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Home , Altiphrynoides, Amietia vertebralis, Amnirana, Amolops larutensis, Amphibian, Amphignathodontidae

THE TREE OF

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 (),2Center for and Conservation, and 3Division of , 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, , and Environmental , 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 , University of the Western Cape, Private Bag X17, Bellville 7535, South ([email protected]); 9Department of Zoology, The Natural History Museum, Cromwell Road, SW7 5BD, UK ([email protected]); 10South 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, ([email protected]); 13Wildlife Research Laboratory, Florida 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 , 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 ᭧ 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 , the Questions, and Taxon Sampling ...... 22 Comparability of Systematic Studies ...... 22 Amphibia ...... 23 ...... 24 ...... 25 and Uraeotyphlidae ...... 25 ...... 25 ...... 25 ``'' ...... 25 ...... 26 ...... 26 Hynobiidae ...... 28 Cryptobranchidae ...... 28 ...... 28 Rhyacotritonidae ...... 30 Amphiumidae ...... 30 ...... 30 ...... 36 Dicamptodontidae ...... 37 ...... 37 Anura ...... 38 ``Primitive'' ...... 41 Ascaphidae ...... 41 Leiopelmatidae ...... 41 Discoglossidae and ...... 44 ``Transitional'' Frogs ...... 45 ...... 46 ...... 47 ...... 47 ...... 48 Pelobatidae and Scaphiopodidae ...... 49 ...... 49 ...... 49 ``Advanced'' FrogsÐNeobatrachia ...... 50 ``'' ...... 52 Heleophrynidae ...... 52 and Nasikabatrachidae ...... 53 , , and Rheobatrachidae ...... 53 ``'' ...... 56 ``Ceratophryinae'' ...... 57 ``Cycloramphinae'' ...... 59

3 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Eleutherodactylinae ...... 60 ...... 61 ``Telmatobiinae'' ...... 61 ``Hemiphractinae'' ...... 62 ...... 63 ...... 63 Dendrobatidae ...... 63 Allophrynidae ...... 64 Centrolenidae ...... 64 ...... 64 ...... 64 ...... 65 Phyllomedusinae ...... 65 Bufonidae ...... 67 ...... 70 , Astylosternidae, and ...... 72 Arthroleptidae ...... 72 Astylosternidae ...... 72 Hyperoliidae ...... 74 Hemisotidae ...... 74 ...... 75 Scaphiophyrninae ...... 77 and Genyophryninae ...... 77 Brevicipitinae ...... 78 ...... 78 Dyscophinae ...... 79 Melanobatrachinae ...... 79 ...... 80 Phrynomerinae ...... 80 ``Ranidae'' ...... 81 Ceratobatrachinae ...... 81 Conrauinae ...... 83 Dicroglossinae ...... 83 Lankanectinae ...... 89 Micrixalinae ...... 90 Nyctibatrachinae ...... 91 Petropedetinae, Phrynobatrachinae, and ...... 91 Ptychadeninae ...... 93 ``Raninae'' ...... 93 Ranixalinae ...... 106 and ...... 106 Results ...... 110 Sequence Length Variation and Notes on Analysis ...... 110 Topological Results and Discussion ...... 111 Outgroup Relationships ...... 112 Amphibia () and ...... 112 Gymnophiona ...... 113 Caudata ...... 114 Hynobiidae and Cryptobranchidae ...... 114 Sirenidae and Proteidae ...... 116 Rhyacotritonidae and Amphiumidae ...... 116 Plethodontidae ...... 116 Salamandridae ...... 117 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 5

Dicamptodontidae and Ambystomatidae ...... 118 Anura ...... 118 Ascaphidae and Leiopelmatidae ...... 118 Pipidae and Rhinophrynidae ...... 119 Discoglossidae and Bombinatoridae ...... 121 Pelobatoidea ...... 121 ...... 121 Heleophrynidae ...... 121 Hyloidea, excluding Heleophrynidae ...... 121 Sooglossidae and Nasikabatrachidae ...... 122 Myobatrachidae, Limnodynastidae, and Rheobatrachidae ...... 123 ``Leptodactylidae'' ...... 123 ``Telmatobiinae'' ...... 123 ``Hemiphractinae'' ...... 127 Eleutherodactylinae and Brachycephalidae ...... 127 ``Leptodactylinae'' ...... 127 ``Ceratophryinae'' ...... 128 ``Cycloramphinae'' and Rhinodermatidae ...... 128 Centrolenidae and Allophrynidae ...... 130 Brachycephalidae ...... 130 Rhinodermatidae ...... 130 Dendrobatidae ...... 130 Hylidae ...... 130 Bufonidae ...... 131 Ranoidea ...... 131 Microhylidae and Hemisotidae ...... 131 Arthroleptidae, Astylosternidae, and Hyperoliidae ...... 134 Ranidae, Mantellidae and Rhacophoridae ...... 134 Mantellidae and Rhacophoridae ...... 141 A Taxonomy of Living ...... 141 Taxonomic Accounts ...... 147 Amphibia Gray, 1825 ...... 164 Gymnophiona MuÈller, 1832 ...... 165 Rhinatrematidae Nussbaum, 1977 ...... 165 Stegokrotaphia Cannatella and Hillis, 1993 ...... 166 Ichthyophiidae Taylor, 1968 ...... 166 Caeciliidae Ra®nesque, 1814 ...... 167 Batrachia Latreille, 1800 ...... 168 Caudata Fischer von Waldheim, 1813 ...... 169 Cryptobranchoidei Noble, 1931 ...... 170 Cryptobranchidae Fitzinger, 1825 ...... 170 Hynobiidae Cope, 1859 ...... 170 Diadectosalamandroidei new taxon ...... 171 Hydatinosalamandroidei new taxon ...... 171 Perennibranchia Latreille, 1825 ...... 172 Proteidae Gray, 1825 ...... 172 Sirenidae Gray, 1825 ...... 173 Treptobranchia new taxon ...... 173 Ambystomatidae Gray, 1850 ...... 174 Salamandridae Goldfuss, 1820 ...... 174 Plethosalamandroidei new taxon ...... 175 Rhyacotritonidae Tihen, 1958 ...... 176 Xenosalamandroidei new taxon ...... 176 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Amphiumidae Gray, 1825 ...... 176 Plethodontidae Gray, 1850 ...... 177 Anura Fischer von Waldheim, 1813 ...... 178 Leiopelmatidae Mivart, 1869 ...... 179 Lalagobatrachia new taxon ...... 180 Xenoanura Savage, 1973 ...... 181 Pipidae Gray, 1825 ...... 182 Rhinophrynidae GuÈnther, 1859 ``1858'' ...... 183 Sokolanura new taxon ...... 183 Costata Lataste, 1879 ...... 184 Fitzinger, 1843 ...... 184 Bombinatoridae Gray, 1825 ...... 185 Acosmanura Savage, 1973 ...... 185 Anomocoela Nicholls, 1916 ...... 186 Pelodytoidea Bonaparte, 1850 ...... 187 Pelodytidae Bonaparte, 1850 ...... 187 Scaphiopodidae Cope, 1865 ...... 187 Pelobatoidea Bonaparte, 1850 ...... 188 Pelobatidae Bonaparte, 1850 ...... 188 Megophryidae Bonaparte, 1850 ...... 188 Neobatrachia Reig, 1958 ...... 189 Heleophrynidae Noble, 1931 ...... 190 Pthanobatrachia new taxon ...... 190 Hyloides new taxon ...... 191 Sooglossidae Noble, 1931 ...... 191 Notogaeanura new taxon ...... 192 Australobatrachia new taxon ...... 193 Batrachophrynidae Cope, 1875 ...... 193 Myobatrachoidea Schlegel, 1850 ...... 194 Limnodynastidae Lynch, 1971 ...... 194 Myobatrachidae Schlegel, 1850 ...... 195 Nobleobatrachia new taxon ...... 196 Peters, 1862 ...... 196 Meridianura new taxon ...... 196 Brachycephalidae GuÈnther, 1858 ...... 197 Cladophrynia new taxon ...... 201 Cryptobatrachidae new ...... 201 Tinctanura new taxon ...... 201 Boulenger, 1882 ...... 202 Athesphatanura new taxon ...... 202 Hylidae Ra®nesque, 1815 ...... 202 Hylinae Ra®nesque, 1815 ...... 203 Pelodryadinae GuÈnther, 1858 ...... 204 Phyllomedusinae GuÈnther, 1858 ...... 205 Leptodactyliformes new taxon ...... 205 Diphyabatrachia new taxon ...... 205 Centrolenidae Taylor, 1951 ...... 206 Leptodactylidae Werner, 1896 (1838) ...... 207 Chthonobatrachia new taxon ...... 208 Tschudi, 1838 ...... 208 Hesticobatrachia new taxon ...... 209 Bonaparte, 1850 ...... 210 Agastorophrynia new taxon ...... 210 Cope, 1865 ...... 211 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 7

Thoropidae new family ...... 211 Dendrobatidae Cope, 1865 ...... 211 Bufonidae Gray, 1825 ...... 213 Ranoides new taxon ...... 223 Allodapanura new taxon ...... 224 Microhylidae GuÈnther, 1858 (1843) ...... 224 Asterophryinae GuÈnther, 1858 ...... 227 Cophylinae Cope, 1889 ...... 227 Dyscophinae Boulenger, 1882 ...... 228 Gastrophryninae Fitzinger, 1843 ...... 228 Melanobatrachinae Noble, 1931 ...... 229 Microhylinae GuÈnther, 1858 (1843) ...... 229 Laurent, 1946 ...... 230 Afrobatrachia new taxon ...... 231 Xenosyneunitanura new taxon ...... 231 Bonaparte, 1850 ...... 231 Hemisotidae Cope, 1867 ...... 232 Laurentobatrachia new taxon ...... 232 Hyperoliidae Laurent, 1943 ...... 232 Arthroleptidae Mivart, 1869 ...... 233 Natatanura new taxon ...... 234 Dubois, 1987 ``1986'' ...... 235 Victoranura new taxon ...... 235 Boulenger, 1884 ...... 236 Telmatobatrachia new taxon ...... 236 Micrixalidae Dubois, Ohler, and Biju, 2001 ...... 237 Ametrobatrachia new taxon ...... 237 Africanura new taxon ...... 237 Phrynobatrachidae Laurent, 1940 ...... 237 Pyxicephaloidea Bonaparte, 1850 ...... 238 Noble, 1931 ...... 238 Bonaparte, 1850 ...... 240 Saukrobatrachia new taxon ...... 241 Anderson, 1871 ...... 241 Dicroglossinae Anderson, 1871 ...... 241 Occidozyginae Fei, Ye, and Huang, 1991 ``1990'' ...... 243 Aglaioanura new taxon ...... 243 Rhacophoroidea Hoffman, 1932 (1858) ...... 243 Mantellidae Laurent, 1946 ...... 244 Rhacophoridae Hoffman, 1932 (1858) ...... 245 Ranoidea Ra®nesque, 1814 ...... 247 Blommers-SchloÈsser, 1993 ...... 247 Ranidae Ra®nesque, 1814 ...... 248 Acknowledgments ...... 255 References ...... 257 Appendix 1. Voucher and DNA Locus Information ...... 292 Appendix 2. Accession Numbers for Genbank Sequences Used in This Study ...... 322 Appendix 3. Base-Pair Length of 28S Fragment ...... 324 Appendix 4. Branch Lengths, Bremer Support, and Jackknife Values by Branch ...... 326 Appendix 5. DNA Sequence Transformations for Selected Branches/Taxa ...... 330 Appendix 6. Nomenclatural Notes ...... 355 Appendix 7. New and Revived Combinations and Clari®cations Regarding Taxonomic Content ...... 359 ABSTRACT

The evidentiary basis of the currently accepted classi®cation of living amphibians is dis- cussed and shown not to warrant the degree of authority conferred on it by use and tradition. A new taxonomy of living amphibians is proposed to correct the de®ciencies of the old one. This new taxonomy is based on the largest phylogenetic analysis of living Amphibia so far accomplished. We combined the comparative anatomical character evidence of Haas (2003) with DNA sequences from the mitochondrial transcription unit H1 (12S and 16S ribosomal RNA and tRNAValine genes, ഠ 2,400 bp of mitochondrial sequences) and the nuclear genes histone H3, rhodopsin, tyrosinase, and seven in absentia, and the large ribosomal subunit 28S (ഠ 2,300 bp of nuclear sequences; ca. 1.8 million base pairs; xÅ ϭ 3.7 kb/terminal). The dataset includes 532 terminals sampled from 522 representative of the global diversity of amphibians as well as seven of the closest living relatives of amphibians for outgroup com- parisons. The primary purpose of our taxon sampling strategy was to provide strong tests of the monophyly of all ``family-group'' taxa. All currently recognized nominal families and subfam- ilies were sampled, with the exception of Protohynobiinae (Hynobiidae). Many of the currently recognized genera were also sampled. Although we discuss the monophyly of genera, and provide remedies for nonmonophyly where possible, we also make recommendations for future research. A parsimony analysis was performed under Direct Optimization, which simultaneously op- timizes nucleotide (alignment) and tree costs, using the same set of assumptions throughout the analysis. Multiple search algorithms were run in the program POY over a period of seven months of computing time on the AMNH Parallel Computing Cluster. Results demonstrate that the following major taxonomic groups, as currently recognized, are nonmonophyletic: Ichthyophiidae (paraphyletic with respect to Uraeotyphlidae), Caecili- idae (paraphyletic with respect to Typhlonectidae and Scolecomorphidae), (paraphyletic with respect to Sirenidae), Leiopelmatanura (paraphyletic with respect to Asca- phidae), Discoglossanura (paraphyletic with respect to Bombinatoridae), (par- aphyletic with respect to Neobatrachia), Pipanura (paraphyletic with respect to Bombinatoridae and Discoglossidae/Alytidae), Hyloidea (in the sense of containing Heleophrynidae; paraphy- letic with respect to Ranoidea), Leptodactylidae (polyphyletic, with Batrachophrynidae form- ing the sister taxon of Myobatrachidae ϩ Limnodynastidae, and broadly paraphyletic with respect to Hemiphractinae, Rhinodermatidae, Hylidae, Allophrynidae, Centrolenidae, Brachy- cephalidae, Dendrobatidae, and Bufonidae), Microhylidae (polyphyletic, with Brevicipitinae being the sister taxon of Hemisotidae), Microhylinae (poly/paraphyletic with respect to the remaining non-brevicipitine microhylids), Hyperoliidae (para/polyphyletic, with Leptopelinae forming the sister taxon of Arthroleptidae ϩ Astylosternidae), Astylosternidae (paraphyletic with respect to Arthroleptinae), Ranidae (paraphyletic with respect to Rhacophoridae and Man- tellidae). In addition, many subsidiary taxa are demonstrated to be nonmonophyletic, such as (1) with respect to Brachycephalus; (2) (sensu Dubois, 1992), which is polyphyletic, with various elements falling far from each other on the tree; and (3) , with respect to several nominal bufonid genera. A new taxonomy of living amphibians is proposed, and the evidence for this is presented to promote further investigation and data acquisition bearing on the evolutionary history of amphibians. The taxonomy provided is consistent with the International Code of Zoological Nomenclature (ICZN, 1999). Salient features of the new taxonomy are (1) the three major groups of living amphibians, /Gymnophiona, /Caudata, and frogs/Anura, form a monophyletic group, to which we restrict the name Amphibia; (2) Gymnophiona forms the sister taxon of Batrachia (salamanders ϩ frogs) and is composed of two groups, Rhinatrematidae and Stegokrotaphia; (3) Stegokrotaphia is composed of two families, Ichthyophiidae (including Uraeotyphlidae) and Caeciliidae (including Scolecomorphidae and Typhlonectidae, which are regarded as sub- families); (4) Batrachia is a highly corroborated monophyletic group, composed of two taxa, Caudata (salamanders) and Anura (frogs); (5) Caudata is composed of two taxa, Cryptobran- choidei (Cryptobranchidae and Hynobiidae) and Diadectosalamandroidei new taxon (all other salamanders); (6) Diadectosalamandroidei is composed of two taxa, Hydatinosalamandroidei

8 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 9

new taxon (composed of Perennibranchia and Treptobranchia new taxon) and Plethosala- mandroidei new taxon; (7) Perennibranchia is composed of Proteidae and Sirenidae; (8) Trep- tobranchia new taxon is composed of two taxa, Ambystomatidae (including Dicamptodonti- dae) and Salamandridae; (9) Plethosalamandroidei new taxon is composed of Rhyacotritonidae and Xenosalamandroidei new taxon; (10) Xenosalamandroidei is composed of Plethodontidae and Amphiumidae; (11) Anura is monophyletic and composed of two , Leiopelmatidae (including Ascaphidae) and Lalagobatrachia new taxon (all other frogs); (12) Lalagobatrachia is composed of two clades, Xenoanura (Pipidae and Rhinophrynidae) and Sokolanura new taxon (all other lalagobatrachians); (13) Bombinatoridae and Alytidae (former Discoglossidae) are each others' closest relatives and in a called Costata, which, excluding - tidae and Xenoanura, forms the sister taxon of all other frogs, Acosmanura; (14) Acosmanura is composed of two clades, Anomocoela (ϭ Pelobatoidea of other authors) and Neobatrachia; (15) Anomocoela contains Pelobatoidea (Pelobatidae and Megophryidae) and Pelodytoidea (Pelodytidae and Scaphiopodidae), and forms the sister taxon of Neobatrachia, together form- ing Acosmanura; (16) Neobatrachia is composed of two clades, Heleophrynidae, and all other neobatrachians, Phthanobatrachia new taxon; (17) Phthanobatrachia is composed of two major units, Hyloides and Ranoides; (18) Hyloides comprises Sooglossidae (including Nasikabatrach- idae) and Notogaeanura new taxon (the remaining hyloids); (19) Notogaeanura contains two taxa, Australobatrachia new taxon and Nobleobatrachia new taxon; (20) Australobatrachia is a clade composed of Batrachophrynidae and its sister taxon, Myobatrachoidea (Myobatrach- idae and Limnodynastidae), which forms the sister taxon of all other hyloids, excluding soog- lossids; (21) Nobleobatrachia new taxon, is dominated at its base by frogs of a treefrog morphotype, several with intercalary phalangeal cartilagesÐHemiphractus (Hemiphractidae) forms the sister taxon of the remaining members of this group, here termed Meridianura new taxon; (22) Meridianura comprises Brachycephalidae (former Eleutherodactylinae ϩ Brachy- cephalus) and Cladophrynia new taxon; (23) Cladophrynia is composed of two groups, Cryp- tobatrachidae (composed of and , previously a fragment of the poly- phyletic Hemiphractinae) and Tinctanura new taxon; (24) Tinctanura is composed of Am- phignathodontidae ( and , another fragment of the polyphyletic Hem- iphractinae) and Athesphatanura new taxon; (25) Athesphatanura is composed of Hylidae (Hylinae, Pelodryadinae, and Phyllomedusinae, and excluding former Hemiphractinae, whose inclusion would have rendered this taxon polyphyletic) and Leptodactyliformes new taxon; (26) Leptodactyliformes is composed of Diphyabatrachia new taxon (composed of Centrolen- idae [including Allophryne] and Leptodactylidae, sensu stricto, including and relatives) and Chthonobatrachia new taxon; (27) Chthonobatrachia is composed of a refor- mulated Ceratophryidae (which excludes such genera as and and includes other taxa, such as ) and Hesticobatrachia new taxon; (28) Hesti- cobatrachia is composed of a reformulated Cycloramphidae (which includes ) and Agastorophrynia new taxon; (29) Agastorophrynia is composed of Bufonidae (which is par- tially revised) and Dendrobatoidea (Dendrobatidae and Thoropidae); (30) Ranoides new taxon forms the sister taxon of Hyloides and is composed of two major monophyletic components, Allodapanura new taxon (microhylids, hyperoliids, and allies) and Natatanura new taxon (ranids and allies); (31) Allodapanura is composed of Microhylidae (which is partially revised) and Afrobatrachia new taxon; (32) Afrobatrachia is composed of Xenosyneunitanura new taxon (the ``strange-bedfellows'' Brevicipitidae [formerly in Microhylidae] and Hemisotidae) and a more normal-looking group of frogs, Laurentobatrachia new taxon (Hyperoliidae and Arthroleptidae, which includes Leptopelinae and former Astylosternidae); (33) Natatanura new taxon is composed of two taxa, the African Ptychadenidae and the worldwide Victoranura new taxon; (34) Victoranura is composed of Ceratobatrachidae and Telmatobatrachia new taxon; (35) Telmatobatrachia is composed of Micrixalidae and a worldwide group of ranoids, Ametrobatrachia new taxon; (36) Ametrobatrachia is composed of Africanura new taxon and Saukrobatrachia new taxon; (37) Africanura is composed of two taxa: Phrynobatrachidae (, including Dimorphognathus and Phrynodon as synonyms) and Pyxice- phaloidea; (38) Pyxicephaloidea is composed of Petropedetidae (, , Arthro- leptides, and ), and Pyxicephalidae (including a number of African genera, e.g. [including Afrana], , , , and ); and (39) Saukrobatrachia new taxon is the sister taxon of Africanura and is composed of Dicro- 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

glossidae and Aglaioanura new taxon, which is, in turn, composed of Rhacophoroidea (Man- tellidae and Rhacophoridae) and Ranoidea (Nyctibatrachidae and Ranidae, sensu stricto). Many generic revisions are made either to render a monophyletic taxonomy or to render a taxonomy that illuminates the problems in our understanding of phylogeny, so that future work will be made easier. These revisions are: (1) placement of and Lineatriton (Caudata: Plethodontidae: ) into the synonymy of , to render a monophy- letic Pseudoeurycea; (2) placement of Haideotriton (Caudata: Plethodontidae: Spelerpinae) into the synonymy of Eurycea, to render a monophyletic Eurycea; (3) placement of Nesomantis (Anura: Sooglossidae) into the synonymy of , to assure a monophyletic Sooglossus; (4) placement of Cyclorana and (Anura: Hylidae: Pelodryadinae) into , but retaining Cyclorana as a subgenus, to provide a monophyletic Litoria; (5) partition of ``Lim- nodynastes'' (Anura: Limnodynastidae) into and to render mono- phyletic genera; (6) placement of , , and Vanzolinius (Anura: Leptodac- tylidae) into Leptodactylus, to render a monophyletic Leptodactylus; (7) partition of ``Eleuth- erodactylus'' (Anura: Brachycephalidae) into ,``Eleutherodactylus'', ``Euhyas'', ``Pelorius'', and Syrrhophus to outline the taxonomic issues relevant to the of this nominal taxon to other nominal genera; (8) partition of ``Bufo'' (Anura: Bufonidae) into a number of new or revived genera (i.e., Amietophrynus new , , Chaunus, Cran- opsis, new genus, Epidalea, new genus, , Pelto- phryne, , new genus; Pseudepidalea new genus, , Rhi- nella, new genus); (9) placement of the monotypic Spinophrynoides (Anura: Bufonidae) into the synonymy of (formerly monotypic) to make for a more informative taxonomy; (10) placement of the Bufo taitanus group and Stephopaedes (as a subgenus) into the synonymy of Mertensophryne (Anura: Bufonidae); (11) placement of Xe- nobatrachus (Anura: Microhylidae: Asterophryinae) into the synonymy of to render a monophyletic Xenorhina; (12) transfer of a number of species from to (Microhylidae: Cophylinae) to render a monophyletic Plethodontohyla; (13) placement of Schoutedenella (Anura: Arthroleptidae) into the synonymy of ; (14) transfer of Dimorphognathus and Phrynodon (Anura: Phrynobatrachidae) into the synonymy of Phrynobatrachus to render a monophyletic Phrynobatrachus; (15) placement of Afrana into the synonymy of Amietia (Anura: Pyxicephalidae) to render a monophyletic taxon; (16) place- ment of Chaparana and Paa into the synonymy of (Anura: Dicroglossidae) to render a monophyletic genus; (17) recognition as genera of Ombrana and Annandia (Anura: Dicrog- lossidae: Dicroglossinae) pending placement of them phylogenetically; (18) return of Phry- noglossus into the synonymy of to resolve the paraphyly of Phrynoglossus (Anura: Dicroglossidae: Occidozyginae); (19) recognition of new genus for palpe- bralis to resolve the polyphyly of ``''; (20) synonymy of ``Chirixalus'' with Chi- romantis to resolve the paraphyly of ``Chirixalus''; (21) recognition of the genus , composed of the former subgenera of Rana, Babina and Nidirana (Anura: Ranidae); (22) recognition of the genera , , Nasirana, , Pterorana, Pul- chrana, and Sanguirana, formerly considered subgenera of Rana (Anura: Ranidae), with no special relationship to Rana (sensu stricto); (23) consideration of (Anura: Ranidae), formerly a subgenus of Rana, as a genus, with Rugosa as a synonym; (24) recognition of (Anura: Ranidae) as a genus, with and most species of former Chal- corana included in this taxon as synonyms; (25) recognition of (Anura: Ranidae) as a genus and its content rede®ned; (26) redelimitation of to include as synonyms Eburana and (both former subgenera of Rana); (27) recognition of (An- ura: Ranidae) for all species of North American ``Rana'' not placed in Rana sensu stricto (Aquarana, Pantherana, Sierrana, Trypheropsis, and Zweifelia considered synonyms of Lith- obates); (28) redelimitation of the genus Rana as monophyletic by inclusion as synonyms Amerana, Aurorana, Pseudoamolops, and Pseudorana, and exclusion of all other former sub- genera; (29) redelimitation of the genus (Anura: Ranidae), formerly a subgenus of Rana, with and Tylerana included as synonyms. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 11

INTRODUCTION gun to change in the 2000s with the infusion of signi®cant amounts of molecular evidence Amphibians (caecilians, frogs, and sala- into the discussion of large-scale amphibian manders) are a conspicuous component of diversi®cation. But, although recent molec- the world's vertebrate fauna. They currently ular studies have been very illuminating include 5948 recognized species with repre- (e.g., Biju and Bossuyt, 2003; Darst and sentatives found in virtually all terrestrial and Cannatella, 2004; Faivovich et al., 2005; freshwater , in all but the coldest and Roelants and Bossuyt, 2005; San Mauro et driest regions or the most remote oceanic is- al., 2005), so far they have not provided the lands. The number of recognized species of general roadmap for future research that a amphibians has grown enormously in recent larger and more detailed study could provide. , about a 48.2% increase since 1985 Among the three major taxonomic com- (Frost, 1985, 2004, unpubl. data). This ponents of amphibian diversity, caecilians growth re¯ects the increasing ease of col- appear to have been the focus of the most lecting in remote locations and a signi®cant signi®cant study of large-scale evolutionary growth of active scienti®c communities in a history (Gower et al., 2002; Gower and Wil- few megadiverse countries. Unfortunately, kinson, 2002; M. Wilkinson et al., 2002; M. the rapid increase in knowledge of amphib- Wilkinson et al., 2003; San Mauro et al., ian species diversity is coincident with a 2004; M.H. Wake et al., 2005), although this massive and global decline in amphibian may be an artifact of the relatively small size populations (Alford and Richards, 1999; of the group (173 species currently recog- Houlahan et al., 2000; Young et al., 2001; nized) and the few, mostly coordinated, S.N. Stuart et al., 2004) due to a diversity of workers. Salamanders are the best-known factors, including loss and fragmen- group at the species level, but tation (Green, 2005; Halliday, 2005) but also phylogenetic work has largely focused on the possibly due to global environmental chang- generic and infrageneric levels of investiga- es (Donnelly and Crump, 1998; Blaustein tion (e.g., Zhao, 1994; Titus and Larson, and Kiesecker, 2002; Heyer, 2003; Licht, 1996; Highton, 1997, 1998, 1999; GarcõÂa- 2003) and such proximate causes as emerg- ParõÂs and Wake, 2000; Highton and Peabody, ing infectious diseases (Collins and Storfer, 2000; Jockusch et al., 2001; Parra-Olea and 2003). Wake, 2001; Jockusch and Wake, 2002; Par- Understanding of amphibian evolutionary ra-Olea et al., 2002; Steinfartz et al., 2002; history has not kept pace with knowledge of Parra-Olea et al., 2004; Sites et al., 2004), amphibian species diversity. For all but a few although there have been several important groups, there is only a rudimentary evolu- efforts at an overall synthesis of morpholog- tionary framework upon which to cast the ical and molecular evidence (Larson and theories of cause, predict which lineages are Dimmick, 1993; Larson et al., 2003; Wiens most likely to go extinct, or even compre- et al., 2005). hend the amount of genetic diversity being Research on has also lost (Lips et al., 2005). Indeed, it is arguable focused primarily on generic and infragener- whether our general understanding of frog ic studies (e.g., Graybeal, 1997; Cannatella phylogenetics has progressed substantially et al., 1998; Mendelson et al., 2000; Sheil et beyond the seminal works of the late 1960s al., 2001; Channing et al., 2002a; Dawood et to early 1980s (Inger, 1967; Kluge and Far- al., 2002; Faivovich, 2002; Glaw and ris, 1969; J.D. Lynch, 1971, 1973; Farris et Vences, 2002; Pramuk, 2002; Cunningham al., 1982a). The major advances in frog tax- and Cherry, 2004; Drewes and Wilkinson, onomy in the 1980s and 1990s were domi- 2004; B.J. Evans et al., 2004; Pauly et al., nated by nomenclatural and largely litera- 2004; Crawford and Smith, 2005; Matsui et ture-based phenotypic sorting (e.g., Dubois, al., 2005), and broader discussions of frog 1980, 1981, 1984b; Laurent, 1986; Dubois, phylogenetics have been predominantly nar- 1987 ``1986'', 1992) that provided other rative rather than quantitative (e.g., Canna- workers with digestible ``chunks'' to discuss tella and Hillis, 1993; Ford and Cannatella, and evaluate phylogenetically. This has be- 1993; Cannatella and Hillis, 2004). Illumi- 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 nating large-scale studies have appeared re- taxonomy consistent with phylogeny that cently (Biju and Bossuyt, 2003; Haas, 2003; will serve as a general road map for further Darst and Cannatella, 2004; Roelants and research. That such a diverse group of biol- Bossuyt, 2005; Van der Meijden et al., 2005; ogists (see list of authors) would be willing Faivovich et al., 2005). Nevertheless, a study to set aside their legitimate philosophical dif- of a broad sampling of amphibians, based on ferences to produce this work demonstrates a large number of terminals, has not been the seriousness of the need. We hope that by attempted to date. providing considerable new data and new hy- A serious impediment in amphibian biol- potheses of relationship that we will engen- ogy, and systematics generally, with respect der efforts to test our phylogenetic hypothe- to advancing historically consistent taxono- ses and generate new ones. Regardless, the mies, is the social conservatism resulting in days are over of construing broad conclu- the willingness of many taxonomists to em- sions from small analyses of small numbers brace, if only tacitly, paraphyletic groupings, of taxa using small amounts of molecular or even when the evidence exists to correct morphological data. We also think that the them. The reason for this is obvious. Rec- time is past for authoritarian classi®cations, ognizing paraphyletic groups is a way of de- rich in special pleading and weak on evi- scribing trees in a linear way for the purpose dence (e.g., Dubois, 1992; Delorme et al., of telling great stories and providing favored 2005; Dubois, 2005). In short, we hope that characters a starring role. Because we think this publication will help change the nature that storytelling re¯ects a very deep element of the conversation among scientists regard- of human communication, many systema- ing amphibian systematics, moving it away tists, as normal storytelling humans, are un- from the sociologically conservative to the willing to discard paraphyly. Unfortunately, scienti®cally conservative. As noted by Can- the great stories of science, those popular natella and Hillis (2004: 444), the need for with the general public and some funding ``scaling up'' the rate of data collection is agencies, almost never evidence careful anal- certainly evident (e.g., compare the eviden- ysis of data and precise reasoning or lan- tiary content of Cannatella and Hillis, 1993, guage. And, for much of its history, system- with Cannatella and Hillis, 2004). atics focused on great narrative stories about Nevertheless, even if we are successful in ``adaptive radiations'' and ``primitive'', providing a roadmap for future work, this ``transitional'', and ``advanced'' groups rath- will not assure the health of amphibian sys- er than the details of phylogeny. These sto- tematics. Clearly, the task of understanding ries were almost always about favored char- the evolution and ecological, morphological, acters (e.g., pectoral girdle , repro- and taxonomic diversity of amphibians is ductive modes) within a sequence of para- massive, yet funding remains insuf®cient to phyletic groupings to the detriment of a full maintain a healthy amphibian systematics and detailed understanding of evolutionary commmunity. Further, the institutional, inter- history. institutional, national and international infra- When one deconstructs the existing tax- structure needed to promote the systematics onomy of frogs, for example, one is struck research program needs to be greatly en- by the number of groups delimited by very hanced with respect to state-of-the-art collec- small suites of characters and the special tion facilities, digital libraries of all relevant pleading for particular characters that under- systematic literature, interoperable collection lies so much of the taxonomic reasoning. databases, and associated GIS and mapping- Factoring in the systematic philosophy at the related capacity, supercomputers and the im- time many of these groups were named, both proved analytical software to drive them, re- the origin of the problems and the illogic of motely accessible visualization instrumenta- perpetuating the status quo become apparent. tion and specimen images, and enhanced Our goal in this study is to provide rem- data-aquisition technology, including mas- edies for the problems noted above, by way sive through-put DNA sequencing, in addi- of performing a large phylogenetic analysis tion to already-identi®ed personnel, training, across all living amphibians and providing a and ®nancial needs related to exploring life 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 13

