos. 1307: T → CPos. 1392: T → APos. 1422: G → APos. 1441: C → AtRNA valinePos. 59: C → TPos. 69: G → A16SPos. 76: Gap → APos. 254: T → CPos. 279: Gap → GPos. 367: T → GapPos. 383: T → CPos. 736: T → CPos. 742: T → CPos. 836: A → TPos. 1052: Gap → CPos. 1078: Gap → APos. 1174: T → GapPos. 1348: Gap → APos. 1534: C → TPos. 1544: G → APos. 1718: C → APos. 1737: T → APos. 1842: A → CPos. 1883: C → APos. 1948: G → CPos. 1985: A → CPos. 1997: G → APos. 2027: A → TPos. 2054: C → TPos. 2086: C → TPos. 2110: A → CPos. 2112: T → CPos. 2205: T → CPos. 2208: A → CPos. 2266: T → C
Pos. 2310: A → TPos. 2376: A → GPos. 2422: C → TPos. 2584: A → TPos. 2600: G → APos. 2616: A → TPos. 2768: C → TPos. 2789: C → TPos. 2802: T → APos. 2866: A → CPos. 2913: T → CPos. 2946: G → GapPos. 2997: C → TPos. 3127: T → APos. 3138: A → GPos. 3146: T → APos. 3249: A → GPos. 3277: G → APos. 3295: T → CPos. 3297: T → APos. 3431: C → T28SPos. 1467: G → GapPos. 1476: Gap → CRhodopsinPos. 135: C → TPos. 145: C → TPos. 202: T → CPos. 291: G → TPos. 292: T → ASIAPos. 72: T → CPos. 199: G → APos. 292: G → CPos. 382: G → APos. 397: T → A
Scinax ruber clade
Cytochrome bPos. 241: A → T12SPos. 42: T → CPos. 250: A → CPos. 600: A → GPos. 642: Gap → APos. 650: T → CPos. 654: T → CPos. 673: C → GPos. 729: T → APos. 759: A → CPos. 869: T → CPos. 958: A → GPos. 1135: G → APos. 1144: T → APos. 1153: T → GPos. 1509: A → GtRNA valinePos. 95: C → T16SPos. 58: C → TPos. 107: T → APos. 427: T → APos. 517: A → TPos. 726: A → CPos. 1203: T → APos. 1246: T → CPos. 1337: T → CPos. 1438: Gap → APos. 1529: A → CPos. 1530: A → TPos. 1614: A → G
Pos. 1795: T → CPos. 1874: T → CPos. 1951: A → TPos. 2029: T → CPos. 2030: G → APos. 2163: Gap → TPos. 2267: A → TPos. 2672: A → TPos. 2674: T → CPos. 2719: T → CPos. 2753: T → CPos. 3027: A → CPos. 3054: T → CPos. 3075: T → APos. 3166: C → TPos. 3516: A → T28SPos. 381: Gap → TPos. 393: Gap → GPos. 395: Gap → CPos. 396: Gap → CPos. 415: Gap → GRhodopsinPos. 120: C → TPos. 160: C → TPos. 244: G → T
Sphaenorhynchus
Cytochrome bPos. 17: A → TPos. 26: T → APos. 36: A → CPos. 45: C → TPos. 49: C → TPos. 68: C → TPos. 115: C → TPos. 214: C → TPos. 244: A → TPos. 289: C → TPos. 331: C → TPos. 356: T → CPos. 381: C → T12SPos. 36: A → TPos. 68: A → CPos. 196: G → APos. 206: A → TPos. 251: G → APos. 256: C → TPos. 319: A → GPos. 343: T → CPos. 484: A → TPos. 521: T → APos. 672: C → APos. 743: A → TPos. 760: C → TPos. 765: G → APos. 769: G → APos. 821: C → TPos. 965: T → APos. 1052: A → TPos. 1113: C → GapPos. 1123: Gap → APos. 1233: G → APos. 1245: T → CPos. 1332: G → APos. 1343: CT → GapPos. 1366: C → TPos. 1374: C → TPos. 1409: G → APos. 1412: C → A
Pos. 1415: Gap → TPos. 1426: C → TPos. 1496: T → CPos. 1539: T → APos. 1651: A → TPos. 1761: T → CtRNA valinePos. 39: C → TPos. 56: T → CPos. 74: A → TPos. 75: A → T16SPos. 12: C → TPos. 255: Gap → GPos. 548: A → TPos. 771: T → APos. 780: C → APos. 818: C → APos. 827: T → CPos. 829: C → TPos. 860: G → APos. 902: T → APos. 950: C → TPos. 1010: G → APos. 1019: C → TPos. 1204: Gap → APos. 1331: T → CPos. 1347: Gap → APos. 1389: T → CPos. 1419: T → APos. 1443: T → GPos. 1458: A → TPos. 1503: C → TPos. 1510: A → TPos. 1800: Gap → TPos. 1826: C → APos. 1839: T → APos. 1883: C → APos. 1948: GT → APos. 2019: A → TPos. 2043: A → TPos. 2086: C → TPos. 2098: G → GapPos. 2108: Gap → TPos. 2120: A → TPos. 2544: G → APos. 2551: A → TPos. 2672: A → TPos. 2975: A → TPos. 3059: A → CPos. 3081: A → GPos. 3145: T → APos. 3166: C → APos. 3277: G → APos. 3299: C → TPos. 3313: T → APos. 3425: C → TPos. 3455: G → ARAG-1Pos. 15 C → APos. 44 A → GPos. 47 C → TPos. 58 A → GPos. 64 A → GPos. 89 T → CPos. 164 G → APos. 194 G → APos. 248 A → CPos. 272 C → TPos. 275 C → TPos. 317 T → C
214 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 332 G → APos. 357 C → TPos. 365 T → CPos. 371 A → CPos. 386 T → APos. 398 A → TPos. 410 A → CPos. 425 G → ARhodopsinPos. 45: C → TPos. 93: C → TPos. 160: C → TPos. 220: G → TPos. 224: G → TPos. 235: C → TPos. 272: C → TPos. 290: C → TPos. 308: T → APos. 314: C → TTyrosinasePos. 7: T → APos. 10: A → CPos. 11: C → TPos. 20: T → APos. 72: C → TPos. 139: A → GPos. 183: A → GPos. 197: C → TPos. 209: T → CPos. 210: A → GPos. 211: C → APos. 276: T → CPos. 288: C → TPos. 291: G → CPos. 294: C → TPos. 330: T → CPos. 333: A → GPos. 375: T → CPos. 376: T → GPos. 393: C → TPos. 406: C → APos. 408: A → CPos. 423: T → CPos. 424: G → APos. 447: A → GPos. 472: Gap → TPos. 478: G → GapPos. 481: T → CPos. 508: T → CPos. 513: G → A
Xenohyla
Cytochrome bPos. 6: C → TPos. 10: A → GPos. 32: A → GPos. 49: C → APos. 52: A → TPos. 88: C → TPos. 108: A → GPos. 137: C → TPos. 164: C → TPos. 170: A → TPos. 199: G → APos. 201: T → CPos. 229: C → TPos. 262: A → CPos. 271: C → TPos. 277: T → CPos. 290: C → TPos. 292: T → C
Pos. 313: A → GPos. 322: C → APos. 353: C → TPos. 362: T → GPos. 402: T → CPos. 411: T → CPos. 420: A → T12SPos. 13: G → APos. 140: C → TPos. 148: C → APos. 159: C → APos. 171: G → TPos. 320: A → CPos. 379: A → GPos. 455: C → TPos. 499: Gap → TPos. 528: A → GPos. 547: C → GapPos. 600: A → GPos. 645: Gap → APos. 654: T → GapPos. 683: A → CPos. 733: T → GPos. 774: A → GPos. 815: T → CPos. 817: G → APos. 927: A → TPos. 1043: C → TPos. 1094: A → TPos. 1278: G → APos. 1303: A → GPos. 1321: T → APos. 1376: A → GPos. 1399: A → GPos. 1434: T → CPos. 1441: C → APos. 1549: AT → CPos. 1649: A → TPos. 1670: C → TtRNA valinePos. 46: T → GPos. 75: A → CPos. 76: A → GPos. 86: T → C16SPos. 130: C → TPos. 191: Gap → TPos. 192: C → APos. 259: A → GPos. 272: T → CPos. 323: C → APos. 336: C → APos. 427: T → APos. 434: A → TPos. 472: T → APos. 498: T → CPos. 543: T → CPos. 654: T → CPos. 668: T → APos. 718: T → CPos. 742: T → CPos. 755: A → CPos. 767: A → TPos. 775: T → GPos. 784: C → APos. 792: A → GPos. 821: T → APos. 854: T → CPos. 900: T → CPos. 908: T → C
Pos. 1025: T → CPos. 1160: T → CPos. 1174: T → APos. 1183: A → CPos. 1246: T → CPos. 1259: T → CPos. 1272: T → APos. 1274: A → GPos. 1309: Gap → CPos. 1324: A → GPos. 1436: T → GPos. 1443: T → CPos. 1534: C → APos. 1741: C → TPos. 1896: A → CPos. 1925: A → TPos. 2027: A → TPos. 2028: G → CPos. 2089: C → TPos. 2112: T → CPos. 2115: A → TPos. 2130: C → TPos. 2154: T → APos. 2161: A → TPos. 2229: T → GPos. 2240: A → GPos. 2262: AG → TPos. 2616: A → CPos. 2618: T → CPos. 2664: A → TPos. 2694: T → CPos. 2801: Gap → TPos. 2865: Gap → CPos. 2956: A → GPos. 3018: T → APos. 3027: T → CPos. 3056: T → CPos. 3086: T → APos. 3234: T → CPos. 3426: A → GPos. 3454: T → CPos. 3487: T → C
Hylini
Cytochrome bPos. 29: Gap → CPos. 31: A → CPos. 33: C → GapPos. 57: C → APos. 90: T → APos. 101: C → TPos. 126: G → TPos. 127: C → TPos. 137: T → APos. 353: C → T12SPos. 117: A → CTPos. 140: C → TPos. 148: C → TPos. 432: T → APos. 455: C → TPos. 528: A → TPos. 594: A → GPos. 595: A → TPos. 716: A → TPos. 1008: T → CPos. 1094: A → TPos. 1211: T → CPos. 1265: C → TPos. 1417: G → TPos. 1422: G → A
Pos. 1649: A → TPos. 1670: C → APos. 1678: A → CtRNA valinePos. 39: C → A28SPos. 150: A → TPos. 153: T → CPos. 342: C → GPos. 400: T → GapPos. 454: G → CPos. 516: G → CPos. 563: G → CPos. 569: A → TPos. 863: G → GapPos. 870: G → GapPos. 876: A → GapPos. 1003: Gap →GPos. 1017: Gap →TPos. 1080: Gap →GPos. 1103: G → CRAG-1Pos. 1: C → TPos. 101: T → CPos. 114: T → APos. 115: C → GPos. 143: G → CPos. 151: A → CPos. 169: T → CPos. 178: A → GPos. 194: G → APos. 359: A → CPos. 374: G → ARhodopsinPos. 12: A → GPos. 220: G → APos. 271: C → GPos. 281: T → CPos. 287: C → TSIAPos. 280: C → TPos. 349: T → ATyrosinasePos. 6: G → APos. 33: T → APos. 86: A → CPos. 160: T → CPos. 175: A → 4Pos. 180: 4 → TPos. 394: C → APos. 454: C → GPos. 455: T → C
Acris
Cytochrome bPos. 40: A → GPos. 63: C → GapPos. 68: C → TPos. 125: T → CPos. 137: A → CPos. 152: T → CPos. 155: A → CPos. 201: A → CPos. 207: C → TPos. 223: A → CPos. 302: A → TPos. 306: A → CPos. 311: A → CPos. 362: A → TPos. 369: A → TPos. 387: T → A
2005 215FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 396: A → CPos. 408: C → T12SPos. 142: C → TPos. 206: A → CPos. 229: Gap → APos. 233: Gap → APos. 332: A → CPos. 457: T → APos. 547: A → TPos. 670: A → TPos. 778: A → CPos. 841: A → TPos. 864: C → TPos. 908: A → CPos. 927: A → TPos. 965: C → TPos. 1245: T → APos. 1307: T → APos. 1332: G → APos. 1423: A → TPos. 1568: A → TPos. 1649: T → CPos. 1674: A → CtRNA valinePos. 6: G → APos. 39: A → TPos. 105: C → T16SPos. 172: A → CPos. 188: A → CPos. 272: G → APos. 443: C → GPos. 452: T → CPos. 459: T → CPos. 498: T → CPos. 517: T → CPos. 784: C → TPos. 848: C → APos. 853: A → GPos. 914: A → CPos. 950: C → APos. 1077: T → APos. 1235: C → APos. 1265: A → TPos. 1325: A → GapPos. 1375: Gap → CPos. 1480: T → GapPos. 1673: A → TPos. 1699: A → TPos. 1807: A → CPos. 1819: A → TPos. 1870: A → CPos. 1949: A → TPos. 1965: T → APos. 2030: G → APos. 2072: T → CPos. 2080: A → GPos. 2122: T → GPos. 2179: C → TPos. 2203: T → CPos. 2233: T → APos. 2244: T → CPos. 2250: C → TPos. 2253: A → TPos. 2381: T → CPos. 2422: C → TPos. 2577: G → TPos. 2676: A → TPos. 2787: T → APos. 2889: A → C
Pos. 2982: T → CPos. 3140: C → TPos. 3280: A → TRAG-1Pos. 5 G → APos. 71 G → APos. 125 G → APos. 170 G → APos. 359 C → TPos. 422 T → CRhodopsinPos. 21: C → TPos. 54: A → GPos. 60: T → CPos. 166: C → TPos. 211: G → ASIAPos. 160: T → CPos. 247: C → GPos. 259: A → TPos. 268: A → GPos. 397: T → ATyrosinasePos. 9: T → CPos. 10: A → CPos. 20: T → CPos. 25: T → CPos. 28: T → CPos. 34: T → CPos. 43: T → CPos. 73: A → GPos. 102: A → GPos. 127: T → CPos. 150: C → TPos. 167: G → APos. 168: T → CPos. 169: G → CPos. 180: T → CPos. 195: A → GPos. 204: A → CPos. 215: G → CPos. 231: A → CPos. 254: A → GPos. 307: A → GPos. 312: T → CPos. 315: A → GPos. 321: C → GPos. 330: T → CPos. 352: C → TPos. 384: A → GPos. 387: T → CPos. 407: A → GPos. 447: A → GPos. 453: T → CPos. 491: C → APos. 493: T → GPos. 496: A → GPos. 507: T → C
Anotheca spinosa
Cytochrome bPos. 10: A → TPos. 24: G → APos. 52: A → TPos. 53: G → APos. 57: A → CPos. 91: T → CPos. 155: A → GPos. 167: T → CPos. 170: A → TPos. 173: A → G
Pos. 183: C → TPos. 189: C → TPos. 212: T → APos. 213: G → APos. 214: C → TPos. 217: A → GPos. 247: C → TPos. 248: C → APos. 249: T → CPos. 256: A → TPos. 290: C → TPos. 299: T → CPos. 316: C → TPos. 353: T → CPos. 359: A → GPos. 363: A → GPos. 376: A → GPos. 408: C → TPos. 417: G → C12SPos. 9: T → CPos. 22: G → APos. 45: C → TPos. 83: G → APos. 88: C → TPos. 102: T → CPos. 174: A → GPos. 271: C → APos. 313: C → TPos. 347: T → CPos. 383: G → APos. 389: T → CPos. 391: A → TPos. 414: C → APos. 422: Gap → TPos. 423: Gap → TPos. 432: A → GapPos. 455: C → TPos. 460: T → CPos. 480: A → GPos. 489: G → GapPos. 512: T → APos. 600: T → GPos. 602: A → TPos. 608: A → GPos. 611: A → GPos. 654: T → CPos. 661: A → GPos. 686: G → APos. 726: A → TPos. 788: G → APos. 797: T → CPos. 802: C → TPos. 900: T → GPos. 928: A → GPos. 958: A → GPos. 963: Gap → TPos. 995: T → CPos. 1002: C → TPos. 1023: T → CPos. 1043: C → TPos. 1095: C → TPos. 1096: G → APos. 1100: G → APos. 1101: T → GPos. 1113: T → CPos. 1177: C → TPos. 1194: T → CPos. 1211: T → CPos. 1215: A → CPos. 1265: T → C
Pos. 1273: A → GPos. 1274: T → CPos. 1278: G → APos. 1296: C → TPos. 1317: C → TPos. 1366: C → TPos. 1378: G → APos. 1385: G → APos. 1408: A → GPos. 1427: T → CPos. 1434: T → CPos. 1439: C → APos. 1449: G → APos. 1544: Gap → TPos. 1585: A → TPos. 1645: T → CPos. 1649: T → CPos. 1651: A → TPos. 1664: A → GPos. 1670: T → CPos. 1674: A → CPos. 1693: C → TPos. 1699: Gap → TtRNA valinePos. 46: T → CPos. 55: A → GPos. 99: G → A16SPos. 94: C → TPos. 118: C → TPos. 130: C → TPos. 206: T → CPos. 272: G → APos. 536: C → TPos. 559: T → CPos. 566: T → CPos. 610: T → CPos. 611: A → GPos. 668: T → CPos. 699: A → GapPos. 712: A → GapPos. 742: T → APos. 771: T → CPos. 818: C → TPos. 821: A → TPos. 876: A → GPos. 952: A → GPos. 959: G → APos. 1022: A → CPos. 1025: T → CPos. 1085: A → GapPos. 1121: Gap → TPos. 1140: A → TPos. 1183: A → GPos. 1223: T → CPos. 1246: T → CPos. 1278: C → TPos. 1292: T → GPos. 1510: A → GPos. 1529: A → GPos. 1548: T → CPos. 1600: C → APos. 1607: C → TPos. 1635: Gap → CPos. 1710: T → APos. 1718: T → CPos. 1727: A → CPos. 1813: T → CPos. 2032: A → GapPos. 2039: G → APos. 2059: A → C
