THE AMPHIBIAN TREE OF LIFE DARREL R. FROST,1 TARAN GRANT,1,4 JULIAÂ N FAIVOVICH,1,4 RAOUL H. BAIN,1,2 ALEXANDER HAAS,5 CEÂ LIO F.B. HADDAD,6 RAFAEL O. DE SAÂ ,7 ALAN CHANNING,8 MARK WILKINSON,9 STEPHEN C. DONNELLAN,10 CHRISTOPHER J. RAXWORTHY,1 JONATHAN A. CAMPBELL,11 BORIS L. BLOTTO,12 PAUL MOLER,13 ROBERT C. DREWES,14 RONALD A. NUSSBAUM,15 JOHN D. LYNCH,16 DAVID M. GREEN,17 AND WARD C. WHEELER3 1Division of Vertebrate Zoology (Herpetology),2Center for Biodiversity and Conservation, and 3Division of Invertebrate Zoology, American Museum of Natural History (DRF: [email protected]; TG: [email protected]; JF: [email protected]; RHB: [email protected]; CJR: [email protected]; WCW: [email protected]); 4Department of Ecology, Evolution, and Environmental Biology, Columbia University, New York, NY 10027; 5Biocenter Grindel and Zoological Museum Hamburg, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany ([email protected]); 6Departamento de Zoologia, Instituto de BiocieÃncias, Universidade Estadual Paulista (UNESP), Caixa Postal 199, 13506-900 Rio Claro, SaÄo Paulo, Brazil ([email protected]); 7Department of Biology, University of Richmond, Richmond, VA 23173-0001 ([email protected]); 8Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, South Africa ([email protected]); 9Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK ([email protected]); 10South Australia Museum, Evolutionary Biology Unit, North Terrace, Adelaide 5000, South Australia ([email protected]); 11Department of Biology, University of Texas at Arlington, TX 76019-0001 ([email protected]); 12Division HerpetologõÂa, Museo Argentino de Ciencias Naturales ``Bernardino Rivadavia'', Angel Gallardo 470, 1405 Buenos Aires, Argentina ([email protected]); 13Wildlife Research Laboratory, Florida Fish and Wildlife Conservation Commission, 4005 South Main Street, Gainesville, FL 32601-9075 ([email protected]); 14Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, CA 94103-3009 ([email protected]); 15Museum of Zoology and Department of Ecology and Evolutionary Biology, University of Michigan, 1109 Geddes Avenue, Ann Arbor, MI 48109-1079 ([email protected]); 16Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Apartado 7495, BogotaÂ, Colombia ([email protected]); 17Redpath Museum, McGill University, 859 Sherbrooke Street West, Montreal, Quebec H3A 2K6, Canada ([email protected]) BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 297, 370 pp., 71 ®gures, 5 tables, 7 appendices Issued March 15, 2006 Copyright q American Museum of Natural History 2006 ISSN 0003-0090 CONTENTS Abstract ....................................................................... 8 Introduction ................................................................... 11 Materials and Methods ......................................................... 13 Conventions and Abbreviations ................................................ 13 General Analytical Approach: Theoretical Considerations ......................... 13 Taxon Sampling ............................................................. 14 Character Sampling .......................................................... 15 Laboratory Protocols ......................................................... 17 Sequence Preanalysis: Heuristic Error Checking ................................. 17 Molecular Sequence Formatting ............................................... 17 Analytical Strategy .......................................................... 18 Review of Current Taxonomy, the Questions, and Taxon Sampling .................. 22 Comparability of Systematic Studies ........................................... 22 Amphibia ................................................................... 23 Gymnophiona ............................................................... 24 Rhinatrematidae ........................................................... 25 Ichthyophiidae and Uraeotyphlidae .......................................... 25 Scolecomorphidae ......................................................... 25 Typhlonectidae ............................................................ 25 ``Caeciliidae'' ............................................................. 25 Caudata .................................................................... 26 Sirenidae ................................................................. 26 Hynobiidae ............................................................... 28 Cryptobranchidae .......................................................... 28 Proteidae ................................................................. 28 Rhyacotritonidae .......................................................... 30 Amphiumidae ............................................................. 30 Plethodontidae ............................................................ 30 Salamandridae ............................................................ 36 Dicamptodontidae ......................................................... 37 Ambystomatidae .......................................................... 37 Anura ...................................................................... 38 ``Primitive'' Frogs ........................................................... 41 Ascaphidae ............................................................... 41 Leiopelmatidae ............................................................ 41 Discoglossidae and Bombinatoridae .......................................... 44 ``Transitional'' Frogs ......................................................... 45 Pipoidea .................................................................. 46 Rhinophrynidae ........................................................... 47 Pipidae ................................................................... 47 Pelobatoidea .............................................................. 48 Pelobatidae and Scaphiopodidae ............................................. 49 Pelodytidae ............................................................... 49 Megophryidae ............................................................. 49 ``Advanced'' FrogsÐNeobatrachia ............................................. 50 ``Hyloidea'' ................................................................. 52 Heleophrynidae ........................................................... 52 Sooglossidae and Nasikabatrachidae ......................................... 53 Limnodynastidae, Myobatrachidae, and Rheobatrachidae ....................... 53 ``Leptodactylidae'' ......................................................... 56 ``Ceratophryinae'' ......................................................... 57 ``Cycloramphinae'' ........................................................ 59 3 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 Eleutherodactylinae ........................................................ 60 Leptodactylinae ........................................................... 61 ``Telmatobiinae'' .......................................................... 61 ``Hemiphractinae'' ......................................................... 62 Brachycephalidae .......................................................... 63 Rhinodermatidae .......................................................... 63 Dendrobatidae ............................................................. 63 Allophrynidae ............................................................. 64 Centrolenidae ............................................................. 64 Hylidae .................................................................. 64 Hylinae .................................................................. 64 Pelodryadinae ............................................................. 65 Phyllomedusinae .......................................................... 65 Bufonidae ................................................................ 67 Ranoidea ................................................................... 70 Arthroleptidae, Astylosternidae, and Hyperoliidae ............................. 72 Arthroleptidae ............................................................. 72 Astylosternidae ............................................................ 72 Hyperoliidae .............................................................. 74 Hemisotidae .............................................................. 74 Microhylidae .............................................................. 75 Scaphiophyrninae .......................................................... 77 Asterophryinae and Genyophryninae ......................................... 77 Brevicipitinae ............................................................. 78 Cophylinae ............................................................... 78 Dyscophinae .............................................................. 79 Melanobatrachinae ......................................................... 79 Microhylinae .............................................................. 80 Phrynomerinae ............................................................ 80 ``Ranidae'' ................................................................ 81 Ceratobatrachinae .......................................................... 81 Conrauinae ............................................................... 83 Dicroglossinae ...........................................................