on this planet and maintaining large research GENERAL ANALYTICAL APPROACH: collections (Q.D. Wheeler et al., 2004; Page THEORETICAL CONSIDERATIONS et al., 2005). There has been the salutary de- velopment of additional support in the train- CHOICE OF PHYLOGENETIC METHOD: All ing of systematists (e.g., Rodman and Cody, phylogenetic methods minimize the number 2003) and important successes in increasing of character transformations required to ex- systematics capacity in a few megadiverse plain the observed variation. Unweighted countries (e.g., Brazil; see de Carvalho et al., (equally weighted) parsimony analysis min- 2005), but it is also clear that increased re- imizes hypothesized transformations global- search support is needed to assure another ly, whereas the assumptions (expressed as generation of evolutionary capable differential probabilities or costs) about the of the detailed anatomical work to document evolutionary process or perceived impor- how organisms have changed and diversi®ed tance of different classes of transformations through time. But, especially in this time of employed in statistical (maximum-likeli- increasing optimization of university hiring hood, Bayesian analysis) and weighted par- and retention policies on the ability of faculty simony methods minimize certain classes of to garner extramural funding, additional transformations at the expense of others. Op- funding is needed to make sure that jobs ex- erational considerations aside (e.g., tree- ist for the systematists that are being trained. space searching capabilities), disagreements ABOUT THE COLLABORATION: This collabora- between the results of unweighted parsimony tion was undertaken with the knowledge that analysis and the other methods are due to the everyone involved would have to compro- increased patristic distance required to ac- mise on deeply held convictions regarding commodate the additional assumptions. For the nature of evidence, methods of analysis, this study, we chose to analyze the data un- and what constitutes a reasonable assump- der the minimal assumptions of unweighted tion, as well as the nature of taxonomic no- parsimony. Given the size and complexity of menclature. Nevertheless, all data are provid- our dataset, an important advantage of par- ed either through GenBank or from http:// simony algorithms (whether weighted or un- research.amnh.org/herpetology/down- weighted) is that thorough analysis could be loads.html, and we expect several of the achieved in reasonable times given currently coauthors to deal in greater detail with the available hardware and software. problems and taxonomic hypotheses noted in NUCLEOTIDE HOMOLOGY AND THE TREAT- this paper, on the basis of even greater MENT OF INSERTIONS/DELETIONS (INDELS): The amounts of data with various taxonomic method of inferring nucleotide homology units within Amphibia and from their own (i.e., alignment) and insertions/deletions (in- points of view. We are unanimous in thinking dels) and the treatment of indels in evaluat- that the capacity for systematic work needs ing phylogenetic hypotheses are critically to be expanded, and given existing university important in empirical studies. A given da- hiring and retention practices, this expansion taset aligned according to different criteria or can only take place through enhanced fund- under different indel treatments may strongly ing. support contradictory solutions (e.g., W.C. Wheeler, 1995; Morrison and Ellis, 1997). Many workers infer indels as part of their MATERIALS AND METHODS procedure to discover nucleotide homology CONVENTIONS AND ABBREVIATIONS but then either treat the inferred indels as nu- cleotides of unknown identity by converting Commands used in computer programs are gaps into missing data or eliminate gap-con- italicized. Tissues are referenced in appendix taining column vectors altogether, because 1 with the permanent collection number for they are believed to be unreliable or because the voucher specimen or, if that is unavail- the method of phylogenetic analysis does not able, the tissue-collection number or ®eld- allow them (Swofford et al., 1996). Others voucher number. (See appendix 1 for acro- argue that indels provide evidence of phy- nyms.) logeny but believe, we think incorrectly, that 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 sequence alignment and tree evaluation are pology combination, and search strategies logically independent and must be performed must take into consideration the extent of the separately (e.g., Simmons and Ochoterena, heuristic shortcuts applied at both levels. The 2000; Simmons, 2004). details of our analyses are discussed below We treat indels as evidentially equivalent under Heuristic Homology Assessment and to any other kind of inferred transformation Heuristic Tree Searching, with the general and as a deductively inferred component of approach being to increase the rigor at both the explanation of DNA sequence diversity levels as the overall search progresses. In any observed among the sampled terminals. Fur- heuristic analysis, a balance is sought where- thermore, because nucleotides lack the struc- by the algorithmic shortcuts speed up anal- tural and/or developmental complexity nec- ysis enough to permit a suf®ciently large and essary to test their homology separately, hy- diverse sample of trees and alignments to potheses of nucleotide homology can be discover the global optimum during ®nal re- evaluated only in reference to a topology ®nement, but not so severe that the sampling (Grant and Kluge, 2004; see also Frost et al., is so sparse or misdirected that the global 2001). In recognition of these considerations, optimum is not within reach during ®nal re- we assessed nucleotide homology dynami- ®nement. Ideally, indicators of search ade- cally by optimizing observed sequences di- quacy (e.g., multiple independent minimum- rectly onto competing topologies (Sankoff, length hits, stable consensus; see Goloboff, 1975; Sankoff et al., 1976), thereby heuris- 1999; Goloboff and Farris, 2001; Goloboff tically evaluating competing hypotheses by et al., 2003) should be employed to judge the simultaneous searching of tree space. This is adequacy of analysis, as is now reasonable achieved using Direct Optimization (W.C. in parsimony analysis of large prealigned da- Wheeler, 1994, 1996, 1998, 1999; Phillips et tasets (e.g., as performed by the software al., 2000; W.C. Wheeler, 2000, 2001, 2002, package TNT; Goloboff et al., 2003). How- 2003a, 2003b, 2003c) as implemented in the ever, current hardware and software limita- computer program POY (W.C. Wheeler et tions make those indicators unreachable in al., 1996±2003). reasonable amounts of time for our dataset Determination of nucleotide homology is analyzed under Direct Optimization. The ad- treated as an optimization problem in which equacy of our analysis may only be judged the optimal scheme of nucleotide homologies intuitively in light of the computational effort for a given topology is that which requires and strategic use of multiple algorithms de- the fewest transformations overallÐthat is, signed for large datasets. that which minimizes patristic distance, thus providing the most parsimonious explanation TAXON SAMPLING of the observed diversity. Determining the optimal alignment for a given topology is The 532 terminals (re¯ecting 7 outgroup NP-complete1 (Wang and Jiang, 1994). For species, 522 ingroup species [with three re- even a minuscule number of sequences, the dundancies]) included in our analysis are number of possible alignments is staggering- given in appendix 1. Because this study is ly large (Slowinski, 1998), making exact so- predominantly molecular, outgroup sampling lutions impossible for any contemporary da- was restricted to the closest living relatives taset, and heuristic algorithms are required to of living amphibians and did not include fos- render this problem tractable. sil taxa. These included two , two Phylogenetic analysis under Direct Opti- turtles, one crocodylian, one squamate, and mization, therefore, addresses two nested a as the root. Our study was not NP-complete problems. POY searches si- designed to identify the sister taxon of tet- multaneously for the optimal homology/to- rapods, and our use of a coelacanth instead of a ®sh was due to expediency and not a decided preference for any particular hy- 1 The notion of NP-completeness extends from formal complexity theory. But, we can regard NP-complete pothesis of relationship. problems as those problems for which there is no prac- The remaining 525 terminals were sam- tical way to determine or verify an exact solution. pled from the three orders of living amphib- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 15 ians. Our general criteria were (1) availabil- rior ceratohyla emargination) were excluded ity of tissues and/or sequences on GenBank, from our analysis because POY is unable to and (2) representation of taxonomic diversi- address noninteger transformations. We did ty. Although taxonomic rank per se is mean- include Haas' transformation 102 (presence/ ingless, our taxon sampling was guided to a absence of larval ) which he excluded large degree by generic diversity. Experience from analysis because of dif®culty in scoring suggested that this ``genus-level'' sampling absences; its inclusion did not alter his ®nal would thoroughly sample the diversity of liv- topology and provided us the opportunity to ing amphibians. The median number of spe- incorporate known occurence of larval ribs cies per genus for living taxa is only three, in our ®nal hypothesis. something that we think has to do with hu- Of the 81 frog and 4 salamander species man perception of similarity and difference, in Haas' (2003) study, our study overlaps in not evolutionary processes. Some genera 41 anurans and 2 caudates. We did not com- (e.g., Eleutherodactylus, ca. 605 species) are bine into one virtual taxon morphology from so large and/or diverse that directed subsam- one species and DNA sequences from anoth- pling of species groups was required to eval- er, even if putatively closely related. Al- uate likely paraphyly (e.g., with respect to though that would have allowed us to incor- ). porate more (and potentially all) morpholog- Summarizing, our sample constituted ical data, and in some cases it probably about 8.8% of all species of Recent amphib- would not have affected our results detri- ians currently recognized, with approximate- mentally, because of our general skepticism ly the same proportion of species diversity regarding the current understanding of am- sampled from each . Of the ca. 467 Re- 2 phibian relationships we were unwilling to cent amphibian genera , 326 (69.8%) are rep- assume the monophyly of any group prior to resented in our sample. We targeted 17 spe- the analysis. cies of caecilians, representing 16 genera of DNA SEQUENCES: In light of the differing all 6 family groups. Among salamanders we levels of diversity included in this study, we sampled 51 species from 42 genera of all 10 sought to sample loci of differing degrees of families. The bulk of our ingroup sample fo- variability (i.e., rates). From the mitochon- cused on frogs, with 437 terminals targeted. drial genome, we targeted the mitochondrial The remaining 457 terminals represent 454 H-strand transcription unit 1 (H1), which in- anuran species from ca. 269 genera and 32 Valine anuran families. A more extensive discussion cludes the 12S ribosomal, tRNA , and of the terminals and the rationale behind their 16S ribosomal sequences, yielding approxi- choice is presented under ``Review of Cur- mately 2,400 base pairs (bp) generated in 5± rent Taxonomy''. 7 overlapping fragments. We also targeted the nuclear coding genes histone H3 (328 bp), rhodopsin (316 bp), tyrosinase (532 CHARACTER SAMPLING bp), seven in absentia (397 bp), and the nu- MORPHOLOGY: The 152 transformation se- clear 28S ribosomal gene (ca. 700 bp), giving ries of morphology were incorporated di- a total of approximately 2,300 bp of nuclear rectly from Haas (2003). Of his original 156 DNA. Primers used in PCR ampli®cation and transformations, the gap-weighted morpho- cycle-sequencing reactions (and respective metric transformations 12 (relative larval citations) are given in table 1. When possi- thickness), 83 (cornua trabeculae pro- ble, terminals for which we were unable to portions), 116 (ratio of anterior ceratohyal generate all fragments were augmented with processes), and 117 (relative depth of ante- sequences from GenBank (see appendices 1, 2) under the assumption that the tissues were 2 The estimate of the number of amphibian genera, for actually conspeci®c. The amount of se- purposes of these comparisons, rests on our perception quence/terminal varied (®g. 1) with a range of common usage. Because we arbitrarily treated many from 490 bp ( limborgi) to 4,790 nominal subgenera (e.g., Clinotarsus, Hydrophylax, Lithobates) as genera, our working number of genera, (Eleutherodactylus pluviacanorus), and the for purposes of this manuscript, is considerably larger. mean being 3,554 bp (see appendix 1). 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

TABLE 1 Primers Used for Various Loci in This Study and Their Published Sources

Gene Primer name Direction Primer sequence (5Ј to 3Ј) Source 12S MVZ59 Forward ATAGCACTGAAAAYGCTDAGATG Graybeal, 1997 MVZ50 Reverse TYTCGGTGTAAGYGARAKGCTT Graybeal, 1997 12S A-L Forward AAACTGGGATTAGATACCCCACTAT Goebel et al., 1999 12S F-H Reverse CTTGGCTCGTAGTTCCCTGGCG Goebel et al., 1999 12S L1 Forward AAAAGCTTCAAACTGGGATTAGATACCCCACTAT Feller and Hedges, 1998 L13 Forward TTAGAAGAGGCAAGTCGTAACATGGTA Feller and Hedges, 1998 tRNAVal tRNAVal-H Reverse GGTGTAAGCGARAGGCTTTKGTTAAG Goebel et al., 1999 16S AR Forward CGCCTGTTTATCAAAAACAT Palumbi et al., 1991 BR Reverse CCGGTCTGAACTCAGATCACGT Palumbi et al., 1991 Wilkinson2 Reverse GACCTGGATTACTCCGGTCTGA J.A. Wilkinson et al., 1996 Titus I Reverse GGTGGCTGCTTTTAGGCC Titus and Larson, 1996 L2A Forward CCAAACGAGCCTAGTGATAGCTGGTT Hedges, 1994 H10 Reverse TGATTACGCTACCTTTGCACGGT Hedges, 1994 Rhodopsin Rhod1A Forward ACCATGAACGGAACAGAAGGYCC Bossuyt and Milinkovitch, 2000 exon 1 Rhod1C Reverse CCAAGGGTAGCGAAGAARCCTTC Bossuyt and Milinkovitch, 2000 Rhod1D Reverse GTAGCGGAAGAARCCTTCAAMGTA Bossuyt and Milinkovitch, 2000 Tyrosinase TyrC Forward GGCAGAGGAWCRTGCCAAGATGT Bossuyt and Milinkovitch, 2000 exon 1 TyrG Reverse TGCTGGCRTCTCTCCARTCCCA Bossuyt and Milinkovitch, 2000 Histone H3 H3F Forward ATGGCTCGTACCAAGCAGACVGC Colgan et al., 1999 H3R Reverse ATATCCTTRGGCATRATRGTGAC Colgan et al., 1999 28S 28SV Forward AAGGTAGCCAAATGCCTCATC Hillis and Dixon, 1991 28SJJ Reverse AGTAGGGTAAAACTAACCT Hillis and Dixon, 1991 Seven in SIA1 (T3) Forward TCGAGTGCCCCGTGTGYTTYGAYTA Bonacum et al., 2001 absentiaa SIA2 (T7) Reverse GAAGTGGAAGCCGAAGCAGSWYTGCATCAT Bonacum et al., 2001 a Primers SIA1 and SIA2 were used with the universal T3 and T7 primers following Bonacum et al. (2001).

Fig. 1. Number of DNA base-pairs per terminal. For speci®c terminal data see appendix 1. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 17

LABORATORY PROTOCOLS tice, we generated sequences in forward and reverse directions. The ca. 2400 bp of H1 Whole cellular DNA was extracted from were generated in 5±7 overlapping frag- frozen and ethanol-preserved tissues ( or ments, which allowed further sequence con- muscle) using either phenol-chloroform ex- ®rmation. We also compared the sequences traction methods or the Qiagen DNeasy kit generated for multiple extractions of the following manufacturer's guidelines. PCR same tissues, as well as multiple specimens ampli®cation was carried out in 25 ␮l reac- of the same species. Using Sequencher (Gene tions using Amersham Biosciences puRe Taq Codes) we selected all edited sequences for Ready-To-Go Beads. The standard PCR pro- a given locus and used the ``assemble inter- gram consisted of an initial denaturing step actively'' option to establish the threshold at of 3 minutes at 94ЊC, 35±40 cycles of 1 min- which a given sequence would align with ute at 94ЊC, 1 minute at 45±62ЊC, and 1±1.5 any other sequence, which allowed identical minutes at 72ЊC, followed by a ®nal exten- and nearly identical sequences to be isolated sion step of 6 minutes at 72ЊC. PCR-ampli- for inspection. We compared questionable se- ®ed products were cleaned using the AR- quences with those of con®rmed identity and RAYIT kit (TeleChem International) on a sequences in GenBank. Beckman Coulter Biomek 2000 robot. Cycle- The sequences that passed these tests were sequencing using BigDye Terminators v. 3.0 then aligned using ClustalX (Thompson et (Applied Biosystems) were run in 8 ␮l re- al., 1997). The resulting alignments and actions, and this was followed by isopropa- neighbor-joining trees for each partition were nol-ethanol precipitation and sequencing on examined to detect aberrant sequences and either an ABI 3700 or ABI 3730XL auto- formatting errors (e.g., reverse-comple- mated DNA sequencer. Sequences were ed- ments). Finally, preliminary phylogenetic ited in Sequencher (Gene Codes). analyses were performed, and the resulting Given the magnitude and complexity of topologies were used to identify terminals this project (over 8,500 sequences were gen- that required further scrutiny. Extreme vari- erated), the potential for errors to accumulate ance from expected position suggested the from a variety of sources (e.g., mislabeled possibility of error and caused us to perform vials, contamination, mispipetting, incorrect experiments to con®rm sequence identities. naming of ®les) was a serious concern. We We clarify that no sequence was eliminated took several measures to avoid errors. Tis- solely because it did not ®t our prior notions sues, stock solutions (including DNA ex- of relationships. Rather, the topologies were tracts), and diluted working solutions were used heuristically to single out terminals/se- stored separately. Extractions were done at quences for reexamination. different times in batches of no more than 30 Once sequence identities were con®rmed, samples. Filtered tips were used to manipu- sequences derived from the independent late stock DNA extracts. Multichannel pi- DNA extractions were merged. With a few pettes were used whenever possible, and all exceptions noted later, those from conspeci®c PCR cleaning was done using a Beckman specimens were merged into chimeras (with Coulter Biomek 2000 robot. We extracted polymorphisms coded as ambiguities) to re- 100 tissues twice independently and se- duce the number of terminals in the analysis, quenced at least one locus of each to con®rm but all sequences are deposited separately in sequence identity, and we distributed multi- GenBank (appendix 1). ple specimens of 10 species among different batches and generated all sequences for each to con®rm species identi®cations and se- MOLECULAR SEQUENCE FORMATTING quence identities and detect errors. To allow integration of incomplete se- quence fragments (particularly those obtained SEQUENCE PREANALYSIS: from GenBank; see Taxon Sampling Strategy HEURISTIC ERROR CHECKING and Character Sampling Strategy, above), ac- Numerous steps were taken to detect er- celerate cladogram diagnosis, and reduce rors in DNA sequences. As is standard prac- memory requirements under Iterative Pass 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

TABLE 2 were separated by inserting ampersands (&). Summary of DNA Sequence Data Thus, the multiple fragments of the mtDNA cluster remain in the same ®le and order. The Number of terminals for resulting POY-formatted ®les can be ob- Number which a gene tained from http://research.amnh.org/herpe- of sequence was tology/downloads.html or from the authors. Sequence fragments available

Mitochondrial ribosomal ANALYTICAL STRATEGY cluster 25 532 28S 5 343 We analyzed all data simultaneously using Histone H3 2 378 the program POY (W.C. Wheeler et al., Rhodopsin 2 375 1996±2003) v. 3.0.11a (released May 20, Seven in absentia (SIA) 4 302 2003) run on the AMNH Parallel Computing Tyrosinase 3 202 Cluster. We visualized results using Winclada (Nixon, 1999±2002) and performed addition- al searches of implied alignments by - Optimization, we broke complete sequences ing NONA (Goloboff, 1993±1999) from into contiguous fragments. (This also im- Winclada (see below). proves the performance of POY's implemen- HEURISTIC HOMOLOGY ASSESSMENT: Nu- tation of the parsimony ratchet; see Heuristic merous algorithms of varying degrees of ex- Tree Searching, below.) We did so sparingly, haustiveness have been proposed to optimize however, as these breaks constrain homology unaligned data on a given topology. Our assessment by prohibiting nucleotide com- search strategy employed three Direct Opti- parisons across fragments, that is, it is as- mization algorithms. In order of increasing sumed that no nucleotides from fragment X exhaustiveness and execution time, these are homologous with any nucleotides from were Fixed States Optimization (W.C. fragment Y. As the number of breaks increas- Wheeler, 1999), Optimization Alignment es, so too does the risk of overly constraining (W.C. Wheeler, 1996), and Iterative Pass Op- the analysis and failing to discover the glob- timization (W.C. Wheeler, 2003a). As an in- ally optimal solution(s). dication of the magnitude of the problem of We, therefore, inserted as few breaks as analyzing this 532-terminal dataset, execu- were necessary to maximize the amount of tion time for a single random-addition se- sequence data included, minimize the inser- quence Wagner build (RAS), without swap- tion of terminal N's, and attain maximum- ping, on a 1.7 GHz Pentium 4 Dell Inspiron length fragments of about 500 bases (table 2650 running WindowsXP was 2.69 hours 2). Breaks were placed exclusively in highly under Fixed States and 3.26 hours under Op- conserved regions (many of which corre- timization Alignment. spond to commonly used PCR primers), as Although Fixed States Optimization was recovery of such highly invariable regions is proposed as a novel means of conceptualiz- largely alignment-method independent and ing DNA-sequence homology (W.C. Wheel- the inserted breaks do not prevent discovery er, 1999), we employed it here simply as a of global optima. These highly conserved re- heuristic shortcut. Because Fixed States is so gions were identi®ed via preliminary much faster than the Optimization Alignment ClustalX (Thompson et al., 1997) alignments algorithm, it allowed us to sample more thor- under default parameters. Except for their oughly the universe of trees. (The speed-up usefulness in placing fragments derived from for multiple replicates is actually much great- different PCR primers and detecting errors er than noted earlier for a random-addition (see Sequence Preanalysis, above), these pre- sequence Wagner build, as generating the ini- liminary alignments were used solely for the tial state set is the slowest step in Fixed purpose of identifying conserved regions; States analysis.) The trees obtained in Fixed they did not otherwise inform or constrain States analyses were then used as starting our phylogenetic analysis. Once appropriate points for further analysis under Optimiza- conserved regions were identi®ed, fragments tion Alignment. The potential exists for the 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 19 globally optimal tree (or trees that would Iterative Pass Optimization (W.C. Wheeler, lead to the global optimum when swapped 2003a). under a more exhaustive optimization algo- See table 3 for a summary of general rithm) to be rejected from the pool of can- searching techniques. Initial runs used the didates under the heuristic. To minimize this approxbuild heuristic to speed up building of risk, we also generated a smaller pool of can- starting trees, but the resulting trees required didate trees under Optimization Alignment. much more subsequent re®nement, nullifying The resulting 10 optimal and near-optimal the initial speed-up. Remaining analyses candidate trees were then submitted to ®nal were therefore run without approxbuild. We evaluation and re®nement under Iterative conducted searches without slop or check- Pass optimization using iterativelowmem to slop, both of which increase the pool of trees reduce memory requirements. (For details on examined by swapping suboptimal trees tree-searching algorithms see Heuristic Tree found during the search. Although these Searching, below.) steps can be highly effective, initial trials We did not employ exact during most showed they were too time-consuming for searches, although we did use that command the dataset (especially under Iterative Pass, in the ®nal stages of analysis. To verify where they would also be most relevant). lengths reported in POY, we output the im- A variant of Goloboff's (1999) tree drift- plied alignment (W.C. Wheeler, 2003b) and ing was also used to escape local optima. Al- binary version of the optimal topology in though it is based loosely on Goloboff's al- Hennig86 format with phastwinclad®le and gorithm, the implementation in POY differs opened the resulting ®le in Winclada (Nixon, signi®cantly in the way it accepts candidate 1999±2002), following the procedure of trees during the search (see Goloboff, 1999, Frost et al. (2001). Because each topology for his accept/reject calculation). In POY, the may imply a different optimal alignment, probability of accepting a candidate tree that when multiple optimal topologies were ob- is equal to or worse than the current optimum tained we examined them separately by in- (better trees are always accepted) is given by putting each as a separate ®le using topo®le. 1/(n ϩ c Ϫ b), where c is the length of the Examination of the implied alignments, candidate topology, b is the length of the cur- whether formatted as Hennig ®les or as stan- rent optimum (best), and n is a user-speci®ed dard alignments (impliedalignment), grants factor that decreases the probability of ac- another opportunity to detect errors in for- cepting a suboptimal tree, effectively allow- matting or sequencing (e.g., reverse comple- ing the user to control the ease with which ments; see Sequence Preanalysis, above). the search will drift away from the current HEURISTIC TREE SEARCHING: Ef®cient optimum (we used the default of 2). search strategies for large datasets are to a The parsimony ratchet (Nixon, 1999) was certain degree dataset-dependent (Goloboff, proposed for analysis of ®xed matrices. Giv- 1999), and, as discussed above, common in- en that there are no prespeci®ed column vec- dicators of suf®ciency are unrealistic given tors to be reweighted under dynamic homol- current technological limitations. Therefore, ogy, the original approach had to be modi- rather than apply a simple, prede®ned search ®ed. In the current version of POY, the ratch- strategy (e.g., 100 random-addition sequence et is programmed to reweight randomly Wagner builds ϩ TBR branch swapping), we selected DNA fragments. Our data were di- employed a variety of tree-searching algo- vided into 41 fragments (see table 2), so rithms in our analysis, spending more com- ratchetpercent 15 randomly reweighted 7 puting time on those that proved most fruit- fragments, regardless of their length or rela- ful. Tree fusing (Goloboff, 1999) and TBR tive position. In our analyses we reweighted swapping were performed at various points 15±35% of the fragments and applied throughout the analysis, and optimal trees weights of 2±8ϫ. from different searches were pooled for ®nal As a complementary approach, we also tree fusing and TBR swapping, all of which performed quick searches (few random-ad- was re®ned by submitting optimal topologies dition sequence Wagner builds ϩ SPR) under to swapping and ratcheting (see below) under indel, transversion, and transition costs of 3: 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

TABLE 3 Summary of Tree-Searching Methods Combined in Overall Search Strategy See the text for more detailed explanations and references. Different runs combined multiple procedures, and all runs included SPR and/or TBR re®nement.

Searching method Description of procedure RAS Random addition sequence Wagner builds Constrained RAS As above, but constrained to agree with an input group inclusion matrix derived from the consensus of topologies within 100±150 steps of optimum Subset RAS Separate analysis of subsets of 10±20 taxa; resulting arrangements used to de®ne starting trees for further analysis of complete data set Tree drifting Tree drifting as programmed in POY, using TBR swapping; control factor ϭ 2 (default) Ratcheting (fragment) Ratcheting as programmed in POY, with 15±35% of DNA fragments selected randomly and weighted 2±8 times, saving 1 minimum-length tree per replicate Ratcheting (indel, tv, ts) Ratcheting approximated by applying relative indel- transversion-transition weights of 311, 131, and 113, saving all minimum length trees Constrained ratcheting (fragment) As above, but beginning with the current optimum input as a starting tree and constrained to agree with an input group inclusion matrix derived from the consensus of topologies within 100±150 steps of present optimum Tree fusing Standard tree fusing followed by TBR branch swapping, with the maximum number of fusing pairs left unconstrained Manual rearrangement Manual movement of branches of current optimum Ratcheting (original) of ®nal implied Parsimony ratchet of ®xed matrix, as implemented in Winclada alignment

1:1, 1:3:1, and 1:1:3 and included the result- inclusion matrix (Farris, 1973) in the pro- ing topologies in the pool of trees submitted gram Jack2Hen (W.C.Wheeler, unpublished; to tree-fusing and re®nement under equal available at http://research.amnh.org/sci- weights, following the general procedure of comp/projects/poy.php), and then employing d'Haese (2003). Reweighting in this method constraint in the subsequent searches. To cal- is not done stochastically and therefore dif- culate the consensus we included trees within fers from both Nixon's (1999) original and 100±150 steps of the current optimum, the POY's implementation of the ratchet. How- goal being to collapse enough branches for ever, because it weights sets of transforma- swapping to be effective, but only enough tions drawn from throughout the entire da- branches to make for signi®cant speed-ups of taset, it is likely to capture different patterns RAS ϩ swapping, while still allowing dis- in the data and may be a closer approxima- covery of optimal arrangements within the tion to the original ratchet than POY's im- polytomous groups (see Goloboff, 1999: plementation. Both approaches attempt to es- 420). This is effectively a manual approxi- cape local optima. mation of Goloboff's (1999) consensus- We also performed constrained searches based sectorial search procedure, the main by using Winclada to calculate the strict con- difference being that we collapsed branches sensus of trees within an arbitrary number of based only on tree length and not relative ®t steps of the present optimal, saving the to- difference (Goloboff, 1999; Goloboff and pology as a tree®le, constructing the group- Farris, 2001). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 21