216 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 2071: T → CPos. 2130: T → CPos. 2138: Gap → CPos. 2156: C → TPos. 2169: A → GPos. 2170: A → GPos. 2217: T → CPos. 2229: T → CPos. 2262: G → APos. 2267: A → GPos. 2280: A → TPos. 2364: A → GapPos. 2392: A → GPos. 2518: A → GPos. 2524: T → CPos. 2544: A → GPos. 2615: A → GPos. 2655: A → GPos. 2660: C → TPos. 2694: T → CPos. 2719: T → CPos. 2768: T → CPos. 2787: T → CPos. 2960: T → GPos. 3014: T → CPos. 3036: T → APos. 3039: A → GPos. 3071: A → GPos. 3094: T → CPos. 3281: A → CPos. 3282: T → CPos. 3297: C → TPos. 3321: A → GPos. 3324: T → CPos. 3363: Gap → TPos. 3380: A → TPos. 3419: G → APos. 3458: T → CPos. 3462: C → TRAG-1Pos. 11 A → GPos. 77 T → CPos. 89 T → CPos. 149 A → CPos. 179 A → GPos. 185 G → TPos. 263 G → APos. 293 G → APos. 395 C → TRhodopsinPos. 28: A → TPos. 57: C → TPos. 89: A → TPos. 93: C → APos. 114: C → TSIAPos. 21: A → GPos. 292: G → CTyrosinasePos. 0: A → TPos. 6: A → TPos. 46: G → CPos. 97: G → APos. 124: T → CPos. 139: A → CPos. 164: A → GPos. 194: A → GPos. 357: C → TPos. 418: G → APos. 426: C → APos. 484: T → C
Pos. 526: C → TPos. 537: T → G
Bromeliohyla bromeliacia
Cytochrome bPos. 62: T → CPos. 63: C → TPos. 76: C → TPos. 116: C → TPos. 149: A → TPos. 161: A → GPos. 183: A → TPos. 186: A → GPos. 241: A → GPos. 283: C → TPos. 286: C → TPos. 331: T → GapPos. 359: A → GPos. 369: A → CPos. 381: C → TPos. 405: CT → APos. 420: A → G12sPos. 28: A → GPos. 180: C → TPos. 206: A → TPos. 310: T → CPos. 370: T → CPos. 412: T → CPos. 421: A → CPos. 438: A → TPos. 452: T → CPos. 454: T → CPos. 455: T → CPos. 512: T → CPos. 541: A → TPos. 627: Gap → TPos. 631: Gap → GPos. 641: A → CPos. 736: C → TPos. 1002: C → TPos. 1094: T → CPos. 1136: A → GPos. 1187: C → TPos. 1233: G → APos. 1374: C → TPos. 1407: G → APos. 1429: C → TPos. 1455: T → CPos. 1469: Gap → TPos. 1496: T → CPos. 1639: Gap → TtRNA valinePos. 2: A → TPos. 55: A → CPos. 74: C → GPos. 94: C → TPos. 98: A → G16SPos. 8: A → GPos. 111: Gap → APos. 152: Gap → TPos. 162: C → TPos. 383: A → GPos. 459: T → GPos. 589: C → TPos. 618: A → GPos. 640: A → TPos. 648: G → APos. 736: T → CPos. 754: A → G
Pos. 763: G → APos. 780: C → TPos. 788: C → TPos. 890: G → APos. 941: G → APos. 948: C → TPos. 979: C → TPos. 1085: A → GPos. 1187: A → TPos. 1211: A → GapPos. 1235: C → GapPos. 1289: A → GPos. 1314: A → GPos. 1328: A → GPos. 1332: G → APos. 1468: A → TPos. 1561: C → TPos. 1574: A → TPos. 1596: A → TPos. 1673: A → GPos. 1757: CT → APos. 1799: A → TPos. 1861: C → TPos. 1925: A → TPos. 1975: A → CPos. 1985: A → TPos. 2029: T → CPos. 2063: A → GPos. 2094: A → GPos. 2110: A → GPos. 2151: C → TPos. 2201: Gap → CPos. 2202: Gap → APos. 2233: T → CPos. 2312: T → APos. 2423: A → CPos. 2450: C → TPos. 2620: C → TPos. 2655: A → GPos. 2697: T → CPos. 2753: A → GPos. 2856: T → CPos. 3010: C → TPos. 3143: A → GPos. 3199: A → TPos. 3291: A → GPos. 3303: C → TPos. 3310: A → TPos. 3320: G → APos. 3324: T → CPos. 3326: T → CPos. 3425: C → TPos. 3455: G → APos. 3458: T → C28sPos. 561: C → APos. 812: C → ARAG-1Pos. 179: A → GRhodopsinPos. 9: C → TPos. 95: C → TPos. 154: C → TPos. 244: G → TPos. 268: A → TPos. 279: C → ASIAPos. 133: C → APos. 256: A → GTyrosinasePos. 106: C → T
Pos. 112: C → TPos. 127: T → CPos. 176: A → GPos. 197: T → APos. 254: A → GPos. 263: C → TPos. 267: C → TPos. 269: C → TPos. 297: C → TPos. 299: A → CPos. 300: A → GPos. 354: G → C
Charadrahyla
Cytochrome bPos. 128: A → TPos. 274: T → CPos. 353: T → C12SPos. 183: A → GPos. 328: G → APos. 337: C → TPos. 454: T → CPos. 571: A → CPos. 1002: C → TPos. 1010: T → APos. 1061: T → CPos. 1113: C → GapPos. 1116: C → TPos. 1236: T → CPos. 1343: T → CPos. 1383: C → APos. 1441: C → APos. 1461: C → TtRNA valinePos. 61: C → T16SPos. 2: T → CPos. 130: C → TPos. 452: T → CPos. 770: Gap → CPos. 780: C → GapPos. 1174: T → CPos. 1265: A → GPos. 1328: A → GPos. 1330: C → TPos. 1332: G → TPos. 1643: T → APos. 1673: A → CPos. 1817: A → TPos. 1861: C → APos. 1870: A → CPos. 2253: A → GPos. 2268: C → APos. 2461: C → TPos. 2569: T → CPos. 2616: A → TPos. 2694: T → CPos. 2780: A → GapPos. 2802: T → APos. 2913: T → CPos. 3010: C → APos. 3111: G → APos. 3118: C → APos. 3130: A → TPos. 3154: A → GPos. 3389: C → TPos. 3425: C → TSIAPos. 175: C → TPos. 376: T → C
2005 217FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
TyrosinasePos. 86: C → APos. 102: A → GPos. 172: A → GPos. 432: T → C
Duellmanohyla
Cytochrome bPos. 229: C → TPos. 277: T → CPos. 292: T → CPos. 313: T → CPos. 366: T → CPos. 376: T → CPos. 399: T → C12SPos. 13: A → GPos. 22: G → APos. 131: T → GapPos. 272: A → GPos. 414: C → TPos. 427: C → TPos. 726: A → GapPos. 1010: T → APos. 1333: A → CPos. 1405: A → GPos. 1408: A → GPos. 1412: C → TPos. 1427: T → CPos. 1431: T → CPos. 1500: Gap → A16SPos. 172: A → CPos. 234: A → GapPos. 272: G → TPos. 376: T → CPos. 755: T → APos. 949: T → CPos. 1160: T → CPos. 1699: A → CPos. 1826: T → CPos. 2164: Gap → TPos. 2308: C → TPos. 2551: A → GPos. 2742: T → APos. 2787: A → GapPos. 3054: A → GPos. 3222: T → CPos. 3249: T → CPos. 3375: T → CPos. 3487: T → CRAG-1Pos. 35 A → GRhodopsinPos. 28: A → TSIAPos. 175: C → TPos. 241: G → ATyrosinasePos. 70: T → GPos. 517: T → CPos. 527: A → G
Ecnomiohyla
Cytochrome bPos. 49: C → TPos. 136: C → TPos. 152: T → CPos. 176: C → TPos. 183: A → TPos. 189: C → TPos. 229: C → T
Pos. 369: A → T12SPos. 102: T → CPos. 164: C → APos. 322: C → TPos. 427: C → T16SPos. 1118: A → CPos. 1183: T → APos. 1230: Gap → TPos. 1235: C → APos. 1491: A → TPos. 1585: A → GPos. 1741: C → TPos. 1870: A → TPos. 1925: A → TPos. 2069: T → APos. 2089: A → CPos. 2190: C → GapPos. 2229: T → CPos. 2441: C → TPos. 2457: G → APos. 3056: C → APos. 3173: G → APos. 3222: T → GapPos. 3249: T → CPos. 3285: G → APos. 3315: A → TRAG-1Pos. 242 G → ARhodopsinPos. 28: A → TPos. 281: C → TPos. 316: A → G
Exerodonta
Cytochrome bPos. 71: A → CPos. 115: C → TPos. 232: C → APos. 247: C → TPos. 256: A → TPos. 334: A → GPos. 411: T → C12SPos. 227: T → CPos. 322: C → APos. 370: A → GPos. 392: T → CPos. 594: G → TPos. 673: T → CPos. 760: C → APos. 1376: A → GPos. 1392: T → APos. 1585: A → GPos. 1601: T → APos. 1602: T → APos. 1636: A → TPos. 1649: T → CPos. 1651: A → G16SPos. 151: Gap → TPos. 284: Gap → APos. 434: A → TPos. 443: C → TPos. 755: A → TPos. 758: A → CPos. 853: A → TPos. 890: G → APos. 948: C → TPos. 1153: A → C
Pos. 1160: T → CPos. 1351: Gap → APos. 1389: T → APos. 1468: A → TPos. 1534: C → TPos. 1587: T → CPos. 1817: A → TPos. 1819: A → TPos. 1839: T → CPos. 2024: G → APos. 2067: A → TPos. 2130: T → APos. 2263: C → TPos. 2615: A → CPos. 2676: A → TPos. 2780: A → CPos. 2812: T → APos. 2929: C → TPos. 2946: G → GapPos. 3081: A → TPos. 3234: A → TPos. 3290: A → GPos. 3426: G → TPos. 3439: T → CPos. 3454: C → A28SPos. 544: G → CRAG-1Pos. 32 A → GPos. 128 T → CPos. 155 T → CPos. 248 A → GPos. 293 G → APos. 413 T → CRhodopsinPos. 232: C → TPos. 259: C → TPos. 281: C → TPos. 299: T → CSIAPos. 81: C → TPos. 106: C → APos. 205: T → CPos. 211: C → TPos. 331: G → APos. 376: T → CTyrosinasePos. 4: C → TPos. 105: G → CPos. 179: A → GPos. 257: C → TPos. 327: T → CPos. 514: G → C
Exerodonta sumichrastigroup
Cytochrome bPos. 24: G → APos. 45: C → TPos. 126: T → CPos. 153: C → TPos. 167: C → TPos. 176: C → TPos. 189: C → TPos. 198: T → CPos. 214: C → TPos. 217: A → GPos. 241: A → GPos. 296: C → TPos. 331: C → TPos. 366: T → C
Pos. 367: C → TPos. 380: C → TPos. 383: T → CPos. 417: A → C12SPos. 24: A → GPos. 115: A → TPos. 159: A → TPos. 231: T → CPos. 241: T → CPos. 333: C → APos. 452: T → CPos. 541: T → CPos. 558: A → CPos. 594: T → CPos. 641: A → GPos. 733: A → TPos. 778: A → GPos. 869: C → TPos. 1043: C → TPos. 1211: C → TPos. 1549: A → TPos. 1625: T → C16SPos. 107: C → APos. 234: T → CPos. 472: C → APos. 656: T → CPos. 708: A → GPos. 718: T → CPos. 736: T → CPos. 758: C → TPos. 774: T → GPos. 780: C → TPos. 834: A → GPos. 862: T → CPos. 914: A → TPos. 1574: A → TPos. 1699: A → CPos. 1727: A → TPos. 1826: T → CPos. 2057: T → CPos. 2069: T → CPos. 2200: T → CPos. 2248: A → GPos. 2381: T → APos. 2587: T → CPos. 2615: C → TPos. 2715: Gap → TPos. 2982: T → CPos. 2995: A → CPos. 3010: C → TPos. 3087: Gap → CPos. 3125: T → CPos. 3209: T → CPos. 3230: T → ARAG-1Pos. 392 T → ARhodopsinPos. 9: C → TPos. 36: G → APos. 133: C → TPos. 167: A → CSIAPos. 21: A → GPos. 205: C → ATyrosinasePos. 420: A → G
Hyla
Cytochrome b
218 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 11: C → APos. 67: C → TPos. 250: T → CPos. 417: G → A12SPos. 560: Gap → TPos. 778: A → GPos. 814: C → TPos. 1101: T → CPos. 1346: T → CPos. 1549: A → Gap16SPos. 543: T → CPos. 780: C → APos. 1103: A → TPos. 1246: T → APos. 1436: T → APos. 1713: Gap → APos. 1925: T → CPos. 2267: A → TPos. 2787: T → GapPos. 2812: T → APos. 2858: Gap → CRhodopsinPos. 316: A → GTyrosinasePos. 73: G → APos. 484: T → CPos. 493: T → C
Hyla arborea group
Cytochrome bPos. 145: C → TPos. 271: C → T12SPos. 13: A → GPos. 34: T → CPos. 152: T → APos. 169: T → CPos. 268: T → CPos. 455: C → TPos. 880: C → GPos. 1039: Gap → TPos. 1043: C → APos. 1365: Gap → CPos. 1561: A → T16SPos. 664: A → TPos. 718: T → CPos. 1057: T → CPos. 1389: T → APos. 1402: A → CPos. 1799: A → CPos. 2089: A → CPos. 2820: A → CPos. 2856: T → APos. 3239: A → GapPos. 3282: T → CRAG-1Pos. 4 C → TPos. 380 C → TPos. 395 C → TRhodopsinPos. 93: C → APos. 245: G → TPos. 262: G → ATyrosinasePos. 29: C → APos. 31: A → CPos. 120: A → TPos. 186: C → A
Pos. 245: C → TPos. 315: G → A
Hyla cinerea group
Cytochrome bPos. 53: G → APos. 108: G → APos. 125: T → CPos. 149: A → TPos. 183: A → TPos. 214: C → TPos. 286: C → TPos. 338: T → CPos. 366: T → C12SPos. 597: T → CPos. 1105: A → TPos. 1113: T → CPos. 1422: G → APos. 1589: C → A16SPos. 107: C → APos. 177: Gap → CPos. 272: G → APos. 517: T → CPos. 521: A → TPos. 1603: T → CPos. 1699: T → CPos. 1874: T → CPos. 2058: C → APos. 2156: C → APos. 2161: A → TPos. 2280: A → TRAG-1Pos. 83 A → GPos. 284 A → GSIAPos. 250: C → TPos. 20: T → GPos. 257: C → TPos. 506: G → A
Hyla eximia group
Cytochrome bPos. 153: T → CPos. 167: T → CPos. 238: T → C16SPos. 162: T → CPos. 566: T → CPos. 1376: T → CPos. 1842: A → GPos. 2107: A → GPos. 2620: C → T
Hyla versicolor group
Cytochrome bPos. 49: C → GPos. 103: C → TPos. 108: G → APos. 161: A → GPos. 214: C → TPos. 286: C → TPos. 289: C → TPos. 313: C → TPos. 356: C → TPos. 383: A → TPos. 417: A → C12SPos. 285: A → TPos. 352: G → APos. 560: T → Gap
Pos. 565: T → CPos. 571: A → CPos. 738: A → TPos. 1278: G → APos. 1579: A → GapPos. 1671: Gap → T16SPos. 521: A → TPos. 573: A → TPos. 822: A → TPos. 906: T → CPos. 965: C → TPos. 1092: A → GPos. 1167: A → TPos. 1203: T → CPos. 1223: C → TPos. 1718: C → TPos. 1741: C → APos. 1749: T → CPos. 1799: A → TPos. 1807: A → GPos. 1925: C → TPos. 2057: C → TPos. 2089: A → TPos. 2177: C → TPos. 2233: T → CPos. 2280: A → GPos. 2508: T → CPos. 2997: C → TPos. 3010: C → TPos. 3140: C → TPos. 3389: C → TPos. 3524: C → TPos. 3538: G → ARhodopsinPos. 316: G → ASIAPos. 211: C → TTyrosinasePos. 120: A → T
Isthmohyla
Cytochrome bPos. 68: T → CPos. 91: T → CPos. 167: T → CPos. 247: C → TPos. 265: C → TPos. 376: A → C12SPos. 117: C → TPos. 169: T → APos. 348: T → CPos. 550: T → APos. 565: T → CPos. 726: A → TPos. 1002: C → TtRNA valinePos. 4: A → G16SPos. 323: C → TPos. 355: T → CPos. 559: T → CPos. 580: T → CPos. 611: A → GPos. 1015: T → APos. 1657: Gap → CPos. 1727: A → CPos. 1865: C → TPos. 2043: A → TPos. 2107: A → G
Pos. 2112: CT → APos. 2156: C → TPos. 2208: AT → CPos. 2217: T → CPos. 2544: A → GPos. 2776: A → GapPos. 2820: A → TPos. 2982: T → CPos. 3056: C → TPos. 3111: G → APos. 3154: A → GPos. 3173: T → CPos. 3237: T → APos. 3281: A → GPos. 3282: T → CRhodopsinPos. 93: C → TPos. 124: C → A
Megastomatohyla mixe
Cytochrome bPos. 10: C → APos. 46: C → TPos. 49: C → APos. 57: A → GPos. 101: T → APos. 109: G → APos. 125: T → CPos. 137: C → TPos. 145: C → TPos. 164: C → TPos. 168: A → GPos. 184: A → CPos. 204: C → APos. 214: C → TPos. 223: A → CPos. 238: T → APos. 241: A → CPos. 292: T → CPos. 302: A → GPos. 327: C → APos. 338: T → CPos. 366: T → CPos. 383: A → TPos. 414: T → APos. 415: C → T12SPos. 34: T → CPos. 61: C → TPos. 68: A → CPos. 69: C → TPos. 110: A → GapPos. 183: A → TPos. 269: A → TPos. 326: A → GPos. 339: T → CPos. 434: G → APos. 440: A → GPos. 451: T → CPos. 452: T → APos. 480: A → GPos. 521: T → CPos. 558: A → GapPos. 579: Gap → GPos. 619: G → TPos. 646: C → TPos. 650: T → GPos. 726: A → CPos. 736: C → APos. 738: A → CPos. 769: G → A
2005 219FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 821: C → TPos. 835: T → CPos. 869: C → TPos. 908: A → CPos. 989: A → GPos. 1036: C → TPos. 1043: C → APos. 1048: T → CPos. 1067: C → TPos. 1089: T → CPos. 1105: A → GapPos. 1122: T → CPos. 1128: T → CPos. 1144: T → CPos. 1244: A → GPos. 1245: T → CPos. 1278: G → APos. 1327: A → GPos. 1366: C → TPos. 1392: T → APos. 1405: A → GPos. 1408: A → GPos. 1412: C → TPos. 1418: A → CPos. 1427: T → CPos. 1431: T → CPos. 1439: C → TPos. 1455: T → CPos. 1537: Gap → TPos. 1561: A → CPos. 1568: A → GPos. 1651: A → TPos. 1664: A → TtRNA valinePos. 39: A → GPos. 46: T → CPos. 59: C → TPos. 69: G → APos. 76: AG → T16SPos. 0: G → APos. 107: C → TPos. 130: C → APos. 234: A → TPos. 294: A → GapPos. 323: C → TPos. 376: A → TPos. 418: T → CPos. 600: Gap → APos. 630: A → TPos. 636: T → CPos. 665: A → TPos. 702: A → GPos. 731: T → CPos. 784: C → APos. 807: G → APos. 818: C → TPos. 839: C → TPos. 876: A → GPos. 906: T → APos. 908: T → APos. 954: G → APos. 968: G → APos. 1012: T → GapPos. 1092: T → APos. 1153: A → CPos. 1167: T → CPos. 1228: A → GapPos. 1253: Gap → CPos. 1259: T → CPos. 1310: C → A
Pos. 1331: A → TPos. 1332: G → APos. 1405: T → APos. 1480: C → APos. 1544: A → GPos. 1572: T → CPos. 1574: A → TPos. 1614: A → GPos. 1727: A → TPos. 1740: Gap → TPos. 1768: A → GapPos. 1861: C → TPos. 1865: C → APos. 1928: C → TPos. 1932: T → CPos. 1977: G → CPos. 1997: G → APos. 2054: C → APos. 2068: C → TPos. 2070: A → CPos. 2107: A → CPos. 2110: A → TPos. 2130: T → APos. 2177: C → TPos. 2298: C → TPos. 2398: A → CPos. 2450: C → TPos. 2544: A → GPos. 2588: C → TPos. 2605: G → APos. 2669: A → GPos. 2683: Gap → APos. 2691: C → TPos. 2787: T → GapPos. 2812: T → CPos. 2906: T → APos. 2946: G → TPos. 2990: A → GapPos. 2997: C → GapPos. 3004: A → TPos. 3014: T → GapPos. 3036: T → CPos. 3127: T → CPos. 3143: A → GPos. 3145: T → CPos. 3236: Gap → TPos. 3268: C → TPos. 3278: C → TPos. 3287: T → CPos. 3298: G → APos. 3385: T → CPos. 3426: G → APos. 3454: C → TPos. 3486: A → GPos. 3545: A → G28sPos. 205: A → GPos. 569: T → CPos. 779: Gap → CPos. 780: Gap → CPos. 1073: G → APos. 1078: C → ARAG-1Pos. 117: A → GPos. 137: T → CPos. 155: T → CPos. 179: A → GRhodopsinPos. 28: A → TPos. 39: A → GPos. 135: C → T
Pos. 205: C → TPos. 219: C → TPos. 256: C → TSIAPos. 6: G → APos. 127: A → GPos. 211: C → TPos. 235: G → APos. 259: A → GPos. 322: C → TPos. 373: C → TTyrosinasePos. 19: T → GPos. 26: C → APos. 96: A → GPos. 133: AC → GPos. 179: A → GPos. 191: A → GPos. 261: A → GPos. 307: A → GPos. 308: A → CPos. 394: A → GPos. 456: G → CPos. 484: T → CPos. 511: G → A
Plectrohyla
Cytochrome bPos. 45: C → TPos. 189: C → TPos. 238: T → APos. 310: A → CPos. 402: T → C12SPos. 9: T → CPos. 502: Gap → CPos. 601: A → GPos. 650: T → CPos. 1113: C → TPos. 1118: C → TPos. 1579: A → G16SPos. 188: A → TPos. 459: T → APos. 654: T → APos. 906: T → APos. 1085: T → APos. 1103: T → CPos. 1741: C → TPos. 1826: T → CPos. 1948: T → CPos. 2039: G → APos. 2154: C → APos. 2229: T → APos. 2267: A → TPos. 2820: A → TPos. 2985: T → CPos. 3326: T → C28SPos. 525: G → GapRAG-1Pos. 362 C → APos. 395 T → CSIAPos. 3: T → CPos. 142: C → GPos. 148: C → TTyrosinasePos. 37: C → TPos. 58: G → APos. 86: C → A
Pos. 172: A → GPos. 300: A → GPos. 324: G → APos. 416: A → GPos. 475: C → T
Plectrohyla bistincta group
Cytochrome bPos. 101: T → AGPos. 204: A → TPos. 319: C → TPos. 356: T → C12SPos. 295: A → CPos. 427: C → TPos. 512: T → APos. 1002: C → TPos. 1005: C → A16SPos. 517: T → CPos. 1219: A → TPos. 1223: C → TPos. 1618: Gap → TPos. 1649: T → APos. 2190: C → ARAG-1Pos. 74 T → C
Plectrohyla guatemalensisgroup
Cytochrome bPos. 3: C → TPos. 111: C → TPos. 131: A → GPos. 229: C → TPos. 244: A → TPos. 271: C → TPos. 383: T → CPos. 396: A → G12SPos. 180: C → TPos. 251: G → APos. 646: C → TPos. 751: A → CPos. 1270: A → GPos. 1307: T → CPos. 1321: T → APos. 1521: G → APos. 1659: C → TtRNA valinePos. 110: T → A16SPos. 12: C → TPos. 355: C → TPos. 498: T → CPos. 691: G → APos. 767: A → GPos. 839: C → TPos. 1092: T → APos. 1458: T → CPos. 1718: T → CPos. 2057: T → CPos. 2240: A → GPos. 2854: Gap → CPos. 3010: C → TPos. 3027: T → CPos. 3242: Gap → T28SPos. 891: C → T
Pseudacris
Cytochrome b
220 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 36: A → CTPos. 39: T → CPos. 71: A → CPos. 153: C → TPos. 189: C → TPos. 253: T → CPos. 274: T → CPos. 319: C → TPos. 327: C → APos. 354: A → GPos. 383: A → T12SPos. 556: Gap → APos. 838: A → TPos. 843: T → CPos. 1113: C → TtRNA valinePos. 59: C → TPos. 69: G → A16SPos. 664: A → TPos. 1085: T → APos. 1211: A → GapPos. 1223: C → GapPos. 1561: C → TPos. 1710: T → CPos. 2025: C → TPos. 2112: T → APos. 2836: Gap → TPos. 3035: C → TPos. 3208: Gap → T28SPos. 464: G → TPos. 525: G → GapPos. 909: C → ATyrosinasePos. 191: A → GPos. 283: G → TPos. 324: G → APos. 382: A → CPos. 396: A → C
Ptychohyla
Cytochrome bPos. 111: C → T12SPos. 1113: C → TPos. 1422: A → G16SPos. 308: A → GPos. 1298: C → TPos. 1532: C → TPos. 1953: T → APos. 2143: Gap → TPos. 2785: T → CPos. 3278: C → TPos. 3298: G → A
Smilisca
Cytochrome bPos. 119: CT → APos. 198: T → APos. 274: T → CPos. 280: C → TPos. 330: C → TPos. 383: A → TPos. 411: T → C12SPos. 206: A → GPos. 231: T → APos. 244: A → TPos. 330: C → T
Pos. 336: G → APos. 386: A → TPos. 566: Gap → APos. 600: T → APos. 729: T → APos. 880: C → APos. 1101: T → CPos. 1316: A → TPos. 1330: A → TPos. 1614: T → GapPos. 1632: Gap → CPos. 1678: T → CtRNA valinePos. 39: A → G16SPos. 443: T → APos. 853: A → GPos. 906: T → CPos. 1103: A → TPos. 1235: A → TPos. 1468: A → CPos. 1743: Gap → TPos. 2117: A → TPos. 2267: A → TPos. 2820: A → CPos. 2849: A → TPos. 3127: T → CSIAPos. 232: A → GPos. 292: G → A
Tlalocohyla
Cytochrome bPos. 90: A → TPos. 128: A → TPos. 158: C → TPos. 244: A → TPos. 259: C → TPos. 322: A → CPos. 330: C → TPos. 381: C → TPos. 387: T → A12SPos. 40: C → TPos. 61: C → APos. 131: T → CPos. 231: T → APos. 297: T → CPos. 348: T → CPos. 353: G → APos. 378: G → APos. 437: C → TPos. 521: T → GPos. 528: A → CPos. 582: C → APos. 970: A → CPos. 1001: A → CPos. 1122: T → APos. 1265: T → CPos. 1346: T → APos. 1417: T → CPos. 1531: C → Gap16SPos. 188: A → CPos. 323: C → APos. 336: C → APos. 399: A → TPos. 459: T → APos. 589: C → TPos. 634: T → APos. 718: T → C
Pos. 777: A → CPos. 853: A → TPos. 874: T → CPos. 906: T → CPos. 982: A → GPos. 1024: C → TPos. 1183: T → APos. 1275: G → APos. 1376: T → GapPos. 1451: T → CPos. 1458: T → APos. 1870: A → TPos. 1874: T → APos. 1878: C → APos. 1928: C → APos. 1948: T → APos. 1994: A → TPos. 2008: C → TPos. 2059: A → TPos. 2069: T → APos. 2151: T → CPos. 2200: T → APos. 2398: A → TPos. 2587: T → CPos. 2785: A → CPos. 2913: T → CPos. 2975: A → TPos. 3056: C → APos. 3118: C → TPos. 3260: C → TPos. 3291: A → GPos. 3318: T → CPos. 3357: A → TPos. 3428: C → TPos. 3429: T → CPos. 3451: A → GRhodopsinPos. 185: C → APos. 262: G → APos. 296: T → GPos. 300: C → TSIAPos. 3: T → CPos. 289: A → CPos. 340: A → GTyrosinasePos. 0: A → CPos. 6: A → GPos. 26: C → GPos. 49: C → TPos. 118: T → CPos. 155: C → GPos. 170: A → CPos. 171: G → CPos. 177: A → CPos. 254: A → GPos. 327: T → CPos. 453: T → CPos. 466: A → C
Triprion
Cytochrome bPos. 2: C → TPos. 23: T → CPos. 39: C → TPos. 68: T → CPos. 79: A → GPos. 88: C → TPos. 94: C → TPos. 126: T → GPos. 127: T → C
Pos. 137: C → TPos. 140: A → CPos. 145: C → TPos. 149: A → GPos. 153: T → CPos. 158: C → TPos. 164: C → TPos. 176: C → TPos. 180: C → TPos. 204: C → APos. 241: A → GPos. 265: C → TPos. 286: C → TPos. 311: A → GPos. 367: C → TPos. 399: C → T12SPos. 13: A → GPos. 38: T → CPos. 41: T → CPos. 70: C → TPos. 117: C → APos. 149: Gap → APos. 169: T → APos. 227: T → CPos. 326: A → GPos. 352: A → GPos. 532: Gap → CPos. 547: T → CPos. 550: T → CPos. 565: T → CPos. 582: T → GPos. 716: A → CPos. 733: T → APos. 738: A → GPos. 778: A → GPos. 814: C → TPos. 864: C → GapPos. 869: C → TPos. 880: C → TPos. 1105: C → TPos. 1122: T → CPos. 1362: T → CPos. 1402: A → TPos. 1405: A → GPos. 1423: A → CPos. 1431: T → CPos. 1440: T → APos. 1441: C → TPos. 1549: A → CPos. 1561: A → TPos. 1761: C → TPos. 1762: G → AtRNA valinePos. 4: A → GPos. 59: C → TPos. 69: G → APos. 90: A → GPos. 101: A → GPos. 108: T → C16SPos. 162: T → GPos. 434: A → TPos. 517: T → CPos. 543: T → CPos. 630: A → GPos. 656: T → CPos. 726: A → TPos. 780: C → TPos. 839: T → CPos. 890: A → G
2005 221FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 914: A → GPos. 1103: A → GPos. 1174: T → CPos. 1259: T → CPos. 1436: T → APos. 1443: T → CPos. 1480: T → CPos. 1650: Gap → CPos. 1737: C → TPos. 1741: C → APos. 1817: A → GPos. 1948: T → CPos. 1953: T → CPos. 2063: A → GPos. 2089: A → TPos. 2142: C → APos. 2587: T → CPos. 2613: A → GPos. 2849: A → GPos. 2913: T → CPos. 2985: T → CPos. 3081: A → TPos. 3209: G → APos. 3249: C → TPos. 3326: C → TRAG-1Pos. 68: A → CPos. 107: A → GPos. 122: A → GPos. 302: C → TSIAPos. 48: C → TPos. 106: C → ATyrosinasePos. 12: C → TPos. 22: G → APos. 49: C → TPos. 248: G → TPos. 268: C → TPos. 312: C → TPos. 394: A → CPos. 401: G → APos. 416: G → APos. 432: C → TPos. 450: C → GPos. 456: G → CPos. 481: G → TPos. 493: T → CPos. 506: G → APos. 514: G → A
Hylini 1 Lophiohylini
Cytochrome bPos. 14: A → TPos. 97: A → CPos. 310: C → A12SPos. 231: A → TPos. 761: T → CPos. 890: A → CPos. 965: T → CPos. 1116: A → C16SPos. 272: T → GPos. 383: T → APos. 848: A → CPos. 906: A → TPos. 1223: Gap → CPos. 1796: Gap → APos. 1817: T → APos. 2057: A → CT
Pos. 2151: T → CPos. 2618: T → APos. 2787: Gap → T28SPos. 249: C → GapPos. 327: T → GapSIAPos. 331: A → GTyrosinasePos. 315: C → APos. 429: T → CPos. 481: T → G
Lophiohylini
Cytochrome bPos. 3: C → APos. 36: A → CPos. 265: C → APos. 268: A → CPos. 302: A → C12SPos. 24: A → TPos. 285: A → CPos. 593: T → CPos. 672: C → APos. 771: C → TPos. 817: G → APos. 908: A → TPos. 1589: T → AtRNA valinePos. 6: G → APos. 105: C → T16SPos. 138: Gap → ACPos. 573: A → TPos. 736: T → APos. 758: A → TPos. 814: T → APos. 836: A → CPos. 878: C → TPos. 978: G → APos. 1160: T → APos. 1174: T → CPos. 1419: T → CPos. 1470: Gap → CPos. 1710: T → CPos. 1789: Gap → TPos. 1842: A → GapPos. 1880: A → TPos. 1953: T → APos. 2041: A → CPos. 2091: T → CPos. 2110: A → TPos. 2156: A → CPos. 2525: A → TPos. 2694: T → APos. 2889: A → CPos. 3004: A → CPos. 3297: T → A28SPos. 390: C → TPos. 532: Gap → APos. 544: G → CRAG-1Pos. 89 T → CPos. 233 A → GPos. 248 A → CPos. 317 T → CPos. 341 C → TPos. 371 A → TSIA