Using constraint ®les generated in the tic shortcuts employed in POY to speed up same way, we also input the current optimum optimization did not prevent discovery of as a starting point for ratcheting. This strat- global optima. It also ensures that users of egy avoids spending time on RAS builds of other programs will be able to duplicate our the unconstrained parts of the tree (which results given our alignment. tend to be highly suboptimal) and seeks to PARALLEL COMPUTING: All POY runs were escape local optima in the same way as un- parallelized across 95 or 64 processors of the constrained ratcheting, discussed earlier. AMNH 256-processor Pentium 4 Xeon 2.8 However, there is a tradeoff in that the ar- GHz Parallel Computing Cluster. Initial anal- rangements may be less diverse (and there- yses divided replicates among 5 sets of 19 fore unable to ®nd global optima) but are processors using controllers, that is, 5 repli- likely to be, on average, closer to the opti- cates were run simultaneously, each paralle- mum score than those examined through lized across 19 processors. Although that RAS. strategy may lead to a more ef®cient parallel As a further manual approximation of sec- implementation of POY (Janies and Wheeler, torial searches, we analyzed subsets of taxa 2001), a shortcoming is that catchslaveout- separately by de®ning reduced datasets with put, which saves all intermediate results to terminals ®les that listed only the targeted the standard error ®le, is disabled when con- terminals. More rigorous searches (at least trollers is in use. Consequently, crashes (e.g., 100 RAS ϩ TBR for each of the reduced due to HVAC failures and overheating) or datasets) of these reduced datasets were then maintenance reboots result in the irrecover- performed, and the results were used to spec- able loss of days or weeks of analysis. To ify starting topologies for additional search- avoid this problem in subsequent runs, we ing of the complete dataset. parallelized each replicate across all proces- As a ®nal attempt to discover more par- sors and ran replicates serially, which al- simonious solutions in POY, we also re- lowed recovery from interrupted runs by in- arranged branches of current optima manu- putting the intermediate results as starting ally. As a general search strategy this would points. obviously be highly problematic, if for no SUPPORT MEASURES: We calculated support other reason than that it would bias results. using the implied alignment of the optimal However, we performed this step primarily hypothesis. That is, the values reported re- to ensure that the ``received wisdom'' was ¯ect the degree of support by the hypothe- evaluated explicitly in our analysis. Our pro- sized transformation series and not by the cedure was to open the current optimum in data per se. It is preferable to evaluate sup- Winclada, target taxa whose placement was port based on the unaligned data, as that pro- strongly incongruent with current taxonomy, vides a more direct assessment of evidential and move them to their expected positions ambiguity. (That is, it is possible for a clade (or place them in polytomies, depending on to appear strongly supported given a partic- the precision of the expectations). The re- ular alignment, but for support to dissolve sulting topology was saved as a tree®le that when an alternative alignment is considered, was read into POY as a starting topology for meaning that the support by the data them- diagnosis and re®nement (e.g., swapping, selves is ambiguous.) We based support mea- tree-fusing). In this way we were sure that sures on the implied alignment because (1) the more heterodox aspects of our results it is much less time-consuming than support were not due simply to failing to evaluate the calculation under dynamic homology, and orthodox alternatives in our searches. we preferred to concentrate computational We analyzed the ®nal implied alignment resources on searches for the optimal solu- obtained in the ®nal searches under Iterative tion; and (2) these values are directly com- Pass Optimization (i.e., the optimal solution parable to those reported in the majority of found through all searching in POY) by car- phylogenetic studies, which derive support rying out 10 independent ratchet runs of 200 values from a single, ®xed alignment. iterations each, using the default reweight- To estimate Bremer values (Bremer, 1994), ings (Nixon, 1999). This ensured that heuris- we output the implied alignment and optimal 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 trees in Hennig86 format using phastwin- have not attempted to provide comparable clad®le, converted it to NEXUS format in characters among the taxa because such a de- Winclada, and then generated a NEXUS in- scription has yet to be accomplished in a de- verse-constraints batch ®le in PRAP (K. tailed way (but see J.D. Lynch, 1973, and MuÈller, 2004), which was analyzed in Laurent, 1986, for general attempts) and is PAUP* 4.0 (Swofford, 2002). Given time outside the scope of this study. A general constraints, tree searches for the Bremer study would obviously change both the de- analysis were super®cial, consisting of only limitation of the characters and the levels of 2 RAS ϩ TBR per group. Jackknife frequen- generality. cies were calculated from 1000 replicates of 1 RAS per replicate without TBR swapping; COMPARABILITY OF SYSTEMATIC STUDIES jackknife analysis was performed by spawn- ing NONA from Winclada. Throughout the review of current taxono- my that follows, we make only passing ref- REVIEW OF CURRENT TAXONOMY, erence to the various analytical techniques THE QUESTIONS, AND TAXON used by various authors. There are two rea- SAMPLING sons for this. Not only is a deep review of In this section we review the existing tax- techniques of phylogenetic inference beyond onomy of living amphibians and explain the scope of this paper, but it probably would which species we sampled and what the jus- be impossible for us to put together a quorum ti®cations were for this sampling3. We also of authors to support any view beyond that examine the evidentiary basis of the current it is monophyletic taxa that we are attempt- taxonomy in an attempt to evaluate which ing to apprehend. parts provide a scienti®c template on which Our main concerns regard the repeatability to interpret evolutionary patterns and trends, of systematic analyses and that readers un- and which parts form an arbitrary and mis- derstand that many, if not most, of the anal- leading structure that are merely anointed by yses cited in this section are not rigorously time and familiarity or, worse, by authority. comparable. In morphological studies it is The canonical issue is obviously monophyly, common practice to report on individual so the question becomes: Does our taxonomy transformation series and the logic behind re¯ect evolutionary (i.e., monophyletic) treating these transformations as additive or groups? And, regardless of that answer, what nonadditive or whether these transformations is the evidentiary basis of the claims that can be polarized individually or not. This have been made about amphibian relation- makes these analyses repeatable because ships? Can we sample taxa in such a way as workers can duplicate data as well as ana- to test those claims? In this section we have, lytical conditions. where practical, provided speci®c evidence DNA sequence studies, however, have from the published record as it bears on these tended not to provide the information nec- questions. The reader should bear in mind essary for independent workers to repeat that much of the current taxonomy rests on analyses, regardless of the accessibility of the subjective notions of overall similarity and original sequence data. In most cases, au- the relative importance of certain characters thors align their sequences manually (which to speci®c Linnaean ranks. Even where is necessarily idiosyncratic and nonrepeata- knowledge claims derive from phylogenetic ble, even if one uses models of secondary analysis, the evidence can be highly contin- structure to help). In cases where alignment gent on a speci®c phylogenetic context. We is done under algorithmic control, it is com- mon to not cite the indel, transversion, and 3 We do not address literature that appeared after 1 transition costs that went into the alignment, August 2005 (although we do address electronically rendering these alignments unrepeatable. available ``in-press'' articles that had not yet appeared Also, many authors ``correct'' alignments by in hard-copy form by that date). This decision will have excluded some important literature, but the date is well without explaining what this means or after the submission date of the manuscript (29 May what these corrections were, further remov- 2005) and a practical end-point was needed. ing alignment from the sphere of repeatabil- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 23 ity. (This ``correction'' almost always means being the smallest taxon enclosing the living that the trees become longer.) crown groups (cf. de Blainville, 1816; Gray, Of concern, at least for the AMNH au- 1825; de Queiroz and Gauthier, 1992; Can- thors, is that the assumptions of alignment natella and Hillis, 1993, 2004). We concur may not be consistent with the assumptions with authors who restrict application of the of analysis. For instance, an author may have name Amphibia to the living crown groups, aligned sequences using one transversion: so for this study we use the terms ``Amphib- transition cost ratio but subsequently ana- ia'' and ``Lissamphibia'' interchangeably. lyzed those data under an evolutionary mod- Testing lissamphibian monophyly and the el that makes entirely different assumptions relationships among the three crown groups about these relative costs. If the alignment of amphibians was and continues to be method is not adequately speci®ed, as is daunting because morphologically the groups common in published works (e.g., Pauly et are mutually very divergent and temporally al., 2004), one is at a loss to know what com- distant from each other and from nonamphi- ponent of the ultimate tree structure is due to bian relatives. Furthermore, testing lissam- the assumptions of alignment or to the as- phibian monophyly may be outside the abil- sumptions of analysis. To illuminate the un- ity of this study to address inasmuch as the derlying incomparability of many molecular major controversy has to do with the phylo- studies, we have provided in the relevant ®g- genetic structure of various fossil groups. ure legends, and where this information can Most authors regard Lissamphibia as a taxon be gleaned from the publication, the align- imbedded in (e.g., Estes, ment costs and whether the sequence was ex- 1965; Trueb and Cloutier, 1991; Lombard cluded for being ``unalignable'' (generally and Sumida, 1992) whereas others regard meaning that the authors did not like the frogs to be temnospondyls and salamanders number of gaps required to align the se- and caecilians to be lepospondyls (Carroll quences), the amount of sequence and from and Currie, 1975; Carroll et al., 1999; Car- what genes, and the kind of analysis (parsi- roll, 2000a; J.S. Anderson, 2001). Laurin mony, Bayesian, or maximum-likelihood), (1998a, 1998b, 1998c) regarded Lissamphi- and, if some general model of nucleotide bia to be completely within , evolution was assumed, what that model but more recent work (e.g., Ruta et al., 2003) was. Because we are alarmed by the lack of returned a monophyletic Lissamphibia to the explicitness in the literature regarding under- temnospondyls. (See Lebedkina, 2004, and lying assumptions, we urge editors to require Schoch and Milner, 2004, for extensive re- that these pieces of information to be includ- views of the alternative views of phylogeny ed in any works that pass over their desks. of modern amphibian groups.) Because none Having provided this preface to our review of these paleontological studies adequately of current taxonomy as a caveat for readers, addressed living diversity, we hope that fu- we now embark on a peregrination through ture work will integrate data presented here the evidentiary basis of current amphibian with fossil taxa as part of the resolution of taxonomy. the problem. Regardless of the consideration of fossil AMPHIBIA taxa, the choice of Recent outgroups for analysis is clearly based on knowledge of the For the purposes of this paper, we are con- relationships of major tetrapod groups. A cerned with amphibians not as the ®ctional coelacanth () represents a near-rel- ``transitional'' group from ®shes to , ative of , and among tetrapods, sev- but as the taxon enclosing the extant crown eral amniotes (Mammalia: Didelphis and Ga- clades Gymnophiona (caecilians), Caudata zella; Testudines: Pelomedusa and Chelydra; (salamanders), and Anura (frogs), together Diapsida: Iguana and Alligator) represent the forming Lissamphibia of Gadow (1901) and nearest living relatives of amphibians. Al- most recent authors (e.g., Milner, 1988, 1993, though our choice of outgroups is made spe- 1994; Ruta et al., 2003; Schoch and Milner, ci®cally to root the ingroup tree, our choice 2004) or Amphibia in the restricted sense of of terminals will allow weak tests of the var- 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 3. Currently accepted view of relation- ships among families based on Nuss- baum and Wilkinson (1989), Hedges and Maxson (1993), M. Wilkinson and Nussbaum (1996), Gower et al. (2002), and M. Wilkinson et al. (2002). Quotation marks denote nonmonophyletic Fig. 2. Four phylogenetic hypotheses of tet- taxa. rapod relationships. A, Gauthier et al. (1988a, 1988b); B, Rieppel and de Braga (1996); C, Zar- doya and Meyer (1998); D, Hedges and Poling (1999). phyletic with respect to caecilians (although Laurin himself considered this conclusion implausible). Previously published molecular data placed salamanders as the sister taxon ious hypotheses of relationships. of either caecilians (Larson, 1991; Feller and The alternative relationships suggested by Hedges, 1998) or frogs (Iordansky, 1996; various authors is large, and an extensive dis- Zardoya and Meyer, 2000, 2001; San Mauro cussion of these alternatives is outside the et al., 2004; Roelants and Bossuyt, 2005; San scope of this paper. Nevertheless, we show Mauro et al., 2005). The latter arrangement four topologies in ®gure 2. The most popular is most favored by morphologists (e.g., Trueb tree of amniote groups among paleontolo- and Cloutier, 1991). Additional tests using gists is shown in ®gure 2A and re¯ects the morphological data of the relative placement preferred topology of Gauthier et al. (1988a, of the living lissamphibians will require eval- 1988b), although some authors suggested, uation of fossils, such as also on the basis of morphological evidence, (McGowan and Evans, 1995; Milner, 2000; that turtles are the sister taxon of lepidosaurs, Gardner, 2001, 2002) and the putative Me- with and mammals successively sozoic and Tertiary caecilians, salamanders, more distantly related (Rieppel and de Braga, and frogs (Estes, 1981; Jenkins and Walsh, 1996; ®g. 2B). This position, however, was 1993; Shubin and Jenkins, 1995; SanchõÂz, disputed by M. Wilkinson et al. (1997). Also 1998; Carroll, 2000a; Gao and Shubin, 2001, relevant to our study, some recent DNA se- 2003). quence studies have found turtles to form the sister taxon of archosaurs (Zardoya and Mey- GYMNOPHIONA er, 1998; Iwabe et al., 2005; ®g. 2C), and others found turtles to be the sister taxon of Caecilians (6 families, 33 genera, 173 spe- archosaurs to the exclusion of lepidosaurs, cies) are found almost worldwide in tropical with mammals outside this group (Hedges terrestrial, semiaquatic, and aquatic habitats. and Poling, 1999; Mannen and Li, 1999; ®g. A reasonably well-corroborated cladogram 2D). Our data will provide a weak test of exists for at least the major groups of cae- these alternatives. cilians (Nussbaum and Wilkinson, 1989; Assuming lissamphibian monophyly, the Hedges and Maxson, 1993; M. Wilkinson relationships among the three major groups and Nussbaum, 1996, 1999; Gower et al., of living lissamphibians remain controver- 2002; M. Wilkinson et al., 2002; ®g. 3). Tax- sial. On the basis of a parsimony analysis of on sampling has not been dense and taxo- morphological data, Laurin (1998a, 1998b, nomic assignments are almost certain to 1998c) suggested that salamanders are para- change with the addition of new taxa and ev- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 25 idence. Nevertheless, it appears that most group Uraeotyphlidae is supported by the caecilian taxa are monophyletic, with the ex- morphological character of the tentacle being ception of ``Ichthyophiidae'' with respect to positioned below the naris. Our sole sample Uraeotyphlidae (Gower et al., 2002) and of this taxon is narayani. ``Caeciliidae'', which includes most of the SCOLECOMORPHIDAE (2 GENERA,6SPECIES): diversity (93 species; 54% of all caecilians) The East African Scolecomorphidae was and which is paraphyletic with respect to Ty- suggested to form the sister taxon of ``Cae- phlonectidae (M.H. Wake, 1977; M. Wilkin- ciliidae'' ϩ Typhlonectidae (Nussbaum, son, 1991) and possibly with respect to Sco- 1979), but because this suggestion was based lecomorphidae (M.H. Wake, 1993; M. Wil- on one of the early phylogenetic studies of kinson et al., 2003). caecilians, the sampling over which this gen- The following taxa were sampled: eralization was made was small. Subsequent RHINATREMATIDAE (2 GENERA,9SPECIES): A studies from mtDNA (M. Wilkinson et al., South American group, Rhinatrematidae is 2003) and morphology (M.H. Wake, 1993; hypothesized to be the sister taxon of re- M. Wilkinson, 1997) suggested that Scole- maining caecilians and is clearly composed comorphidae, like Typhlonectidae, is imbed- of the most generally plesiomorphic living ded within ``Caeciliidae''. The monophyly of caecilians (Nussbaum, 1977, 1979; Duellman Scolecomorphidae is well-corroborated by and Trueb, 1986; San Mauro et al., 2004). morphology (Nussbaum, 1979; M. Wilkin- They retain a tail (a plesiomorphy) but share son, 1997). Nevertheless, we sampled mem- the putatively derived characters of high bers of each of the two nominal genera (Cro- numbers of secondary annuli, having an os taphatrema tchabalmbaboensis and Scole- basale, and lacking the fourth ceratobranchi- comorphus vittatus), both as a test of scole- al. We sampled one species each of the two comorphid monophyly and to help optimize nominal genera ( bivittatum and molecular characters for the family to the ap- sp.) to optimize characters for propriate branch4. the family appropriately and to test the TYPHLONECTIDAE (5 GENERA,14SPECIES): monophyly of this group. The South American Typhlonectidae is a ICHTHYOPHIIDAE (2 GENERA,39SPECIES) morphologically well-corroborated taxon of AND URAEOTYPHLIDAE (1 GENUS,5SPECIES): secondarily aquatic caecilians (M.H. Wake, Tropical Asian Ichthyophiidae was hypothe- 1977; Nussbaum, 1979; M. Wilkinson, sized to form the sister taxon of Uraeotyph- 1991), clearly derived out of ``Caeciliidae''. lidae (M. Wilkinson and Nussbaum, 1996; Although there are several nominal genera of San Mauro et al., 2004), or to include Uraeo- typhlonectids, because of the highly apo- typhlidae (cf. Gower et al., 2002), or, cur- morphic nature and highly corroborated rently less corroborated, to be the sister taxon monophyly of the taxon we sampled only Ty- of Uraeotyphlidae plus stegokrotaphic cae- phlonectes natans. cilians (i.e., ``Caeciliidae'' ϩ Scolecomor- ``CAECILIIDAE'' (21 GENERA, 100 SPECIES): phidae ϩ Typhlonectidae; Nussbaum, 1979; Duellman and Trueb, 1986). The morpholog- 4 A minor but controversial issue is exposed here ical diagnosis of Ichthyophiidae is contingent among the coauthors. Throughout the text, ``phyloge- netic tree'' and ``cladogram'' are used interchangeably, on whether Uraeotyphlus is within or outside although there is good reason to consider the latter to be of Ichthyophiidae, but the presence of an- the operational basis of the former (Platnick, 1977). A gulate annuli anteriorly in ichthyophiids re- related issue is that we prefer the nomenclature of trans- mains an apomorphy among these phyloge- formation series containing characters (e.g., Wiley, netic hypotheses. We have sampled one 1981; Grant and Kluge, 2004), rather than the more op- erational terminology of characters containing character striped (Ichthyophis sp.) that is states. Character transformations happen through time not suspected to be close to Uraeotyphlus along lineages (i.e., along branches in the tree, therby and one unstriped Ichthyophis (I. cf. penin- rendering the notion of branch length). We use the term sularis), which we suspect (M. Wilkinson ``branch'' rather than the more operational ``node'' (a term from computer science, not biology). In other and D.J. Gower, unpubl. data) to be phylo- words, we attempt to use evolutionary terms rather than genetically close to Uraeotyphlus. Monophy- the operational equivalents that have enjoyed consider- ly of the endemic and monotypic Indian able usage. Frost bears responsibility for this decision. 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

This nominal taxon can be diagnosed only in Gao and Shubin (2001) provided a parsi- the sense of being coextensive with the mony analysis of DNA sequences and mor- ``higher'' caecilians (Stegokrotaphia of Can- phology (including relevant fossils) suggest- natella and Hillis, 1993) in lacking a tail, ing that Sirenidae is not the sister taxon of having a stegokrotaphic , and not being the remaining salamander families, but the either a scolecomorphid or typhlonectid. We sister taxon of Proteidae (®g. 5). Otherwise, chose taxa from within the pantropical ``Cae- their results were largely congruent with ciliidae'' that on the basis of previously pub- those of Larson and Dimmick (1993). The lished results (M.H. Wake, 1993; M. Wilkin- exemplars used for their family-group tree son et al., 2003) we predicted would illumi- were not provided nor were the distribution nate the paraphyly of ``Caeciliidae'' with re- of morphological characters suf®ciently de- spect to the presumptively derivative groups tailed to allow us to include their data. Fur- Typhlonectidae and Scolecomorphidae. We ther, Larson et al. (2003), on the basis of mo- sampled: uluguruensis (Afri- lecular data alone (the data themselves not ca), tentaculata (South America), presented or adequately described beyond oaxacae (), noting that they are from nuclear rRNA and ramaswanii (), seraphini mtDNA sequences), suggested the tree (Africa), squalostoma (Africa), Hy- shown in ®gure 6. Larson et al. (2003) also pogeophis rostratus (Seychelles), Schisto- noted that phylogenetic analysis of most metopum gregorii (Africa), and morphological characters, other than those hardyi (South America). associated with production, do not support the monophyly of their Sal- CAUDATA amandroidea (sensu Duellman and Trueb, Salamanders (10 families, 62 genera, 548 1986; all salamander families other than Sir- species) are largely Holarctic and Neotropi- enidae, Hynobiidae, and Cryptobranchidae). cal and are the best known amphibian group, Although we address salamander phylogeny even though their phylogeny is notoriously through the application of a large amount of problematic because of the confounding ef- molecular data, we did not address the mor- fects of paedomorphy on interpreting their phological data set presented by Larson and morphology by (Larson et al., 2003; Wiens Dimmick (1993) and Gao and Shubin (2001, et al., 2005). Apparently independent pae- 2003) because of the lack of correspondence domorphic lineages include Cryptobranchi- between our exemplars and theirs and be- dae, Proteidae, and Sirenidae, as well as var- cause this would have required reconciliation ious lineages within Ambystomatidae and of these data with the frog morphology data Salamandridae. Larson et al. (2003) provided we did include, an undertaking that is outside an extensive discussion of salamander sys- the scope of this study. tematics, offering detailed discussion of the Most recently, Wiens et al. (2005) provid- existing issues, although much of the sup- ed an analysis that included additional char- porting evidence was not disclosed. Until re- acters of morphology and the addition of data cently, Larson and Dimmick (1993) provided from RAG-1 DNA sequences (®g. 7). These the received wisdom on salamander relation- authors presented results from different ana- ships based on a combined analysis of mor- lytical approaches (e.g., maximum-likeli- phology (29 transformation series) and mol- hood, Bayesian, parsimony). We illustrate ecules (177 informative sites from rRNA se- only the parsimony analysis of morphology quences; ®g. 4). The branch associated with ϩ molecules, which most closely approxi- in their tree (all sala- mates our own assumption set. A paper by manders excluding Sirenidae, Cryptobran- San Mauro et al. (2005) provided substan- chidae, and Hynobiidae) is corroborated pri- tially similar results using the RAG-1 gene marily by a number of morphological char- also used by Wiens et al. (2005). acters that are functionally related to the se- SIRENIDAE (2 GENERA,4SPECIES): Sirenidae cretion of a spermatophore (Sever, 1990; is a North American, pervasively paedomor- Sever et al., 1990; Sever, 1991a, 1991b, phic taxon, whose members are obligately 1992, 1994). aquatic and possess large external and 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 27

Fig. 4. Relationships of salamanders suggested by Larson and Dimmick (1993). Families are noted on right. and were employed as outgroups. Consensus of 40 equally-parsimonious trees (length ϭ 460, ci ϭ 0.59). Data are 32 morphological and 177 molecular (nu rDNA) character transformations (from Larson, 1991). The method of DNA alignment was not speci®ed. Gaps were excluded as evidence. lack pelvic girdles and hind limbs as well as morphological similarities (such as external . Only two genera ( and Pseu- gills and reduced maxillae) shared with other dobranchus) are recognized. Sirenidae has obligate paedomorphs have been more-or- been considered the sister taxon of the re- less universally considered by authors to be maining salamanders by most authors be- convergent. Nevertheless, Gao and Shubin cause of its lack of internal fertilization (this (2001), on the basis of an analysis of living is assumed on the basis of its lacking sper- and fossil taxa, concluded that sirenids are matophore-producing glands and not on any the sister taxon of proteids (®g. 5). Wiens et observation regarding its reproductive be- al. (2005) suggested, on the basis of a par- havior) and its primitive closure mech- simony analysis of DNA sequences and mor- anism (Larson and Dimmick, 1993). Other phology, that sirenids are the sister taxon of 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 5. Tree of salamander families from Gao and Shubin (2001; fossil terminals pruned) based on a parsimony analysis of nu rRNA sequence data from Larson and Dimmick (1993) and 60 morphological transformation series (length ϭ Fig. 6. Relationships of salamander families 402; ci ϭ 0.549; ri ϭ 0.537). The sequence align- suggested by Larson et al. (2003) on the basis of ment method was not disclosed. Indels (i.e., gaps) undisclosed nu rRNA and mtDNA sequence data. were treated as evidence. all other salamanders (®g. 7), although their ), are not bounded phylogenetically Bayesian analysis placed Sirenidae as the sis- by these taxa, so our analysis will not pro- ter taxon of Salamandroidea, with Crypto- vide a rigorous test of hynobiid monophyly. branchoidea outside the inclusive group. We CRYPTOBRANCHIDAE (2 GENERA,3SPECIES): selected representatives of each nominal ge- Cryptobranchids are a Holarctic group rep- nus: Siren lacertina, S. intermedia, and Pseu- resented by only three species in two genera, dobranchus striatus. Cryptobranchus (eastern North America) and HYNOBIIDAE (7 GENERA,46SPECIES): The (eastern temperate Asia). We includ- Asian Hynobiidae and Asian and North ed all three species, Cryptobranchus allegan- American Cryptobranchidae are usually con- iensis, Andrias davidianus, and A. japonicus, sidered each others' closest relatives because to test the monophyly of Andrias and opti- they share the putatively plesiomorphic con- mize ``family'' evidence to the appropriate dition of external fertilization and have the branch. The monophyly of Cryptobranchidae m. pubotibialis and m. puboischiotibialis is not seriously in doubt as these giant, ob- fused to each other (Noble, 1931; Larson et ligately paedomorphic salamanders are high- al., 2003; Wiens et al., 2005). Hynobiids ly apomorphic in many ways, such as in have aquatic larvae and transformed adults, lacking gills or functional , and instead and they retain angular bones in the lower respiring across the extensive surface. jaw (presumed plesiomorphies). Morpholog- Like Hynobiidae and Sirenidae (presum- ical evidence in support of monophyly of this ably), cryptobranchids lack internal fertiliza- group are septomaxilla absent (also absent in tion. plethodontids and ambystomatids), ®rst hy- PROTEIDAE (2 GENERA,6SPECIES): Protei- pobranchial and ®rst ceratobranchial fused dae is another obligate paedomorphic per- (also in ), second ceratobranchial ennibranch clade considered to be monophy- in two elements, and palatal dentition re- letic because of its loss of the maxillae (also placed from the posterior of the vomer (also very reduced in sirenids, apparently indepen- in ambystomatids; Larson and Dimmick, dently) and the basilaris complex of 1993). Our selection of hynobiid taxa was (also lost in sirenids, plethodontids, and restricted to sibiricus and Batra- some salamandrids), and because it has other chuperus pinchoni. Larson et al. (2003) sug- characters associated with paedomorphy, gested that Onychodactylus, especially, and such as lacking a m. rectus abdominis (No- several genera that we could not obtain (e.g., ble, 1931). Unlike sirenids, cryptobranchids, 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 29

Fig. 7. Relationships of salamanders suggested by Wiens et al. (2005). Families are noted on right. Results re¯ect a parsimony analysis of 326 character transformations of morphology (221 parsimony- informative), and DNA sequences from nu rRNA (212 bp from Larson, 1991; 147 parsimony-infor- mative) and RAG-1 (1,530 bp; 624 parsimony-informative). Sequence alignment was made using Se- quencher (Gene Codes Corp.). Morphological characters identi®ed as paedomorphic were treated as unknown for adult morphology and in some cases hypothetical terminals were related-species chimaeras of composite molecular and morphological data. Molecular transformations were weighted equally in analysis. Inferred insertion-deletion events were coded as binary characters separate from the nucleotide sequence characters and indel-required gaps within sequences were coded as missing. The tree was rooted on Gymnophiona ϩ Anura. 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

and hynobiids, but like other salamander ERA, 374 SPECIES): Plethodontidae includes families, proteids employ internal fertiliza- the large majority of salamander species, tion through use of a spermatophore (Noble, with most being in North America, Central 1931). In our analysis, we included two spe- America, and South America, with Speleo- cies of (of North America), N. cf. mantes found in Mediterranean Europe and beyeri and N. maculosus, but were unsuc- Karsenia found in the Korean Peninsula cessful in amplifying DNA of the only other (Min et al., 2005). The monophyly of the genus, Proteus (which is found only in the group is not seriously questioned, as its western Balkans). Nevertheless, Trontelj and members share a number of morphological Goricki (2003) did study Proteus and pro- synapomorphies such as nasolabial grooves vided molecular evidence consistent with the in transformed adults and the absence of monophyly of Proteidae, and Wiens et al. lungs (found in other groups as well; Larson (2005), also reporting on both Necturus and and Dimmick, 1993). Starting in 2004, and Proteus, subsequently provided strong evi- while this project was in progress, under- dence in favor of its monophyly. standing of the evolution of Plethodontidae RHYACOTRITONIDAE (1 GENUS,4SPECIES): moved into a dynamic state of ¯ux with the Western North American Rhyacotriton was publication of a series of important studies originally placed in its own subfamily within addressing substantial amounts of DNA se- Ambystomatidae (Tihen, 1958) but was quence data and morphology (Chippindale et shown to be distantly related to ambysto- al., 2004; Mueller et al., 2004; Macey, 2005; matines by Edwards (1976), Sever (1992), Wiens et al., 2005). Before 2004, - and Larson and Dimmick (1993), who con- tid phylogeny appeared to be reasonably well sidered it to be a family distinct from Am- understood (D.B. Wake, 1966; D.B. Wake bystomatidae. Wiens et al. (2005) consid- and Lynch, 1976; J.F. Lynch and Wake, ered, on the basis of their parsimony analy- 1978; D.B. Wake et al., 1978; Maxson et al., sis, that Rhyacotritonidae is the sister taxon 1979; Larson et al., 1981; Maxson and Wake, of Amphiumidae ϩ Plethodontidae. Good 1981; Hanken and Wake, 1982; J.F. Lynch et and Wake (1992) provided the most recent al., 1983; D.B. Wake and Elias, 1983; Lom- revision. Rhyacotritonidae retains a reduced bard and Wake, 1986; D.B. Wake, 1993; ypsiloid cartilage and has at least one apo- Jackman et al., 1997; GarcõÂa-ParõÂs and Wake, morphy associated with the glandular struc- 2000; Parra-Olea et al., 2004) with the group ture of the (Sever, 1992). Inasmuch as putatively composed of two monophyletic the four species are seemingly very closely subfamilies (®g. 8), Desmognathinae and related and morphologically very similar, we Plethodontinae, although the morphological sampled only Rhyacotriton cascadae, al- evidence for any suprageneric group other though this leaves the taxon's monophyly un- than Desmognathinae and Bolitoglossini (a tested. tribe in Plethodontinae as then de®ned) was AMPHIUMIDAE (1 GENUS,3SPECIES): The equivocal. of eastern North America have Desmognathines (2 genera, 20 species; reduced limbs and are obligate aquatic pae- ϩ Phaeognathus) as tradi- domorphs. They have internal fertilization tionally understood share nine morphological and a suite of morphological features that are characters suggested to be synapomorphies associated with spermatophore formation and (Schwenk and Wake, 1993; Larson et al., internal fertilization. Some authors have as- 2003), although at least some of them may sociated Amphiumidae with Plethodontidae be manifestations of a single transformation (sharing fused maxillae and reproductive be- having to do with the unique method of jaw havior patterns; e.g., Salthe, 1967; Larson closure: (1) heavily ossi®ed and strongly ar- and Dimmick, 1993) and recent molecular ticulated skull and ; (2) dorsoven- studies place them here as well (Wiens et al., trally ¯attened, wedge-like head; (3) modi- 2005). The three species are very similar and ®ed anterior trunk vertebrae; (4) enlarged share many apomorphies, so we restricted dorsal spinal muscles; (5) hindlimbs larger our sampling to . than forelimbs; (6) stalked occipital con- PLETHODONTIDAE (4 SUBFAMILIES,27GEN- dyles; (7) enlarged quadratopectoralis mus- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 31

Fig. 8. Composite tree of hypothesized relationships among Plethodontidae as inferred from 1966± 2004 literature; subfamilies and tribes noted on the right (D.B. Wake, 1966; D.B. Wake and Lynch, 1976; J.F. Lynch and Wake, 1978; D.B. Wake et al., 1978; Maxson et al., 1979; Larson et al., 1981; Maxson and Wake, 1981; Hanken and Wake, 1982; J.F. Lynch et al., 1983; D.B. Wake and Elias, 1983; Lombard and Wake, 1986; D.B. Wake, 1993; Jackman et al., 1997; GarcõÂa-ParõÂs and Wake, 2000; and Parra-Olea et al., 2004). Quotation marks denote nonmonophyletic taxa. cles; (8) modi®ed atlas; and (9) presence of Plethodontinae were included three nominal atlantomandibular ligaments. Most species tribes: Hemidactyliini, Plethodontini, and have a biphasic life history, but at least some Bolitoglossini. species have either nonfeedling larvae or di- Hemidactyliini (5 genera, 33 species) was rect development (Tilley and Bernardo, the only putative plethodontine group with 1993). Plethodontinae in the pre-2004 sense free-living larvae and transformation into (®g. 8) did not have strong morphological adults (although this is shared with most des- evidence in support of its monophyly, al- mognathines). Lombard and Wake (1986) though Lombard and Wake (1986) suggested suggested that Hemidactylium is the sister that possessing three embryonic or larval taxon of Stereochilus ϩ (Eurycea, Gyrino- epibranchials is synapomorphic. Within philus, and ) but provided only 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 a single morphological character (parietal breakage specializations was synapomorphic with a distinct ventrolateral shelf) in support and for the supergenus exclud- of the monophyly of this group. ing they suggested that fused Plethodontini (3 genera, 62 species), as maxillae was a synapomorphy. traditionally understood, was a heteroge- As noted above, in 2004±2005 three stud- neous assemblage composed of Plethodon, ies appeared that transformed our under- Aneides, and the more distant (1 standing of plethodontid relationships nominal species, but likely containing many (Mueller et al., 2004; Chippindale et al., species under any meaningful de®nition of 2004; Macey, 2005). Although there are that term; see Highton, 1998). Lombard and three relevant analyses, there are only two Wake (1986) suggested two morphological data sets. Mueller et al. (2004; ®g. 9) pre- characters in support of the monophyly of sented a Bayesian analysis of complete this group (radii expanded and fused to bas- mtDNA genomes; this data set was reana- ibranchial, and presence of a posterior max- lyzed by parsimony and extensively dis- illary facial lobe). cussed by Macey (2005; ®g. 10). Another As traditionally viewed (before 2004), data set and analysis of combined morphol- Bolitoglossini (15 genera, 222 species) rep- ogy and DNA sequence evidence was pro- resented a highly-speciose group in the New vided by Chippindale et al. (2004; ®g. 11). World tropics and west-coastal North Amer- All three studies suggested strongly not ica, with isolated representation in Mediter- only that Plethodontinae (as traditionally un- ranean Europe. The group was characterized derstood) is paraphyletic with respect to Des- by having a projectile , although this mognathinae, but that the traditional view of also appears in other plethodontids. plethodontid relationships was largely mis- Lombard and Wake (1986) proposed a taken, presumably due in part to the special (nonparsimonious) scenario in which they pleading for particular characters that under- suggested 10 synapomorphies of Bolitoglos- pinned the older system of subfamilies and sini, all associated with the structure and tribes. Mueller et al. (2004) found that all function of the tongue. They regarded the su- three of the traditionally recognized pletho- pergenus (Hydromantes ϩ dontine tribes, Bolitoglossini, Hemidactyli- ) to be the sister taxon of the ini, and Plethodontini, are polyphyletic. supergenus Bolitoglossa ϩ supergenus Ba- Chippindale et al. (2004) found Hemidacty- trachoseps (containing solely Batrachoseps) liini and Plethodontini to be polyphyletic, based on two synapomorphies. Elias and with Bolitoglossini insuf®ciently sampled to Wake (1983) discussed phylogeny within test its monophyly rigorously. Macey (2005; Bolitoglossini and suggested the topology ®g. 10) also rejected the monophyly of Bol- Hydromantes [including Speleomantes] ϩ itoglossini and Hemidactyliini, in his reanal- (Batrachoseps (Nyctanolis ϩ other bolito- ysis of the data of Mueller et al. (2004). glossine genera)). Synapomorphies given by Mueller et al. (2004; ®g. 9) placed Hemidac- Elias and Wake (1983) for Bolitoglossini are tylium as the sister taxon of Batrachoseps (a (1) urohyal lost; (2) radii fused to the basi- bolitoglossine) and the remaining hemidac- branchial; (3) long epibranchials relative to tyliines as the sister of a group of bolito- the ceratobranchials; (4) second ceratobran- glossines (excluding Hydromantes and Spe- chial modi®ed for force transmission; (5) leomantes). Chippindale et al. (2004; ®g. 11) presence of a cylindrical muscle complex considered Hemidactylium to be the sister around the tongue; (6) juvenile otic capsule taxon of all other bolitoglossines and hemi- con®guration. The synapomophry for Batra- dactyliines, and the remaining hemidactyli- choseps ϩ Nyctanolis ϩ other bolitoglossine ines to form the sister taxon of Hemidacty- genera was reduction in number of caudos- lium ϩ bolitoglossines (Hydromantes and acral vertebrae to two. For the supergenus Speleomantes not analyzed). Bolitoglossa (Nyctanolis ϩ other genera of Chippindale et al (2004; ®g. 11) provided bolitoglossines, excluding Batrachoseps and a new taxonomy, recognizing a newly for- supergenus Hydromantes), they suggested mulated Plethodontinae (including Pletho- that having the tail base with complex of dontini and Desmognathinae of the older tax- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 33