Pos. 36: T → CPos. 145: A → GPos. 148: C → TPos. 211: C → TPos. 235: G → APos. 373: C → TTyrosinasePos. 0: AC → TPos. 4: C → GPos. 212: G → APos. 268: C → TPos. 333: A → TPos. 372: T → CPos. 396: A → G
Aparasphenodon brunoi
Cytochrome bPos. 42: T → CPos. 71: A → GPos. 76: C → APos. 97: C → TPos. 105: A → GPos. 108: G → APos. 152: C → TPos. 164: C → TPos. 167: C → TPos. 183: C → TPos. 214: C → TPos. 223: A → TPos. 235: A → GPos. 247: C → TPos. 250: C → TPos. 296: C → TPos. 327: C → TPos. 330: C → TPos. 331: C → TPos. 367: C → TPos. 396: A → GPos. 399: C → TPos. 414: C → TPos. 415: C → TPos. 420: C → A12sPos. 10: A → GPos. 66: G → APos. 68: A → CPos. 159: C → GPos. 183: A → CPos. 194: T → CPos. 206: A → TPos. 227: C → TPos. 452: T → CPos. 521: T → CPos. 550: C → TPos. 594: A → CPos. 646: C → TPos. 693: G → APos. 864: C → GapPos. 965: C → TPos. 970: A → CPos. 1043: C → TPos. 1270: A → GPos. 1392: T → APos. 1402: G → CPos. 1599: C → TPos. 1636: C → TPos. 1670: C → T16SPos. 1085: T → CPos. 1310: C → TPos. 1325: C → T
Pos. 1468: A → GPos. 1470: C → TPos. 1718: T → CPos. 1737: T → CPos. 1795: C → TPos. 1874: C → APos. 1878: T → CPos. 1928: T → CPos. 1998: A → CPos. 2027: A → GPos. 2028: G → APos. 2029: T → CPos. 2063: A → GPos. 2072: T → CPos. 2130: C → TPos. 2151: C → TPos. 2203: A → CPos. 2240: A → GPos. 2253: A → GPos. 2395: A → GPos. 2566: C → TPos. 2568: C → TPos. 2844: G → CPos. 2856: A → TPos. 2866: A → TPos. 2982: C → TPos. 2990: C → TPos. 3014: T → CPos. 3032: T → CPos. 3127: T → CPos. 3189: G → A
Argenteohyla siemersi
Cytochrome bPos. 3: C → TPos. 17: A → GPos. 57: A → GPos. 63: C → TPos. 76: C → TPos. 108: G → CPos. 109: G → APos. 128: A → GPos. 186: A → GPos. 223: A → GPos. 238: T → APos. 239: C → TPos. 292: T → APos. 311: A → GPos. 319: C → TPos. 334: G → APos. 344: A → GPos. 378: T → CPos. 396: A → TPos. 408: A → T12SPos. 36: A → GPos. 133: Gap → APos. 141: Gap → TPos. 159: C → APos. 285: C → APos. 492: Gap → TPos. 493: Gap → TPos. 672: A → CPos. 673: C → TPos. 880: A → TPos. 1010: T → CPos. 1084: T → CPos. 1309: A → TPos. 1346: T → CPos. 1392: T → CPos. 1413: A → T
222 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 1422: G → APos. 1589: A → CPos. 1674: C → TtRNA valinePos. 72: G → APos. 98: G → A16SPos. 0: G → APos. 11: C → TPos. 138: A → TPos. 172: A → CPos. 308: A → GPos. 356: T → CPos. 452: T → CPos. 726: A → CPos. 827: T → CPos. 831: A → TPos. 839: C → TPos. 1022: G → APos. 1049: T → CPos. 1062: T → CPos. 1278: T → CPos. 1443: T → CPos. 1458: A → GPos. 1510: A → GPos. 1649: T → CPos. 1795: C → APos. 1874: C → TPos. 1944: A → GPos. 2016: T → CPos. 2024: G → APos. 2091: C → TPos. 2125: T → CPos. 2187: A → GPos. 2203: A → TPos. 2233: T → CPos. 2250: C → APos. 2263: C → TPos. 2267: T → CPos. 2285: A → GPos. 2345: C → TPos. 2392: A → TPos. 2416: G → APos. 2450: C → TPos. 2728: Gap → APos. 2889: C → GapPos. 3039: A → GPos. 3059: A → CPos. 3116: Gap → GPos. 3199: A → GPos. 3285: G → APos. 3397: Gap → G28SPos. 262: C → GapPos. 505: G → GapPos. 532: A → TRAG-1Pos. 6: T → CPos. 128 T → GPos. 183 C → GPos. 324 G → CPos. 380 C → TSIAPos. 205: T → GTyrosinasePos. 91: C → TPos. 284: C → TPos. 312: T → CPos. 324: G → APos. 339: C → APos. 351: C → T
Pos. 416: A → G
Corythomantis greenengi
Cytochrome bPos. 10: A → CPos. 45: C → TPos. 60: A → GPos. 67: C → TPos. 71: A → CPos. 126: G → APos. 127: C → TPos. 155: A → GPos. 211: C → TPos. 268: T → APos. 274: C → TPos. 327: C → APos. 381: C → TPos. 393: C → TPos. 411: T → C12SPos. 18: T → CPos. 24: T → CPos. 34: T → CPos. 152: A → CPos. 250: A → TPos. 440: A → GPos. 452: T → CPos. 454: T → CPos. 460: T → APos. 576: A → TPos. 593: C → TPos. 683: A → GPos. 729: T → CPos. 775: G → APos. 814: C → TPos. 864: C → TPos. 875: C → APos. 965: C → TPos. 1048: T → CPos. 1144: T → CPos. 1194: A → CPos. 1297: T → CPos. 1307: T → CPos. 1315: A → CPos. 1316: A → GPos. 1376: A → GPos. 1509: A → GPos. 1585: A → T16SPos. 3: A → GPos. 107: T → APos. 138: A → CPos. 220: C → TPos. 411: C → APos. 452: T → APos. 543: T → CPos. 606: C → APos. 622: Gap → CPos. 625: T → CPos. 634: T → GapPos. 665: A → CPos. 667: A → GPos. 668: T → CPos. 736: A → TPos. 751: C → APos. 758: T → CPos. 777: A → TPos. 787: T → CPos. 844: A → TPos. 863: C → TPos. 890: G → A
Pos. 997: C → TPos. 1025: T → CPos. 1049: T → GapPos. 1080: Gap → APos. 1085: T → CPos. 1174: C → APos. 1261: G → TPos. 1267: Gap → GPos. 1451: T → GPos. 1530: A → GPos. 1614: A → TPos. 1620: T → CPos. 1654: A → TPos. 1705: T → APos. 1741: T → APos. 1817: A → GapPos. 1844: Gap → CPos. 1865: T → APos. 1893: G → APos. 1952: T → CPos. 2025: C → TPos. 2034: A → GPos. 2041: CT → GPos. 2043: A → TPos. 2058: A → CPos. 2068: T → CPos. 2107: A → TPos. 2120: A → TPos. 2122: T → CPos. 2130: C → TPos. 2154: T → CPos. 2156: C → TPos. 2190: A → GPos. 2229: A → TPos. 2525: T → APos. 2812: T → APos. 2872: Gap → CPos. 3014: T → APos. 3032: T → CPos. 3125: A → T28SPos. 563: G → TPos. 569: A → TRAG-1Pos. 59 G → APos. 110 C → TPos. 131 G → APos. 197 T → CPos. 201 A → CPos. 233 G → APos. 398 A → CRhodopsinPos. 9: C → TPos. 135: C → TPos. 219: C → TSIAPos. 27: G → APos. 100: A → TPos. 151: C → APos. 241: G → ATyrosinasePos. 58: G → APos. 144: G → CPos. 230: C → TPos. 257: C → TPos. 270: G → APos. 321: C → TPos. 342: C → TPos. 360: G → TPos. 435: C → TPos. 450: C → T
Pos. 475: C → T
Itapotihyla langsdorffii
Cytochrome bPos. 13: A → CPos. 14: T → APos. 17: A → CPos. 24: G → APos. 36: C → GPos. 63: C → TPos. 91: T → CPos. 115: C → TPos. 158: C → GPos. 227: A → TPos. 241: A → CPos. 244: A → TPos. 259: C → TPos. 277: T → CPos. 296: C → TPos. 299: T → CPos. 319: C → TPos. 327: C → TPos. 334: A → CPos. 344: A → GPos. 363: A → GPos. 383: A → TPos. 396: A → CPos. 420: A → G12SPos. 18: T → CPos. 26: A → GPos. 33: A → GPos. 36: A → TPos. 148: C → TPos. 169: A → CPos. 206: A → GPos. 297: T → APos. 452: T → CPos. 541: C → APos. 594: A → GPos. 615: T → CPos. 650: T → CPos. 672: A → GPos. 683: A → GPos. 743: A → TPos. 761: C → TPos. 880: A → TPos. 989: A → GPos. 1008: T → CPos. 1010: T → APos. 1048: T → APos. 1118: C → APos. 1269: T → CPos. 1321: T → CPos. 1343: T → CPos. 1392: T → CPos. 1422: G → APos. 1429: C → APos. 1439: C → TPos. 1631: Gap → TPos. 1649: A → GapPos. 1761: T → C16SPos. 172: A → CPos. 347: C → TPos. 355: C → APos. 376: A → TPos. 498: T → CPos. 548: A → CPos. 634: A → CPos. 753: A → T
2005 223FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 757: A → GPos. 848: C → APos. 863: C → TPos. 1004: G → APos. 1077: T → GPos. 1211: A → GPos. 1228: A → CPos. 1246: T → APos. 1259: T → APos. 1278: T → CPos. 1419: C → APos. 1443: T → GPos. 1468: A → GPos. 1727: A → GapPos. 1823: T → CPos. 1932: T → APos. 1949: A → GPos. 2043: A → CPos. 2059: A → CPos. 2089: C → TPos. 2110: T → CPos. 2112: T → APos. 2115: A → TPos. 2130: C → TPos. 2156: C → TPos. 2176: C → TPos. 2525: T → CPos. 2669: A → TPos. 2844: T → APos. 2901: T → CPos. 2956: A → GPos. 2997: C → GPos. 3010: T → APos. 3014: T → APos. 3018: T → CPos. 3032: T → CPos. 3239: A → GPos. 3287: T → A28SPos. 312: C → TPos. 1482: G → CPos. 1483: T → GRAG-1Pos. 4 C → TPos. 44 A → GPos. 71 G → APos. 173 C → TPos. 311 C → TPos. 357 C → TRhodopsinPos. 316: G → ATyrosinasePos. 104: G → APos. 148: C → TPos. 203: G → APos. 270: G → APos. 297: C → TPos. 399: A → TPos. 444: G → APos. 459: C → TPos. 471: C → T
Nyctimantis rugiceps
Cytochrome bPos. 10: A → CPos. 39: C → TPos. 45: C → TPos. 53: G → APos. 57: A → TPos. 62: G → APos. 91: T → C
Pos. 101: T → APos. 140: A → GPos. 145: T → GPos. 150: T → CPos. 161: A → GPos. 170: A → CPos. 189: C → TPos. 201: T → CPos. 213: G → CPos. 217: T → APos. 232: A → CPos. 238: T → GPos. 262: A → CPos. 268: T → APos. 274: C → TPos. 290: C → TPos. 299: T → CPos. 316: C → TPos. 347: A → GPos. 365: T → CPos. 366: T → CPos. 376: T → CPos. 383: A → GPos. 390: A → GPos. 394: A → GPos. 408: A → G16SPos. 996: A → GPos. 1103: T → CPos. 1153: C → TPos. 1228: A → GPos. 1242: T → CPos. 1246: T → GPos. 1443: T → GapPos. 1503: A → GPos. 1574: A → TPos. 1602: T → CPos. 1607: T → GPos. 1654: A → GapPos. 1727: A → TPos. 1766: Gap → CPos. 1826: C → TPos. 1894: T → CPos. 1925: C → TPos. 1960: A → GPos. 2030: G → APos. 2034: A → GPos. 2091: C → APos. 2094: A → GPos. 2115: A → GPos. 2117: T → CPos. 2200: A → GPos. 2229: A → GPos. 2233: T → GPos. 2265: T → CPos. 2279: T → APos. 2288: A → GPos. 2315: A → GPos. 2461: C → TPos. 2508: T → CPos. 2518: A → CPos. 2525: T → CPos. 2544: G → APos. 2564: T → CPos. 2570: A → TPos. 2616: A → GPos. 2643: A → GPos. 2694: A → GPos. 2737: Gap → TPos. 2889: C → APos. 2937: A → T
Pos. 2966: C → TPos. 2975: A → GPos. 2997: C → TPos. 3081: A → TPos. 3106: T → APos. 3138: A → GPos. 3230: C → TPos. 3239: A → TPos. 3272: Gap → TPos. 3285: G → TPos. 3326: C → TPos. 3406: T → CPos. 3425: C → TPos. 3486: A → G
Osteocephalus
12SPos. 46: A → TPos. 386: A → TPos. 521: T → CPos. 646: T → CPos. 673: T → GapPos. 675: A → CPos. 1327: A → GPos. 1579: G → TtRNA valinePos. 98: G → A16SPos. 367: T → APos. 498: T → APos. 576: A → TPos. 616: A → GPos. 1015: T → APos. 1741: T → CPos. 1813: T → APos. 1839: C → APos. 1847: A → TPos. 1948: C → APos. 2019: A → TPos. 2041: C → APos. 2058: A → TPos. 2059: A → CPos. 2080: A → CPos. 2117: T → CPos. 2244: T → APos. 2308: T → CPos. 2450: C → TPos. 2508: T → CPos. 2509: C → TPos. 2538: C → TPos. 2780: A → TPos. 3018: T → ARAG-1Pos. 127 T → C
Osteopilus
Cytochrome bPos. 127: C → TPos. 128: A → CTPos. 201: A → CPos. 299: T → C12SPos. 36: A → GPos. 110: A → CPos. 131: T → APos. 368: A → GPos. 592: C → TPos. 781: A → CPos. 864: C → TPos. 1245: C → TPos. 1683: C → A16S
Pos. 580: A → CPos. 625: T → GPos. 780: C → APos. 839: C → APos. 853: A → TPos. 1353: A → TPos. 1470: C → APos. 1607: T → CPos. 1789: T → APos. 1823: T → CPos. 2089: C → APos. 2233: T → CPos. 2820: A → TPos. 3054: T → CPos. 3516: A → CRAG-1Pos. 58 A → GPos. 230 C → TPos. 266 C → TPos. 404 T → CPos. 425 G → ATyrosinasePos. 43: T → CPos. 127: T → CPos. 157: C → TPos. 218: G → APos. 294: C → TPos. 357: T → CPos. 381: C → TPos. 396: G → APos. 444: G → TPos. 526: C → T
Phyllodytes
Cytochrome bPos. 10: A → CPos. 33: C → TPos. 46: C → TPos. 48: Gap → APos. 53: G → GapPos. 68: C → APos. 71: A → CPos. 79: A → CPos. 111: C → TPos. 136: C → TPos. 183: A → TPos. 199: G → APos. 201: A → TPos. 214: C → TPos. 223: A → TPos. 241: A → TPos. 244: A → CPos. 246: A → GPos. 290: C → TPos. 331: CT → APos. 402: T → CPos. 403: G → APos. 414: T → A12SPos. 22: G → APos. 34: T → CPos. 45: T → CPos. 46: A → TPos. 61: C → APos. 251: G → APos. 278: C → TPos. 281: A → CPos. 293: G → APos. 313: C → TPos. 322: C → APos. 348: T → C
224 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 427: C → APos. 455: C → GapPos. 480: A → GPos. 521: T → CPos. 547: A → GapPos. 601: A → CPos. 619: G → APos. 621: A → GPos. 670: A → GPos. 816: A → TPos. 864: C → TPos. 1061: T → APos. 1128: T → CPos. 1135: G → APos. 1177: C → TPos. 1187: C → TPos. 1233: G → APos. 1316: A → GPos. 1383: C → APos. 1405: A → GPos. 1431: T → CPos. 1509: A → GPos. 1602: T → APos. 1640: T → CPos. 1664: A → CPos. 1692: G → AtRNA valinePos. 56: T → CPos. 61: C → TPos. 76: A → GPos. 77: A → TPos. 94: C → T16SPos. 2: T → CPos. 12: C → APos. 130: C → APos. 367: AT → CPos. 383: A → CPos. 452: T → CPos. 483: A → TPos. 536: C → TPos. 579: A → GPos. 592: T → CPos. 623: Gap → CPos. 625: T → CPos. 658: A → GapPos. 668: T → GapPos. 691: G → APos. 780: C → TPos. 784: C → TPos. 941: G → CPos. 960: G → APos. 968: G → APos. 969: A → GPos. 1009: G → APos. 1085: T → APos. 1140: A → TPos. 1223: C → APos. 1259: T → CPos. 1289: A → GPos. 1480: T → CPos. 1585: A → GPos. 1592: G → CPos. 1603: T → CPos. 1673: A → CPos. 1699: A → CPos. 1839: T → CPos. 1925: A → TPos. 1928: C → GPos. 2030: G → APos. 2058: A → C
Pos. 2139: Gap → CPos. 2268: C → APos. 2284: G → APos. 2312: T → APos. 2319: T → CPos. 2441: C → GapPos. 2450: C → TPos. 2457: G → APos. 2477: C → TPos. 2484: T → CPos. 2496: G → APos. 2503: C → APos. 2518: A → GPos. 2551: A → GPos. 2568: C → TPos. 2652: T → CPos. 2711: C → TPos. 2800: Gap → CPos. 2856: T → APos. 2982: T → APos. 3039: A → TPos. 3041: G → APos. 3127: T → CPos. 3166: C → GapPos. 3173: G → APos. 3189: G → APos. 3267: Gap → TPos. 3277: G → APos. 3299: C → TPos. 3439: T → CPos. 3447: G → APos. 3449: C → APos. 3458: T → C28SPos. 260: Gap → CPos. 261: Gap → CPos. 283: Gap → TPos. 318: Gap → APos. 440: Gap → GPos. 441: Gap → GPos. 484: C → GPos. 738: Gap → GPos. 741: Gap → CPos. 925: Gap → GPos. 1048: Gap → GRhodopsinPos. 10: G → APos. 279: C → APos. 296: T → CPos. 311: C → TSIAPos. 106: C → TPos. 250: C → TTyrosinasePos. 22: G → APos. 43: T → CPos. 52: C → TPos. 73: A → GPos. 103: G → APos. 127: T → CPos. 157: C → APos. 177: C → TPos. 209: T → CPos. 215: G → APos. 233: C → APos. 256: G → APos. 282: A → CPos. 285: C → TPos. 343: G → APos. 345: G → TPos. 490: G → A
Pos. 507: T → GPos. 529: C → TPos. 537: T → G
Tepuihyla edelcae
12SPos. 26: A → GPos. 231: T → APos. 251: G → APos. 449: Gap → CPos. 451: T → CPos. 452: T → CPos. 454: T → CPos. 528: A → TPos. 594: A → GPos. 696: A → GPos. 875: C → APos. 970: A → CPos. 989: A → GPos. 999: T → CPos. 1010: T → CPos. 1144: T → CPos. 1211: T → CPos. 1309: C → GPos. 1441: T → CPos. 1602: T → APos. 1640: T → CPos. 1683: C → T16SPos. 56: C → GapPos. 162: C → TPos. 220: T → APos. 336: C → APos. 376: A → TPos. 383: C → GPos. 411: C → TPos. 589: T → CPos. 630: A → GPos. 668: T → APos. 671: A → GPos. 718: T → CPos. 736: A → GPos. 742: T → APos. 754: A → GPos. 755: A → GPos. 758: T → CPos. 814: A → TPos. 890: G → APos. 996: A → GPos. 1012: T → CPos. 1018: T → CPos. 1057: C → TPos. 1082: Gap → CPos. 1160: A → CPos. 1174: C → TPos. 1328: A → GPos. 1419: C → TPos. 1436: T → CPos. 1470: C → TPos. 1534: T → CPos. 1699: A → TPos. 1705: T → GPos. 1727: A → TPos. 1783: C → TPos. 1796: A → TPos. 1799: A → TPos. 1823: T → APos. 1880: T → CPos. 1883: C → TPos. 1894: T → CPos. 2039: G → A
Pos. 2063: A → GPos. 2067: A → GPos. 2154: T → CPos. 2177: C → TPos. 2217: T → CPos. 2395: A → TPos. 2533: T → CPos. 2564: T → CPos. 2566: T → CPos. 2616: A → GPos. 2676: A → GPos. 3010: T → APos. 3027: T → CPos. 3056: T → CPos. 3077: T → CPos. 3111: G → APos. 3138: A → GPos. 3140: C → TPos. 3166: C → TPos. 3222: C → TPos. 3234: T → APos. 3287: T → CPos. 3295: T → CPos. 3298: G → APos. 3439: T → CRAG-1Pos. 419: C → T
Trachycephalus
Cytochrome bPos. 36: C → APos. 173: A → CPos. 201: A → CPos. 247: C → T12SPos. 169: A → TPos. 838: A → TPos. 880: A → CPos. 1636: C → GapPos. 1674: C → T16SPos. 130: C → APos. 188: AT → CPos. 718: T → APos. 836: C → APos. 1259: T → CPos. 1443: T → CPos. 1470: C → APos. 1534: T → CPos. 2229: A → CPos. 2525: T → CPos. 2753: A → CPos. 2827: Gap → TPos. 3287: T → CRAG-1Pos. 6 T → CPos. 125 G → APos. 242 G → ARhodopsinPos. 96: T → CPos. 154: C → APos. 160: C → TPos. 209: C → TPos. 220: G → ATyrosinasePos. 70: T → APos. 251: C → TPos. 351: C → TPos. 456: G → APos. 490: G → APos. 497: C → A
2005 225FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 511: G → A
Phyllomedusinae
Cytochrome bPos. 11: A → CPos. 53: G → APos. 71: A → TPos. 78: C → GapPos. 88: C → TPos. 126: G → APos. 134: C → TPos. 136: C → TPos. 204: C → T12SPos. 13: G → APos. 22: G → A
Pos. 83: A → G
Pos. 235: A → T
Pos. 278: C → T
Pos. 281: A → T
Pos. 293: G → A
Pos. 313: C → A
Pos. 389: T → A
Pos. 528: A → T
Pos. 602: A → C
Pos. 641: AT → G
Pos. 658: A → G
Pos. 672: C → A
Pos. 771: C → T
Pos. 775: G → A
Pos. 814: C → T
Pos. 817: G → A
Pos. 838: A → T
Pos. 900: T → A
Pos. 965: T → C
Pos. 1166: Gap → C
Pos. 1321: T → C
Pos. 1392: T → C
Pos. 1439: C → T
Pos. 1568: A → T
tRNA valine
Pos. 57: C → T
Pos. 76: A → C
16S
Pos. 403: Gap → A
Pos. 427: T → C
Pos. 579: A → G
Pos. 616: A → C
Pos. 626: Gap → C
Pos. 664: A → T
Pos. 758: A → T
Pos. 807: G → A
Pos. 861: A → C
Pos. 890: A → G
Pos. 904: A → T
Pos. 948: T → C
Pos. 1167: A → T
Pos. 1758: Gap → A
Pos. 1799: A → T
Pos. 1819: A → T
Pos. 1948: T → C
Pos. 2045: A → C
Pos. 2069: A → T
Pos. 2070: A → C
Pos. 2127: G → A
Pos. 2154: T → A
Pos. 2161: A → T
Pos. 2179: C → T
Pos. 2203: A → T
Pos. 2396: Gap → C
Pos. 2669: A → T
Pos. 2768: C → T
Pos. 2906: T → C
Pos. 2953: A → G
Pos. 3041: G → A
Pos. 3097: A → C
Pos. 3126: A → T
Pos. 3145: T → A
Pos. 3222: T → C
Pos. 3234: A → Gap
Pos. 3285: G → A
Pos. 3292: C → T
Pos. 3429: T → C
Pos. 3451: A → G
28S
Pos. 223: Gap → G
Pos. 232: Gap → G
Pos. 327: T → G
Pos. 505: G → T
Pos. 558: G → T
Pos. 564: Gap → A
Pos. 825: Gap → C
Pos. 876: A → C
Pos. 934: Gap → T
Rhodopsin
Pos. 10: A → G
Pos. 16: A → T
Pos. 18: G → T
Pos. 93: C → T
Pos. 105: G → A
Pos. 145: C → T
Agalychnis
12S
Pos. 73: T → C
Pos. 159: C → T
Pos. 164: T → C
Pos. 277: A → T
Pos. 550: T → A
Pos. 558: T → C
Pos. 599: T → C
Pos. 864: C → T
Pos. 1144: T → C
Pos. 1321: C → T
Pos. 1539: C → T
tRNA valine
Pos. 39: T → C
16S
Pos. 220: T → A
Pos. 479: Gap → A
Pos. 536: C → T
Pos. 904: T → C
Pos. 2151: T → C
Pos. 2312: T → A
Pos. 3097: C → T
Pos. 3111: G → A
Pos. 3290: A → T
Cruziohyla calcarifer
Cytochrome b
Pos. 10: A → C
Pos. 13: A → C
Pos. 33: C → T
Pos. 64: C → T
Pos. 105: A → C
Pos. 108: A → C
Pos. 115: C → T
Pos. 116: C → T
Pos. 137: T → A
Pos. 144: C → T
Pos. 152: T → C
Pos. 158: C → G
Pos. 183: C → A
Pos. 202: G → A
Pos. 214: C → T
Pos. 265: C → T
Pos. 271: C → T
Pos. 290: C → T
Pos. 306: A → G
Pos. 313: A → G
Pos. 327: C → T
Pos. 334: A → C
Pos. 347: A → C
Pos. 362: A → C
Pos. 411: T → C
Pos. 415: C → T
12S
Pos. 61: C → T
Pos. 73: T → C
Pos. 88: C → T
Pos. 98: T → Gap
Pos. 142: C → T
Pos. 152: A → C
Pos. 173: G → A
Pos. 206: A → T
Pos. 231: A → Gap
Pos. 280: C → T
Pos. 291: G → A
Pos. 319: A → T
Pos. 359: A → T
Pos. 370: A → T
Pos. 391: C → T
Pos. 512: T → C
Pos. 521: T → C
Pos. 550: T → A
Pos. 599: T → C
Pos. 613: C → T
Pos. 646: T → C
Pos. 673: C → T
Pos. 692: T → C
Pos. 729: T → A
Pos. 776: G → A
Pos. 812: C → T
Pos. 862: Gap → T
Pos. 869: C → A
Pos. 875: C → G
Pos. 908: A → C
Pos. 1002: C → T
Pos. 1061: T → C
Pos. 1116: A → T
Pos. 1144: T → C
Pos. 1194: A → C
Pos. 1307: T → A
Pos. 1408: A → C
Pos. 1600: Gap → C
Pos. 1670: T → C
tRNA valine
Pos. 3: A → T
Pos. 11: G → A
Pos. 54: C → T
16S
Pos. 3: A → Gap
Pos. 4: A → C
Pos. 14: Gap → C
Pos. 51: Gap → C
Pos. 107: A → C
Pos. 118: A → C
Pos. 172: A → C
Pos. 294: A → G
Pos. 347: C → T
Pos. 355: C → T
Pos. 443: T → A
Pos. 452: T → C
Pos. 539: Gap → A
Pos. 544: Gap → C
Pos. 573: A → Gap
Pos. 586: Gap → C
Pos. 592: T → C
Pos. 634: A → G
Pos. 665: T → G
Pos. 668: T → A
Pos. 692: A → T
Pos. 718: T → C
Pos. 763: G → A
Pos. 767: A → T
Pos. 784: C → T
Pos. 788: C → T
Pos. 798: T → C
Pos. 827: T → C
Pos. 920: G → A
226 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
APPENDIX 5(Continued)
Pos. 972: T → A
Pos. 1008: A → C
Pos. 1049: T → A
Pos. 1153: A → T
Pos. 1174: T → A
Pos. 1235: C → T
Pos. 1242: C → T
Pos. 1468: T → A
Pos. 1480: T → C
Pos. 1491: A → C
Pos. 1572: T → C
Pos. 1631: T → C
Pos. 1783: C → T
Pos. 1817: T → C
Pos. 1823: T → C
Pos. 1826: T → A
Pos. 1887: T → C
Pos. 1925: A → G
Pos. 2016: T → C
Pos. 2018: T → A
Pos. 2039: G → A
Pos. 2064: C → T
Pos. 2208: A → C
Pos. 2217: T → A
Pos. 2250: C → T
Pos. 2379: Gap → T
Pos. 2391: Gap → C
Pos. 2406: T → C
Pos. 2450: C → T
Pos. 2517: G → A
Pos. 2566: T → C
Pos. 2581: C → T
Pos. 2605: G → A
Pos. 2616: A → T
Pos. 2652: T → C
Pos. 2664: A → T
Pos. 2671: Gap → C
Pos. 2711: C → T
Pos. 2741: Gap → C
Pos. 2856: T → A
Pos. 2913: T → C
Pos. 2985: T → C
Pos. 3010: T → A
Pos. 3032: T → A
Pos. 3048: A → T
Pos. 3071: A → T
Pos. 3083: T → C
Pos. 3088: A → G
Pos. 3195: Gap → T
Pos. 3199: A → C
Pos. 3303: C → T
Pos. 3310: A → T
Pos. 3316: A → T
Pos. 3320: G → A
Pos. 3357: A → T
Pos. 3431: C → T
Pos. 3432: A → G
Pos. 3458: T → C
Pos. 3499: C → T
Pos. 3516: A → G
28S
Pos. 207: Gap → G
Pos. 208: C → T
Pos. 238: G → Gap
Pos. 306: T → G
Pos. 373: C → T
Pos. 390: C → T
Pos. 509: G → Gap
Pos. 511: C → T
Pos. 516: G → A
Pos. 653: C → T
Rhodopsin
Pos. 6: T → C
Pos. 9: C → T
Pos. 314: C → T
Hylomantis
Cytochrome b
Pos. 33: C → T
Pos. 98: C → T
Pos. 126: A → G
Pos. 170: T → C
Pos. 204: T → C
Pos. 250: T → A
Pos. 274: C → T
Pos. 316: T → C
Pos. 402: T → C
12S
Pos. 83: G → A
Pos. 251: G → A
Pos. 528: T → A
Pos. 875: C → T
Pos. 1549: A → C
tRNA valine
Pos. 98: A → G
16S
Pos. 50: Gap → C
Pos. 252: Gap → T
Pos. 347: C → T
Pos. 399: T → Gap
Pos. 452: T → C
Pos. 459: T → C
Pos. 625: T → C
Pos. 774: T → A
Pos. 1160: T → A
Pos. 1242: C → T
Pos. 1278: T → C
Pos. 1334: A → T
Pos. 1737: T → C
Pos. 1783: C → T
Pos. 2112: T → Gap
Pos. 2605: G → A
Pos. 2616: A → T
Pos. 2866: A → Gap
Pos. 2875: A → Gap
Pos. 2906: C → Gap
Pos. 2946: A → C
Pos. 3277: G → A
Pos. 3375: T → C
Rhodopsin
Pos. 305: C → T
Pachymedusa dacnicolor
12S
Pos. 28: A → G
Pos. 33: A → G
Pos. 49: C → T
Pos. 117: A → T
Pos. 307: G → A
Pos. 370: A → G
Pos. 381: C → T
Pos. 386: A → T
Pos. 414: C → T
Pos. 427: A → G
Pos. 434: G → A
Pos. 451: T → C
Pos. 480: A → G
Pos. 519: Gap → G
Pos. 601: C → T
Pos. 646: T → C
Pos. 658: G → A
Pos. 672: A → C
Pos. 733: T → A
Pos. 736: A → T
Pos. 750: C → T
Pos. 780: C → T
Pos. 835: T → C
Pos. 908: A → C
Pos. 1105: A → T
Pos. 1128: T → C
Pos. 1153: T → G
Pos. 1166: C → T
Pos. 1187: C → T
Pos. 1233: G → A
Pos. 1307: T → C
Pos. 1317: A → G
Pos. 1426: T → C
Pos. 1429: C → A
Pos. 1439: T → C
Pos. 1509: A → G
Pos. 1622: Gap → G
Pos. 1630: A → T
Pos. 1761: T → C
tRNA valine
Pos. 6: G → A
Pos. 34: Gap → T
Pos. 55: A → C
Pos. 84: T → C
Pos. 95: C → T
Pos. 105: C → T
16S
Pos. 94: A → Gap
Pos. 130: A → G
Pos. 272: T → A
Pos. 323: C → T
Pos. 493: Gap → G
Pos. 513: Gap → A
Pos. 580: A → G
Pos. 654: T → A
Pos. 663: T → A
Pos. 667: A → T
Pos. 772: Gap → A
Pos. 777: C → Gap
Pos. 807: A → G
Pos. 818: C → T
Pos. 872: C → T
Pos. 959: A → G
Pos. 983: G → A
Pos. 1034: G → Gap
Pos. 1049: T → A
Pos. 1167: T → A
Pos. 1208: Gap → A
Pos. 1268: C → T
Pos. 1281: T → C
Pos. 1370: Gap → T
Pos. 1371: Gap → T
Pos. 1468: T → C
Pos. 1491: A → T
Pos. 1699: T → A
Pos. 1855: A → T
Pos. 2007: A → T
Pos. 2033: A → T
Pos. 2041: A → T
Pos. 2049: G → A
Pos. 2054: T → C
Pos. 2060: C → T
Pos. 2070: C → A
Pos. 2105: A → G
Pos. 2107: C → T
Pos. 2552: T → C
Pos. 2606: A → G
Pos. 2717: Gap → T
Pos. 2797: Gap → C
Pos. 2844: A → T
Pos. 2913: T → G
Pos. 3071: A → T
Pos. 3083: T → C
Pos. 3121: A → T
Pos. 3181: G → A
Pos. 3189: G → T
Pos. 3199: A → T
Pos. 3249: G → Gap
Pos. 3260: C → Gap
Pos. 3266: C → T
Pos. 3268: C → T
Pos. 3291: A → T
2005 227FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
APPENDIX 5(Continued)
Pos. 3439: T → C
Pos. 3504: C → T
Pos. 3507: C → T
SIA
Pos. 93: C → T
Pos. 220: G → A
Phasmahyla
Cytochrome b
Pos. 10: A → C
Pos. 36: A → T
Pos. 99: T → C
Pos. 137: T → A
Pos. 173: A → T
Pos. 207: C → T
Pos. 248: C → T
Pos. 250: T → A
Pos. 280: C → T
Pos. 356: C → T
Pos. 362: A → T
Pos. 380: C → T
12S
Pos. 61: C → T
Pos. 69: C → T
Pos. 142: C → T
Pos. 173: G → A
Pos. 281: T → A
Pos. 319: A → T
Pos. 432: T → A
Pos. 518: Gap → C
Pos. 547: A → T
Pos. 600: T → C
Pos. 835: T → C
Pos. 861: Gap → C
Pos. 908: A → C
Pos. 989: A → T
Pos. 1036: C → T
Pos. 1048: T → A
Pos. 1058: Gap → T
Pos. 1164: Gap → T
Pos. 1233: G → A
Pos. 1454: G → A
Pos. 1549: A → T
Pos. 1645: T → C
tRNA valine
Pos. 59: C → T
Pos. 69: G → A
Pos. 82: C → T
16S
Pos. 162: C → T
Pos. 343: Gap → T
Pos. 355: C → T
Pos. 580: A → C
Pos. 668: T → A
Pos. 718: T → A
Pos. 751: C → T
Pos. 761: T → C
Pos. 780: C → T
Pos. 822: A → T
Pos. 861: C → A
Pos. 1085: T → A
Pos. 1324: A → T
Pos. 1510: A → T
Pos. 1534: A → Gap