Fig. 9. Tree of Plethodontidae by Mueller et al. (2004), with the traditional taxonomic assignments (Desmognathinae ϩ tribes of Plethodontinae; Wake, 1966) placed on the right, with taxonomic fragments numbered for clarity. The generic taxonomy was updated to re¯ect name changes of former luschani to Lyciasalamandera (Veith and Steinfartz, 2004) and Hydromantes italicus to Speleomantes. The results re¯ect a Bayesian analysis of entire mt DNA genomes (number of informative sites not stated, but analyzed fragments totalled 14,040 bp), with control region and ambiguously alignable region excluded. Sequences were aligned with default costs of GCG v. 10.3 (Accelrys, San Diego; cost of 8 for gap creation and extension cost of 2) and subsequently adjusted manually. It was not stated whether gaps were treated as evidence or as missing data. onomy). The sister taxon of Plethodontinae Hemidactylium (Hemidactyliinae) as the sis- was not named in their taxonomy, the com- ter taxon of the remaining plethodontids. ponent parts being named Hemidactyliinae Clearly, the analyses of mtDNA-sequence (for Hemidactylium alone), Spelerpinae (for data by Mueller et al. (2004) and Macey the remainder of the old Hemidactyliini), and (2005) and of nuDNA, mtDNA, and mor- Bolitoglossinae (identical in content to the phology by Chippindale et al. (2004) 5 are old Bolitoglossini, these authors not having studied Hydromantes sensu lato). Mueller et 5 The Bayesian analysis of plethodontid relationships presented by Min et al. (2005) was based on a subset of al. (2004), followed by Macey (2005), the data provided by Chippindale et al. (2004) and showed that Hydromantes (in the sense of Wiens et al. (2005), with the addition of sequences of including Speleomantes) is not imbedded in Karsenia koreana and Hydromantes brunus. Because Bolitoglossini, as previously supposed, but is that analysis rests on a smaller amount of data than the earlier studies and was performed solely to place the imbedded in Plethodontinae. Macey (2005) newly-discovered Karsenia (as the sister taxon of Anei- arrived at the same taxonomy as Chippindale des ϩ desmognathines), we restrict our comments about et al. (2004), although Macey (2005) placed this paper to the placement of Karsenia. 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 10. Parsimony tree of Plethodontidae by Macey (2005), a reanalysis of entire mt DNA genome sequence data provided by Mueller et al. (2004). On right are the traditional taxonomy and Macey's revised subfamilial taxonomy, which is substantially identical to that suggested by Chippindale et al. (2004; ®g. 11). The generic taxonomy is updated to re¯ect name changes of former Salamandra luschani (Veith and Steinfartz, 2004) and Hydromantes italicus.

strongly discordant with previous (and more pinae of Chippindale et al., 2004, and Macey, limited) morphological and molecular re- 2005): Eurycea wilderae and sults. Because of the timing of the appear- porphyriticus. ance of these papers, our selection of taxa Of the new Plethodontinae (composed of was chosen to address the older, more tradi- former Desmognathinae, Plethodontini, and tional view but may provide a weak test of supergenus Hydromantes of Bolitoglossini) the new view of plethodontid phylogeny and we sampled broadly. We included one spe- taxonomy. cies of western Plethodon, P. dunni, and one We included in our analysis Hemidacty- species of eastern Plethodon, P. jordani.We lium scutatum (Hemidactyliinae) as well as also included Aneides hardii and Ensatina the more ``typical'' hemidactyliines (Speler- eschscholtzii. Mueller et al. (2004), based on 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 35

Fig. 11. Tree of Plethodontidae suggested by Chippindale et al. (2004) based on parsimony analysis of 104 transformation series of morphology and 1,493 informative sites of nu DNA (RAG-1) and mt DNA (cytochrome c and ND4a). On the right (left to right) are the old taxonomy of plethodontids and the taxonomy recommended by Chippindale et al. (2004). Sequences were aligned manually with only single-codon indels; gaps were considered missing data in the analysis. analysis of mtDNA, rejected the monophyly Plethodon and Aneides to be rather distantly of Plethodontini, placing Ensatina as the sis- related. We bracketed the diversity (Titus and ter taxon of desmognathines. (In a parsimony Larson, 1996) of desmognathines (the pre- analysis of the same data, Macey, 2005, 2004 Desmognathinae) by sampling Phaeog- placed Ensatina as the sister taxon of Hydro- nathus hubrichti, Desmognathus quadrama- mantes.) The monophyly of Plethodon,in culatus, and D. wrighti. Of the supergenus particular, is controversial, with some authors Hydromantes, formerly in Bolitoglossini, we (e.g., Larson et al., 1981; Mahoney, 2001) sampled Hydromantes platycephalus and ®nding the western species to be closer to Speleomantes italicus. Aneides to the exclusion of eastern species, Of Bolitoglossinae we sampled 11 of the and others (e.g., Chippindale et al., 2004; 14 nominal genera: supergenus Batrachoseps Mueller et al., 2004; Macey, 2005) ®nding (B. attenuatus and B. wrightorum), and su- 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 12. Salamandrid relationships suggested by Titus and Larson (1995) based on a parsimony analysis of 44 morphological character transformations and 431 informative sites of ca. 1.8 kb of the 12S and 16S mt rRNA and tRNAVal fragments of mtDNA. Sequence alignment was done using MALIGN (W.C. Wheeler and Gladstein, 1992) with equal weighting of transversions and transitions and a gap penalty cost of 6. Sequence data and morphology in parsimony analysis had equal costs and gaps were treated as evidence. The tree was rooted on Eurycea ϩ Phaeognathus; tree length ϭ 2,081. Generic names are updated to re¯ect the naming of (Veith and Steinfartz, 2004) and the partition of into Mesotriton, (not studied by Titus and Larson, 1995), and Triturus (GarcõÂa-ParõÂs et al., 2004b). pergenus Bolitoglossa (Bolitoglossa rufes- in addition to two cranial characters (pres- cens, alvarezdeltoroi, Dendrotri- ence of a frontosquamosal arch and fusion of ton rabbi, , Lineatriton lineo- the premaxillaries [reversed in ϩ lus, abscondens, unifor- , and Chioglossa]). mis, townsendi, Pseudoeurycea Titus and Larson (1995) provided a phy- conanti, and sp.). logenetic tree on the basis of a study of mt SALAMANDRIDAE (18 GENERA,73SPECIES): rRNA and morphology data (®g. 12). Scholz Salamandridae is found more-or-less (1995; ®g. 13) obtained similar results on the throughout the Holarctic, with the bulk of its basis of morphology and courtship behavior. phylogenetic and species diversity in tem- Zacj and Arntzen (1999) also reported on perate Eurasia. Salamandrids are character- phylogenetics of Triturus, showing (as did ized by strongly keratinized skin in adults Titus and Larson, 1995) that it is composed (except for the strongly aquatic ), of two groups: (1) Triturus vulgaris ϩ Tri- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 37

written, but we chose taxa that should allow the basic structure of salamandrid phylogeny to be elucidated. To bracket this suggested topology with appropriate taxonomic sam- ples we chose asper, crocatus, viridescens, Pach- ytriton brevipes, sp., Pleu- rodeles waltl, Salamandra salamandra, Tar- icha sp., Triturus cristatus, and . DICAMPTODONTIDAE (1 GENUS,4SPECIES): The North American Dicamptodon is related to Ambystomatidae (Larson and Dimmick, 1993; ®g. 4) and, like them, some popula- tions are neotenic (Nussbaum, 1976). Like other salamandroid salamanders they have internal fertilization and a suite of morpho- Fig. 13. Consensus of salamandrid relation- logical features associated with forming and ships suggested by Scholz (1995) based on a par- collecting . Dicamptodon dif- simony analysis of 27 character transformations fers from Ambystomatidae in glandular fea- of morphology and behavior. Triturus in Scholz's tures of the cloaca and in attaining a large sense included what is now Lissotriton, Mesotri- size, but is considered by most workers as ton, and Triturus (GarcõÂa-ParõÂs et al., 2004b). the sister taxon of Ambystomatidae (e.g., Larson et al., 2003Юg. 6; Wiens et al., turus marmoratus species groups; (2) and 2005Юg. 7). We sampled both Dicampto- Triturus cristatus group, but not addressing don aterrimus and D. tenebrosus. its polyphyly. Steinfartz et al. (2002) report- AMBYSTOMATIDAE (1 GENUS,31SPECIES): ed on salamandrid phylogeny and substanti- North American Ambystomatidae is a mor- ated the polyphyly of Triturus and of Mer- phologically compact family having internal tensiella. Subsequently (and appearing after fertilization via a spermatophore and the this analysis was completed), GarcõÂa-ParõÂs et suite of morphological characters that sup- al. (2004b) partitioned the polyphyletic ``Tri- port this attribute. Some populations exhibit turus'' into three genera (Triturus, Lissotri- neotenic aquatic adults. ton, and Mesotriton), based on the sugges- The last summary of phylogeny within the tions that (1) Triturus, sensu stricto (Triturus group based on explicit evidence was pre- cristatus ϩ T. marmoratus species groups) is sented by Shaffer et al. (1991; see also Lar- most closely related to Euproctus; (2) Me- son et al., 2003), who provided a cladogram sotriton (Triturus alpestris) is the sister taxon based on 32 morphological transformation of a group composed of Cynops, Parame- series and 26 allozymic transformation se- sotriton, and Pachytriton); and (3) Lissotri- ries. The dichotomy in this tree is be- ton (Triturus vulgaris species group) is of tween Ambystoma gracile ϩ A. maculatum uncertain relationship to the other compo- ϩ A. talpoideum on one hand, and all other nents, but does not form a monophyletic species of Ambystoma, on the other. We were group with either Mesotriton or Triturus. unable to obtain any of these three species, GarcõÂa-ParõÂs et al. (2004a: 602) also sug- but we did sample Ambystoma cingulatum, gested that ongoing molecular work (evi- A. mexicanum and A. tigrinum. Ambystoma dence undisclosed), will show Euproctus to mexicanum and A. tigrinum are very closely be paraphyletic and that Triturus vittatus will related, and A. cingulatum is distantly related not be included within Triturus, the oldest to them. This is a weaker test of monophyly available name for this taxon being Omma- than we would have liked because it does not totriton Gray, 1850. include A. gracile, A. maculatum,orA. tal- We could not address these ®nal issues, poideum. Further, Larson et al. (2003) sug- these appearing well after the manuscript was gested that, in addition to A. gracile, A. ma- 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 culatum, and A. talpoideum, A. jeffersonian- the extent of character con¯ict within their um, A. laterale, A. macrodactylum, and A. data set was never adequately exposed. More opacum were likely to be outside of the taxa recently, Haas (2003; ®g. 15) provided a dis- bracketed by our species, although the evi- cussion of frog evolution, based primarily on dence for this was not presented. new larval characters. Haas did, however, ex- clude several of the adult characters included ANURA by Ford and Cannatella (1993) as insuf®- ciently characterized or assayed. More re- Frogs (32 families, ca. 372 genera, 5227 cently, important discussions of phylogeny species) constitute the vast majority (88%) of have been made in the context of DNA se- living species of amphibians and the bulk of quence studies (Roelants and Bossuyt, their genetic, physiological, ecological, and 2005Юg. 16; San Mauro et al., 2005Юg. morphological diversity. Despite numerous 17) that will be cited throughout our review. studies that point towards its de®ciencies The monophyly of frogs (Anura) relative (e.g. Kluge and Farris, 1969; Lynch, 1973; to other living amphibians has not been gen- Sokol, 1975, 1977; Duellman and Trueb, erally questioned6 (although the universality 1986; Ruvinsky and Maxson, 1996; Maglia, of this taxon with respect to some fossil an- 1998; Emerson et al., 2000; Maglia et al., tecedent taxa has (e.g., Grif®ths, 1963; Ro- 2001; Scheltinga et al., 2002; Haas, 2003; cÃek, 1989, 1990), and the number of mor- Roelants and Bossuyt, 2005; San Mauro et phological characters corroborating this al., 2005; Van der Meijden et al., 2005), the monophyly is largeÐe.g., (1) reduction of current classi®cation continues in many of its vertebrae to 9 or fewer; (2) atlas with a single parts to re¯ect sociological conservatism and centrum; (3) hindlimbs signi®cantly longer the traditional preoccupation with groupings than forelimbs, including elongation of ankle by subjective impressions of overall similar- bones; (4) fusions of radius and ulna and tib- ity; special pleading for characters consid- ia and ®bula; (5) fusion of caudal vertebral ered to be of transcendent importance; and segments into a urostyle; (6) fusion of hyob- notions of ``primitive'', ``transitional'', and ranchial elements into a hyoid plate; (7) pres- ``advanced'' groups instead of evolutionary ence of keratinous jaw sheaths and kerato- propinquity. Understanding of frog relation- donts on larval mouthparts; (8) a single me- ships remains largely a tapestry of con¯icting dian spiracle in the , a characteristic of opinion, isolated lines of evidence, unsub- the Type III (consideration of this as stantiated assertion, and unresolved paraphy- a synapomorphy being highly contingent on ly and polyphyly. Indeed, the current taxon- the preferred overall cladogram); (9) skin omy of frogs is based on a relatively small with large subcutaneous lymph spaces; and sampling of species and in many cases the (10) two m. protractor lentis attached to , putative morphological characteristics of ma- based on very narrow taxon sampling (Saint- jor clades within Anura are overly-general- Aubain, 1981; Ford and Cannatella, 1993). ized, overly-interpreted, and rei®ed through Haas (2003) suggested (®g. 15) an addi- generations of literature reviews (e.g., Ford tional 20 synapomorphies from larval mor- and Cannatella, 1993), of which this review phology: (1) paired venae caudalis lateralis is presumably guilty as well. This general short; (2) operculum fused to abdominal lack of detailed understanding of anuran re- wall; (3) m. geniohyoideus origin from cer- lationships has been exacerbated by the ex- atobranchials I±II; (4) m. interhyoideus pos- plosive discovery of new species in the past terior absent; (5) larval jaw depressors orig- 20 years. inate from palatoquadrate; (6) ramus maxil-

Currently, the most widely cited review of laris (cranial nerve V2) medial to the m. le- frog phylogeny is Ford and Cannatella (1993; ®g. 14), which provided a narrative 6 RocÏek and Vesely (1989) suggested a diphyletic or- discussion of the evidence for a novel view igin of Anura based on a hypothesized nonhomology between the rostral plate of pipoid larvae and the cornua of frog phylogeny without providing all of trabeculae of other anuran larvae. The developmental the underlying data from which this discus- homology of these structures was later established (Ols- sion was largely derived. The result was that son and Hanken, 1996; de Sa and Swart, 1999). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 39

Fig. 14. Narrative tree of relevant anuran taxa by Ford and Cannatella (1993). A branch subtending Hylidae ϩ Pseudidae in the original ®gure is collapsed per errata distributed with reprint. An asterisk was used by these authors to denote a metataxon, and quotation marks to denote nonmonophyly.

vator manidbulae longus; (7) ramus hypobranchiale; (12) processus urobranchial- mandibularis (cranial nerve V3) anterior (dor- is short, not reaching beyond the hypobran- sal) to the m. levator mandibulae longus; (8) chial plates; (13) commisura proximalis I ramus mandibularis (cranial nerve V3) ante- present; (14) commisura proximalis II pre- rior (dorsal) to the externus group; (9) car- sent; (15) commisura proximalis III present; tilago labialis superior (suprarostral cartilage) (16) ceratohyal with diarthrotic articulation present; (10) two perilymphatic foramina; present, medial part broad; (17) cleft between (11) hypobrachial skeletal parts as planum hyal arch and I closed; (18) 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 41 ligamentum cornuquadratum present; (19) sacral vertebrae (Ford and Cannatella, 1993), ventral valvular velum present; (20) branchi- both considered plesiomorphic within Anu- al traps present. Haas also suggested ra7. Ascaphus has an intromittant organ (apo- that the following were synapomorphies not morphic) in males and a highly modi®ed tor- mentioned as such by Ford and Cannatella rent-dwelling tadpole. The vertebrae are am- (1993): (1) inguinal; (2) vertical phicoelous and ectochordal (Nicholls, 1916; pupil shape; (3) overlapping scapula Laurent, 1986), presumably plesiomorphic at anteriorly; and (4) cricoid cartilage as a this level of generality. Our sampled species closed ring. for this taxon is Ascaphus truei, one of the two closely-related species. ``PRIMITIVE'' FROGS LEIOPELMATIDAE (1 GENUS,4SPECIES): Iso- We ®rst address the groups that are some- lated in New Zealand, Leiopelmatidae, like times referred to collectively as Archaeoba- Ascaphidae, is a generally very plesiomorph- trachia (Duellman, 1975) and traditionally ic group of frogs. Nevertheless, it possesses are considered ``primitive'', even though the apomorphies, such as ventral inscription ribs, component taxa have their own apomorphies found nowhere else among frogs (Noble, and the preponderance of evidence suggests 1931; Laurent, 1986; Ford and Cannatella, strongly that they do not form a monophy- 1993). Unlike Ascaphus, Leiopelma does not letic group (Roelants and Bossuyt, 2005; San have feeding larvae (Archey, 1922; Altig and Mauro et al., 2005). McDiarmid, 1999; Bell and Wassersug, ASCAPHIDAE (1 GENUS,2SPECIES): Ford and 2003). As in Ascaphidae, the vertebrae are Cannatella (1993) considered North Ameri- amphicoelous and ectochordal with a persis- can Ascaphus (Ascaphidae) to be the sister tent notochord (Noble, 1924; Ritland, 1955) taxon of all other frogs (®g. 14), although on and both vocal sacs and vocalization are ab- the basis of allozyme study by Green et al. sent (Noble and Putnam, 1931). (1989) and, more recently, Roelants et al. Ford and Cannatella (1993) suggested that (2005; ®g. 16) and San Mauro et al. (2005; Leiopelmatidae is the nearest relative of all ®g. 17), on the basis of evidence from DNA other frogs (excluding Ascaphidae) and listed sequences, suggested that Ascaphidae ϩ ®ve synapomorphies in support of this Leiopelmatidae forms a monophyletic group. grouping (their Leiopelmatanura): (1) elon- BaÂez and Basso (1996) presented a phylo- gate arms on the ; (2) loss of the as- genetic analysis designed to explore the re- cending process of the palatoquadrate; (3) lationships of the fossil anurans Vieraella sphenethmoid ossifying in the anterior posi- and with the extant taxa As- tion; (4) exit of the root of the facial nerve caphus, Leiopelma, Bombina, Alytes, and from the braincase through the facial fora- . Despite their restricted taxon men, anterior to the auditory capsule, rather sampling, their results also support the than via the anterior acoustic foramen into monophyly of Ascaphus ϩ Leiopelma, al- the auditory capsule; (5) palatoquadrate ar- though the authors considered their evidence weak for reasons of dif®culty in evaluating 7 Ritland (1955) suggested the possibility that the m. characters. caudalipuboischiotibialis is not homologous with the Green and Cannatella (1993) did not ®nd tail-wagging muscles of salamanders but instead, an ac- ϩ cessory coccygeal head of the m. semimembranosus. In a monophyletic Ascaphus Leiopelma. As- that case, the character would be judged to be a syna- caphus and Leiopelma share the presence of pomorphy of Ascaphus ϩ Leiopelma, rather than a sym- a m. caudalipuboischiotibialis and nine pre- plesiomorphy shared by those taxa.

← Fig. 15. Anuran relationships suggested by Haas (2003). Consensus of 144 equally parsimonious trees discovered by parsimony analysis of 151 character-transformation series (excluding his morpho- metric characters 12, 83, 116, and 117, as well as 102) of larval and adult morphology and reproductive mode (ci ϭ 0.31; ri 0.77). Taxonomy is updated to re¯ect subsequent publications. 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 16. Tree of amphibians provided by Roelants and Bossuyt (2005). This tree re¯ects a maximum- likelihood analysis of 3,963 aligned positions (2,022 variable and 1,788 parsimony-informative) of three protein-coding nuDNA genes (ca. 555 bp of RAG-1, ca. 675 bp of CXCR-4, ca. 1280 bp of NCX-1) and ca. 1940 bp of the mitochondrial genome (part of 16S and tRNAMet, and all of tRNALeu, tRNAIle, ND-1, and tRNAGln). Alignment was done initially using ClustalX (Thompson et al., 1997; presumably applying default cost functions) followed by a probabilistic method implemented in the program ProAlign (LoÈytynoja and Milinkovitch, 2003) and, in the case of 16S and tRNA seqments, subsequently modi®ed manually, guided by models of secondary structure for Xenopus. Gaps were treated as missing data and ambiguously aligned sequences were excluded. The model of evolution assumed was GTR ϩ ⌫ϩI. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 43

Fig. 17. Tree of amphibians provided by San Mauro et al. (2005). This tree re¯ects a maximum- likelihood analysis of 1,368 bp of the nuclear protein-coding gene RAG-1, assuming the GTR ϩ⌫ϩ I substitution model (as suggested by ModelTest v. 3.6; Posada and Crandall, 1998). Sequence alignment was made manually with only one gap excluded from analysis. ticulates with the braincase via a pseudobasal respect to salamanders; the other three char- process rather than a basal process. acters were likely polarized on the assump- Characters 4 (facial nerve exit) and 5 (pal- tion that Ascaphus is plesiomorphic and the atoquadrate articulation) are polarized with sister taxon of remaining frogs, thereby pre- 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 supposing the results, although this was not caudalipuboischiotibialis. In addition, stated. With respect to character 1 (the tri- Abourachid and Green (1999) noted that al- radiate sternum), the parsimony cost of this though Leiopelma and Ascaphus do hop, transformation on the overall tree is identical they swim with alternating sweeps of the if Ascaphidae and Leiopelmatidae are sister hind legs (the presumably plesiomorphic taxa and Bombinatoridae and Discoglossidae condition), unlike those in Bombianura, are sister taxa. The remaining characters, 2 which swim with coordinated thrusts of the and 3, were not discussed with respect to out- hind limbs, a likely synapomorphy. groups or reversals in the remainder of Ford Bombinatoridae was considered (Ford and and Cannatella's tree, implying that they are Cannatella, 1993) to have as synapomorphies unreversed and unique. (1) expanded ¯ange of the quadratojugal, and With Ascaphus, Leiopelma shares the apo- (2) presence of endochondral ossi®cations in morphy of columella not present (N.G. Ste- the hyoid plate (both unreversed). We sam- phenson, 1951). Haas (2003) did not include pled four species of Bombina: B. bombina, Leiopelma in his analysis of exotrophic lar- B. microdeladigitora, B. orientalis, and B. val morphology because of their endotrophy. variegata. The genus may be monophyletic, We included in our analysis Leiopelma ar- but no rigorous phylogenetic study has been cheyi and L. hochstetteri, which bracket the performed so far, and paraphyly of Bombina phylogenetic diversity of Leiopelmatidae with respect to remains an open (E.M. Stephenson et al., 1974), although it is question. We could not obtain tissues of Bar- not suf®cient to test hypotheses of the evo- bourula so its phylogenetic position will re- lution of direct development (exoviviparity main questionable. Bombina has aquatic in this case; Thibaudeau and Altig, 1999) feeding , but larvae of Barbourula within Leiopelma. are unknown and are suspected to be endo- DISCOGLOSSIDAE8 (2 GENERA,12SPECIES) trophic (Altig and McDiarmid, 1999). Dis- AND BOMBINATORIDAE (2 GENERA,10SPECIES): coglossidae (sensu stricto) also has free-liv- Ford and Cannatella (1993; ®g. 14) suggest- ing aquatic tadpoles (Boulenger, 1892 ed that Bombina ϩ Barbourula forms the sis- ``1891''; Altig and McDiarmid, 1999). ter taxon of all other frogs, exclusive of Leio- Ford and Cannatella (1993; ®g. 14) also pelmatidae and Ascaphidae, although recent posited a taxon, Discoglossanura, composed molecular evidence (Roelants and Bossuyt, of Discoglossidae (sensu stricto) and the re- 2005; ®g. 16) placed Bombinatoridae and maining frogs (exclusive of Ascaphidae, Discoglossidae in the familiar position of sis- Leiopelmatidae, and Bombinatoridae) which ter taxa. they suggested to be monophyletic on the ba- Ford and Cannatella's (1993) arrangement sis of two synapomorphies: (1) bicondylar (®g. 14; i.e., paraphyly of Bombinatoridae ϩ sacrococcygeal articulation; and (2) epister- Discoglossidae) required a partition of the num present. Monophyly of Discoglossidae traditionally recognized Discoglossidae (sen- (sensu stricto) was supported by their pos- su lato) to place Bombina and Barbourula in session of (1) V-shaped parahyoid bones their own family, Bombinatoridae. In their (also in Pelodytes) and (2) a narrow epipubic system, Bombinatoridae ϩ its sister taxon cartilage plate. (all frogs excluding Leiopelmatidae and As- Haas (2003; ®g. 15) presented a cladogram caphidae) was named Bombianura. Bombi- that is both deeply at variance with the re- anura is corroborated by four synapomor- lationships suggested by Ford and Cannatella phies: (1) fusion of the halves of the sphe- (1993) and, at least with respect to this part nethmoid; (2) reduction to eight presacral of their cladogram, consistent with the mo- vertebrae; (3) loss of the m. epipubicus (re- lecular evidence presented by Roelants and gained in Xenopus); and (4) loss of the m. Bossuyt (2005; ®g. 16). Haas (2003) pre- sented six morphological synapomorphies of Discoglossidae ϩ Bombinatoridae (as Dis- 8 SanchõÂz (1998) and Dubois (2005) noted that the name with priority for this taxon is Alytidae. However, coglossidae, sensu lato) and rejected Discog- to re¯ect the relevant literature we retain the name Dis- lossidae (sensu Ford and Cannatella) as par- coglossidae in this section. aphyletic, placing Alytes as the sister taxon 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 45 of the remaining members of Discoglossidae Ford and Cannatella (1993; ®g. 14) sug- ϩ Bombinatoridae. Synapomorphies of gested this group, Mesobatrachia, to be Haas' Discoglossidae are: (1) origin of m. monophyletic and composed of Pipoidea intermandibularis restricted to the medial (Pipidae ϩ Rhinophrynidae) and Pelobato- face of the cartilago meckelii; (2) larval m. idea (Pelobatidae [including Scaphiopodidae] levator mandibulae externus present as two ϩ Megophryidae ϩ Pelodytidae). They pro- bundles (profundus and super®cialis); (3) vided four synapomorphies for their Meso- posterior processes of pars alaris double; (4) batrachia: (1) closure of the frontoparietal cartilaginous roo®ng of the cavum cranii fontanelle by juxtaposition of the frontopa- present only as taenia traversalis; (5) verte- rietal bones (not in Pelodytes or ); (2) bral centra formation epichordal; and (6) pro- partial closure of the hyoglossal sinus by the cessus urobranchialis absent. Synapomor- ceratohyals; (3) absence of the taenia tecti phies suggested by Haas (2003; ®g. 15) for medialis; and (4) absence of the taenia tecti Discoglossidae, excluding Alytes are (1) epi- transversum. dermal melanocytes forming an orthogonal Pugener et al. (2003) rejected Mesobatra- pattern; (2) advertisement call inspiratory; chia and suggested three synapomorphies for and (3) pupil an inverted drop-shape (trian- a clade composed of all frogs excluding pi- gular). Of Discoglossidae (sensu stricto), we poids. (This statement is based on Pugener sampled one species of Alytes (A. obstetri- et al.'s, 2003, ®gure 12; they provided no cans) and two species of Discoglossus (D. comprehensive list of synapomorphies.) galganoi and D. pictus). Discoglossidae and Haas (2003; ®g. 15), in contrast, suggested Bombinatoridae show opisthocoelous and a number of characters that placed Pipoidea epichordal vertebrae according to Mookerjee as the sister taxon of all frogs except Asca- (1931), Grif®ths (1963), and Haas (2003). phidae (although he did not study Leiopel- Kluge and Farris (1969: 23) suggested that ma). This is consistent with the molecular vertebral development in studies of San Mauro et al. (2005; ®g. 17). is perichordal, although Haas (2003) reported Haas' characters also placed Pelobatoidea (as it as epichordal. represented by his exemplars) as a paraphy- Roelants and Bossuyt (2005; ®g. 16) and, letic series of Spea,(Pelodytes, Heleophry- with denser taxon sampling, San Mauro et ne), and Pelobates ϩ ϩ Lepto- al. (2005; ®g. 17) provided substantial brachium, ``between'' Discoglossidae (sensu amounts of DNA evidence suggesting lato) and Limnodynastes on a pectinate tree. strongly that Bombinatoridae ϩ Discoglos- This is inconsistent with the results of Roe- sidae forms a monophyletic group, thereby lants and Bossuyt (2005). Larval characters rejecting Discoglossanura, Leiopelmatanura, suggested by Haas (2003) to support the and Bombianura of Ford and Cannatella group of all frogs exclusive of Ascaphidae (1993). and Pipoidea are (1) m. mandibulolabialis present; (2) upper jaw cartilages powered by ``TRANSITIONAL'' FROGS jaw muscles; (3) larval m. levator mandibu- lae externus main portion inserts in upper The following few groups traditionally jaw cartilages; (4) insertion of the larval m. have been considered ``transitional'' from the levator mandibulae internus in relation to jaw primitive to advanced frogs, even though one articulation lateral; (5) m. levator mandibulae component taxon in particular, Pipidae, is longus super®cialis and profundus in two highly apomorphic in several ways. The bundles; (6) processus anterolateralis of cris- monophyly of this collection of families was ta parotica present; (7) processus muscularis supported by some authors (e.g., Ford and present; (8) distal end of cartilago meckeli Cannatella, 1993; GarcõÂa-ParõÂs et al., 2003), with stout dorsal and ventral processes form- but recent morphological (e.g., Haas, 2003; ing a shallow articular fossa; and (9) liga- Pugener et al., 2003) and DNA sequence ev- mentum mandibulosuprarostrale present. idence (Roelants and Bossuyt, 2005; San GarcõÂa-ParõÂs et al. (2004b; ®g. 18) pre- Mauro et al., 2005) does not support its sented mtDNA sequence evidence for the monophyly. monophyly of Mesobatrachia although their 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 18. Tree of Pelobatoidea and outgroups of GarcõÂa-ParõÂs et al. (2003) based on 1,000 bp of two mitochondrial genes: cytochrome c and 16S rRNA. The sequences were aligned using Clustal X (Thomp- son et al., 1997) using default costs then manually modi®ed based on published secondary-structure models of the 16S gene. Gaps were treated as missing data and data were analyzed under the assumption of the GTR ϩ⌫substitution model, as suggested by ModelTest 3.06 (Posada and Crandall, 1998). The tree was rooted on Ascaphus montanus ϩ A. truei. Quotation marks denote nonmonophyly. outgroup sampling (which was limited to As- changing places, with San Mauro et al.'s caphus truei, A. montanus, Discoglossus gal- (2005) placement of Pipoidea agreeing with ganoi, and Rana iberica) provided only a that of Haas (2003). minimal test of this proposition. Even more PIPOIDEA: Pipoidea (Pipidae ϩ Rhino- recently, on the basis of more DNA sequence phrynidae) is clearly well corroborated as evidence and better sampling, Roelants and monophyletic but not clearly resolved with Bossuyt (2005; ®g. 16) and San Mauro et al. respect to its rather dense fossil record. Ford (2005; ®g. 17) found ``Mesobatrachia'' to and Cannatella (1993; ®g. 14) considered Pi- have its elements in a paraphyletic series poidea to be supported by ®ve morphological with respect to Neobatrachia. Roelants and synapomorphies: (1) lack of mentomeckelian Bossuyt (2005) found (Ascaphidae ϩ Leio- bones; (2) absence of lateral alae of the par- pelmatidae) ϩ (Discoglossoidea ϩ (Pipoidea asphenoid; (3) fusion of the frontoparietals ϩ (Pelobatoidea ϩ Neobatrachia))) and San into an azygous element; (4) greatly enlarged Mauro et al. (2005) found Ascaphidae ϩ otic capsule; and (5) tadpole with paired spi- Leiopelmatidae as the sister taxon of Pipo- racles and lacking keratinized jaw sheaths idea ϩ (Discoglossoidea ϩ (Pelobatidae ϩ and keratodonts (Type I tadpole). Haas Neobatrachia)). In other words, their substan- (2003) added a substantial number of larval tial difference is in Discoglossoidea (ϭ Bom- characters: (1) eye position lateral; (2) oper- binatoridae ϩ Discoglossidae) and Pipoidea cular canal and spiracles paired; (3) insertion 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 47 of m. levator arcuum branchialium reaching phy of Rhinophrynidae at this level of gen- medially and extending on proximal parts of erality. ceratobranchial IV; (4) m. constrictor bran- PIPIDAE (5 GENERA,30SPECIES): South chialis I absent; (5) m. levator mandibulae American and African Pipidae is a highly internus shifted anteriorly; (6) m. levator apomorphic group of bizarre, highly aquatic mandibulae longus originates exclusively species. Ford and Cannatella (1993) provided from arcus subocularis; (7) posterolateral 11 characters in support of its monophyly: projections of the crista parotica with expan- (1) lack of a quadratojugal; (2) presence of sive ¯at chondri®cations; (8) arcus subocu- an epipubic cartilage; (3) unpaired epipubic laris with a distinct processus lateralis pos- muscle; (4) free ribs in larvae; (5) fused ar- terior projecting laterally from the posterior ticulation between the coccyx and the sa- palatoquadrate; (9) articulation of cartilago crum; (6) short, stocky scapula; (7) elongate labialis superior with cornu trabeculae fused septomaxillary bones; (8) ossi®ed pubis; (9) into rostral plate; and (10) forelimb erupts a single median palatal opening of the eus- out of limb pouch, outside of peribranchial tachian tube; (10) organs in the space. In addition, recent DNA sequence adults; and (11) loss of tongue. BaÂez and data (Roelants and Bossuyt, 2005; ®g. 16) Trueb (1997) added to this list (fossil taxa strongly support a monophyletic group of pruned by us for purposes of this discussion): Rhinophrynidae ϩ Pipidae. (1) the possession of an optic foramen with RHINOPHRYNIDAE (1 GENUS,1SPECIES): a complete bony margin formed by the sphe- Tropical North American and Central Amer- nethmoid; (2) anterior ramus of the pterygoid ican Rhinophrynus dorsalis is a burrowing arises near the anteromedial corner of the frog with a number of apomorphies with re- otic capsule; (3) parasphenoid fused at least spect to its nearest living relative, Pipidae: partially with the overlying braincase; (4) vo- (1) of the distal condyle of the femur mer without an anterior process if the bone into lateral and medial condyles; (2) modi- is present; (5) mandible bears a broad-based, bladelike coronoid process along its postero- ®cation of the prehallux and distal phalanx medial margin; (6) sternal end of the cora- of the ®rst digit into a spade for digging; (3) coid not widely expanded; (7) anterior ramus tibiale and ®bulare short and stocky, with of pterygoid dorsal with respect to the max- distal ends fused; and (4) an elongate atlantal illa; and (8) premaxillary alary processes ex- neural arch. In addition to the previous char- panded dorsolaterally. Haas (2003) provided acters provided by Ford and Cannatella 11 additional larval characters: (1) origin of (1993; ®g. 14), Haas (2003; ®g. 15) provided the m. subarcualis rectus II±IV placed far lat- (1) larval m. geniohyoideus absent; (2) larval erally; (2) anterior insertion of m. subarcualis m. levator mandibulae externus present in rectus II±IV on ceratohyal III; (3) commis- two bundles (profundus and super®cialis); surae craniobranchiales present; (4) arcus su- (3) ramus mandibularis (cranial nerve V3) bocularis round in cross section; (5) one peri- posterior (ventral) to m. levator mandibulae lymphatic foramen; (6) vertebral centra for- externus group; (4) endolymphatic spaces mation epichordal; (7) processus urobran- extend into more than half of the vertebral chialis absent; and (8) ventral valvular velum canal (presacral vertebrae 4 or beyond); (5) absent, as well as these additional characters branchial food traps divided crescentrically; of the adult: (9) advertisement call without (6) cricoid ring with dorsal gap; and (7) uro- air¯ow; (10) pupil shape round; and (11) branchial process very long. Available DNA pectoral girdle pseudo®rmisternal. sequence data (e.g., Roelants and Bossuyt, On the basis of morphology, Cannatella 2005) also suggest strongly that Rhinophry- and Trueb (1988; ®g. 19A) considered the nus is the sister taxon of Pipidae. We sam- generic relationships to be Xenopus ϩ (Sil- pled the single species in this taxon, Rhino- urana ϩ ((Hymenochirus ϩ Pseudhymeno- phrynus dorsalis.BaÂez and Trueb (1997) not- chirus) ϩ )). Subsequently, de Sa and ed that Rhinophrynus also has amphicoelous Hillis (1990; ®g. 19B), on the basis of a com- ectochordal vertebrae, as in Ascaphidae and bined analysis of morphology and mtDNA, Leiopelmatidae, which may be a synapomor- proposed the arrangement Hymenochirus 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

postzygopophyses bear sulci and ridges, with the prezygopophyses covering the lateral margin of the postzygopophysis; and (4) an- terior process of the pterygoid laminae. They also suggested the following synapomorphies for Pipinae (Pipa ϩ Hymenochirus) (fossil taxa pruned for purposes of this discussion): (1) wedge-shaped skull; (2) vertebrae with parasagittal spinous processes; (3) anterior position of the posterior margin of the par- asphenoid; (4) possession of short coracoids broadly expanded at their sternal ends. In ad- dition, they noted other characters of more ambiguous placement that optimize on this stem in this topology. Recent DNA sequence Fig. 19. Trees of intergeneric relationships data (Roelants and Bossuyt, 2005; ®gs. 16, within Pipidae: A, Analysis of Cannatella and 19D), however, suggest a topology of Pipa Trueb (1988) based on 94 character transforma- ϩ (Hymenochirus ϩ (Xenopus ϩ )). tions of morphology and 7 ingroup taxa (4 species We sampled three species of Dactylethri- of Pipa collapsed and Pseudhymenochirus consid- nae (Africa): Silurana tropicalis, Xenopus ered a synonym of Hymenochirus in our ®gure for laevis, and X. gilli. From Pipinae (South clarity of discussion). Monophyly of Pipidae was America and Africa) we sampled Hymeno- assumed as well as the sister-taxon relationship of chirus boettgeri, Pipa pipa, and P. carvalhoi. Rhinophrynidae, with pelobatoids accepted as the second taxonomic outgroup (no tree statistics pro- According to the cladogram provided by vided). B, Analysis of de Sa and Hillis (1990) Trueb and Cannatella (1986), inclusion of ei- based on 1.486kb of sequence from nuclear 18S ther Pipa parva or P. myersi would have and 28S rDNA and the morphological data from bracketed the phylogenetic diversity of Pipa Cannatella and Trueb (1988). Sequences were somewhat better, although our sampling was aligned manually and analyzed under equally adequate to test pipine (weakly), dactyleth- weighted parsimony; gaps were not treated as ev- rine, and pipid monophyly, and the place- idence. The tree was rooted on Spea (tree length ment of Pipidae among other frogs. counting only informative characters ϭ 81, ci ϭ 0.74). C, Parsimony tree of BaÂez and Pugener PELOBATOIDEA: Pelobatoidea (Megophryi- (2003) based on 49 characters of adult morphol- dae, Pelobatidae, Pelodytidae, and Scaphio- ogy, outgroups and fossils pruned for graphic pur- podidae) has also been the source of consid- poses (the effect of this pruning on the number of erable controversy. Haas (2003; ®g. 15) did characters being relevant is not known). The tree not recover the group as monophyletic (see was rooted on Rhinophrynus, Discoglossus, and the earlier discussion under Mesobatrachia), Ascaphus. (The length of original tree ϭ 93, ci ϭ although Ford and Cannatella (1993) sug- 0.677.) D, Relevant section of tree from Roelants gested that synapomorphies include the pres- and Bossuyt (2005). See ®gure 16 for information ence of a palatine process of the and on alignment and analysis. ossi®cation of the sternum into a bony style. Gao and Wang (2001) found Pelobatoidea to (Xenopus ϩ Silurana), and this was further be more closely related to Discoglossidae on corroborated by BaÂez and Trueb (1997) and the basis of a limited analysis of fossil taxa. BaÂez and Pugener (2003; who found [Hy- But, GarcõÂa-ParõÂs et al. (2003; ®g. 18) sug- menochirus ϩ Pipa] ϩ [Xenopus ϩ Silur- gested that Pelobatoidea is the sister taxon of ana]; ®g. 19C), and suggested the following Pipoidea on the basis of a maximum-likeli- synapomorphies for Dactylethrinae (Xenopus hood analysis of mtDNA evidence, although ϩ Silurana; fossil taxa pruned for this dis- their outgroup structure was insuf®cient to cussion): (1) scapula extremely reduced; (2) provide a strong test of this proposition. margins of olfactory foramina cartilaginous; (This position was effectively rejected by re- (3) articular surfaces of the vertebral pre- and cent molecular evidence [Roelants and Bos- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 49 suyt, 2005; San Mauro et al., 2005; ®gs. 16, calcaneum (also found in some Centroleni- 17].) dae; SanchõÂz and de la Riva, 1993) and Maglia (1998) also provided an analysis of placed them in their Pelobatoidea as did Pelobatoidea, but because she constrained GarcõÂa-ParõÂs et al. (2003; ®g. 18). Haas the monophyly of this group we are not sure (2003), however, recovered Pelodytes in a how to interpret the distribution of her mor- polytomy with , Neobatrachia phological evidence. Pugener et al. (2003) and Megophrys ϩ Pelobates ϩ Leptobrach- provided a cladogram based on morphology ium. We sampled Pelodytes punctatus as our in which Pelobatoidea was recovered as exemplar of Pelodytidae. Larvae in pelody- monophyletic (and imbedded within Neoba- tids are also typical free-living exotrophs trachia), but the underlying data were not (Altig and McDiarmid, 1999). provided. Roelants and Bossuyt (2005; ®g. MEGOPHRYIDAE (11 GENERA, 129 SPECIES): 16) suggested on the basis of DNA evidence Ford and Cannatella (1993; ®g. 14) diag- that Pelobatoidea is the sister taxon of Neo- nosed Megophryidae as having (1) a com- batrachia, a result that is consistent with the plete or nearly complete absence of cerato- older view of Savage (1973; cf. Noble, hyals in adults; (2) intervertebral cartilages 1931). Dubois (2005) most recently treated with an ossi®ed center; and (3) paddle- all pelobatoids as a single family composed shaped tongue. Haas (2003; ®g. 15) recov- of four subfamilies, but this was merely a ered a group consisting of the megophryids change in Linnaean rank without a concom- (Leptobrachium and Megophrys being his itant change in understanding phylogenetic exemplars) and Pelobates but did not resolve history. the megophryids sensu stricto. Evidence for PELOBATIDAE (1 GENUS,4SPECIES) AND this megophryid ϩ Pelobates clade is: (1) SCAPHIOPODIDAE (2 GENERA,7SPECIES): Ford distal anterior labial ridge and keratodont- and Cannatella (1993; ®g. 14) diagnosed Pe- bearing row very short and median; (2) vena lobatidae (including Scaphiopodidae in their caudalis dorsalis present; (3) anterior inser- sense) on the basis of (1) fusion of the joint tion of the m. subarcualis rectus II±IV on between the sacrum and urostyle; (2) exos- ceratobranchial III; (4) m. mandibulolabialis tosed frontoparietals; and (3) presence of a superior present; (5) adrostral cartilage very metatarsal spade supported by a well-ossi®ed large and elongate; and (6) cricoid ring with prehallux. As noted earlier, Haas (2003; ®g. a dorsal gap. 15) did not recover Pelobatidae (sensu lato) Dubois (1980) and Dubois and Ohler as monophyletic, instead placing Spea phy- (1998) suggested that megophryids form two logenetically far from Pelobatidae, more dis- subfamilies based on whether the larvae have tant than Heleophryne. More recently, funnel-shaped oral discs (Megophryinae), an GarcõÂa-ParõÂs et al. (2003; ®g. 18) provided apomorphy, or nonmodi®ed oral discs (Lep- molecular data suggesting that Pelobatidae tobrachiinae), a plesiomorphy. Megophryi- and Scaphiopodidae are not each other's nae includes Atympanophrys, Brachytarso- closest relatives. These results were aug- phrys, Megophrys, , and Xen- mented by the DNA sequence studies of ophrys. Their Leptobrachiinae includes Lep- Roelants and Bossuyt (2005) and San Mauro tobrachella, , Leptobrachium, et al. (2005), both of which supported Sca- , , and Vibrissaphora. De- phiopodidae as the sister taxon of Pelodyti- lorme and Dubois (2001) presented a con- dae ϩ (Pelobatidae ϩ Megophryidae) (®gs. sensus tree (®g. 20) based on 54 transfor- 16, 17). All species have typical exotrophic mation series of morphology (not including aquatic larvae (Altig and McDiarmid, 1999). Vibrissaphora). This tree suggests that Me- We sampled , gophryinae ( being their couchii, and S. holbrooki from Scaphiopod- exemplar) is deeply imbedded within a par- idae, and and P. cultripes aphyletic Leptobrachiinae (the remaining from Pelobatidae. megophryid exemplars being of this nominal PELODYTIDAE (1 GENUS,3SPECIES): Ford subfamily); that Scutiger is composed of a and Cannatella (1993; ®g. 14) diagnosed Pe- paraphyletic subgenus Scutiger and a mono- lodytidae as having a fused astragalus and phyletic subgenus Aelurophryne), and that 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 20. Consensus of 12 equally parsimonious trees of selected members of Megophryidae of Delorme and Dubois (2001), rooted on Scaphiopus and Pelodytes. Underlying data were 54 transfor- mation of morphology, rooted on Pelodytes and Scaphiopus (ci ϭ 0.581; ri ϭ 0.713). Although the tree and a list of the underlying character transformation were provided, no association was made between the character transformations and taxa or particular branches on the tree, rendering the analysis practi- cally unrepeatable. Nominal subfamilies are noted on the right.

Oreolalax is composed of a paraphyletic sub- were feae, Leptobrachium genus Oreolalax and a monotypic subgenus chapaense, L. hasselti, Leptolalax bourreti, Aelurolalax. Megophrys nasuta, Ophryophryne hansi, O. Within megophryines, Xie and Wang microstoma, major (formerly X. (2000) noted con¯ict between isozyme and lateralis). We were unable to obtain samples karyological data regarding the monophyly of Atympanophrys, , Oreola- of Brachytarsophrys, and also noted that lax, Scutiger, and Vibrissaphora, so, al- Atympanophrys is only dubiously diagnos- though we are con®dent that our sampling able from Megophrys or Xenophrys. They will allow phylogenetic generalizations to be also suggested that Xenophrys may not be made regarding the family, most of the prob- diagnosable from Megophrys. lems within the group (e.g., the questionable Lathrop (1997) suggested that, among monophyly of Leptobrachium, Leptolalax, nominal leptobrachiines, Leptolalax has no Megophrys, Scutiger, and Xenophrys) will identi®ed apomorphies. Xie and Wang remain unanswered. (2000) noted that Oreolalax is diagnosable from Scutiger on the basis of unique maxil- ``ADVANCED'' FROGSÐNEOBATRACHIA lary teeth and that Vibrissaphora has apo- Neobatrachia9 includes about 96% of ex- morphies (e.g., keratinized spines along the tant frogs and is a poorly understood array lips of adults), although the effect of recog- of apparently likely paraphyletic groups with nizing Oreolalax and Vibrissaphora on the apomorphic satellites. So, at this juncture in monophyly of Scutiger has not been evalu- our discussion the quantity of evidence sug- ated. Similarly, the monophyly of Lepto- brachium is undocumented. 9 There is controversy regarding the appropriate name The species sampled for DNA sequences of this taxon. It is addressed in appendix 6. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 51

Fig. 21. Bayesian tree of anuran exemplars of Biju and Bossuyt (2003), with particular reference to Neobatrachia. Underlying data are two mtDNA fragments, covering part of 12S rRNA, complete t- RNAVal, and part of 16S rRNA. In addition, one fragment of the nuclear genome: exon 1 of rhodopsin, single exon of RAG-1, and exon 2 of CXCR-4, for a total of 2,325 bp of sequence. Alignment was made using Clustal X (Thompson et al., 1997), alignment costs not disclosed, with ambiguous sections excluded and gaps excluded as evidence. Model of nucleotide substitution assumed for analysis was GTR ϩ⌫ϩI. gested by authors to support major groups, prephylogenetic systematics, Neobatrachia and the quality of published taxonomic rea- also has within it its own nominally ``prim- soning drops signi®cantly to the realm of itive'' groups aggregated on plesiomorphy grouping by overall similarity and special (e.g., Leptodactylidae), as well as its own pleading for particularly favored characters. nominally ``transitional'' and ``advanced'' Like the larger-scale Archeobatrachia (prim- groups (e.g., Ranidae and Rhacophoridae, itive frogs), Mesobatrachia (transitional Arthroleptidae and Hyperoliidae). Further, frogs), and Neobatrachia (advanced frogs) of the unwillingness of the systematics com- 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 munity to change taxonomies in the face of lossidae, and, presumably Rheobatrachidae evidence is best illustrated here. For exam- as well. Darst and Cannatella (2004; ®g. 22) ple, Brachycephalidae was shown to be im- redelimited Hyloidea as the descendants of bedded within the leptodactylid taxon the most recent common ancestor of Eleuth- Eleutherodactylinae, but the synonymy was erodactylini, Bufonidae, Centrolenidae, Hy- not made by Darst and Canntella (2004), and linae, Phyllomedusinae, Pelodryadinae, and Leptopelinae was shown to be more closely Ceratophryinae, thereby excluding Heleo- related to Astylosternidae than to hyperoliine phrynidae, Limnodynastidae, Myobatrachi- hyperoliids by Vences et al. (2003c), but was dae, Rheobatrachidae, and Sooglossidae (and retained by those authors in an explicitly par- by implication, presumably Nasikabatrachi- aphyletic Hyperoliidae. dae) from Hyloidea10. For this discussion, we Ford and Cannatella (1993; ®g. 14) sug- retain the older, more familiar de®nition of gested ®ve characters in support of the Hyloidea as all neobatrachians excluding the monophyly of Neobatrachia: (1) (neo)palatine ranoids. bone present; (2) fusion of the third distal HELEOPHRYNIDAE (1 GENUS,6SPECIES): carpal to other carpals; (3) complete separa- South African Heleophryne was considered tion of the m. sartorius from the m. semiten- by Ford and Cannatella (1993; ®g. 14) to be dinosus; (4) presence of an accessory head a member of Neobatrachia and Hyloidea. of the m. adductor longus; and (5) absence The synapomorphy of Heleophrynidae sug- of the parahyoid bone. In addition, Haas gested by these authors includes only ab- (2003; ®g. 15) presented the following larval sence of keratinous jaw sheaths in exotrophic characters (but see Heleophrynidae): (1) up- free-living larvae. Haas (2003; ®g. 15), in per lip papillation with broad diastema; (2) contrast, placed Heleophrynidae outside cartilage of the cavum cranii forms tectum Neobatrachia in a pectinate relationship parietale; (3) secretory ridges present; and among ``pelobatoids'' or as the sister taxon (4) pupil horizontally elliptical. The charac- ter of central importance historically to the 10 Because Darst and Cannatella (2004) used Limno- recognition of this taxon is the (neo)palatine dynastes and Heleophryne as outgroups to root the re- mainder of the tree, it was not possible for them to have bone, a character not without its own contro- obtained a tree in which traditional Hyloidea is mono- versy. phyletic so their statement (p. 46) that ``the placement ``HYLOIDEA'': The worldwide Hyloidea, of some basal neobatrachian clades (Heleophrynidae, for which no morphological synapomorphy Myobatrachidae, and Sooglossidae) remains uncertain'' is actually an assumption of their phylogenetic analysis. has been suggested, was long aggregated on Uncited by Darst and Cannatella (2004), Biju and Bos- the basis of its being ``primitive'' with re- suyt (2003) differentiated between ``Hyloidea'' sensu spect to the ``more advanced'' Ranoidea, al- lato (the traditional view of Hyloidea) and Hyloidea sen- though molecular evidence under certain an- su stricto, which they considered to be monophyletic and alytical methods and assumptions supports which, like the concept of Darst and Cannatella (2004), excluded Myobatrachidae, Limnodynastidae, Heleo- its monophyly (Ruvinsky and Maxson, 1996; phrynidae, Sooglossidae, and Nasikabatrachidae. Anoth- Feller and Hedges, 1998). Hyloidea is de- er issue is that Ford and Cannatella (1993) and Canna- ®ned by the plesiomorphic (at least within tella and Hillis (2004) de®ned the name Hylidae to apply Neobatrachia) possession of arciferal pecto- cladographically to the hypothetical ancestor of Hemi- phractinae, Hylinae, Pseudinae (now part of Hylinae), ral girdles (coracoids not fused) and simple and Pelodryadinae, and all of its descendants. However, procoelous vertebrae, although descriptions Darst and Cannatella (2004) implied that their Hylidae of both characters have been highly rei®ed was rede®ned to exclude Hemiphractinae. This rede®- through repetition and idealization. More re- nition would be necessary to keep content and diagnosis cently, Biju and Bossuyt (2001: ®g. 21) sug- as stable as possible with respect to the traditional use of the term ``Hylidae'', because without this kind of re- gested on the basis of a DNA sequence anal- de®nition in a system that aspires to precision, the pre- ysis that Hyloidea, as traditionally viewed, is tense of precision is lost. For example, the cladographic paraphyletic with respect to Ranoidea, but de®nition of Hylidae by Ford and Cannatella (1993) and within ``Hyloidea'' is a monophyletic group Cannatella and Hillis (2004) applied to the cladogram of Darst and Cannatella (2004) would require that the largely coextensive with ``Hyloidea'', but ex- following be included within Hylidae: Brachycephali- cluding Heleophrynidae, Limnodynastidae, dae, Leptodactylidae, Bufonidae, Centrolenidae, Den- Myobatrachidae, Nasikabatrachidae, Soog- drobatidae, and, likely, Rhinodermatidae. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 53 of Pelobates, Leptobrachium, and Mego- We sampled one species each of the nom- phrys. Heleophryne is included at this level inal sooglossid genera (Nesomantis thomas- in Haas' analysis by having (1) m. tympan- seti and Sooglossus sechellensis). Of Nasi- opharyngeus present; (2) m. interhyoideus kabatrachidae we sampled posterior present; (3) m. diaphragmatoprae- sahyadrensis as well as sequences attributed cordialis present; (4) m. constrictor bran- by Dutta et al. (2004) only to an unnamed chialis I present; and (5) interbranchial sep- species of Nasikabatachidae, also from the tum IV musculature with the lateral ®bers of . Although Dutta et al. did not the m. subarcualis rectus II±IV invading the name their species as new, they explicitly septum, and lacking the characters listed by treated it as distinct from N. sahyadrensis Haas for Neobatrachia. In addition, the ver- (Dutta et al., 2004: 214), and we therefore tical pupil and ectochordal vertebrae tie he- follow their usage. (Our statement that Na- leophrynids to myobatrachines, and non-neo- sikabatrachidae contains two species rests on batrachians. Recent DNA sequence evidence this assertion, although any clear diagnosis (San Mauro et al., 2005; ®g. 17) strongly of the second has yet to be cogently provid- supports Heleophrynidae as the sister taxon ed.) All species of Sooglossidae that are of all other neobatrachians (although Biju known are endotrophic according to Thibau- and Bossuyt, 2003, also on the basis of mo- deau and Altig (1999). Sooglossus sechellen- lecular evidence as well had suggested that sis has free tadpoles that are carried on the Heleophrynidae is the sister taxon of Lim- back of the mother. The tadpoles are likely nodynastidae ϩ Myobatrachidae). endotrophic, but this is not de®nitely known We sampled Heleophryne purcelli and H. (R.A. Nussbaum, personal obs.). Dutta et al. regis. These species are likely close relatives (2004) reported exotrophic tadpoles occur- (Boycott, 1982) so broader sampling (to have ring in fast-¯owing streams for their un- included H. rosei, whose isolation on Table named species of Nasikabatrachidae. Mountain near Cape Town suggests a likely LIMNODYNASTIDAE (8 GENERA,50SPECIES), distant relationship to the other species) MYOBATRACHIDAE (11 GENERA,71SPECIES), would have been preferable. AND RHEOBATRACHIDAE (1 GENUS,2SPECIES): SOOGLOSSIDAE (2 GENERA,4SPECIES) AND Different authors consider this taxonomic NASIKABATRACHIDAE (1 GENUS,2SPECIES): cluster to be one family (Myobatrachidae, Sooglossidae is a putative Gondwanan relict sensu lato) with two or three subfamilies (Savage, 1973) on the Seychelles, possibly (Heyer and Liem, 1976); to be two families, related to myobatrachids as evidenced by Limnodynastidae and Myobatrachidae (Zug sharing with that taxon the plesiomorphy of et al., 2001; Davies, 2003a, 2003b); or to be ectochordal vertebrae (J.D. Lynch, 1973), al- three families, Limnodynastidae, Myoba- though Bogart and Tandy (1981) suggested a trachidae, and Rheobatrachidae (Laurent, relationship with the arthroleptines (a ranoid 1986). Because Rheobatrachidae (Rheoba- group). In fact, the group is plesiomorphic in trachus; Laurent, 1986) was only tentatively many characters, being arciferal (although associated with Myobatrachidae by Ford and having a bony sternum; see Kaplan, 2004, Cannatella (1993), we retain its familial sta- for discussion of the various meanings of tus for clarity of discussion. ``arcifery'') and all statements as to its rela- Limnodynastidae, Myobatrachidae, and tionships, based on morphology, have been Rheobatrachidae are primarily united on the highly conjectural. Biju and Bossuyt (2003; basis of their geographic propinquity on Aus- ®g. 21) suggested on the basis of DNA se- tralia and New (Tyler, 1979; Ford quence evidence that Sooglossidae is the sis- and Cannatella, 1993). And, only one line of ter taxon of the recently discovered Nasika- evidence, that of spermatozoal morphology, batrachus, found in the Western Ghats of has ever suggested that these taxa taken to- South India. Nasikabatrachus has so far had gether are monophyletic (Kwon and Lee, little of its morphology documented. They 1995). Heyer and Liem (1976) provided a also found Sooglossidae ϩ Nasikabatrachi- character analysis that assumed familial and dae to form the sister taxon of all other neo- generic monophyly, but this was criticized batrachians. methodologically (Farris et al., 1982a). 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 55

Rheobatrachinae (Rheobatrachus)isofun- horizontal ones); (4) broad alary process certain position, although Farris et al. (Grif®ths, 1959a), which they found in (1982a) in their reanalysis of Heyer and Myobatrachidae and Rheobatrachus (as well Liem's (1976) data, considered it to be part as in Adenomera, [in the sense of Limnodynastinae. Ford and Cannatella of including and Eupemphix], (1993) subsequently argued that Rheobatra- and ); and (5) divided chinae is more closely related to Myoba- sphenethmoid. trachidae than to Limnodynastidae, although Ford and Cannatella (1993) reported at this suggestion, like the ®rst, rests on highly least one synapomorphy for Limnodynasti- contingent phylogenetic evidence. Moreover, dae: a connection between the m. interman- Myobatrachidae may be related to Sooglos- dibularis and m. submentalis (also found in sidae (J.D. Lynch, 1973) and Limnodynasti- Leptodactylinae and Eleutherodactylinae ac- nae to Heleophrynidae (J.D. Lynch, 1973; cording to Burton, 1998b). Rheobatrachus Ruvinsky and Maxson, 1996), although these was diagnosed by having gastric brooding of views are largely conjectural inasmuch as the larvaeÐan unusual reproductive mode, to character evidence of J.D. Lynch (1973) was say the least. It is tragic that the two species presented in scenario form. are likely now extinct (Couper, 1992). Ford and Cannatella (1993) suggested, on Read et al. (2001) provided a phylogenetic the basis of discussion of characters present- study of myobatrachine frogs (®g. 23) based ed by Heyer and Liem (1976), that Myoba- on mtDNA sequence data that assumed trachidae (Myobatrachinae in their sense and monophyly of the group and used only Lim- presumably including Rheobatrachus) has nodynastes to root the myobatrachine tree. four morphological synapomorphies: (1) The evolutionary propinquity of Limnodyn- presence of notochordal (ectochordal) verte- astes (Limnodynastidae) and brae with intervertebral discs; (2) m. petro- (Myobatrachidae) was supported on the basis hyoideus anterior inserting on the ventral of DNA sequence evidence by Biju and Bos- face of the hyoid; and, possibly, (3) reduction suyt (2003). of the vomers and concomitant reduction of We were able to sample at least one spe- vomerine teeth (J.D. Lynch, 1971). cies for most of the genera of the three nom- Ford and Cannatella (1993) suggested sev- inal families. For Limnodynastidae we sam- eral synapomorphies of Myobatrachidae and pled at least one species for all nominal gen- Sooglossidae to the exclusion of Limnodyn- era: Adelotus brevis, australia- astidae: (1) incomplete cricoid cartilage ring; cus, ¯etcheri, Limnodynastes (2) semitendinosus tendon inserting dorsal to depressus, L. dumerilii, L. lignarius, L. or- the m. gracilis (in myobatrachines excluding natus, L. peronii, L. salmini, Mixophyes car- and Rheobatrachus, which have binensis, sudelli, N. pictus, a ventral trajectory of the tendon; (3) hori- melanoscaphus, sphagni- zontal pupil (except in ; (also ver- cola. Recent authors (e.g., Cogger et al., tical in Rheobatrachus; limnodynastines 1983) have considered Kyarranus to be a primitively have a vertical pupil according to synonym of Philoria, and we follow this. Heyer and Liem, 1976, although several have J.D. Lynch (1971) provided morphological

← Fig. 22. Parsimony tree of Darst and Cannatella (2004) of hyloid frogs and outgroups based on analysis of 12S and 16S fragments of mitochondrial rRNA gene sequences. Sequence alignment was performed using Clustal X (Thompson et al., 1997) under a number of different cost regimes (not disclosed) and then compared with secondary structures and manually manipulated to minimize the number of informative sites under a parsimony criterion. Unalignable regions were excluded and gaps were treated as missing. The number of informative sites was not stated. The tree was rooted on Limnodynastes ϩ Heleophryne. We updated the taxonomy of the terminals and higher taxonomy to correspond with changes made after the paper was published. Use of Euhyas (instead of Eleutherodac- tylus) is our modi®cation to illuminate discussion. 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

characters that are evidence of monophyly of Kyarranus ϩ Philoria (e.g., presence of stub- by ®ngers and concealed tympana as well as direct developmentÐLittlejohn, 1963; De Bavay, 1993; Thibaudeau and Altig, 1999). For Rheobatrachidae, we obtained Rheo- batrachus silus. And, for Myobatrachidae, we obtained at least single representatives of all nominal genera: rotunda, Assa darlingtoni, nimbus, C. signi- fera, victoriana, ni- chollsi, Myobatrachus gouldii, Paracrinia haswelli, bibroni, P. coriacea, ¯ammocaerulea, Taudactylus acutirostris, and Uperoleia laevigata. With exceptions, this taxon selection will not al- low us to comment on generic monophyly, but it will identify major monophyletic groups and questions that will guide future research. All rheobatrachids and most myob- atrachids have endotrophic larvae and vari- ous degrees of direct development (Thibau- deau and Altig, 1999). ``LEPTODACTYLIDAE'' (57 GENERA, 1243 SPECIES): ``Leptodactylidae'' holds the same position in the Americas as Myobatrachidae (sensu lato, as containing Limnodynastidae and Rheobatrachidae) does in AustraliaÐa likely nonmonophyletic hodgepodge ``prim- itive'' holochordal or rarely stegochordal, ar- ciferal, and procoelous neobatrachian group united by geography and not synapomorphy. ``Leptodactylidae'' is currently divided into ®ve subfamilies, some of which are not clearly monophyletic (or consistently diag- nosable) and some of which may be poly- Fig. 23. Parsimony tree of Crinia, Geocrinia, phyletic (Ruvinsky and Maxson, 1996; Haas, and allied myobatrachids, of Read et al. (2001). 2003; Darst and Cannatella, 2004; Faivovich Data were of mtDNA: approximately 621 bp (266 et al., 2005; San Mauro et al., 2005; ®gs. 17, variable) from the 12S rRNA region and 677 bp (383 variable) of ND2. Sequence alignment of 22, 24). 12S and ND2 were done under ClustalX (Thomp- J.D. Lynch (1971, 1973) considered lep- son et al., 1997) with gap opening and extension todactylids to be divided into four subfami- costs set at 50, and transversion: transition cost lies, on the basis of both synapomorphy and ratio set at 2. Ambiguously alignable regions were symplesiomorphy: (1) Ceratophryinae (for excluded. In analysis, transversion:transition costs and ); (2) Elo- were set at 2. It was not stated whether gaps were siinae (ϭ Hylodinae of other authors; for treated as evidence but we infer that gaps were , , and ); treated as missing data. Branches marked with an (3) Leptodactylinae (for , Edalor- asterisk were collapsed in the original publication hina, , Leptodactylus [including because of low bootstrap support. Adenomera], Limnomedusa, Lithodytes, Par- atelmatobius, Physalaemus [including En- gystomops and Eupemphix], , and Pseudopaludicola); and (4) Telmatobi- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 57 inae, aggregated on the basis of plesiomor- closely related to at least some component of phy. Within his Telmatobiinae Lynch de®ned Limnodynastidae (Lechriodus) than to other ®ve tribes, each aggregated on a variable ba- South American ``leptodactylids''. Another sis of synapomorphy and symplesiomorphy: leptodactylid satellite is Brachycephalidae, a Telmatobiini (Batrachophrynus, Caudiver- small monophyletic taxon, likely the sister bera, Telmatobius, and ); Also- taxon of (Leptodactylidae: dini (, [including Al- Eleutherodactylinae) based on digit reduction sodes], Hylorina, and ); Odonto- (Izecksohn, 1988; Giaretta and Sawaya, phrynini (, Odontophry- 1998). Similarly, Rhinodermatidae (Rhinod- nus, and Proceratophrys); Grypiscini erma) is a small group that is likely also a (, , and Za- telmatobiine leptodactylid (Barrio and Rin- chaenus); and Eleutherodactylini (Eleuther- aldi de Chieri, 1971; Lavilla and Cei, 2001), odactylus, Euparkerella, , and Is- differing from them in having partial or com- chnocnema, as well as several other genera plete larval development within the male vo- subsequently placed in the synonymy of cal sac and, except for Eupsophus, in having Eleutherodactylus), with being endotrophic larvae (Formas et al., 1975; Al- left incertae sedis. Subsequently, Heyer tig and McDiarmid, 1999). (1975) provided a preliminary clustering Laurent (1986) provided the subfamilial (based on the nonphylogenetic monothetic taxonomy we employ for discussion (his ar- subset method of Sharrock and Felsenstein, rangement being the formalization of the 1975) of the nominal genera within the fam- groupings tentatively recommended by Hey- ily that assumed monophyly of both the fam- er, 1975). He recognized Ceratophryinae (in ily and the constituent genera (see Farris et the larger sense of including J.D. Lynch's al., 1982a, for criticism of the approach) in Odontophrynini, transferred from Telmato- which Heyer identi®ed, but did not recognize biinae), Telmatobiinae (including calyptoce- formally, ®ve units that were recognized sub- phallelines and excluding J.D. Lynch's sequently (Laurent, 1986) as Ceratophryinae, Eleutherodactylini), Cycloramphinae (as Eleutherodactylinae, Cycloramphinae, Lep- Grypiscinae, including Grypscini and Elosi- todactylinae, and Telmatobiinae. J.D. Lynch inae of J.D. Lynch), Eleutherodactylinae, and (1978b) revised the genera of Telmatobiinae, Leptodactylinae. where he recognized three tribes: Telmato- ``CERATOPHRYINAE'' (6 GENERA,41SPE- biini (, , Batrachophry- CIES): Reig (1972) and Estes and Reig (1973) nus, Eupsophus, Hylorina, , suggested that the leptodactylid subfamily Limnomedusa, Somuncuria, and Telmato- Ceratophryinae was ``ancestral'', in some bius), Calyptocephalellini (Caudiverbera and sense, to Bufonidae, although others rejected Telmatobufo), and Batrachylini (Batrachyla this (e.g., J.D. Lynch, 1971, 1973). Laurent and Thoropa). The justi®cation for this ar- (1986), following Heyer (1975), transferred rangement was partially based on character Macrogenioglottus, Odontophrynus, and argumentation, although plausibility of re- Proceratophrys (J.D. Lynch's tribe Odonto- sults was based on subjective notions of phrynini) into this nominal subfamily, with overall similarity and relative character im- Ceratophrys, , and Lepidobatra- portance. A cursory glance at ®gure 24 (Fai- chus being placed in Ceratophryini. Haas vovich et al., 2005) shows that several of (2003; ®g. 15) presented morphological evi- these groups are nonmonophyletic. dence that Ceratophryini and Odontophry- Burton (1998a) suggested on the basis of nini are not each other's closest relatives (fol- hand muscles (although his character polarity lowing J.D. Lynch, 1971), with Odontophry- was not well supported) that the leptodactyl- nus most closely related to Leptodactylus, id tribe Calyptocephalellini is more closely and the clade Ceratophryini (Lepidobatra- related to the South African Heleophrynidae chus ϩ Ceratophrys) most closely related to than to other South American leptodactylids. hylids, exluding hemiphractines. Duellman San Mauro et al. (2005; ®g. 17) suggested on (2003) treated the two groups as subfamilies, the basis of DNA sequence data that Cau- Odontophryninae and Ceratophryinae, pre- diverbera (Calyptocephalellini) is more sumably following the results of Haas 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 59