Pos. 1606: Gap → A
Pos. 1699: T → Gap
Pos. 1705: T → A
Pos. 1787: C → T
Pos. 1874: T → C
Pos. 1878: C → T
Pos. 1932: T → C
Pos. 1951: A → T
Pos. 1964: C → T
Pos. 2018: T → A
Pos. 2026: A → C
Pos. 2028: G → A
Pos. 2030: G → T
Pos. 2057: A → T
Pos. 2059: A → T
Pos. 2112: T → A
Pos. 2345: C → T
Pos. 2381: T → A
Pos. 2398: C → T
Pos. 2416: G → A
Pos. 2422: C → T
Pos. 2552: T → C
Pos. 2566: T → C
Pos. 2581: C → T
Pos. 2583: A → G
Pos. 2586: C → T
Pos. 2674: C → T
Pos. 2711: C → T
Pos. 2975: A → T
Pos. 3036: T → A
Pos. 3059: A → C
Pos. 3111: G → A
Pos. 3140: C → T
Pos. 3199: A → G
Pos. 3230: T → C
Pos. 3323: A → T
Pos. 3375: T → Gap
Pos. 3484: G → A
Pos. 3491: C → A
Pos. 3551: C → T
Phyllomedusa
12S
Pos. 65: C → A
Pos. 251: G → A
Pos. 313: A → T
Pos. 327: A → T
Pos. 391: C → T
Pos. 434: G → A
Pos. 530: Gap → C
Pos. 864: C → A
Pos. 1066: C → T
Pos. 1130: Gap → C
Pos. 1177: T → Gap
Pos. 1585: T → A
Pos. 1614: A → T
tRNA valine
Pos. 98: A → G
16S
Pos. 348: Gap → A
Pos. 427: C → T
Pos. 483: T → C
Pos. 573: A → T
Pos. 610: T → C
Pos. 634: A → T
Pos. 1025: T → C
Pos. 1077: T → A
Pos. 1153: A → C
Pos. 1274: A → G
Pos. 1737: T → C
Pos. 1819: T → A
Pos. 1897: T → C
Pos. 2045: C → T
Pos. 2094: A → Gap
Pos. 2107: C → A
Pos. 2203: T → C
Pos. 2217: T → C
Pos. 2233: T → A
Pos. 2285: A → G
Pos. 2567: T → C
Pos. 2664: A → T
Pos. 3143: G → A
RAG-1
Pos. 68 A → G
Rhodopsin
Pos. 133: C → T
Pos. 305: C → T
Pos. 314: C → T
SIA
Pos. 148: C → T
Tyrosinase
Pos. 2: T → C
Pos. 444: C → T
Pelodryadinae
Cytochrome b
Pos. 62: G → T
Pos. 164: C → T
12S
Pos. 61: C → A
Pos. 169: A → Gap
Pos. 194: T → C
Pos. 432: T → A
Pos. 619: G → A
Pos. 621: A → G
Pos. 661: A → G
Pos. 880: A → C
Pos. 1174: C → T
Pos. 1303: A → T
Pos. 1317: A → T
Pos. 1670: T → A
tRNA valine
Pos. 98: A → G
16S
Pos. 265: Gap → A
Pos. 459: T → A
Pos. 643: Gap → A
Pos. 821: T → A
Pos. 902: T → A
Pos. 914: C → A
Pos. 959: A → G
Pos. 1077: T → A
Pos. 1085: T → A
Pos. 1235: C → A
Pos. 1324: A → C
Pos. 1447: Gap → C
Pos. 1725: Gap → T
Pos. 1896: A → C
Pos. 1900: C → T
Pos. 1945: G → A
Pos. 2112: T → A
Pos. 2200: T → A
Pos. 2224: A → T
Pos. 2267: T → A
Pos. 2268: C → A
Pos. 2381: T → A
Pos. 2676: A → T
Pos. 2719: T → C
Pos. 2860: Gap → A
Pos. 3086: T → A
Pos. 3181: G → A
Pos. 3260: C → T
Pos. 3274: A → T
Pos. 3280: A → T
Pos. 3297: T → A
RAG-1
Pos. 8 A → G
Pos. 89 T → C
Pos. 134 C → T
Pos. 350 C → T
Pos. 410 C → T
Rhodopsin
Pos. 99: G → A
Pos. 270: A → T
Tyrosinase
Pos. 273: C → G
Pos. 282: A → C
Pos. 449: C → G
Pos. 455: T → C
Pos. 456: G → A
228 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Note added in proof
While this paper was in press, two relevant contributions were published, eachone describing a new species of Hyla. Kohler et al. (2005) described Hyla coffea,and tentatively assigned it to the Hyla microcephala group. For these reasons, weinclude this species in the resurrected genus Dendropsophus, as Dendropsophuscoffea (Kohler, Jungfer, and Reichle, 2005) new comb. We follow the authors intentatively assigning it to the Dendropsophus microcephalus group, increasing thenumber of species currently included in this group to 31. Carvalho e Silva andCarvalho e Silva (2005) described as Hyla eugenioi the species that we included inthis paper as Hyla sp. 1 (aff. H. ehrhardti). Therefore, we include this species inAplastodiscus, as Aplastodiscus eugenioi (Carvalho e Silva and Carvalho e Silva,2005) new comb. Following our results, and the authors’ assigning the species tothe former Hyla albofrenata complex of the H. albomarginata group, we considerit a member of the Aplastodiscus albofrenatus group, increasing to seven the numberof species of the group. Carvalho e Silva and Carvalho e Silva (2005) also noticedthat the species of the former Hyla albofrenata complex of the H. albomarginatagroup (now the Aplastodiscus albofrenatus group) have a red-orange iris, while spe-cies of the former H. albosignata complex (now the Aplastodiscus albosignatusgroup) have a characteristic ring (red externally, gray internally).
ReferencesCarvalho e Silva, A.M.P.T., and S.P. Carvalho e Silva. 2005. New species of the
Hyla albofrenata group, from the states of Rio de Janeiro and Sao Paulo, Brazil(Anura, Hylidae). Journal of Herpetology 38: 73–81.
Kohler, J., K.H. Jungfer, and S. Reichle. 2005. Another new species of small Hyla(Anura, Hylidae) from amazonian sub-andean forest of western Bolivia. Journalof Herpetology 39: 43–50.
2005 229FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
INDEX
Numbers in bold indicate pages where only species groups are mentioned.
Acris barbouri 127crepitans 10, 32, 99, 152, 175gryllus 10, 32, 99, 152, 175
Adenomera sp. 10, 197Agalychnis annae 19, 113, 172
calcarifer 11, 19, 54, 113, 114, 172, 192callidryas 11, 19, 113, 172, 182craspedopus 11, 54, 114, 172litodryas 11, 19, 113, 172, 192moreletii 19, 113, 172saltator 11, 19, 113, 172, 193spurrelli 19, 113, 172
Allophryne ruthveni 12, 14, 194Alsodes gargola 12, 50, 198Amphodus wuchereri 109Anotheca spinosa 10, 32, 41, 74, 99, 152, 175,
215Aparasphenodon bokermanni 38, 108, 152
brunoi 10, 38, 108, 123, 152, 175, 222venezolanus 38, 108, 152
Aplastodiscus albofrenatus 82, 152, 153, 156,159, 163, 205albosignatus 82, 152, 154, 157, 158, 162, 206arildae 82, 153callipygius 82, 154cavicola 82, 154cochranae 10, 41, 78, 82, 152, 175ehrhardti 82, 156flumineus 82, 156ibirapitanga 82, 156leucopygius 82, 158musicus 82, 159perviridis 10, 41, 60, 75, 78, 80, 82, 152, 175,
206sibilatus 82, 162weygoldti 82, 163
Argenteohyla siemersi 10, 38, 73, 152, 175, 222siemersi pederseni 38
Atelognathus patagonicus 12, 198Atelopus varius 12, 195
Batrachyla leptopus 12, 198Bokermannohyla ahenea 83, 152
alvarengai 84, 153astartea 83, 153caramaschii 83, 154carvalhoi 83, 154circumdata 82, 83, 87, 123, 128, 152, 153, 154,
155, 156, 157, 158, 159, 161, 162, 207claresignata 83, 155clepsydra 83, 155feioi 83, 155gouveai 83, 155
hylax 83, 157ibitiguara 84, 157ibitipoca 83, 157izecksohni 83, 157langei 83, 123, 158lucianae 83, 158luctuosa 83, 158martinsi 83, 123, 158nanuzae 83, 123, 159, 159pseudopseudis 83, 84, 123, 153, 157, 161, 162,
207ravida 83, 161saxicola 84, 162sazimai 83, 123, 162
Brachycephalus ephippium 12, 14, 49, 195Bromeliohyla bromeliacia 99, 154, 216
dendroscarta 99, 155Bufo arenarum 12, 195
marmoratus 90
Calamita melanorabdotus 111quadrilineatus 111
Calyptahyla crucialis 39, 73Centrolene prosoblepon 12, 14, 195Centrolenella pulidoi 203Ceratophrys cranwelli 12, 50, 196Charadrahyla altipotens 99, 104, 153
chaneque 100, 104, 155nephila 100, 159taeniopus 100, 162trux 100, 163
Cinclidium granulatum 85Cochranella bejaranoi 12, 14, 196
siren 201Colostethus talamancae 12, 14, 196Cophomantis punctillata 85Corythomantis greeningi 10, 38, 108, 109, 152,
175, 222Crossodactylus schmidti 12, 50, 196Cruziohyla calcarifer 113, 114, 118, 172, 226
craspedopus 113, 114, 172Cryptobatrachus sp. 11, 191Cyclorana australis 12, 19, 194
brevipes 19platicephala 19
Dendrobates auratus 12, 14, 196tinctorius 86
Dendrophryniscus minutus 12, 195Dendropsophus acreanus 91, 152
allenorum 93, 153amicorum 93, 153anataliasiasi 92, 153anceps 91, 123, 153
230 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Dendropsophus (continued)aperomeus 92, 121, 153araguaya 92, 153battersbyi 93, 121, 153berthalutzae 92, 154bifurcus 91, 154bipunctatus 92, 154bogerti 90, 154bokermanni 93, 154branneri 92, 154brevifrons 93, 154cachimbo 92, 154carnifex 90, 154cerradensis 92, 155columbianus 90, 121, 154, 155cruzi 92, 155decipiens 92, 154, 155, 157, 159delarivai 93, 121, 155dutrai 91, 156ebraccatus 91, 118, 156elegans 91, 156elianeae 92, 156garagoensis 90, 91, 121, 128, 156, 159, 160,
163gaucheri 93, 156giesleri 93, 123, 156grandisonae 93, 156gryllatus 92, 156haddadi 92, 157haraldschultzi 93, 157jimi 92, 157joannae 92, 157koechlini 93, 157labialis 91, 121, 123, 158, 159, 160leali 92, 158leucophyllatus 91, 153, 154, 156, 158, 161,
162, 163, 210limai 93, 158luteoocellatus 93, 158marmoratus 91, 152, 156, 158, 159, 162, 211mathiassoni 92, 158melanargyreus 91, 158meridensis 91, 159meridianus 92, 159microcephalus 91, 92, 118, 153, 154, 155, 156,
157, 158, 159, 161, 162, 163, 211microps 93, 159minimus 92, 158, 159, 161minusculus 92, 159minutus 92, 93, 123, 153, 155, 158, 159, 163miyatai 92, 159nadhereri 91, 159nanus 92, 159novaisi 91, 159oliveirai 92, 159padreluna 90, 159
parviceps 93, 153, 154, 156, 157, 158, 159,160, 161, 162, 163, 211
pauiniensis 93, 160pelidna 91, 160phlebodes 92, 119, 160praestans 90, 121, 160pseudomeridianus 92, 160rhea 92, 161rhodopeplus 92, 161riveroi 92, 161robertmertensi 92, 119, 161rossalleni 91, 161rubicundulus 92, 153, 154, 155, 156, 157, 161,
163ruschii 93, 161sanborni 92, 161sarayacuensis 91, 162sartori 92, 119, 162schubarti 93, 162seniculus 91, 123, 162soaresi 91, 162stingi 93, 121, 162studerae 92, 162subocularis 93, 119, 162timbeba 93, 163tintinnabulum 93, 163triangulum 91, 163tritaeniatus 92, 163virolinensis 90, 163walfordi 92, 163werneri 92, 163xapuriensis 93, 163yaracuyanus 93, 121, 163
Duellmanohyla chamulae 32, 69, 100, 152ignicolor 32, 69, 100, 152lythrodes 32, 69, 100, 124, 152rufioculis 10, 32, 100, 124, 152, 175salvavida 32, 69, 100, 152schmidtorum 32, 69, 100, 152soralia 10, 32, 69, 100, 152, 175uranochroa 32, 100, 124, 152
Dydinamipus sjoestedti 12, 195
Ecnomiohyla echinata 100, 156fimbrimembra 100, 124, 156miliaria 100, 124, 159minera 100, 159miotympanum 100, 159phantasmagoria 100, 160salvaje 100, 161thysanota 100, 124, 162tuberculosa 100, 119, 163valancifer 100, 163
Edalorhina perezi 12, 197Eleutherodactylus pluvicanorus 12, 197
thymelensis 12, 197Eupsophus calcaratus 12, 50, 198
2005 231FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
Exerodonta abdivita 101, 152bivocata 101, 154catracha 101, 154chimalapa 101, 155juanitae 101, 157melanomma 101, 158perkinsi 101, 160pinorum 101, 160smaragdina 101, 162sumichrasti 100, 101, 155, 162, 164, 218xera 100, 164
Fejervarya limnocharis 12, 199Flectonotus sp. 11, 17, 191
Garbeana garbei 94Gastrotheca cf. marsupiata 11, 192
cornuta 11, 16, 192coronata 99fissipes 11, 16, 192marsupiata 11, 16nicefori 16ovifera 11, 16plumbea 16pseustes 11, 15, 192riobambae 15
Heleophryne purcelli 12, 13, 196Hemiphractus helioi 11, 16, 49, 192Hemisus marmoratus 12, 196Hyalinobatrachium eurignathum 12, 14, 196
estevesi 85, 152, 203taylori 20
Hyla abdivita 34, 101, 152acreana 29, 152ahenea 23, 152albofrenata 10, 21, 41, 45, 54, 56, 60, 76, 78,
80, 152, 153, 156, 159, 163alboguttata 111, 152albolineata 21albomarginata 10, 12, 21, 22, 24, 26, 41, 45,
54, 56, 57, 59, 60, 61, 75, 76, 78, 85, 86, 87,152, 153, 154, 156, 157, 159, 160, 161, 162,163, 176
albonigra 25, 152, 158albopunctata 10, 21, 22, 26, 45, 56, 59, 60, 61,
75, 76, 85, 86, 152, 158, 159, 161, 176albopunctulata 21, 26, 27, 152albosignata 10, 21, 22, 41, 45, 54, 56, 60, 76,
78, 80, 152, 154, 157, 158, 162, 176albovittata 152, 202alemani 24, 76, 88, 152allenorum 30, 31, 64, 153altipotens 35, 153alvarengai 23, 25, 26, 59, 82, 83, 153alytolylax 27, 153ameibothalame 33, 153americana 153
amicorum 31, 153anataliasiasi 31, 92, 153anceps 11, 31, 65, 90, 91, 153, 186andersonii 11, 32, 35, 36, 66, 72, 102, 153, 178andina 10, 25, 76, 153, 183angustilineata 35, 153annectans 10, 33, 102, 153, 177aperomea 30, 153araguaya 31, 92, 153arborea 10, 32, 33, 34, 66, 72, 101, 102, 124,
125, 127, 153, 155, 157, 159, 161, 162, 163,177, 218
arborescandens 10, 34, 66, 68, 104, 153, 182arboricola 34, 102, 153arenicolor 10, 32, 33, 34, 35, 102, 153, 179arianae 21arildae 10, 21, 22, 153, 176armata 10, 26, 27, 40, 57, 84, 153, 155, 177aromatica 10, 40, 54, 59, 76, 89, 153, 157astartea 10, 23, 74, 153, 179atlantica 25, 88, 153aurantiaca 97auraria 111, 153, 202, 203aurata 94avivoca 10, 32, 36, 102, 153, 185balzani 10, 25, 76, 153, 183battersbyi 31, 153baudinii 106beckeri 25, 87, 153benitezi 11, 26, 60, 86, 153, 186berthalutzae 10, 27, 28, 29, 92, 154, 179bifurca 28, 154biobeba 22, 23bipunctata 10, 29, 31, 90, 154, 181bischoffi 10, 24, 25, 88, 154, 183bistincta 10, 33, 37, 66, 68, 70, 104, 153, 154,