(2003), and this was followed by Dubois to be close to Eleutherodactylini on the basis (2005). Faivovich et al. (2005; ®g. 24) also of overall similarity.) Grypiscines and hylo- found Ceratophryinae to be polyphyletic. We dines differ in (1) the shape of the transverse sampled exemplars from all nominal cerato- processes of the posterior presacral vertebrae, phryid genera except Macrogenioglottus, being short in hylodines and not short in gry- which is similar to Odontophrynus (J.D. piscines; (2) the shape of the facial lobe of Lynch, 1971) and karyologically similar to the maxillae (deep in grypiscines, shallow in Proceratophrys (Silva et al., 2003; Odonto- hylodines); (3) the shape of the nasals (large phrynus not examined in that study) that we and in median contact in grypiscines, small doubt that this will be an important problem. and widely separated in hylodines); and (4) Ceratophryini does have synapomorphies, whether the nasal contacts the frontoparietal for example: (1) transverse processes of an- (contact in grypiscines, no contact in hylo- terior presacral vertebrae widely expanded; dines). We were unable to obtain samples of (2) cranial bones dermosed; and (3) teeth Crossodactylodes, ,orZachaenus, fanglike, nonpedicellate (J.D. Lynch, 1971, but we did obtain at least one species of ev- 1982b), although nominal Odontophrynini ery other nominal genus in the group: Cros- does not have unambiguously synapomor- sodactylus schmidti, Cycloramphus bora- phies, and the group is united on overall sim- ceiensis, , Megaelosia ilarity. All ceratophryids have free-living ex- goeldii, sp., Scythrophrys otrophic larvae (Altig and McDiarmid, sawayae, and . Denser sam- 1999). We sampled three species of Cerato- pling of this particular taxon would have phryini (Ceratophrys cranwelli, Chacophrys been preferable, but what we obtained will pierotti, and ) and test cycloramphine monophyly and its puta- three species of Odontophrynini (Odonto- tive relationship to Dendrobatidae and will phrynus achalensis, O. americanus, and Pro- provide an explicit hypothesis of its internal ceratophrys avelinoi). Our sampling of Pro- phylogenetic structure as the basis of future ceratophrys should have been denser, but this studies. proved a practical impossibility. Duellman (2003) did not accept Laurent's ``CYCLORAMPHINAE'' (10 GENERA,79SPE- (1986) uni®cation of J.D. Lynch's Hylodinae CIES): Haas (2003) suggested that this group and Grypiscini and recognized Hylodinae may be closely related to Dendrobatidae, in (Crossodactylus, Hylodes, and Megaelosia) part supporting the earlier position of Noble as a different subfamily from Cycloramphi- (1926) and J.D. Lynch (1973) that the hylo- nae. Duellman distinguished Hylodinae and dine part of this nominal subfamily (Cros- Cycloramphinae by T-shaped terminal pha- sodactylus, Hylodes, and Megaelosia) is par- langes in Hylodinae and knoblike terminal aphyletic with respect to Dendrobatidae. Fai- phalanges in Cycloramphinae; and glandular vovich et al. (2005; ®g. 24) recovered Cros- pads on the dorsal surface of the digits, ab- sodactylus (their exemplar of this group) as sent in Hylodinae and present in Cycloram- the sister taxon of Dendrobatidae. Laurent phinae. However, neither the particulars of (1986) recognized this subfamily, thus uni- distribution of these characters in the taxa fying J.D. Lynch's (1971, 1973) Grypiscini nor the levels of universality of their appli- and Elosiinae (ϭ Hylodinae), although the cation as evidence was discussed. Duellman evidentiary basis for uniting these was based (2003) also suggested that Hylodinae and on Heyer's (1975) results based on monoth- Cycloramphinae differ in chromosome num- etic subsets, not parsimony. (Note that J.D. bers, with 13 pairs in Cycloramphinae and 3 Lynch, 1971, had considered his Grypiscini pairs in Hylodinae. However, Kuramoto

← Fig. 24. Tree of Hylidae and outgroups from Faivovich et al. (2005), based on 5.5kb sequence from four mitochondrial genes (12S, 16S, tRNAVal, cytochrome c) and ®ve nuclear genes (rhodopsin, tyrosi- nase, RAG-1, seven in absentia, 28S) and analyzed by Direct Optimization in POY under equal cost functions. Gaps were treated as evidence. 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(1990) noted that hylodines in Duellman's that work in the near future can rectify this sense have 11±13 pairs of chromosomes, and with the recognition of major monophyletic cycloramphines in Duellman's sense also groups from within Eleutherodactylus. What have 11±13 pairs, so Duellman's statement is we could sample of the non-Eleutherodac- taken to be an error. tylus eleutherodactyline taxa were Barycho- ELEUTHERODACTYLINAE (13 GENERA, 782 los ternetzi, quixensis, and two SPECIES): The only suggested synapomorphy species of Phrynopus.OfEleutherodactylus of this taxon is direct terrestrial development (sensu lato) we sampled two species of the of large deposited in small clutches North American subgenus Syrrhophus (J.D. Lynch, 1971). The universality of direct (Eleutherodactylus marnocki of the E. mar- development in this group is based on ex- nocki group of J.D. Lynch and Duellman, trapolation from the few species for which 1997, and E. nitidus of the E. nitidus group direct development has been observed; the of J.D. Lynch and Duellman, 1997); one spe- occurrence of large, unpigmented eggs, and cies of the Antillean subgenus Euhyas because free-living larvae are unknown (see (Eleutherodactylus planirostris of the E. ri- cautionary remarks in Thibaudeau and Altig, cordii group of J.D. Lynch and Duellman, 1999). Inasmuch as this taxon contains the 1997); two species of the South American largest vertebrate genus, Eleutherodactylus subgenus Eleutherodactylus (E. binotatus (ca. 600 species) of which the vast majority and E. juipoca, both of the E. binotatus are not represented by genetic samples, this group of J.D. Lynch, 1978a; see also J.D. taxon will remain inadequately sampled for Lynch and Duellman, 1997); and six species some time. There has never been any com- of the Middle American subgenus Craugas- prehensive phylogenetic study of the rela- tor12 (E. bufoniformis of the E. bufoniformis tionships within the group and the likelihood group of J.D. Lynch, 2000, E. alfredi of the of many (or even most) of the non-Eleuth- E. alfredi group of J.D. Lynch, 2000, E. au- erodactylus genera being components of gusti of the E. augusti group of J.D. Lynch, Eleutherodactylus is high. Indeed, Ardila- 2000, E. pluvicanorus of the E. fraudator Robayo (1979) suggested strongly that for group of KoÈhler, 2000, E. punctariolus and the taxon currently referred to as Eleuthero- E. cf. ranoides13 of the E. rugulosus group dactylus (sensu lato) to be rendered mono- of J.D. Lynch, 2000) and E. rhodopis of the phyletic it would need to include Barycholos, E. rhodopis group of J.D. Lynch, 2000). (For Geobatrachus, Ischnocnema, and Phrynopus expediency, all of these are noted in ``Re- (and likely , Phyllonastes, Phy- sults'' in combination with their subgeneric zelaphryne, Holoaden and Euparkerella, and names; e.g., Eleutherodactylus (Syrrhophus) Brachycephalidae [Izecksohn, 1971; Giaretta marnockii is treated as Syrrhophus marnock- and Sawaya, 1998; Darst and Cannatella, ii.) As noted earlier, we expect that Eleuth- 2004; Faivovich et al., 2005]11). Regardless, erodactylus will be found to be paraphyletic many of the nominal eleutherodactyline gen- with respect to a number of other eleuther- era represent rare and extremely dif®cult an- odactyline taxa (e.g., Barycholos, Phrynopus, imals to obtain (e.g., , Dischi- and Ischnocnema) and hope that this selec- dodactylus), so our sampling of this partic- tion will allow some illumination of this. ular taxon is clearly inadequate to address most systematic problems. We could not ob- 12 Craugastor was recently considered to be a genus tain samples of Adelophryne, Atopophrynus, by Crawford and Smith (2005) and we follow that ar- , Euparkerella (even though rangement, although we refer to Craugastor in this sec- it was suggested to be closely related to Bra- tion and elsewhere as part of Eleutherodactylus (sensu lato) for consistency with the immediately relevant lit- chycephalidae), Geobatrachus, Holoaden, erature. Phyllonastes,orPhyzelaphryne. We hope 13 We report this species as Craugastor cf. ranoides, because we discovered late in this project that the vouch- 11 Dubois (2005) noted that if Brachycephalidae is a er specimen was lost. However, the identi®cation in the synonym of Eleutherodactylinae, as suggested by the re- associated ®eld notes was ``Eleutherodactylus rugulo- sults of Darst and Cannatella (2004), the appropriate sus'' and the only member of the rugulosus group (and name for this taxon, within Leptodactylidae, would be of Craugastor) otherwise in that collection and from that Brachycephalinae. region is Craugastor ranoides. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 61

Nevertheless, we are aware that this tiny dense enough to evaluate well the likely par- fraction of the species diversity of Eleuth- aphyly of this taxon with respect to others, erodactylus is insuf®cient to fully resolve the such as Adenomera (Heyer, 1998), being re- phylogeny of this massive taxon and that the stricted to only two of the ®ve nominal spe- value of the results will be in highlighting cies groups. Leptodactylines vary from hav- outstanding problems and providing a basis ing endotrophic larvae, facultatively endotro- for future, more densely sampled studies. phic larvae (Adenomera) to having exotroph- LEPTODACTYLINAE (12 GENERA, 159 SPE- ic, free-living larvae (, CIES): Monophyly of this group is supported Engystomops, Eupemphix, Leptodactylus, by the possession of foam nests (except in Lithodytes, Physalaemus, Pleurodema, Pseu- Limnomedusa [Langone, 1994] and Pseudo- dopaludicola, Vanzolinius; Altig and Mc- paludicola [Barrio, 1954], and in some spe- Diarmid, 1999). cies of Pleurodema [Duellman and Veloso ``TELMATOBIINAE'' (11 GENERA,98SPE- M., 1977]) and the presence of a bony ster- CIES): Telmatobiinae is a similarity grouping num (rather than the cartilaginous sternum of of mostly austral South American frogs. As other leptodactylids; J.D. Lynch, 1971). currently employed, contents of this subfam- However, Haas (2003; ®g. 15) sampled three ily stem from Laurent's (1986) formalization species of Leptodactylinae (Physalaemus bi- of Heyer's (1975) informal grouping. Tel- ligonigerus, , and matobiines are currently arranged in three ) for mostly larval mor- tribes (J.D. Lynch, 1971, 1978b; Burton, phology and found the group to be para- or 1998a): Telmatobiini (Alsodes, Atelognathus, polyphyletic with respect to Odontophrynus, Eupsophus, Hylorina, Insuetophrynus, So- and with Physalaemus14 and Pleurodema muncuria, and Telmatobius); Batrachylini forming, respectively, more exclusive out- (Batrachyla and Thoropa); and Calyptoce- groups of Haas' hylodines and dendrobatids. phalellini (Batrachophrynus, Caudiverbera, In Darst and Cannnatella's (2004) phyloge- and Telmatobufo). All telmatobiines have netic analysis of mtDNA (®g. 22), their lep- aquatic, exotrophic larvae except Eupsophus, todactyline exemplars are monophyletic in which has endotrophic larvae (Altig and the maximum-likelihood analysis of mtDNA, McDiarmid, 1999), and Thoropa, which is but polyphyletic in the parsimony analysis. semiterrestrial (Bokermann, 1965; Wassersug In Faivovich et al.'s (2005; ®g. 24) parsi- and Heyer, 1983; Haddad and Prado, 2005). mony analysis of multiple mtDNA and Batrachylini (in J.D. Lynch's sense of in- nuDNA loci, exemplars of most genera of cluding Thoropa) is diagnosed by having ter- Leptodactylinae obtained as monophyletic, restrial eggs and aquatic to semiterrestrial with the exception of Limnomedusa. There- larvae and T-shaped terminal phalanges. fore, the monophyly of Leptodactylinae is an Laurent (1986) did not (apparently) accept open question. We could not obtain samples J.D. Lynch's (1978b) transferral of Thoropa of Hydrolaetare (or the recently resurrected into Batrachylini, and retained Thoropa in Eupemphix and Engystomops), but we sam- Cycloramphinae following Heyer (1975). pled at least one species of each of the other Calyptocephalellini was most recently dis- nominal leptodactyline genera: Adenomera cussed and diagnosed by Burton (1998a) on hylaedactyla, , Leptodac- the basis of hand musculature, but the char- tylus fuscus, L. ocellatus, Limnomedusa ma- acter argumentation was essentially that of croglossa, Lithodytes lineatus, Physalaemus overall similarity, not synapomorphy. For- gracilis, Pleurodema brachyops, Pseudopa- mas and Espinoza (1975) provided karyolog- ludicola falcipes, and Vanzolinius discodac- ical evidence for the monophyly of Calyp- tylus). Our sampling of Leptodactylus is not tocephallelini (although they did not address Batrachophrynus). Cei (1970) suggested on 14 Nascimento et al. (2005) recently partitioned Phys- the basis of immunology that Calyptoce- alaemus into Physalaemus, Eupemphix, and Engysto- phalellini is phylogenetically distant from mops on the basis of phenetic comparisons. Unfortu- nately, the historical reality of these taxa will remain leptodactylids, being closer to Heleophryni- arguable until a phylogenetic analysis is performed on dae than to any South American leptodactyl- this group. id group. J.D. Lynch (1978b) suggested the 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 following to be synapomorphies of Calyp- the basis of mtDNA evidence that Hemi- tocephalellini (composed of solely Caudiv- phractinae is polyphyletic, with Cryptobatra- erbera and Telmatobufo): (1) occipital artery chus closest to direct-developing eleuthero- enclosed in a bony canal; and (2) very broad dactylines, and Gastrotheca imbedded in an- pterygoid process of the premaxilla. In ad- other group of leptodactylids. Similarly, in dition, (1) a very long cultriform process of the analysis by Faivovich et al. (2005; ®g. the parasphenoid; and (2) presence of a me- 24) of multiple mtDNA and nuDNA loci, dial process on the pars palatina of the pre- hemiphractines do not appear as monophy- maxilla are osteological characters suggested letic. They recovered one clade composed of by J.D. Lynch possibly to unite Batracho- Gastrotheca and Flectonotus, one clade com- phrynus with Caudiverbera and Telmatobu- posed of Stefania and Cryptobatrachus, and fo. they found to form a clade We sampled representatives of two of the with the few included exemplars of Eleuth- genera of Calyptocephallelini (Caudiverbera erodactylinae and Brachycephalidae. Further, caudiverbera and ). We inasmuch as the sole noncontingent synapo- could not sample Batrachophrynus, which morphy of nominal Hemiphractinae, bell- was considered a calyptocephalelline by shaped larval gills, has not been surveyed Burton (1998a), and in some of the clado- widely in direct-developing leptodactylids, grams presented by J.D. Lynch (1978b) Ba- we consider the morphological evidence for trachyophrynus was considered to form the the monophyly of the hemiphractines to be sister taxon of his Calyptocephalellini, so its questionable. absence from our analysis is unfortunate. Faivovich et al. (2005; ®g. 24) transferred ``Telmatobiini'' of J.D. Lynch (1978b) is ``Hemiphractinae'' out of a reformulated Hy- explicitly paraphyletic with respect to Batra- lidae and into ``Leptodactylidae'' on the ba- chylini and as such has no diagnosis other ses that continued inclusion in Hylidae than that of the inclusive clade ``Telmatobi- would render Hylidae polyphyletic; its nom- ini'' ϩ Batrachylini: (1) presence of an outer inal inclusion in ``Leptodactylidae'' did no metatarsal tubercle (dubiously synapomorph- violence to a taxon already united solely by ic), and (2) reduction of imbrication on the plesiomorphy and geography; and placing it neural arches of the vertebrae. Among spe- incertae sedis within Hyloidea was to suggest cies of ``Telmatobiini'' we sampled Alsodes its possible placement outside of the ``lep- gargola, , Eupso- todactylid'' region of the overall tree, which phus calcaratus, , Telma- it is not. ``Hemiphractinae'' is a grouping of tobius jahuira, T. cf. simonsi, and T. sp. Of South American frogs united by (1) brooding Batrachylini, we sampled Batrachyla lepto- of eggs on the female's back, generally with- pus. On this basis we provide a weak test of in a dorsal depression or well-developed telmatobiine relationships with regard to Ba- pouch; (2) possession in the developing lar- trachylini. We were unable to sample any vae of bell-shaped gills (Noble, 1927); and member of Insuetophrynus or Somuncuria. (3) presence of a broad m. abductor brevis ``HEMIPHRACTINAE'' (5 GENERA,84SPE- plantae hallucis (Burton, 2004). Larvae may CIES): Mendelson et al. (2000) provided a be exotrophic and endotrophic among spe- cladogram of Hemiphractinae but assumed cies of Gastrotheca and Flectonotus, and en- its monophyly and its hylid af®nities, as had dotrophic alone in Cryptobatrachus, Hemi- all authors since Duellman and Gray (1983) phractus, and Stefania. Based on Faivovich and Duellman and Hoogmoed (1984). Haas et al.'s (2005) topology (®g. 24), - (2003) suggested (®g. 15), on the basis of shaped terminal phalanges and presence of morphological data, that his examplar of intercalary cartilages between the ultimate Hemiphractinae, Gastrotheca, was far from and penultimate phalanges must be consid- other hylids and imbedded within a heter- ered either convergent with those found in eogeneous group of leptodactylids and ran- Hylidae or plesiomorphically retained in hy- oids. Darst and Cannatella (2004), who ex- lids (and lost in intervening lineages), while amined one exemplar species each of Gas- the proximal head of metacarpal II not be- trotheca and Cryptobatrachus, suggested on tween prepollex and distal prepollex, and the 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 63 larval spiracle sinistral and ventrolateral acters that are possible synapomorphies. Two (Duellman, 2001) are convergent with those species are currently recognized, Rhinoder- in the Phyllomedusinae. Our sampling of ma darwinii and R. rufum. We sampled R. Gastrotheca is not dense enough to allow for darwinii. the detection of the paraphyly suggested by DENDROBATIDAE (CA.11GENERA, 241 SPE- Mendelson et al. (2000). Our sampling pre- CIES): The monophyly of Dendrobatidae has cludes evaluation of paraphyly of any of the been upheld consistently (e.g., Myers and nominal genera. Nevertheless, we did sample Ford, 1986; Ford, 1993; Haas, 1995; Clough at least one species per genus, which allows and Summers, 2000; Vences et al., 2000b), us to test the monophyly of the hemiphrac- but different datasets place Dendrobatidae at tines based on more extensive outgroup sam- various extremes within the neobatrachian pling. Our sampled taxa are Cryptobatrachus clade. It is either nested deeply within hy- sp., Flectonotus sp., Gastrotheca ®ssipes, G. loids and arguably related to cycloramphine cf. marsupiata, Hemiphractus helioi, and leptodactylids (Noble, 1926, 1931; J.D. . Lynch, 1971, 1973; Burton, 1998a; Haas, BRACHYCEPHALIDAE (1 GENUS,8SPECIES): 2003; Faivovich et al., 2005); the sister group This tiny group of diminutive south- to of Telmatobius (Vences et al., 2003b); or southeastern Brazilian species are united by closely related to Hylinae (Darst and Can- (1) the absence through fusion of a distin- natella, 2004). Alternatively, they have been guishable sternum; (2) digital reduction (pos- suggested to be deeply imbedded within ran- sibly homologous with that in Euparkerella oids, usually considered close to arthrolep- and Phyllonastes in Eleutherodactylinae); tids or petropedetids (Grif®ths, 1959b, 1963; and (3) complete ossi®cation of the epicor- Duellman and Trueb, 1986; Ford, 1993; Ford acoid cartilages with coracoids and and Cannatella, 1993; Grant et al., 1997). (Ford and Cannatella, 1993; Kaplan, 2002). Rigorous evaluation of the support for these Brachycephalidae was suggested to be im- contradictory hypotheses is required. bedded within Eleutherodacylinae (Izeck- Ford and Cannatella (1993; ®g. 14) pro- sohn, 1971; Giaretta and Sawaya, 1998), vided the following as synapomorphies of which also shows direct development. Fur- Dendrobatidae: (1) retroarticular process pre- ther, Darst and Cannatella (2004; ®g. 22) sent on the mandible; (2) conformation of the provided molecular data to link this taxon to super®cial slip of the m. depressor mandib- Eleutherodactylinae, but continued its rec- ulae; and (3) cephalic amplexus. They men- ognition despite the demonstrable paraphyly tioned other features suggested by other au- that its recognition requires. Although there thors but that were suspect for one reason or are several named and unnamed species in another. Haas (2003; ®g. 15) considered the the genus, the monophyly of the group is not following to be synapomorphies that nest in question (Kaplan, 2002), and we sampled Dendrobatidae within hylodine leptodactyl- the type species, Brachycephalus ephippium, ids: (1) guiding behavior observed during for this study. courtship; and (2) T- or Y-shaped terminal RHINODERMATIDAE (1 GENUS,2SPECIES): As phalanges. Like most other frogs, most den- noted earlier, the Chilean Rhinodermatidae is drobatids have aquatic free-living tadpoles a likely satellite of a paraphyletic ``Lepto- (with some endotrophy in ), al- dactylidae''; it is like them in having pro- though the parental-care behavior of carrying coelous and holochordal vertebrae. J.D. tadpoles to on the back of one of the Lynch (1973) conjectured that Rhinoderma- parents appears to be synapomorphic (Altig tidae is the sister taxon of Bufonidae, where- and McDiarmid, 1999), although among as Lavilla and Cei (2001) suggested that Rhi- New World anurans it also occurs in Cyclor- noderma is within the poorly-de®ned ``Tel- amphus stejnegeri (Heyer and Crombie, matobiinae'' (``Leptodactylidae''). The only 1979). notable synapomorphy of Rhinodermatidae Taxon sampling was designed to provide is the rearing of tadpoles within the vocal the maximal ``spread'' of phylogenetic vari- sacs of the male, although Manzano and Lav- ation with a minimum number of species: Al- illa (1995) also discussed myological char- lobates femoralis, boulengeri, Co- 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 lostethus undulatus, auratus, nuDNA loci recovered an Allophryne ϩ Cen- trinitatis, Minyobates clau- trolenidae clade nested within a grade of diae, Phobobates silverstonei, and Phyllo- ``Leptodactylidae''. Clearly, the diversity of bates lugubris. We did not sample any rep- opinions on the placement of Centrolenidae resentative of , Cryptophylloba- is great. For our analysis we selected species tes, Nephelobates, nor did we sample either of the three nominal genera: gec- of two generally-not-recognized genera Oop- koideum, C. prosoblepon, be- haga or . On the basis of ongo- jaranoi, and ¯eischman- ing work by T. Grant, we think that all of ni. Larvae of centrolenids are aquatic or bur- these are imbedded within our sampled gen- rowing exotrophs (Altig and McDiarmid, era and their absence does not hamper our 1999). ability to test dendrobatid monophyly and HYLIDAE (48 GENERA, 806 SPECIES): Hyli- place the family in the larger phylogenetic dae, as traditionally recognized, was recently scheme. shown to be polyphyletic (Ruvinsky and ALLOPHRYNIDAE (1 GENUS,1SPECIES): Maxson, 1996; Haas, 2003; Darst and Can- South American Allophryne has been (1) natella, 2004; Faivovich et al., 2005). As an very provisionally associated with Hylidae interim measure to resolve this problem Fai- (J.D. Lynch and Freeman, 1966); (2) asserted vovich et al. (2005) transferred ``Hemiphrac- to be in Bufonidae on the basis of morphol- tinae'' into ``Leptodactylidae'', thereby ogy (Laurent, 1980 ``1979''; Dubois, 1983; restricting Hylidae to the Holarctic and Neo- Laurent, 1986), the evidence for this latter tropical Hylinae, tropical American Phyllo- position not actually presented until much medusinae, and Australo-Papuan Pelodrya- later by Fabrezi and Langone (2000); (3) im- dinae (and thereby formalizing the implica- bedded within Centrolenidae, on the basis of tion of Darst and Cannatella, 2004). morphology (Noble, 1931); or (4) placed as Our notions of hylid relationships extend the sister taxon of Centrolenidae on the basis from the recent revision by Faivovich et al. of mtDNA sequence studies (Austin et al., (2005; ®g. 24), who provided a phylogenetic 2002; Faivovich et al., 2005). Cognoscenti of analysis of multiple mtDNA and nuDNA frogs will marvel at the vastness separating loci. Their study addressed 220 hylid exem- these various hypotheses. Ford and Canna- plar terminals as well as 48 outgroup taxa. tella (1993) noted that Allophryne lacks the For our study, we considered including all interacalary cartilages of hylids and centro- terminals from the Faivovich et al. (2005) lenids and suggested that placement in any study, which would have allowed a more rig- taxon other than Neobatrachia is misleading. orous test, but the increased computational Haas (2003; ®g. 15) did not examine Allo- burden was judged too great for the expected phryne. We sampled the single species, Al- payoff of increased precision within Hylinae. lophryne ruthveni. Larvae are unknown (Al- Our sampling strategy aimed to be suf®cient- tig and McDiarmid, 1999). ly dense to test the position of hylids among CENTROLENIDAE (3 GENERA, 139 SPECIES): other frogs and the monophyly of the major Centrolenidae has long been thought to be clades without unduly exacerbating compu- close to, or the sister taxon of, Hylidae (J.D. tational problems. Lynch, 1973; Ford and Cannatella, 1993; HYLINAE (38 GENERA, 586 SPECIES): Our Duellman, 2001) because of the occurrence sampling structure of Hylinae was guided by of interacalary cartilages between the ulti- the results of Faivovich et al. (2005). Beyond mate and penultimate phalanges. On the ba- their genetic evidence, monophyly of this sis of mostly-larval morphology, Haas subfamily is corrobated by at least one mor- (2003) recovered (weakly) Centrolenidae as phological synapomorphy: tendo super®ci- the sister taxon of all Neobatrachia except for alis digiti V (manus) with an additional ten- Limnodynastes (Limnodynastidae), because don that arises ventrally from the m. palmaris it lacked all characters that Haas' analysis longus (Da Silva In Duellman, 2001). All hy- suggested were synapomorphies of Neoba- lines for which it is known have free-living trachia. The analysis of Faivovich et al. exotrophic larvae (Altig and McDiarmid, (2005; ®g. 24) of multiple mtDNA and 1999). Faivovich et al. (2005) recognized 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 65 four monophyletic tribes within Hylinae: Co- of Pelodryadinae remains unsettled. Haas phomantini (Aplastodiscus, Bokermannohy- (2003), on the basis of six exemplars, recov- la, , Hypsiboas, and Myersiohy- ered the subfamily as paraphyletic with re- la); Hylini (Acris, Anotheca, , spect to hylines and phyllomedusines. Tyler , , Ecnomiohy- (1971c) noted the presence of supplementary la, , , , Megasto- elements of the m. intermandibularis in both matohyla, Pseudacris, Plectrohyla, Ptychoh- Pelodryadinae (apical) and Phyllomedusinae yla, Smilisca [including former Pternohyla], (posterolateral). These characters were inter- , and ); Dendropsophini preted by Duellman (2001) as nonhomolo- (, , , Scar- gous and therefore independent apomorphies thyla, , , and Xeno- of their respective groups. If these conditions hyla); and Lophiohylini (, are homologues as suggested by Faivovich et , , , al. (2005) on the basis of their preferred clad- , Osteocephalus, , ogram, the polarity between the two charac- , , and Trachycephal- ters is ambiguous because either the pelod- us). ryadine or the phyllomedusinae condition In this study we included representatives might be ancestral for Phyllomedusinae ϩ of these four tribes: Cophomantini (Aplas- Pelodryadinae. Because our study aims to todiscus perviridis, Hyloscirtus armatus, H. provide a general phylogenetic structure for palmeri, Hypsiboas albomarginatus, H. amphibians, not to resolve all systematic boans, H. cinerascens (formerly Hypsiboas problems, and in light of ongoing research granosus; see Barrio-AmoroÂs, 2004: 13), H. by S. Donnellan, we have not sampled ``Li- multifasciatus); Dendropsophini (Dendrop- toria'' densely enough to provide a detailed sophus marmoratus, D. minutus, D. nanus, resolution of relationships within Pelodry- D. parviceps, , Pseudis par- adinae. Nevertheless, we sampled densely adoxa, goinorum, , S. enough to provide additional evidence re- ruber, ); Hylini garding the paraphyly of ``Litoria'' with re- (Acris crepitans, Anotheca spinosa, Char- spect to Cyclorana or Nyctimystes. Species adrahyla nephila, Duellmanohyla ru®oculis, sampled in this group are Cyclorana aus- miliaria, Exerodonta chimala- tralis,``Litoria'' aurea,``L.'' freycineti,``L.'' pa, Hyla arbrorea, H. cinerea, Isthmohyla ri- genimaculata,``L.'' inermis,``L.'' lesueurii, vularis, Pseudacris crucifer, P. ocularis, Pty- ``L.'' meiriana,``L.'' nannotis, Nyctimystes chohyla leonhardschultzei, Smilisca phaeota, dayi, and N. pulcher. All pelodryadines ap- Tlalocohyla picta, and ); pear to have free-living exotrophic larvae and Lophiohylini (Argenteohyla siemersi, (Tyler, 1985; Altig and McDiarmid, 1999). Osteocephalus taurinus, Osteopilus septen- PHYLLOMEDUSINAE (7 GENERA,52SPECIES): trionalis, Phyllodytes luteolus, Trachyce- There is abundant morphological and molec- phalus jordani, and T. venulosus). ular evidence for the monophyly and phylo- PELODRYADINAE (3 GENERA, 168 SPECIES): genetic structure of this subfamily of bizarre Knowledge of phylogenetic relationships frogs. Synapomorphies of the group include: among Australo-Papuan hylids is poorly un- (1) vertical pupil; (2) ventrolateral position derstood, beyond the pervasive paraphyly of of the spiracle; (3) arcus subocularis of larval nominal ``Litoria'' with respect to the other chondrocranium with distinct lateral process- two genera, Nyctimystes and Cyclorana (Ty- es; (4) ultralow suspensorium; (5) secondary ler and Davies, 1978; King et al., 1979; Ty- fenestrae parietales; and (6) absence of a pas- ler, 1979; Maxson et al., 1985; Hutchinson sage between the ceratohyal and the cerato- and Maxson, 1987; Haas and Richards, 1998; branchial I (Haas, 2003). Faivovich et al. Haas, 2003; Faivovich et al., 2005). Faivov- (2005) discussed several other characters ich (2005) noted one morphological syna- likely to be synapomorphies of Phyllome- pomorphy of Phyllomedusinae ϩ Pelodry- dusinae. Faivovich et al. (2005) demonstrat- adinae, the presence of a tendon of the m. ed on the basis of molecular data that Cru- ¯exor ossis metatarsi II arising only from ziohyla is the sister taxon of the remaining distal tarsi 2±3. Evidence for the monophyly genera, which are further divided in two 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 67 clades, one containing and Phyl- also noted the possibility that the possession lomedusa, and the other containing the re- of inguinal fat bodies and having a xiphi- maining genera (, , sternum free from the underlying m. rectus Pachymedusa, and ). Our tax- abdominis are synapomorphies of Bufonidae, on sampling re¯ects this understanding: Aga- or some subtaxon of that group. lychnis callidryas, calcarifer, Dubois (1983, 1987 ``1985'') recognized , and vail- ®ve nominal subfamilies, not predicated on lanti. any phylogenetic hypothesis or, seemingly, BUFONIDAE (35 GENERA, 485 SPECIES): Bu- any concern for monophyly (Graybeal and fonidae is a worldwide hyloid clade of non- Cannatella, 1995; Graybeal, 1997). controversial monophyly, although the 35 Graybeal and Cannatella (1995) provided genera for the most part are weakly diag- a discussion of the monophyly of most of the nosed (e.g., Andinophryne, Bufo, Crepido- genera within Bufonidae that is extremely phryne, , and Rhamphophryne). useful. They noted that many bufonid genera Ford and Canntella (1993) suggested the fol- are monotypic and therefore not eligible for lowing synapomorphies for Bufonidae: (1) tests of monophyly: Altiphrynoides Dubois, presence of a Bidder's organ (although ab- 1987 ``1986''; Atelophryniscus McCranie, sent in [Echeverria, 1998] Wilson, and Williams, 1989; Pillai and Truebella [Graybeal and Cannatella, and Yazdani, 1973; Crepidophryne Cope, 1995]); (2) unique pattern of insertion of the 1889; Didynamipus Andersson, 1903, Fros- m. hyoglossus; (3) absence of the m. con- tius Cannatella, 1986; Laurentophryne Tih- strictor posterior (Trewevas, 1933); (4) teeth en, 1960; Mertensophryne Tihen, 1960; Me- absent (also in some basal telmatobiines, Al- taphryniscus SenÄaris, AyarzaguÈena, and Gor- lophryne, some dendrobatids, and some rha- zula, 1994; Tschudi, 1838; cophorids); (5) origin of the m. depressor Schismaderma Smith, 1849; and Spinophry- mandibulae solely from the squamosal and noides Dubois, 1987 ``1986''. associated angle or orientation of the squa- Graybeal and Cannatella (1995) noted that mosal (Grif®ths, 1954; also in several other many genera lack evidence of monophyly: anuransÐsee Manzano et al., 2003); (6) Cope, 1861 ``1860''; Andinophry- presence of an ``otic element'', an indepen- ne Hoogmoed, 1985; Bufo Laurenti, 1768; dent ossi®cation in the temporal region that Noble, 1926; Pedostibes fuses to the otic ramus of the squamosal GuÈnther, 1876 ``1875''; Pelophryne Barbour, (Grif®ths, 1954; also known in two genera 1938; Fitzinger, 1843; Rham- of Ceratophryini, Ceratophrys and Chaco- phophryne Trueb, 1971; Stephopaedes Chan- phrys, but unknown in LepidobatrachusÐ ning, 1979 ``1978''; and Wolterstorf®na Wild, 1997, 1999). Ford and Cannatella Mertens, 1939. Graybeal and Cannatella (1993) considered characters 2±6 to be in- (1995) noted the following genera to show suf®ciently surveyed but likely synapo- evidence of monophyly: Stoliczka, morphic. Da Silva and Mendelson (1999) 1870; DumeÂril and Bibron, 1841;

← Fig. 25. A, Consensus tree of Bufonidae from Graybeal (1997). The tree re¯ects a parsimony anal- ysis of DNA sequence data. Sequences used were primarily of mtDNA gene regions 12S and 16S (total of 1672 bp, aligned, for 50 species), with the addition of the protein coding mtDNA gene cytochrome b (390 bp for 19 species) and the nuDNA protein-coding gene c-mos (280 bp for 7 species). The protein- coding genes were aligned manually according to the amino-acid sequence, while the rDNA sequences were performed manually with reference to assumed secondary structure, with gaps excluded as evi- dence. Length of the component trees is 3,862 steps, ci ϭ 0.305, ri ϭ 0.392. Cunningham and Cherry (2004: 681) noted that her 16S DNA sequences of Bufo garmani (U52746) are Bufo gutturalis; her Bufo vertebralis (U52730) sequences are of B. maculatus, and her B. maculatus sequences are likely not of B. maculatus, but of another species, unidenti®ed by them. B, Consensus tree of Bufonidae from Gray- beal (1997) based on DNA sequence data (from A) and morphological data (undisclosed). 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Capensibufo Grandison, 1980; Dendrophryn- taxon.) She also suggested that Peltophryne iscus JimeÂnez de la Espada, 1871 ``1870''; (the Bufo peltocephalus group) is far from Fitzinger, 1843; Melanophryn- other New World bufonids, that Bufo gar- iscus Gallardo, 1961; Buchholz garizans is far from her other exemplar of and Peters, 1875; Dubois, the B. bufo group (B. bufo), and that the two 1987 ``1986''; Boulenger, members of the B. viridis group (sensu Inger, 1895; Ruiz-Carranza and Her- 1973), B. calamita and B. viridis, are isolated naÂndez-Camacho, 1976; Truebella Graybeal phylogenetically from each other. Neverthe- and Cannatella, 1995; and Poche, less, resolution was not strongly corroborat- 1903. ed. The combined morphology ϩ molecular Graybeal (1997) provided the latest esti- analysis provides less resolution at the base mate of phylogeny within the entire Bufon- of the tree and placed Bufo viridis and B. idae. Unfortunately, although the morpholog- calamita far apart, but it did resolve the Bufo ical results were presented, the morphologi- bufo group as monophyletic (B. bufo and B. cal data matrix and morphological transfor- gargarizans being her exemplars). Beyond mation series were not, though they that, her results do not offer a great deal of presumably are available in her unpublished resolution. Although Graybeal (1997; ®g. 25) dissertation (Graybeal, 1995). Her DNA se- and, more recently Pauly et al. (2004; see quence data and analytical methods are avail- ``Taxonomy of Living Amphibians'') provid- able, however. There have been serious res- ed estimates of bufonid phylogeny and start- ervations published about the quality of ed to delineate the paraphyly of ``Bufo'' Graybeal's 16S sequence data (Harris, 2001; within Bufonidae, taxonomy within ``Bufo'' Cunningham and Cherry, 2004)15 and the pa- remains largely parsed among similarity- per was largely a narrative largely focused based species groups (Blair, 1972b; Cei, on comparing parsimony, maximum-likeli- 1972; Inger, 1972; R.F. Martin, 1972). These hood, and neighbor-joining techniques. For species groups have enjoyed considerable our discussion we present two of her trees popularity and longevity of use, but, with ex- that rest on analytical assumptions similar to ceptions, it is not clear whether their recog- our own: (1) a strict consensus of 82 equally nition continues to be helpful in promoting parsimonious trees based on the unweighted scienti®c progress, inasmuch as no attempt molecular data alone (®g. 25A); and (2) her so far has been made to formulate these combined morphology ϩ molecular tree (®g. groups in phylogenetic terms. 25B). Her molecular results suggest that, of Grandison (1981) provided a phylogenetic the exemplars treated in that particular tree data set for African bufonids that she as- (®g. 25A), Melanophryniscus is the sister sumed were closely related to Didynamipus. taxon of all other bufonids, and Atelopus ϩ Her data were reanalyzed and her tree was Osornophryne form the sister taxon of the corrected by Graybeal and Cannatella remaining bufonids, excluding Melanophryn- (1995), and this tree is presented herein (®g. iscus. (This would suggest that presence of a 26). On the basis of Grandison's (1981) ev- Bidder's organ is not a synapomorphy of Bu- idence, Dubois (1987 ``1985'') partitioned fonidae, but of a smaller component of that former Nectophrynoides into four nominal genera: Spinophrynoides (with aquatic lar- 15 Harris (2001) was unable to duplicate Graybeal's vae), Altiphrynoides (with terrestrial larvae), 16S sequences of Bufo melanostictus and B. calamita, Nectophrynoides (ovoviviparous), and Nim- although her sequences still are most similar to other baphrynoides (viviparous). Graybeal and GenBank sequences of these species. Cunningham and Cannatella's (1995; ®g. 26) reanalysis sug- Cherry (2004) were unable to duplicate most of her 16S sequences for the taxa that Cunningham and Cherry gests, at least on the basis of Grandison's (2004) studied; they suggested widespread sequencing (1981) evidence, that ``Nectophrynoides'' errors in Graybeal's study. Whether the inclusion of bet- (sensu stricto) remains paraphyletic. Necto- ter-quality sequences would change her results is un- phryne and Wolterstorf®na also appear par- known. Nevertheless, the problems with the DNA se- quences and the nondisclosure of the morphological ev- aphyletic in this tree, although Graybeal and idence require that her results not be accepted at face Cannatella (1995) suggested additional char- value. acters in support of the monophyly of Nec- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 69

paedes) form a monophyletic group, that on the basis of larval morphology also includes Mertensophryne. The sister taxon of this Mertensophryne group they suggested is the Bufo angusticeps group, with more distant relatives being the Bufo vertebralis group and . Because of this lack of a corroborated global phylogeny of Bufonidae16, we at- tempted to sample as widely as possible. The nominal bufonid taxa we, unfortunately, were unable to sample are Adenomus, Alti- phrynoides, Andinophryne, Atelophryniscus, several of the species groups of nominal Bufo, Bufoides, Churamiti, Crepidophryne, , Laurentophryne, Leptophryne, Mertensophryne, Metaphryniscus, Nimba- phrynoides, Oreophrynella, Parapelophryne, Pseudobufo, and Truebella. Several of these (e.g., Andinophryne, Bufoides, and Pseudob- Fig. 26. Tree from Graybeal and Cannatella's ufo) are likely imbedded within sampled gen- (1995) parsimony reanalysis of Grandison's (1981) 24 transformation series of morphology for era. African bufonids suggested by Grandison (1981) At least some of the bufonids are descrip- to be related to Didynamipus (length ϭ 79, ci ϭ tively ®rmisternal, such as Atelopus, Dendro- 0.45, ri ϭ 0.68), rooted on a hypothetical ancestor. phryniscus, Melanophryniscus, Oreophrynel- The taxonomy is updated to include generic la, and Osornophryne. Others (Leptophryne) changes made by Dubois (1987 ``1985'') subse- approach this condition (Laurent, 1986; but quent to Grandison's study. see Kaplan, 2004, for discussion of the var- ious meanings of ``®rmisterny''). Some bu- fonids exhibit various kinds of endotrophy: tophryne. This topology may be deeply Altiphrynoides (nidicolous; M.H. Wake, ¯awed, however, because Graybeal's (1997) 1980), Didynamipus (direct development; tree of morphology and molecules (®g. 25) Grandison, 1981), Laurentophryne (direct show that among the exemplars shared with development; Grandison, 1981), Nectophry- the study of Grandison (1981), Altiphryno- ne (nidicolous; Scheel, 1970), Nectophryno- ides, Didynamipus, Nectophrynoides, and ides (oviductal-ovoviviparous; Orton, 1949), Werneria are not necessarily particularly Nimbaphrynoides (viviparous; Lamotte and closely related. Didynamipus, in particular, is Xavier, 1972), Oreophrynella (direct devel- more closely related to Asian Pelophryne opment; McDiarmid and Gorzula, 1989), and and South American Oreophrynella than to Pelophryne (nidicolous; Alcala and Brown, the others in the group addressed by Gran- 1982). Others are also suspected to have en- dison (1981). dotrophic larvae or direct development: Cre- Cunningham and Cherry (2004) provided pidophryne, , Frostius, a DNA sequence study of putatively mono- Metaphryniscus, Osornophryne, Rhampho- phyletic African 20-chromosome Bufo (®g. phryne, Truebella, and Wolterstorf®na (Peix- 27). They suggested that the 20-chromosome oto, 1995; Thibaudeau and Altig, 1999). Un- form a monophyletic group with a re- fortunately, our inability to sample any of versal to 22-chromosomes in the Bufo par- these taxa other than Didynamipus, Necto- dalis group (their exemplars being B. par- dalis and B. pantherinus). They also sug- 16 The study by Pauly et al. (2004) appeared during gested that Stephopaedes and Bufo lindneri the writing phase of this study and therefore did not (a member of the B. taitanus group, long as- in¯uence our taxon sampling. We comment on that pa- sociated with Mertensophryne and Stepho- per in the Taxonomy section. 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 phryne, Nectophrynoides, Osornophryne, Pe- bufonids and pipids; see Kaplan, 1994, 1995, lophryne, Rhamphophryne, and Wolterstorf- 2000, 2001, 2004, for discussion). In addi- ®na prevents us from elucidating the details tion, most ranoids are reported as diplasio- of the evolution of life history in this group coelous, although the de®nitions of amphi- or the considerable morphological variation coely, anomocoely, procoely, diplasiocoely, in bufonid larvae, including such things as as well as ectochordy (ϭ perichordy), epi- ¯eshy accessory respiratory structures on the chordy, holochordy, and stegochordy (ϭ ep- head (e.g., on Stephopaedes, Mertensophry- ichordy) in frogs remains controversial17. ne, Bufo taitanus) and ¯aps on the head Ranoidea contains noncontroversially Ar- (Schismaderma). throleptidae, Astylosternidae, Hemisotidae, Regardless of the taxa we could not in- Hyperoliidae, Mantellidae, Microhylidae, Pe- clude, we were able to sample a worldwide tropedetidae, Ranidae, and Rhacophoridae. selection of 62 bufonid species: Ansonia lon- More controversially included (see above) is gidigitata, A. muelleri, Atelopus ¯avescens, Dendrobatidae, which is placed by various A. spumarius, A. zeteki, Bufo alvarius, B. am- authors within Hyloidea. Haas (2003; ®g. 15) boroensis, B. andrewsi, B. angusticeps, B. did not recover Ranoidea as monophyletic in arenarum, B. cf. arunco, B. asper, Bufo as- his analysis of larval characteristics, instead pinia, B. biporcatus, B. boreas, B. brauni, B. ®nding major ranoid groups (e.g., ranids, bufo, B. camerunensis, B. celebensis, B. cog- rhacophorids, hemisotids ϩ hyperoliids ϩ natus, B. coniferus, B. divergens, B. galeatus, microhylids) interspersed among various hy- B. granulosus, B. guttatus, B. gutturalis, B. loid groups (e.g., Physalaemus, Pleurodema, haematiticus, B. latifrons, B. lemur, B. ma- Odontophrynus ϩ Leptodactylus, and bufon- culatus, B. margaritifer, B. marinus, B. ma- ids). Discussing the evidence that supports zatlanensis, B. melanostictus, B. nebulifer, B. the monophyly of the various ranoid groups punctatus, B. quercicus, B. regularis, B. is extremely dif®cult, partly because of the schneideri, B. spinulosus, B. terrestris, B. highly contingent nature of the evidence and, tuberosus, B. viridis, B. woodhousii, Capen- more commonly because historically the sibufo rosei, C. tradouwi, Dendrophryniscus groups were assembled on the basis of over- minutus, Didynamipus sjostedti, Melano- all similarity or special pleading for speci®c phryniscus klappenbachi, Nectophryne afra, characteristics. N. batesi, , Osor- As understood by most workers, the ques- nophryne guacamayo, Pedostibes hosei, Pe- tions regarding Ranoidea fall into two cate- lophryne brevipes, Rhamphophryne festae, gories: (1) What are the phylogenetic rela- Schismaderma carens, Stephopaedes anotis, tionships within Microhylidae?; and (2) Werneria mertensi, and Wolterstorf®na par- What are the phylogenetic relationships with- vipalmata. This sampling, while not dense in ``Ranidae'' (sensu lato as including all overall given the size of Bufonidae, allows a other ranoid subfamilial and familial taxa). rigorous test of the monophyly and place- The possibility of paraphyly of ``Ranidae'' ment of Bufonidae among anurans, as well (sensu lato) with respect to Microhylidae as a minimal test of the monophyly and re- does not seem to have been considered se- lationships of many groups. Most important, riously. We know of no de®nitive evidence the results, together with the obvious de®- that would reject this hypothesis, although ciencies in taxon sampling, will provide an microhylids predominantly have broadly di- explicit reference point for future, more thor- lated sacral diapophyses, a presumed ple- ough studies of the internal phylogenetic structure of Bufonidae. 17 Several authors (Grif®ths, 1959b, 1963; Tihen, 1965; Kluge and Farris, 1969) considered the amphicoe- RANOIDEA: Ranoidea is an enormous group lous-anomocoelous-procoelous-diplasiocoelous condi- of frogs, arguably monophyletic, grouped tions delimited by Nicholls (1916) to have been over- largely on the basis of one complex morpho- simpli®ed and over-generalized. See Kluge and Farris logical character of the pectoral girdle (i.e., (1969) for discussion, but also see comments by J.D. Lynch (1973: 140) and Haas (2003: 74), who disagreed ®rmisterny, the fusion of the epicoracoid car- with various statements by Kluge and Farris, including tilages), except where considered to be non- their assertion regarding the continuum of variation be- homologous (possibly Dendrobatidae, some tween epichordy and perichordy. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 71

Fig. 27. Implied consensus of two most parsimonious trees of African toads studied by Cunningham and Cherry (2004), showing 22N → 20N transition point and reversal to 22N in the Bufo pardalis group, and alternative placements of Bufo maculatus. The underlying data are sequences from mtDNA (12S, 16S, ND2, and the tRNA genes ¯anking ND2) and nuDNA (ACTC and rhodopsin). Alignment of 12S and 16S were made initially with ClustalX (Thompson et al., 1997), costs not disclosed, and adjusted manually, guided by models of secondary structure. Alignment of coding, tRNA and intron sequences involved so few length variables that these were done manually. Gaps and missing data were treated as unknowns. Outgroups not show in tree: Dendropsophus labialis (Hylidae); Euhyas cuneata (Leptodac- tylidae: Eleutherodactylinae), (Limnodynastidae); Heleophryne natalensis (12S only; Heleophrynidae); H. purcelli (16S only; Heleophrynidae); Nesomantis thomasetti (Sooglossidae); Rana temporaria (Ranidae). siomorphy, and ranoids predominantly have ing microhylids, while noting that any study round sacral diapophyses (Noble, 1931; J.D. of ranoid phylogenetics must address the po- Lynch, 1973), although in the absence of an sition of microhylids within the ranoid explicit cladogram the optimization of this framework. transformation and the number of conver- Within the nonmicrohylid ranoid group, gences is questionable. Nevertheless, we will modern progress in our understanding must restrict our comments to the ranoids, exclud- be dated from the publication of Dubois 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(1981), in which he presented a discussion of peroliidae (Hyperoliinae in Laurent's usage). ranoid nomenclature with reference to the at- Laurent (1986) suggested that the group tendant published morphological diversity of composed of Arthroleptidae, Astylosternidae, Ranidae as then understood. Although non- and Hyperoliidae is distinguished from Ran- phylogenetic in outlook, subsequent papers idae (sensu lato) by having: (1) a cartilagi- by Dubois (1983, 1984b, 1987 ``1985'', nous metasternum without a bony style (pre- 1992) provided workers with phenotypic sumably plesiomorphic at this level of gen- groupings and a working taxonomy that in erality); (2) second carpal free; (3) third dis- earlier manifestations, at least, were useful as tal tarsal free; (4) terminal phalanges rough approximations of phylogenetic generally hooked; (5) pupil usually vertical groups. This approach was criticized for its (usually horizontal in Hyperoliinae, although lack of a phylogenetic rationale and overgen- vertical in someÐe.g., , Heterixal- eralization of characters (Inger, 1996). But us, , ; vertical in Lep- because there was little else with which to topelinae); and (7) metatarsal tubercle absent work, the taxonomies of Dubois have been or poorly developed. None of these charac- in¯uential. The most substantive differences ters is demonstrably synapomorphic. between Dubois' classi®cations (e.g., Du- ARTHROLEPTIDAE (SENSU DUBOIS, 1992; 3 bois, 1992, 2005) and those of other authors GENERA,49SPECIES): Laurent and Fabrezi (e.g., Vences and Glaw, 2001) revolve (1986 ``1985'') provided a discussion of the around category-rank differences, particular- phylogeny of genera within this African tax- ly with respect to the rank and content of on and suggested a relationship of (Arthro- Rhacophoridae (variably including Mantelli- leptis ϩ Coracodichus) ϩ ( ϩ dae as a subfamily, or as Schoutedenella), although the evidence for placed as a subfamily within Ranidae or with this scenario is unclear. Like astylosternines Mantellidae and Rhacophoridae as distinct and hyperoliids, arthroleptids possess a car- families), with the status of the various com- tilaginous sternum, a vertical pupil (horizon- ponents of ``Ranidae'' left as an open ques- tal in most hyperoliines), and a free second tion. With the exception of the recent papers distal carpal, all of which are questionable as by Marmayou et al. (2000) and Roelants et to level of universality and polarity. The al. (2004), which dealt only with Asian taxa, monophyly of this taxon has never been rig- and Van der Meijden et al. (2005), which fo- orously tested by phylogenetic analysis with- cused on an African clade, no comprehensive in a well-sampled larger group although Biju attempt has been made to address the phy- and Bossuyt (2003; ®g. 21), on the basis of logenetics of the entire Ranoidea. a relatively small sampling of frogs found ARTHROLEPTIDAE,ASTYLOSTERNIDAE, AND Hyperoliidae to be polyphyletic, and Vences HYPEROLIIDAE: Arthroleptidae, Astylosterni- et al.'s (2003c; ®gs. 28, 29) analysis of dae, and Hyperoliidae are poorly understood mtDNA suggested that Arthroleptis, Schou- African families that have been joined and tedenella, and Cardioglossa form a clade, ei- separated by various authors (Dubois, 1981; ther as the sister taxon of Astylosternidae ϩ Laurent, 1984b; Dubois, 1987 ``1985'', Leptopelinae, or as the sister taxon of Hy- 1992) and even suggested to be related to at peroliinae. We sampled: Arthroleptis tanneri, least two microhylid subgroups, Scaphio- A. variabilis, Cardioglossa gratiosa, C. leu- phryninae (Laurent, 1951) and Brevicipitinae comystax, Schoutedenella schubotzi, S. syl- (Van der Meijden et al., 2004). Ford and vatica, S. taeniata, and S. xenodactyloides. Cannatella (1993) regarded Arthroleptidae We were unable to sample a member of Cor- (sensu Dubois, 1981; including Astyloster- acodichus (if recognized as distinct from Ar- nidae) as a metataxon (Donoghue, 1985; Es- throleptis). Within Arthroleptidae, Arthrolep- tes et al., 1988; Archibald, 1994), even tis, Schoutedenella, and Coracodichus have though no evidence was suggested to support direct development (Laurent, 1973), but Car- the monophyly of a group composed of Ar- dioglossa have free-living, feeding larva (La- throleptidae and Astylosternidae and as orig- motte, 1961; Amiet, 1989; Altig and Mc- inally proposed was considered to be para- Diarmid, 1999; Thibaudeau and Altig, 1999). phyletic (Laurent, 1951) with respect to Hy- ASTYLOSTERNIDAE (5 GENERA,29SPECIES): 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 73

Fig. 28. Maximum-likelihood tree of hyperoliid, arthroleptid, and astylosternid frogs provided by Vences et al. (2003c). A, Maximum-likelihood analysis of 12S rRNA molecule (187 informative sites) analyzed under a GTR substitution model (cost functions reported) suggested by Modeltest (Posada and Crandall, 1998). Initial alignments under Clustal software, costs not disclosed, and subsequently adjusted manually. Highly variable regions and gaps were excluded as evidence. B, Maximum-likelihood trees based on 138 informative sites of 16S rRNA molecule under a GTR substitution model (cost functions reported) for hyperoliids, arthroleptids, and astylosternids. Initial alignments were made under Clustal, costs not disclosed, and subsequently adjusted manually. Highly variable regions and gaps sites were excluded as evidence.

The African Astylosternidae traditionally has throleptines and hyperoliids, astylosternids been allied with Arthroleptidae and Hyper- have a cartilaginous sternum, a vertical pupil oliidae (see above), although the evidentiary (except in ), and a free sec- justi®cation for this appears to be overall ond distal carpal, all of which are question- similarity rather than synapomorphy. Like ar- able as to level of universality. For our anal- 74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

29) to be more closely related to Astyloster- nidae than to Hyperoliinae. Vences et al. (2003c) further discussed some of the char- acters that Drewes (1984) used in his anal- ysis of the family. Like Hyperoliinae, Lep- topelinae lacks fusion of the second tarsal el- ement and fusion of the second distal carpal (Drewes, 1984; ®g. 30). Channing (1989) re- analyzed the morphological data provided by Drewes (1970; ®g. 30) and provided differ- ent cladistic interpretations of these data; this reanalysis and the underlying characters were Fig. 29. Maximum-likelihood tree of Vences discussed in detail by J.A. Wilkinson and et al. (2003c), based on 566 informative sites of Drewes (2000). All hyperoliids for which it combined fragments of mitochondrial 16S, 12S is known have free-living exotrophic larvae rRNA, and cytochome b, analyzed under the as- (Altig and McDiarmid, 1999). With the ex- sumptions of the Tamura-Nei substitution model ception of a partial revision of (cost functions provided). Gaps were not treated (Wieczorek et al., 1998; Wieczorek et al., as evidence. 2000, 2001), only the intergeneric relation- ships within Hyperoliidae have been ad- dressed phylogenetically (Drewes, 1984; ysis we sampled one species of each nominal Channing, 1989; Richards and Moore, 1998) genus: schioetzi, the presum- and paraphyly of Hyperolius and Kassina re- ably closely related Trichobatrachus robus- main strong possibilities. Our genetic sam- tus, and Leptodactylodon bicolor, pling included four species of the sole genus corrugatus, and gabonicus. in Leptopelinae ( argenteus, L. bo- Vences et al. (2003c; ®gs. 28, 29) suggested cagei, L. sp., and L. vermiculatus). Of Hy- on the basis of mtDNA evidence that Lep- peroliinae we were less complete, as we were topelinae (Hyperoliidae) is either imbedded not able to sample any member of Callixalus, within a paraphyletic Astylosternidae or a Chlorolius, , Kassinula, paraphyletic Arthroleptidae, but they did not Phlyctimantis,orSemnodactylus. Neverthe- express this in the taxonomy. Astylosternus less, we were able to obtain genetic samples and Trichobatrachus have exotrophic aquatic of all remaining genera: spi- larvae; in Scotobleps, the larva is unknown; nosus, Afrixalus fornasinii, A. pygmaeus, Al- and in Leptodactylodon the exotrophic aquat- exteroon obstetricans, sp., H. tri- ic larva has an upturned mouth presumably color, Hyperolius alticola, H. puncticulatus, to feed on the surface ®lm (Amiet, 1970). H. tuberilinguis, , Ne- HYPEROLIIDAE (18 GENERA,2SUBFAMILIES, sionixalus thomensis (transferred back into 250 SPECIES): The African treefrogs of the Hyperolius during the course of this study by family Hyperoliidae are currently divided Drewes and Wilkinson, 2004), into two subfamilies: Hyperoliinae, which is immaculatus, Phlyctimantis leonardi, and united by the presence of a gular gland Tachycnemis seychellensis. (Drewes, 1984), and Leptopelinae, which HEMISOTIDAE (1 GENUS,9SPECIES): Rela- was found by Vences et al. (2003c18; ®gs. 28, tionships of the African taxon Hemisotidae are also unclear (Channing, 1995). Like 18 The various maximum-likelihood trees produced by Rhinophrynus and Brachycephalus, Hemisus this study (Vences et al., 2003c) were not shown. The authors provided trees of (1) 471 bp of the 16S rRNA; lacks a distinguishable sternum. Haas' (2003; (2) combined analysis of 415 bp of the cytochrome b ®g. 15) study of larval morphology found it gene as well as 409 bp of the 12S and 997 bp of the to be the sister taxon of Hyperoliidae among 16S rRNA; and (3) 321 bp of the 12S rRNA gene. Taxon his exemplars. Blommers-SchloÈsser (1993) sampling among the three analyses was quite different and beyond the general conclusion that Hyperoliidae is suggested on the basis of one morphological polyphyletic, this sampling provided low resolution of synapomorphy (median gland) that intergeneric relationships. hemisotines should be united with brevicip- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 75