155, 158, 159, 160, 161, 162, 177bivocata 34, 101, 154boans 10, 21, 22, 23, 26, 54, 56, 57, 59, 60, 61,
75, 86, 87, 88, 154, 155, 156, 158, 160, 161,163, 177
bocourti 34, 102, 154bogerti 27, 154bogotensis 10, 21, 26, 27, 57, 76, 80, 84, 153,
154, 155, 157, 158, 160, 162, 163bokermanni 30, 31, 64, 154branneri 29, 31, 154brevifrons 10, 30, 31, 64, 154, 182bromeliacia 10, 33, 66, 69, 70, 74, 99, 154,
155, 178buriti 25, 87, 154cachimbo 31, 92, 154cadaverina 36caingua 10, 25, 154, 183calcarata 10, 24, 56, 57, 86, 154, 179callipeza 27, 154callipleura 25, 154
232 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Hyla (continued)callipygia 10, 21, 22, 154, 176calthula 10, 33, 154, 177calvicollina 33, 68, 154calypsa 34, 71, 154caramaschii 23, 154carnifex 10, 27, 154, 179carvalhoi 23, 76, 154catharinae 42catracha 34, 101, 154caucana 27, 154cavicola 10, 21, 22, 155, 176celata 33, 154cembra 33, 155cerradensis 31, 92, 155cf. callipleura 25chaneque 35, 155charadricola 33, 68, 155charazani 10, 26, 76, 155, 177chimalapa 11, 35, 100, 155, 184chinensis 32, 33, 102, 155chlorostea 40, 111, 121, 155chryses 33, 68, 155chrysoscelis 32, 36, 102, 155cinerea 10, 32, 33, 35, 36, 66, 101, 102, 155,
156, 162, 178, 218cipoensis 25, 87, 155circumdata 10, 21, 22, 23, 24, 25, 26, 49, 54,
59, 60, 76, 82, 152, 153, 154, 155, 156, 157,158, 159, 161, 162, 179
claresignata 20, 21, 23, 59, 60, 74, 82, 155clepsydra 23, 76, 155columbiana 10, 27, 31, 65, 154, 155colymba 27, 155, 178cordobae 10, 21, 25, 155, 183crassa 33, 155crepitans 10, 21, 22, 23, 25, 56, 61, 85, 87, 155,
177crucialis 109crucifer 36, 105cruzi 29, 155cyanomma 33, 155cyclada 10, 34, 66, 68, 104, 155, 182cymbalum 25, 155dasynota 90debilis 34, 103, 155decipiens 10, 27, 28, 29, 61, 65, 91, 92, 154,
155, 157, 159delarivai 30, 155dendrophasma 35, 66, 68, 100, 106, 155, 185dendroscarta 33, 69, 155dentei 24, 86, 156denticulenta 27, 155dolloi 96, 156, 203dutrai 29, 156ebraccata 10, 28, 61, 65, 156, 181echinata 35, 156
ehrhardti 21, 22, 156elegans 28, 64, 156elianeae 31, 92, 156elongata 29, 31ericae 10, 25, 61, 156, 184euphorbiacea 10, 32, 34, 102, 156, 179exastis 23, 87, 156eximia 10, 32, 33, 34, 35, 36, 49, 66, 72, 101,
102, 124, 125, 153, 154, 156, 157, 160, 162,163, 179, 218
faber 10, 20, 22, 23, 25, 45, 56, 61, 76, 87,156, 178
fasciata 10, 24, 56, 57, 76, 86, 156, 179feioi 23, 156femoralis 10, 32, 33, 36, 49, 66, 102, 156, 185fimbrimembra 35, 156fluminea 21, 156freicanecae 25, 156frontalis 90fuentei 26, 156fusca 111, 156fuscovaria 204garagoensis 20, 27, 28, 31, 65, 66, 74, 90, 156,
159, 163gaucheri 30, 31, 156geographica 10, 21, 24, 26, 54, 56, 57, 59, 60,
61, 75, 76, 85, 86, 88, 89, 154, 155, 156,157, 159, 160, 161, 162, 179
giesleri 28, 30, 31, 63, 156, 182godmani 10, 34, 41, 66, 67, 70, 104, 107, 156,
158, 160, 162goiana 21, 25, 76, 87, 156goinorumgouveai 23, 156graeceae 35, 156grandisonae 30, 31, 156granosa 10, 21, 24, 26, 54, 56, 57, 60, 76, 80,
85, 86, 88, 152, 156, 160, 162, 180gratiosa 10, 33, 66, 101, 102, 157, 178gryllata 29, 156guentheri 11, 25, 157, 184guibei 24haddadi 28, 29, 92, 157hadroceps 39hallowelli 33, 102, 157haraldschultzi 31, 157hazelae 34, 68, 71, 104, 157heilprini 11, 26, 39, 56, 60, 61, 86, 157, 186helenae 111, 157hobbsi 25, 88, 157hutchinsi 24, 86, 157hylax 10, 23, 49, 157, 179hypochondrialis 116hypselops, 157ibirapitanga 22, 157ibitiguara 24, 83, 157ibitipoca 23, 157
2005 233FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
imitator 111, 157immaculata 33, 102, 157inframaculata 111, 157infucata 35, 157infulata 85inparquesi 10, 40, 57, 76, 78, 80, 89, 157, 177insolita 34, 71, 102, 157intermedia 33, 102, 157izecksohni 23, 157jahni 27, 76, 78, 157japonica 10, 32, 33, 66, 72, 102, 124, 157, 177jimi 31, 92, 157joannae 29, 157joaquini 11, 21, 25, 61, 76, 157, 184juanitae 34, 68, 101, 157kanaima 10, 24, 57, 59, 80, 89, 157, 180karenanneae 96, 158, 206koechlini 30, 31, 157labedactyla 33, 68, 158labialis 10, 27, 28, 64, 65, 158, 159, 160, 180lactea 97lancasteri 29, 34, 71, 103, 158lanciformis 10, 22, 86, 158, 176langei 24, 158langsdorffii 22, 109, 158larinopygion 10, 26, 27, 40, 57, 78, 84, 155,
158, 159, 160, 161, 162lascinia 27, 158latistriata 11, 25, 87, 158, 184leali 29, 30, 158lemai 11, 26, 60, 78, 158, 180leonhardschultzei 106leptolineata 11, 25, 87, 158, 184leucocheila 22, 158leucophyllata 10, 27, 30, 63, 64, 65, 66, 154,
156, 158, 161, 162, 163leucopygia 10, 21, 22, 28, 60, 76, 158, 176limai 31, 93, 158lindae 27, 158loquax 34, 104, 104, 158loveridgei 40, 158lucianae 23, 158luctuosa 23, 158lundii 10, 22, 23, 56, 61, 76, 87, 158, 178luteola 109luteoocellata 30, 31, 158lynchi 27, 158marginata 11, 21, 25, 61, 76, 158, 184marianae 39, 158marianitae 11, 21, 25, 158, 184marmorata 11, 27, 28, 29, 31, 63, 64, 66, 152,
156, 158, 159, 162, 181martinsi 21, 24, 25, 54, 59, 82, 158, 181mathiassoni 29, 158maxima 22megapodia 205, 158melanargyrea 29, 158
melanomma 10, 34, 66, 68, 158, 182melanopleura 21, 25, 61, 159melanorabdota 158meridensis 28, 159meridiana 29, 159meridionalis 32, 33, 102, 127, 159microcephala 10, 27, 28, 29, 30, 31, 61, 63, 64,
65, 66, 91, 92, 154, 156, 157, 158, 159, 160,161, 162, 163, 181
microderma 10, 24, 57, 60, 86, 159, 180microps 28, 29, 30, 31, 63, 64, 159miliaria 11, 35, 66, 68, 69, 70, 71, 155, 156,
159, 185minera 35, 159minima 10, 27, 30, 65, 153, 158, 159, 161, 162minuscula 29, 159minuta 10, 27, 29, 30, 31, 64, 65, 93, 155, 159,
182miocenica 127miofloridiana 127miotympanum 10, 33, 34, 35, 68, 69, 70, 71,
100, 152, 153, 154, 155, 157, 158, 159, 160,182
mixe 10, 34, 70, 103, 104, 159, 182mixomaculata 34, 66, 68, 70, 74, 101, 103, 104,
159, 160miyatai 30, 159, 181molitor 111, 159moraviensis 29multifasciata 10, 22, 45, 76, 86, 159, 176musica 21, 159mykter 33, 159nadhereri 29, 64, 65, 159nana 10, 29, 31, 63, 65, 92, 159, 181nanuzae 23, 159nephila 11, 35, 70, 159, 185novaisi 29, 159nubicola 34, 159ocularis 105oliveirai 27, 28, 29, 92, 159ornatissima 24, 88, 159pacha 10, 12, 27, 159, 180pachyderma 33, 159padreluna 28, 65, 66, 159palaestes 25, 61, 76, 160palliata 111, 160, 203palmata 85palmeri 10, 27, 78, 160, 178pantosticta 10, 27, 160, 180pardalis 10, 21, 22, 23, 56, 61, 87, 160, 178parviceps 10, 27, 28, 29, 30, 31, 63, 64, 65, 66,
153, 154, 156, 157, 158, 159, 160, 161, 162,163, 183
pauiniensis 30, 31, 160pelidna 28, 160pellita 34, 160pellucens 10, 21, 22, 24, 56, 87, 160, 176
234 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Hyla (continued)pentheter 33, 160perkinsi 34, 66, 68, 160, 182phaeopleura 25, 87, 160phantasmagoria 35, 69, 160phlebodes 29, 65, 92, 160phyllognatha 27, 76, 160picadoi 33, 34, 160piceigularis 27, 160picta 10, 34, 41, 70, 107, 160, 180pictipes 10, 33, 34, 66, 70, 71, 74, 102, 103,
104, 154, 155, 157, 158, 160, 161, 162, 163picturata 10, 24, 57, 60, 88, 160, 180pinima 40, 62, 94, 160pinorum 10, 34, 68, 100, 101, 160platydactyla 27, 28, 76, 78, 160plicata 34, 102, 160polytaenia 11, 25, 59, 61, 76, 87, 153, 154, 155,
156, 158, 160, 162, 184pombali 24, 88, 160praestans 27, 31, 90, 160prasina 11, 21, 25, 76, 160, 184psarolaima 27, 160psarosema 33, 160pseudomeridiana 29, 161pseudopseudis 10, 21, 23, 24, 25, 54, 59, 82,
83, 157, 161, 162pseudopuma 10, 34, 35, 66, 70, 71, 74, 102,
153, 156, 157, 161, 183ptychodactyla 27, 161pugnax 21, 22, 23, 161pulchella 11, 21, 25, 27, 54, 56, 59, 60, 61, 75,
76, 85, 152, 153, 154, 155, 156, 157, 158,159, 160, 161, 162, 184, 202
pulchrilineata 39pulidoi 86, 161, 203punctata 11, 21, 25, 26, 54, 56, 57, 59, 60, 76,
80, 85, 86, 88, 153, 157, 161, 184quadrilineata 161raniceps 10, 21, 22, 24, 26, 76, 161, 177ravida 23, 161regilla 36, 105rhea 31, 92, 161rhodopepla 10, 29, 31, 161, 181rhodoporus 25rhythmicus 26, 86, 119, 161riojana 11, 21, 25, 161, 184riveroi 30, 161rivularis 10, 34, 35, 70, 71, 103, 161, 183robertmertensi 29, 161robertsorum 33, 161rodriguezi 110roeschmanni 111, 161roraima 10, 24, 57, 60, 86, 161, 180rosenbergi 22, 23, 161rossalleni 28, 30, 161rubeola 25
rubicundula 11, 27, 29, 30, 31, 61, 65, 91, 92,153, 154, 155, 156, 157, 161, 163, 184
rubra 42, 61rubracyla 21, 24, 87, 161rufitela 10, 21, 22, 56, 76, 87, 161, 176ruschii 30, 31, 64, 161sabrina 33, 68, 161salvadorensissalvaje 35, 71, 161sanborni 11, 29, 31, 65, 92, 162, 181sanchiangensis 33, 102, 161sarampiona 27, 162sarayacuensis 28, 65, 162, 181sarda 33, 102, 162sartori 29, 162savignyi 10, 32, 33, 102, 162, 177saxicola 24, 25, 162sazimai 23, 162schubarti 29, 30, 31, 162secedens 24, 25, 162semiguttata 11, 25, 61, 162, 184semilineata 10, 21, 24, 57, 76, 85, 88, 89, 162,
180senicula 10, 29, 64, 65, 90, 162, 181septentrionalis 39, 109, 162sibilata 22, 162sibleszi 10, 24, 60, 76, 162, 180siemersi 108simmonsi 27, 162simplex 33, 102, 162sioplea 33, 162smaragdina 35, 100, 162smithii 10, 34, 107, 162, 180soaresi 29, 64, 65, 162sp. 1 (aff. H. ehrhardti) 10, 22, 60, 175sp. 2, 11, 60, 78, 186sp. 3, 10, 23, 24, 179sp. 4, 10, 23, 49, 179sp. 5 (aff. H. thorectes) 10, 35, 71, 183sp. 6 (aff. H. pseudopseudis) 11, 24, 83, 183sp. 7 (aff. H. semiguttata) 10, 183sp. 8, 11, 60, 185sp. 9 (aff. H. alvarengai) 11, 82, 83, 186spinosa 99squirella 10, 33, 66, 102, 162, 178staufferorum 27, 162stenocephala 25, 87, 162stingi 31, 162strigilata 94studerae 29, 162subocularis 30, 31, 162sumichrasti 11, 35, 66, 68, 100, 101, 155, 162,
163surinamensis 111suweonensis 32, 33, 102, 162swanstoni 127, 162
2005 235FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
taeniopus 11, 35, 66, 68, 70, 99, 100, 153, 155,159, 162, 163, 185
tapichalaca 10, 27, 57, 163, 180thorectes 35, 66, 71, 102, 103, 104, 162thysanota 35, 162tica 35, 163timbeba 30, 31, 162tintinnabulum 31, 163torrenticola 27, 163triangulum 28, 163, 181triseriata 105tritaeniata 31, 92, 163truncata 43, 98trux 35, 70, 163tsinlingensis 33, 102, 163tuberculosa 11, 35, 66, 68, 69, 71, 74, 100, 156,
159, 160, 161, 162, 163uranochroa 100uruguaya 11, 40, 61, 62, 94, 95, 160, 163, 185ussuriensis 33, 102, 163valancifer 35, 163varelae 26, 163vasta 39versicolor 11, 32, 35, 36, 49, 66, 72, 101, 102,
153, 155, 163, 185, 218vigilans 40, 111, 121, 163viridis 101virolinensis 28, 65, 163walfordi 10, 29, 163, 181walkeri 10, 34, 102, 163, 179warreni 41, 111, 119, 121, 163wavrini 23, 88, 163werneri 29, 31, 163weygoldti 10, 21, 22, 163, 176wilderi 39wrightorum 34, 102, 163xanthosticta 35, 163xapuriensis 30, 163xera 11, 35, 100, 163, 185yaracuyana 32, 163zonata 111zeteki 33, 35, 71, 103, 163zhaopingensis 102, 163
Hylella tenera 90Hylomantis aspera 19, 114, 115, 171
buckleyi 115, 173danieli 115, 173granulosa 11, 19, 114, 115, 116, 172, 193hulli 115, 173lemur 115, 173medinai 115, 173psilopygion 115, 173
Hylonomus bogotensis 84Hylopsis platycephalus 97Hyloscirtus albopunctulatus 84, 152
alytolylax 84, 153armatus 83, 84, 153, 155, 207
bogotensis 83, 84, 152, 153, 154, 155, 157,158, 160, 162, 163, 203, 208
callypeza 84, 154caucanus 85, 154charazani 84, 155colymba 84, 155denticulentus 85, 155estevesi 85, 152, 203jahni 85, 157, 203larinopygion 83, 85, 155, 158, 159, 160, 161,
162, 208lascinius 85, 158lindae 85, 158lynchi 85, 158pacha 85, 159palmeri 85, 160pantostictus 85, 160phyllognathus 85, 160piceigularis 85, 160platydactylus 85, 160, 203psarolaimus 85, 160ptychodactylus 85, 161sarampiona 85, 162simmonsi 85, 162staufferorum 85, 162tapichalaca 85, 162torrenticola 85, 163
Hypsiboas albomarginatus 87, 152alboniger 88, 152albopunctatus 86, 123, 124, 152, 154, 155, 156,
157, 158, 159, 161, 208alemani 88, 152andinus 88, 153atlanticus 88, 153balzani 88, 153beckeri 88, 153, 204benitezi 87, 119, 153, 157, 158, 159, 161, 203bischoffi 88, 154boans 89, 118, 154buriti 88, 154caingua 88, 154calcaratus 86, 154callipleura 88, 154cipoensis 88, 155cordobae 88, 155crepitans 87, 123cymbalum 88, 123dentei 86, 123ericae 88, 156exastis 87, 156faber 87, 123, 152, 155, 156, 159, 160, 161,
208fasciatus 86, 156freicanecae 88, 156fuentei 89, 156geographicus 89, 156goianus 88, 156
236 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Hypsiboas (continued)granosus 88, 156guentheri 88, 157heilprini 86, 124, 157hobbsi 88, 157hutchinsi 87, 88, 119, 157hypselops 111joaquini 88, 157lanciformis 86, 158latistriatus 88, 158lemai 87, 158leptolineatus 88, 158leucocheilus 86, 158lundii 87, 158marginatus 88, 158marianitae 88, 158melanopleurus 88, 158microderma 87, 88, 119, 121, 159miliarius 100multifasciatus 86, 159ornatissimus 88, 159palaestes 88, 159pardalis 87, 160pellucens 87, 160, 161, 209phaeopleura 88, 160picturatus 88, 121, 160polytaenius 87, 88, 153, 154, 155, 158, 160,
162pombali 89, 160prasinus 88, 160pugnax 87, 119, 161pulchellus 87, 88, 121, 123, 152, 153, 154, 155,
156, 157, 158, 159, 160, 161, 162, 209pulidoi 87, 161, 204punctatus 88, 152, 153, 156, 157, 159, 160,
161, 162, 209raniceps 86, 124, 161rhythmicus 87, 119, 161riojanus 88, 161roraima 87, 161rosenbergi 87, 119, 161rubracylus 87, 161rufitelus 87, 118, 161secedens 88, 162semiguttatus 88, 162semilineatus 87, 88, 89, 154, 156, 160, 162,
163, 209sibleszi 88, 119, 162stenocephalus 88, 162varelae 89, 163wavrini 89, 163
Isthmohyla angustilineata 103, 153calypsa 103, 154debilis 103, 155graeceae 103, 156infucata 103, 157
insolita 103, 157lancasteri 103, 158picadoi 103, 160pictipes 103, 154, 155, 157, 158, 160, 161, 163pseudopuma 103, 153, 156, 157, 161rivularis 103, 161tica 103, 163xanthosticta 103, 163zeteki 103, 163
Itapotihyla langsdorffii 109, 123, 164, 223
Kaloula conjuncta 12, 199
Leptodactylus ocellatus 12, 197pentadactylus 20
Limnodynastes salmini 12, 13, 199Limnomedusa macroglossa 12, 50, 197Lithodytes lineatus 12, 20, 197Litoria alboguttata 18
americana 111angiana 18arfakiana 12, 18, 53, 194aurea 12, 18, 53, 194becki 18caerulea 12, 18, 194dahlii 18dorsivena 18eucnemis 18freycineti 12, 18, 194infrafrenata 12, 18, 53, 194iris 112jungguy 61lesueuri 61longirostris 112meiriana 12, 18, 53, 194nannotis 18
Lophyohyla piperata 109Lysapsus caraya 42, 93, 163
laevis 11, 42, 93, 164, 186limellum 11, 42, 63, 93, 164, 186
Mantidactylus femoralis 12, 198Megastomatohyla mixe 104, 159, 219
mixomaculata 104, 159nubicola 104, 159pellita 104, 160
Melanophryniscus klappenbachi 12, 195Myersiohyla aromatica 89, 153
inparquesi 89, 157kanaima 89, 157loveridgei 89, 158
Neobatrachus sudelli 12, 13, 199Nyctimantis rugiceps 11, 41, 109, 164, 186, 223Nyctimystes disrupta 18
kubori 12, 18, 194narinosus 12, 18, 194oktediensis 18
2005 237FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
pulcher 12, 18, 194trachydermis 18tyleri 18papua 18zweifeli 53
Odontophrynus americanus 12, 50, 196Ololygon catharinae 204
fuscovaria 203, 204megapodia 203trachythorax 204
Osornophryne guacamayo 12, 195Osteocephalus buckleyi 38, 39, 109, 164
cabrerai 11, 39, 109, 164, 186deridens 39, 109, 164elkejungingerae 39, 109, 121, 164exophthalmus 38, 39, 109, 164fuscifacies 39, 109, 164heyeri 39, 109, 164langsdorffii 11, 39, 72, 73. 108, 164, 186leoniae 39, 109, 121, 164leprieurii 11, 38, 39, 109, 164, 187mutabor 39, 109, 164oophagus 11, 38, 39, 72, 108, 109, 164, 187pearsoni 38, 39, 109, 121, 164planiceps 39, 109, 164rodriguezi 39, 40subtilis 38, 39, 109, 164taurinus 11, 38, 39, 108, 109, 164, 187verruciger 38, 39, 109, 164yasuni 39, 109, 164
Osteopilus brunneus 73, 108, 109, 164crucialis 11, 39, 73, 107, 109, 164, 187dominicensis 11, 39, 108, 109, 164, 187marianae 73, 107, 108, 109, 164pulchrilineatus 73, 109, 164septentrionalis 11, 39, 108, 109, 164, 187vastus 11, 39, 74, 109, 164, 187wilderi 73, 108, 109, 164
Pachymedusa dacnicolor 11, 19, 113, 116, 172,193, 227
Pedostibes hossi 12, 195Pharyngodon petasatus 107Phasmahyla cochranae 12, 19, 20, 116, 172, 193
exilis 19, 20, 116, 172guttata 12, 19, 20, 116, 172, 193jandaia 19, 20, 116, 172
Phrynohyas coriacea 39, 165hadroceps 11, 39, 165, 187imitatrix 39, 165lepida 39, 165mesophaea 11, 39, 74, 111, 165, 187resinifictrix 11, 39, 165, 187venulosa 11, 39, 108, 165, 187
Phrynomedusa appendiculata 116, 172bokermanni 116, 172
fimbriata 116, 172marginata 116, 172vanzolinii 116, 172
Phrynopus sp 12, 49, 197Phyllobates bicolor 12, 196Phyllodytes acuminatus 41, 42, 110, 165
auratus 41, 42, 110, 123, 165brevirostris 41, 42, 110, 165edelmoi 41, 42, 110, 165gyrinaethes 41, 42, 110, 165kautskyi 41, 42, 110, 165luteolus 11, 41, 42, 110, 165, 187melanomystax 41, 42, 110, 165punctatus 41, 42, 110, 165sp. 11, 42, 187tuberculosus 41, 42, 110, 165wuchereri 41, 42, 110, 165
Phyllomedusa aspera 19atelopoides 117, 118, 172ayeaye 117, 172azurea 116baltea 117, 173bicolor 12, 20, 116, 118, 173, 193boliviana 116, 117, 173buckleyi 10, 19, 20, 54, 114, 115, 173burmeisteri 10, 20, 117, 173camba 117, 173centralis 117, 173coelestis 118, 173dacnicolor 115danieli 115, 173distincta 116, 173duellmani 117, 173ecuatoriana 117, 173fimbriata 19guttata 19, 20, 115hulli 114, 173hypochondrialis 12, 20, 116, 117, 172, 173, 193iheringii 116, 117, 173lemur 12, 20, 54, 114, 193medinai 114, 173megacephala 117, 173oreades 117, 173pailona 116palliata 12, 20, 117, 118, 173, 193perinesos 20, 117, 173psilopygion 54, 173rohdei 117, 173sauvagii 116, 117, 173tarsius 12, 20, 116, 117, 173, 193tetraploidea 12, 20, 113, 116, 117, 173, 193tomopterna 12, 20, 118, 173, 193trinitatis 113, 118, 173vaillanti 12, 20, 117, 118, 173, 193venusta 118, 173
Physalaemus cuvieri 12, 198Platymantis sp. 12, 199
238 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Plectrohyla acanthodes 37, 105, 165ameibothalame 105, 153arborescandens 105, 153avia 37, 105, 165bistincta 105, 124, 153, 154, 155, 157, 158,
159, 160, 161, 162, 220calthula 105, 154calvicollina 105, 154celata 105, 155cembra 105, 155charadricola 105, 155chryses 105, 155chrysopleura 37, 105, 165crassa 105, 155cyanomma 105, 155cyclada 105, 155dasypus 37, 105, 165exquisita 37, 105, 165glandulosa 11, 37, 105, 165, 188guatemalensis 11, 37, 104, 105, 124, 165, 166,
188, 220hartwegi 37, 105, 165hazelae 105, 157, 165ixil 37, 105, 165labedactyla 105, 158lacertosa 37, 105, 165matudai 11, 37, 105, 165, 188mykter 105, 159pachyderma 105, 159pentheter 105, 160pokomchi 37, 105, 166psarosema 105, 160psiloderma 37, 105, 166pycnochila 37, 105, 166quecchi 37, 105, 166robertsorum 105, 161sabrina 105, 161sagorum 37, 105, 166sioplea 105, 162tecunumani 37, 105, 166teuchestes 37, 105, 166thorectes 105, 162
Pleurodema brachyops 12, 198Podonectes palmatus 93Proacris mintoni 127Pseudacris brachyphona 36, 106, 166
brimleyi 36, 106, 166cadaverina 11, 36, 106, 166, 188clarkii 36, 106, 166crucifer 11, 36, 105, 166, 188feriarum 36, 106, 166illinoiensis 36, 106, 166maculata 36, 106, 166nigrita 36, 106, 166nordensis 127ocularis 11, 36, 106, 166, 188ornata 36, 106, 166
regilla 11, 36, 106, 166, 188streckeri 36, 106, 166triseriata 11, 36, 106, 166, 188
Pseudis bolbodactyla 42, 94, 166cardosoi 42, 94, 166fusca 42, 94, 166minuta 11, 42, 63, 94, 166, 188paradoxa 11, 42, 63, 93, 94, 108, 166, 188tocantins 42, 94, 166
Pseudopaludicola falcipes 12, 198Pseudophryne bibroni 12, 13, 199Pternohyla dentata 36, 166Pternohyla fodiens 11, 37, 66, 106, 166, 188Ptychohyla acrochorda 37, 106, 166
adipoventris 106dendrophasma 106, 145erythromma 37, 70, 106, 167euthysanota 11, 37, 69, 106, 167, 188hypomykter 11, 37, 69, 106, 167, 189legleri 37, 69, 70, 106, 124, 167leonhardschultzei 11, 37, 69, 106, 167, 189macrotympanum 37, 69, 106, 167panchoi 37, 69, 106, 167salvadorensis 37, 69, 106, 167sanctaecrucis 37, 70, 106, 167sp. 11, 37, 69, 189spinipollex 11, 37, 69, 106, 167, 189zophodes 11, 37, 69, 106, 167, 189
Rana arborea 101bicolorboans 85gryllus 99leucophyllata 90nigrita 105palmipes 20paradoxa 93temporaria 12, 199tinctoria 86venulosa 111
Rhacophorus bipunctatus 12, 199maculatus 202
Scarthyla goinorum 11, 42, 62, 94, 167, 189, 212ostinodactyla 94
Scaphiophryne marmorata 12, 198Schismaderma carens 12, 195Scinax acuminatus 11, 43, 97, 167, 189
agilis 95, 167albicans 95, 167alcatraz 96, 167altae 97, 119, 167alter 74, 97, 167angrensis 95, 167arduous 96, 167argyreornatus 95, 167ariadne 95, 167
2005 239FAIVOVICH ET AL.: PHYLOGENY OF HYLIDAE
aromothyella 95, 167atratus 96, 167auratus 97, 168baumgardneri 97, 168berthae 11, 43, 95, 168, 189blairi 97, 168boesemani 97, 168boulengeri 11, 43, 96, 118, 168, 189brieni 95, 168caldarum 97, 168canastrensis 95, 168cardosoi 97, 168carnevalli 95, 168castroviejoi 97, 121, 168catharinae 11, 42, 43, 74, 95, 123, 167, 168,
169, 170, 189, 213centralis 95, 168chiquitanus 97, 168crospedospilus 97, 168cruentommus 97, 168, 204curicica 97, 168cuspidatus 97, 168danae 97, 119, 168dolloi 97, 156, 203, 204duartei 97, 168elaeochrous 11, 43, 97, 118, 168, 190eurydice 97, 168exiguous 97, 119, 168flavidus 97, 168flavoguttatus 95, 168funereus 97, 169fuscomarginatus 97, 169, 204fuscovarius 11, 43, 96, 97, 121, 169, 190garbei 96, 169granulatus 97, 169hayii 97, 169, 204heyeri 95, 169hiemalis 95, 169humilis 95, 169ictericus 97, 169jolyi 96, 169jureia 95, 169karenanneae 97, 158, 204kautskyi 95, 169kennedyi 96, 169lindsayi 97, 169littoralis 95, 169littoreus 96, 169longilineus 95, 169luizotavioi 95, 169machadoi 95, 169manriquei 97, 121, 169maracaya 97, 169megapodius 96, 169, 204melloi 96, 169nasicus 11, 43, 97, 169, 190nebulosus 96
obtriangulatus 95, 170oreites 97, 121, 170pachycrus 97, 170parkeri 97, 170pedromedinae 96, 170perereca 97, 170, 204perpusillus 42, 74, 96, 99, 167, 169, 170pinima 97, 160proboscideus 96, 170quinquefasciatus 97, 170ranki 95, 170rizibilis 95, 170rostratus 42, 94, 96, 119, 168, 169 170ruber 11, 42, 43, 62, 94, 95, 97, 123, 156, 157,
160, 163, 167, 168, 169, 170, 171, 190, 213similis 97, 170squalirostris 11, 43, 97, 170, 190staufferi 11, 42, 43, 97, 118, 170, 190, 204strigilatus 96, 170sugillatus 96, 170trachythorax 96, 205, 170, 204trapicheiroi 96, 170trilineatus 97, 171uruguayus 96, 97, 123v-signatus 96, 171wandae 97, 171x-signatus 97, 171
Scytopis hebes 111Smilisca baudinii 11, 37, 106, 171, 190
cyanosticta 11, 37, 38, 106, 171, 190daulinia 106dentata 106, 166fodiens 106, 166phaeota 11, 37, 38, 107, 119, 171, 190puma 11, 37, 38, 107, 119, 171, 191sila 37, 107, 119, 171sordida 37, 38, 107, 119, 171
Sphaenorhynchus bromelicola 43, 96, 98, 171carneus 43, 98, 171dorisae 11, 43, 94, 98, 171, 191lacteus 11, 43, 62, 98, 171, 191orophilus 43, 96, 98, 171palustris 43, 98, 171pauloalvini 43, 96, 98, 171planicola 43, 98, 171platycephalus 43, 98, 171prasinus 43, 96, 98, 171surdus 43, 98, 171
Stefania evansi 11, 16, 17, 192ginesi 11, 16, 17scalae 201schuberti 11, 17, 192
Telmatobius sp. 12, 198Tepuihyla aecii 40, 110, 171
celsae 40, 110, 171edelcae 11, 40, 110, 171, 191
240 NO. 294BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Tepuihyla (continued)galani 40, 110, 171luteolabris 40, 110, 171rimarum 40, 110, 171rodriguezi 40, 111, 172talbergae 40, 111, 172
Tetraprion jordani 111Tlalocohyla godmani 107, 156
loquax 107, 158picta 107, 160smithii 107, 162
Trachycephalus atlas 40, 111, 172coriaceus 111, 164hadroceps 111, 164imitatrix 111, 165
jordani 11, 40, 72, 111,172, 191lepidus 111, 165lichenatus 109marmoratus 109mesophaeus 111, 123, 165nigromaculatus 11, 40, 72, 74, 111, 123, 172,
191resinifictrix 111, 165venulosus 111, 118, 165
Trichobatrachus robustus 12, 195Triprion petasatus 11, 38, 107, 172, 191
spatulatus 36, 38, 107, 172
Xenohyla eugenioi 43, 98, 172truncata 11, 43, 98, 123, 172, 191