itine microhylids. That hemisotines and brev- icipitines are quite dissimilar cannot be dis- puted (Channing, 1995; Van Dijk, 2001), and the putative phylogenetic relationship be- tween the two taxa was corroborated via mo- lecular data only recently (Van der Meijden et al., 2004; ®g. 31), although Loader et al. (2004) could not place with con®dence Hem- isus with brevicipitines on the basis of mtDNA sequence evidence. Emerson et al. (2000b) on the basis of mtDNA and a small amount of morphology also allied hemisotids with microhylids, although hemisotids retain a Type IV tadpole unlike the Type II tadpoles of microhylids (or direct development as in brevicipitines). We sampled only Hemisus marmoratus, one species of the single genus, of this morphologically compact taxon. On the basis of the tree of Van der Me- ijden et al. (2004), Dubois (2005) recognized an enlarged family, Brevicipitidae, composed of six subfamilies: Astylosterninae, Arthro- leptinae, Brevicipitinae, Hemisotinae, Hyper- oliinae, and Leptopelinae. For our discussion we maintain the older, more familiar taxon- omy. MICROHYLIDAE (69 GENERA, 432 SPECIES): Microhylidae is a worldwide, arguably well- corroborated taxon (J.D. Lynch, 1973; Star- rett, 1973; Blommers-SchloÈsser, 1975; Sokol, 1975, 1977; Wassersug, 1984; Haas, 2003; but see Van der Meijden et al., 2004 (®g. 31), who suggested that the taxon is paraphyletic with respect to the hemisotines), although the internal relationships of Microhylidae are certainly problematic (Burton, 1986; Zwei- fel, 1986, 2000). The subfamilial taxonomy or taxonomic differentia have not changed Fig. 30. Hyperoliid relationships suggested by materially since the revision by Parker A, Liem (1970) based on 36 dendritic to linear (1934), with the exception of the treatment character transformations of morphology and as- of Phrynomeridae as a subfamily of Micro- suming monophyly of a group composed of hy- hylidae by J.D. Lynch (1973), the inclusion peroliids, mantellids, and rhacophorids, the tree of the Scaphiophryninae by Blommers- rooted on a hypothetical ranid; B, Drewes' (1984) parsimony analysis of 27 morphological character SchloÈsser (1975), and the demonstration of transformations, rooted on a hypothetical ancestor the evolutionary propinquity of Hemisotidae constructed by comparison of a large number of and Brevicipitinae by Van der Meijden et al. ranids, astylosternids, and arthroleptids (Semno- (2004; ®g. 31). Beyond the isolation of brev- dactylus was treated as Kassina weali in this pub- icipitines from other microhylids, the allo- lication.); C, Channing's (1989) parsimony re- zyme data of Sumida et al. (2000a) suggest analysis (with minor modi®cations) of the mor- that the subfamilial de®nitions and generic phological data of Drewes (1984). assignments within nominal Genyophryninae and Asterophryninae may require change. In- deed, Savage (1973) had synonymized the 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 31. Maximum-likelihood tree of various ranoids constructed by Van der Meijden et al. (2004) on the basis of 1,566 bp of the nuclear gene RAG-1. Sequence alignment was not reported. Cost functions of analysis were not provided nor which model of nucleotide evolution (as suggested by ModelTest; Posada and Crandall, 1998) was employed in the analysis. The tree was rooted on Xenopus laevis. We inserted the higher taxonomy on the right to allow easier comparison to other studies dis- cussed in this section. two subfamilies on the basis of their geo- these characters is questionable): (1) absence graphical and morphological similarity. of keratodonts in tadpoles; (2) ventral velum Savage (1973) suggested that Dyscophinae divided medially; (3) glottis fully exposed on is polyphyletic, with the Asian Calluella buccal ¯oor; (4) nares not perforate; and (5) more closely related to asterophryines than secretory ridges of branchial food traps with to the Madagascan Dyscophus. Blommers- only a single row of secretory cell apices. In SchloÈsser (1976) reviewed the controversy addition, adults are characterized as having and retained Dyscophus and Calluella in 2±3 palatal folds (palatal folds also being Dyscophinae. Our taxon sampling allows us found in Hemisus). Van de Meijden et al. to test whether Dyscophinae is monophyletic (2004) suggested on the basis of molecular or diphyletic. evidence that Hemisotidae ϩ Brevicipitinae Ford and Cannatella (1993) identi®ed ®ve is more closely related to Hyperoliidae, Ar- larval synapomorphies for Microhylidae (al- throleptidae, and Astylosternidae than to an though these cannot be documented in line- otherwise monophyletic group of microhy- ages with direct development such as in lids (®g. 31). Therefore, the only articulated brevicipitines, asterophryines, and geny- questions so far regarding the monophyly of ophrynines, so the level of universality of Microhylidae are whether Hemisotidae is im- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 77 bedded within it (see above) or, with Brevi- cipitinae, more closely related to non-micro- hylid groups, although the de®nition, histor- ical reality, and content of the various sub- families are controversial. SCAPHIOPHRYNINAE (2 GENERA,11SPECIES): The Madagascan microhylid subfamily Sca- phiophryninae has no demonstrable synapo- morphies in support of its monophyly, but if its monophyly is assumed it is widely con- sidered to be the sister taxon of the remain- ing Microhylidae. At least some authors (e.g., Dubois, 1992) regard it as a distinct family. Ford and Cannatella (1993) suggest- ed that larval synapomorphies that place it in association with the remaining Microhylidae (at least for those that have larvae) are (1) the possession of a median spiracle in the larvae; (2) ®laments poorly developed or absent; (3) modi®cations of buccal pumping mechanisms (short lever arm on ceratohyal, small buccal ¯oor area); (4) absence of m. suspensoriohyoideus; and (5) lack of sepa- ration of the mm. quadrato-, hyo-, and sus- Fig. 32. Trees of Asterophryinae suggested by pensorioangularis. Parker (1934) reported the A, Zweifel (1972), based on 9 morphological transformation series and showing alternative po- taxon as diplasiocoelus like most other ran- sitions of Barygeny (tree rooted on a generalized oids, although he noted Hoplophryninae primitive hypothetical ancestor); and B, Burton ( ϩ ), Astero- (1986), based on a subjective evaluation of 54 phryinae, and some members of his Micro- transformation series of morphology. This tree re- hylinae (e.g., , Metaphry- lects our understanding of Burton's narrative sum- nella, , ) as procoelous. mary of asterophryine relationships, with the no- Parker (1934) noted that re- menclature updated (Callulops replacing Phryno- tains a complete sphenethmoid, thereby ex- mantis). Quotation marks denote nonmonophyly. cluding it from Microhylidae, which, as he applied the name, included only those taxa where the sphenethmoid is either in two CIES): Zweifel (1972) and Burton (1986) last parts, or, more rarely, not ossi®ed at all. Haas reported on phylogenetics of the Australo- (2003) suggested on the basis of larval mor- phology that Scaphiophryninae is polyphy- Papuan Asterophryinae (®g. 32). Geny- letic, with Scaphiophryne forming the sister ophryninae is also Australo-Papuan but ex- taxon of the remaining microhylids, and Par- tends into the Philippines and Lesser Sundas. adoxophyla more closely related to Phry- No major revision or broad-scale phyloge- nomerinae. Clearly the monophyly of this netic study has appeared since Parker (1934), taxon is controversial, but we, unfortunately, although Burton (1986) did suggest evidence were unable to sample and so that it is paraphyletic with respect to Aster- could not test the monophyly of Scaphio- ophryinae. Sumida et al. (2000a) noted that phryninae. We were able to obtain only a some allozyme evidence suggested that As- representative of the other genus, Scaphio- terophryinae is imbedded within a paraphy- phryne marmorata. Our results regarding the letic Genyophryninae. Savage (1973) consid- Scaphiophryninae will therefore remain in- ered Genyophryninae to be part of Astero- complete. phryinae based on the dubious nature of the ASTEROPHRYINAE (8 GENERA,64SPECIES) procoely±diplasiocoely distinction; that they AND GENYOPHRYNINAE (11 GENERA, 142 SPE- share direct-development; and, in part, that 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 they are both biogeographically centered in sole synapomorphy) that brevicipitines New Guinea. should be united with hemisotids (but see Zweifel (1971) summarized the distinction Channing, 1995, who considered this change between the subfamilies as (1) maxillae often premature at the time because of the other- overlapping the premaxillae and usually in wise trenchant differences between them). contact anteriorly (Asterophryinae; this pre- Van der Meijden et al. (2004; ®g. 31) and sumably is apomorphic), maxillae not over- Loader et al. (2004) provided molecular data lapping the premaxillae (Genyophryninae); in support of a hemisotid±brevicipitine rela- (2) diplasiocoelous (rarely tionship. Of the nominal genera we were un- procoelous; Asterophryinae), procoelous able to sample the monotypic Balebreviceps (Genyophryninae); and 3) tongue subcircu- and . The effect of excluding lar, entirely adherent, often with a median these taxa is unknown, although Loader et furrow and posterior pouch (Asterophryi- al. (2004) recovered Spelaeophryne in a nae), tongue oval, half-free behind, no trace clade with and , to the of a median furrow or pouch (Genyophry- exclusion of . We were able to sam- ninae; shared with Cophylinae). Genyophry- ple at least one species of the remaining ninae and Asterophryinae share direct devel- nominal genera: Breviceps mossambicus, opment (Zweifel, 1972; Thibaudeau and Al- Callulina kisiwamitsu, C. kreffti, and Pro- tig, 1999). Our sampling will allow us to test breviceps macrodactylus. Because there are the hypotheses of relationship so far pub- 13 species of Breviceps and 3 species of Pro- lished and elucidate the possible paraphyly breviceps, we were unable to test rigorously of Genyophryninae. Unfortunately, we could the monophyly of these taxa. sample only one species of Asterophryinae, COPHYLINAE (7 GENERA,41SPECIES): The Callulops slateri, which will not allow us to Madagascan Cophylinae is similar to Dys- test its monophyly. The effect of excluding cophinae and Genyophryninae in retaining representatives of , Barygenys, maxillary and vomerine teeth (except in , , Pherohapsis, Stumpf®a) but differs from Dyscophinae in Xenobatrachus, and Xenorhina is unknown. having procoelous vertebrae and from Dys- Of Genyophryninae, we were able to sam- cophinae and Genyophryninae in having a ple pansa, sp., divided vomer (Parker, 1934); none of the sphagnicola, sp., Geny- characters is demonstrably synapomorphic. ophryne thomsomi, Liophryne rhododactyla, Blommers-SchloÈsser and Blanc (1993) pro- brachypus, and vided a cladogram (®g. 33A) of the genera sp. We were unable to sample Albericus, based on nine morphological characters, in , Oreophryne,orOxydacty- which they suggested that Plethodontohyla la. was paraphyletic and that Platypelis did not BREVICIPITINAE (5 GENERA,22SPECIES): have apomorphies to assure its monophyly. Like the Australo-Papuan Asterophryninae Andreone et al. (2004 ``2005'') recently pro- and Genyophryninae, the African Brevicipi- vided a maximum likelihood analysis of tinae has direct development (Parker, 1934; 1173 bp of mtDNA (®g. 33B), in which he Thibaudeau and Altig, 1999). Parker (1934) documented Plethodontohyla paraphyly. Be- considered the subfamily to be distantly re- cause these sequences became available after lated to all other microhylid taxa and char- our analyses were completed, we did not acterized by the retention of a medially ex- sample , ,or panded vomer. Parker (1934) reported the Rhombophryne. The effect of this on the taxon as diplasiocoelus like most other ran- placement of the subfamily will remain un- oids. The species within the subfamily are known, although we did sample four species closely similar and unlike all other micro- of the four remaining genera: hylids in general habitus, although the mono- montana, Platypelis grandis, Plethodonto- phyly of the group has never been tested rig- hyla sp., and Stumpf®a psologlossa. Unfor- orously. tunately, because of our limited sampling we Blommers-SchloÈsser (1993) suggested (the will not be able to test rigorously the results presence of a median thyroid gland being the of either Blommers-SchloÈsser and Blanc 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 79

Calluella as associated with the direct-devel- oping Asterophryinae and any similarities with Dyscophus as re¯ecting plesiomorphy. We sampled one species each of the two nominal genera: Calluella guttulata and Dys- cophus guineti. MELANOBATRACHINAE (3 GENERA,4SPE- CIES): On the basis of geography alone (East Africa [2 genera] and southern India [1 ge- nus]), one would suspect that this is not a monophyletic taxon. Nevertheless, the three genera share an incomplete auditory appa- ratus (convergent in Balebreviceps [Brevi- cipitinae]; Largen and Drewes, 1989) and fu- sion of the sphenethmoid with the paras- phenoid (Parker, 1934). Savage (1973), fol- lowed by Laurent (1986) and Dubois (2005), placed Melanobatrachus in Microhylinae and retained and Parhoplo- phryne in Hoplophryninae, but did so only Fig. 33. Trees of Cophylinae suggested by A, by discarding absence of the auditory appa- Blommers-SchloÈsser and Blanc (1993), on the ba- ratus and fusion of the sphenethmoid to the sis of nine morphological transformation series, parasphenoid, as convergences, without of- rooted (by implication) on Dyscophinae and Sca- fering speci®c characters that con¯icted with phiophryninae. The ®gure is redrawn with branch- these as synapomorphies. Although we are es collapsed that were unsupported by evidence suspicious of the monophyly of this taxon, in the original; B, Andreone et al. (2004 ``2005''), we stick with the most parsimonious hypoth- based on 1,173 bp of 12S and 16S rRNA mtDNA. esis (monophyly of Melanobatrachinae, sen- This tree is redrawn to note only monophyletic su lato) until alternative evidence emerges. genus-group taxa. Alignment was made using the Clustal option in Sequence Navigator (Applied Apparently based on information provided Biosystems), with cost functions for alignment not for Hoplophryne by Barbour and Loveridge provided. All sections that could not be aligned, (1928) and Noble (1929), Parker (1934) gen- including those with three of more gaps in one or eralized that all members of his Melanoba- more taxa, were excluded from analysis. Whether trachinae lack a free-swimming tadpole, the gaps were treated as unknown or evidence was larvae with `` taking place on not stated. The Tamura-Nei substitution model land, but not in an ''. No reproductive or was selected for maximum-likelihood analysis of developmental data on Parahoplophryne or aligned data. The tree was rooted on Scaphio- Melanobatrachus have been published (Dal- phryne (not shown). Quotation marks around try and Martin, 1997). Thibaudeau and Altig names denotes nonmonophyly. (1999) listed Melanobatrachus and Parho- plophryne as having endotrophic larvae, pre- (1993) or Andreone et al. (2004 ``2005''). sumably because of the earlier statement by Species of Cophylinae have nidicolous lar- Parker (1934). McDiarmid and Altig (1999: vae (Blommers-SchloÈsser and Blanc, 1991; 13), however, listed Hoplophryne as exo- Glaw and Vences, 1994). trophic, because Barbour and Loveridge DYSCOPHINAE (2 GENERA,10SPECIES): The (1928: 256) reported vegetable matter in the Madagascan Dyscophinae is distinguished guts of larvae and because R. Altig examined from most other microhylid subfamilies by AMNH larvae of Hoplophryne and inferred retaining maxillary and vomerine teeth, oth- that they could feed (R.W. McDiarmid, per- erwise known only in Cophylinae and Gen- sonal commun.). Laurent (1986) reported the yophryninae, both of which are procoelous taxon (Parhoplophryne and Hoplophryne in rather than diplasiocoelous (Parker, 1934) as his Hoplophrynnae; Melanobatrachus in his in Dyscophinae. Savage (1973) had regarded Microhylinae) as procoelous, unlike most 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

ed the Old World and New World compo- nents separately, implying some kind of tax- onomic division. This was followed, without discussion, by Dubois (2005), who recog- nized Gastrophryninae for the New World component and Microhylinae for the Old World component. We are not aware of any evidence in support of this arrangement so we retain the old taxonomy. Of the 30 nom- inal genera we were able to sample represen- tatives of 14: fusca, geayei, schirchi, muelleri, ovalis, elegans, G. olivacea, bolivi- ana, pleurostigma, pulchra, heymonsi, Microhyla sp., inornata, Nelsonophryne aequa- torialis, Ramanella obscura, and Synaptur- anus mirandaribeiroi). We were not able to sample Adelastes, Altigius, , , , Glyphog- lossus, Hyophryne, , Metaphry- Fig. 34. Tree of New World microhylids by nella, Myersiella, , Phrynella, Re- Wild (1995) based on a parsimony analysis of 14 lictovomer, , Syncope, and morphological transformations series, outgroups . Most of these appear to be clus- and evidence for ingroup monophyly not speci- tered with sampled taxa. The exclusion of ®ed. Otophryne and Uperodon, however, is partic- ularly regrettable. Our sampling will not al- other ranoids so this may also be synapo- low detailed elucidation of the evolution of morphic. Unfortunately, we were able to life-history strategies. Adelastes, Altigius, sample only Hoplophryne rogersi and so will Gastrophrynoides, Hyophryne, Kalophrynus not be able to comment on the monophyly (nidicolous), Myersiella (direct develop- of Melanobatrachinae. ment), Phrynella, (nidicolous), MICROHYLINAE (30 GENERA, 133 SPECIES): and Syncope (nidicolous) have endotrophic The American and tropical Asian Microhy- larvae that exhibit (or are suspected to ex- linae have free-swimming tadpoles (except hibit) various degrees of truncation of larval for a few species, such as Myersiella mi- development (Thibaudeau and Altig, 1999). crops, that have direct development; Izeck- That we lack representatives of about half of sohn et al., 1971). Although microhylines these is lamentable, but our results will pro- can be morphologically characterized, they vide an explicit starting point for future, have no known synapomorphies, and their more detailed studies. The remaining genera monophyly is deeply suspect. According to have exotrophic larvae of typical microhylid Parker (1934), maxillary and vomerine teeth morphology (Altig and McDiarmid, 1999). are absent (as in several other extra-Mada- PHRYNOMERINAE (1 GENUS,5SPECIES): The gascar subfamilies); the vomer is much re- African Phrynomerinae is diagnosable from duced and usually divided; the sphenethmoid Microhylinae solely by possessing intercala- is divided or absent; and the vertebrae are ry cartilages between the ultimate and pen- diplasiocoelous (or rarely procoelous). Wild ultimate phalanges (Parker, 1934). Like most (1995) provided a cladogram of New World other ranoids it is diplasiocoelous. Of this genera (®g. 34), but this assumed that the small taxon we sampled bifas- New World group is monophyletic and was ciatus. Phrynomantis typically has aquatic, unclear about the outgroup(s) used to polar- exotrophic microhylid larvae (Altig and ize the transformations. Laurent (1986) treat- McDiarmid, 1999). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 81

``RANIDAE'' (CA.54GENERA, 772 SPECIES): Ceratobatrachus). Ceratobatrachinae repre- Ranidae is a large ranoid taxon, that is likely sents the direct-developing part of - paraphyletic with respect to Mantellidae and inae sensu Noble (1931) and Platymantinae RhacophoridaeÐat least on the basis of mo- of later authors (e.g., Savage, 1973; Laurent, lecular evidence (Vences and Glaw, 2001; 1986). Those taxa formerly included in Roelants et al., 2004; Van der Meijden et al., Cornuferinae or Platymantinae that exhibit 2005). Ford and Cannatella (1993; ®g. 14) unforked omosterna and/or free-living tad- suggested that the group is paraphyletic, or, poles (what are now , Huia, Mer- at least, that it does not have recognized syn- istogenys, , Hylarana [sensu lato], apomorphies. Nevertheless, Haas (2003; ®g. and ) are now placed in Raninae 15) suggested the following to be synapo- or Micrixalinae. Batrachylodes is inferred to morphies for Ranidae, excluding other ran- have direct development (Noble, 1931; oids: (1) cartilaginous roo®ng of the cavum Brown, 1952; Duellman and Trueb, 1986; cranii present as taenia transversalis and me- Thibaudeau and Altig, 1999), but unlike oth- dialis; (2) free basihyal present; and (3) ®r- er members of Ceratobatrachinae, Batrachy- misterny (convergent elsewhere in Haas' lodes has an entire omosternum (rather than tree). being forked). Noble (1931) regarded Batra- Laurent (1986) included the mantellines chylodes as derived from his Cornufer (ϭ and rhacophorids in his Ranidae, a content ) and, by inference, exhibiting di- that allows at least two other characters (dis- rect development. Because of the character tinctly notched tongue and bony sternal con¯ict of omosternum shape and life-his- style) to be considered as possible synapo- tory, Brown (1952) regarded Batrachylodes morphies (Ford and Cannatella, 1993). as related either to ``Hylarana'' (exotrophic, (These are, however, incongruent with char- entire omosternum) or to the Ceratobatra- acters suggested by Haas, 2003). chus group (direct-developing, forked omos- Dubois and coauthors (Dubois, 1992; Du- ternum). Laurent (1986) treated Batrachylo- bois and Ohler, 2001; Dubois et al., 2001; des as a member of Raninae, although Bou- Dubois, 2005) suggested a taxonomy of 11± lenger (1920) had noted the intraspeci®c 14 subfamilies of uncertain monophyly or re- plasticity of omosternum shape, the only ev- lationship with respect to each other. For dis- idence supporting placement of Batrachylo- cussion, we recognize Dubois' subfamilies, des in Raninae. This arrangement was ac- except as noted. As discussed by Inger cepted by Dubois (1987 ``1985''), although (1996), the diagnostic features supporting subsequently, Dubois (2005) transferred Ba- Dubois' (1992) classi®cation at the time of trachylodes out of Raninae and into Cerato- that writing frequently re¯ected overgener- batrachinae, presumably on the basis of the alized and postfacto approximations for clus- direct development. Our analysis should pro- ters that were aggregated with overall simi- vide more evidence on the placement of this larity, not synapomorphy, as the organizing taxon. principle. The relationships suggested by this Dubois (1992) recognized Ceratobatrachi- taxonomy (and Dubois, 2005, as well) can ni within his Dicroglossinae, but later (Du- be at variance with evidence of monophyly, bois et al., 2001) considered it to be a sub- notably evidence from DNA sequences (Em- family, of unclear relationship to Dicroglos- erson and Berrigan, 1993; Bossuyt and Mil- sinae. Even later, Dubois (2003) stated, on inkovitch, 2000; Emerson et al., 2000b; Mar- the basis of unpublished molecular data, that mayou et al., 2000; Biju and Bossuyt, 2003; Ceratobatrachini is a tribe within Dicroglos- Roelants et al., 2004), so this taxonomy re- sinae. Van der Meijden et al. (2005) present- quires careful evaluation. ed DNA sequence evidence that Ceratoba- CERATOBATRACHINAE (6 GENERA,81SPE- trachus is outside of Dicroglossinae, and on CIES): Ceratobatrachinae is composed of di- that basis (Dubois, 2005) once again em- rect-developing species found from western braced the subfamilial rank Ceratobatrachi- China (i.e., ) to the Indo-Australian nae. Roelants et al. (2004; ®g. 35), in a study archipelago (Batrachylodes, Discodeles, Pal- of predominantly Indian taxa, provided mo- matorappia, Platymantis, and the monotypic lecular evidence that suggest that Ingerana is 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 83 in Occidozyginae, rather than in Ceratobatra- neous group of southern African ranoids, in- chinae, although Dubois (2005), without dis- cluding Afrana, , Natalobatra- cussion, did not accept this. chus, Petropedetes, Pyxicephalus, Strongy- Of this group we sampled Batrachylodes lopus, and Tomopterna. Kosuch et al. (2001; vertebralis, Discodeles guppyi, Ceratobata- ®gs. 38, 39), on a relatively small amount of chus guentheri, Platymantis pelewensis, P. evidence, had previously placed Conraua al- weberi, and Ingerana baluensis. Thus, we ternatively as either the sister taxon of Lim- only lack Palmatorappia from this group19. nonectes (based on 16S alone) or as the sister Although we obviously cannot test the taxon of Tomopterna ϩ Cacosternum (based monophyly of these individual genera (ex- on combined 12S and 16S). The latter result cept Platymantis), our taxon sampling is ad- was suggestive of the more complete results equate to test the monophyly of the inclusive of Van der Meijden et al. (2005). Although group. characters have not been suggested that are CONRAUINAE (1 GENUS,6SPECIES): Until clearly synapomorphic, the group is morpho- the recent publication by Dubois (2005), this logically compact and monophyly is likely. genus (Conraua) had been placed on the ba- Of the six species we sampled two: Conraua sis of overall similarity in a monotypic tribe, robusta and C. goliath. Conrauini, in Dicroglossinae (Dubois, 1992). DICROGLOSSINAE (12 GENERA, 152 SPECIES): Conrauini was proposed (Dubois, 1992) for Recounting the taxonomic history of Dicrog- the West African genus Conraua, the diag- lossinae is dif®cult inasmuch as it was orig- nostic characters being the retention of a inally formed on the basis of overall similar- free-living tadpole stage (plesiomorphic), ity, and the content has varied widely, even with a larval keratodont formula of 7±8/6± by the same authors. Only recently has its 11 (see Dubois, 1995, for the de®nition of concept begun to be massaged by phyloge- keratodont formula) and lateral line not re- netic evidence. Dubois (1987 ``1985'', 1992) tained into adulthood (plesiomorphic). Van diagnosed Dicroglossinae (in the sense of in- der Meijden et al. (2005; ®g. 36), on the ba- cluding Conrauinae and excluding Paini) as sis of DNA sequence data, showed that Con- having the omosternum moderately or raua is not close to Dicroglossinae but the strongly bifurcate at the base and the nasals sister taxon to a taxonomically heteroge- usually large and in contact with each other and with the frontoparietal, although none of 19 The status of Liurana Dubois, 1987, is unclear. Du- these characters is demonstrably synapo- bois (1987 ``1985'') named Liurana as a subgenus of morphic. The most recent taxonomy of Di- Ingerana (Ceratobatrachinae) but, without discussing evidence, Dubois (2005: 4) subsequently considered croglossinae (Dubois, 2005) recognized four Liurana to be a synonym of Taylorana (ϭ Limnonectes, tribes: Dicroglossini (for , Fejer- Dicroglossinae). Similarly, Dubois (2005), with minimal varya, Hoplobatrachus, , Nan- discussion, placed Annandia Dubois, 1992, in his tribe nophrys, and ), Limnonectini Limnonectini, although he had named this taxon as a subgenus of Paa, in his Paini. Because these statements (for Limnonectes, as well as some taxa con- are not associated with evidence, they do not merit fur- sidered by most authors to be synonyms of ther discussion. Limnonectes), Occidozygini (for Occidozyga

← Fig. 35. One of 24 most parsimonious trees of ranoids of Roelants et al. (2004) that corresponds, except for branches marked with an asterisk (*), to their maximum-likelihood tree, based on 698 infor- mative sites out of 1,895 bp of: (1) 750 bp covering part of 12S rRNA gene, complete tRNAVal gene, and part of the 16S rRNA gene; (2) 550 bp of the 16S rRNA gene; (3) ca. 530 bp of exon 1 of the nuclear tyronsinase gene; (4) ca. 315 bp of exon 1 of the rhodopsin gene; (5) ca. 175 bp of exon 4 of the nuclear rhodopsin gene. Alignment was made using the programs SOAP v. 1.0 (LoÈytynoja and Milinkovitch, 2000) and ClustalX (Thompson et al., 1997). Cost functions were not speci®ed, and alignment was subsequently adjusted manually. Sequence segments considered to be ambiguously aligned were excluded from analysis (508 bp). Substitution model assumed for analysis was GTR ϩ ⌫ϩ I. It was not stated whether gaps were treated as missing data or as evidence. 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 36. Maximum likelihood tree of exemplars of Ranoidea, with a focus on African taxa, by Van der Meijden et al. (2005), based on mt DNA (12S and 16S rRNA) and nu DNA (RAG-1, RAG-2, rhodopsin), for 2,995 bp of sequence. Alignment was made using ClustalW (Thompson et al., 1994), with costs not disclosed and gaps and highly variable sites excluded from analysis. The model assumed for maximum-likelihood analysis was TrN ϩ I ϩ G. The tree was rooted on an hierarchical outgroups (not shown in original) composed of Latimeria, Homo, Gallus, Lyciasalamandra, Alytes (2 spp.), Aga- lychnis, and Litoria. The ``southern African clade'' represents Pyxicephalinae as subsequently redelim- ited by Dubois (2005). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 85 and Phrynoglossus), and Paini (for Chapar- have expected given that the species of ana, Nanorana, and ). Sphaerotheca were long placed in Tomopter- Dicroglossini was diagnosed by Dubois na (Pyxicephalinae). Roelants et al. (2004; (1992; in the sense of including Occidozy- ®g. 35) also placed in Dicrog- ginae) as retaining a free-living tadpole (ple- lossinae (by implication) on the basis of siomorphic) and having a lateral line system mtDNA and nuDNA evidence, substantiating that usually is retained into adulthood (pre- the earlier assessment by Kosuch et al. sumably apomorphic, but not present in Oc- (2001; ®gs. 38) which was made on less ev- cidozyga, sensu stricto). As conceived by idence. It was previously assigned to Ranix- Dubois (1992), the taxon contained Euphlyc- alini by Dubois (1987 ``1985'') and to Di- tis, Occidozyga, and Phrynoglossus. Fei et al. croglossini by Dubois et al. (2001). Dubois (1991 ``1990'') and, subsequently, Dubois et et al. (2001: 55) implied on the basis of var- al. (2001) on the basis of published and un- ious published and unpublished mtDNA data published molecular evidence (Marmayou et that Euphlyctis (formerly in his Dicroglossi- al., 2000Юg. 37; Kosuch et al., 2001Юgs. ni), , Hoplobatrachus, Minervar- 38; Delorme et al., 2004Юg. 40) placed Oc- ya, Nannophrys, and Sphaerotheca (formerly cidozyga and Phrynoglossus in the subfamily in his Limnonectini) should be included in a Occidozyginae, and transferred without dis- reconstituted Dicroglossini. cussion into Dicroglossini Fejervarya and Delorme et al. (2004; ®g. 40) demonstrat- Hoplobatrachus (from Limnonectini) and edÐas had Roelants et al. (2004; ®g. 35)Ð Sphaerotheca (from Tomopterninae), and that is phylogenetically distant Nannophrys (from Ranixalinae). from Limnonectes. Grosjean et al. (2004), building on the ear- Of these taxa we sampled rather broadly: lier work of Kosuch et al. (2001) suggested ; Fejervarya cancri- on the basis of several mtDNA and nuDNA vorus, F. kirtisinghei, F. limnocharis, and F. loci that Euphlyctis is the sister taxon of Ho- syhadrensis; Hoplobatrachus occipitalis and plobatrachus with Fejervarya, Sphaerothe- H. rugulosus; Limnonectes acanthi, L. grun- ca, Nannophrys, and Limnonectes forming niens, L. heinrichi, L. kuhlii, L. limborgi (for- more distant relations, a result that is consis- merly Taylorana limborgi), L. poilani, and L. tent with the tree of Roelants et al. (2004; visayanus; ; Sphaer- ®g. 35). otheca breviceps and S. pluvialis. On the ba- Dubois (1992) also recognized a tribe sis of this sampling we should be able to Limnonectini diagnosed nearly identically evaluate the reality of this taxon and, at least with Conrauini (Conrauinae of this review), to some degree, the monophyly of the con- differing only in the larval keratodont for- tained genera. mula of 1±5/2±5, which is arguably plesiom- Occidozygini is a tropical Asian group of orphic. Nominal genera contained in this arguable position. Marmayou et al. (2000; group occur from tropical Africa to tropical ®g. 40) presented mtDNA evidence that Oc- Asia with most taxonomic diversity being in cidozyga and Phrynoglossus are not within Asia: Hoplobatrachus, Limnonectes, and Fe- Dicroglossinae but are outside of a clade jervarya (which was considered a subgenus composed of Rhacophoridae and other mem- of Limnonectes at the time). In addition Mar- bers of a paraphyletic Ranidae. Fei et al. mayou et al. (2000; ®g. 37) and Delorme et (1991 ``1990'') had already transferred Oc- al. (2004; ®g. 40) suggested on the basis of cidozyga (sensu lato) out of Dicroglossinae mtDNA evidence that Sphaerotheca (former- and into its own subfamily on the basis of ly in Tomopterninae; Dubois, 1987 ``1985'') larval characters and this evidence supported and Taylorana (now a synonym of Limno- the view that Dicroglossinae, as previously nectes; originally considered to be a member conceived, is polyphyletic. Roelants et al.'s of Limnonectini [Dubois, 1987 ``1985''] but (2004) greater sampling of Asian ranoids subsequently transferred to Ceratobatrachi- suggested that Ingerana (nominally in Cer- nae by Dubois, 1992) are in Limnonectini. atobatrachinae) is in this clade and together Sphaerotheca, therefore, is likely not to be form the sister taxon of a reformulated Di- closely related to Tomopterna, as one would croglossinae (®g. 35), which together are the 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 37. Consensus of two equally parsimonious trees from Marmayou et al. (2000) of exemplars of Ranidae and Rhacophoridae (Ranidae: Rhacophorinae in their usage) based on 305 bp (151 infor- mative sites) of 12S mtDNA, aligned using the program MUST (Philippe, 1993) and subsequently manually modi®ed with reference to secondary structure models. Cost functions for alignment were not stated, nor whether gaps were treated as missing data or as evidence (ci ϭ 0.382, ri ϭ 0.429). Tree rooted on Eleutherodactylus cuneatus (ϭ Euhyas cuneata). sister taxon of a clade composed of Mantel- sampled Phrynoglossus baluensis, P. boreal- lidae, Rhacophoridae, and Raninae. No Af- is, P. martensii, and Occidozyga lima. rican taxa were examined by Marmayou et Paini is a montane Asian tribe diagnosed al. (2000; ®g. 37), Roelants et al. (2004; ®g. among ranids by having an unforked omos- 35), or Delorme et al. (2004; ®g. 40), so the ternum (and was therefore formerly included relative position and monophyly of Occidoz- in Raninae by Dubois, 1987 ``1985'', 1992) yginae and Dicroglossinae needed to be fur- and males having black, keratinous ventral ther elucidated. This issue was addressed by spines (presumably a synapomorphy with Van der Meijden et al. (2005; ®g. 36), who Nanorana; Jiang et al., 2005: 357). Paini ac- did analyze Asian and African taxa simulta- cording to Dubois (1992) was composed of neously and found Occidozygya lima to sit two genera, each with four subgenera: genus within their Dicroglossinae. Dubois (2005), Chaparana with subgenera Annandia, Cha- on the strength of the evidence produced by parana, Feirana, and Ombrana; genus Paa Van der Meijden et al. (2005), returned Oc- with subgenera Eripaa, Gynandropaa, Paa, cidozyginae to Dicroglossinae as a tribe. We and Quasipaa. Dubois et al. (2001), citing 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 87

Fig. 38. Neighbor-joining tree of ranoid exemplars of Kosuch et al. (2001), which ``agreed well'' with the consensus of four equally parsimonious trees (ci ϭ 0.51). Underlying data were 572 bp of aligned 16S mtDNA sequences of which 221 are parsimony-informative. Alignment was done manually using Sequencher (Applied Biosystems). Indels were treated as missing data. Taxon assignments on the right re¯ect the taxonomy as it existed at the time. 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

composed of ``Chaparana'', several species of ``Paa'', and Nanorana, characterized by spines forming two patches on the chest (save C. quadranus, the type of subgenus Feirana, which does not have spines on the chest); and (2) Group 2, composed of ``Paa'' species as- sociated previously with the subgenera Quas- ipaa Dubois, 1992 (P. robertingeri), and one species nominal of the genus Chaparana, sub- genus Feirana Dubois, 1992 (Paa yei). The second group is characterized by having spines as a single group, more or less over the entire venter, but this characteristic is suf®- ciently variable among subgroups as not to be diagnostic practically except in the not-Na- norana group sense. These authors recom- mended that the generic name Quasipaa be applied to Group 2, but for unstated reasons Fig. 39. Neighbor-joining tree of ranoid ex- hesitated to resolve taxonomically the non- emplars of Kosuch et al. (2001). Underlying data monophyly of Chaparana and Paa in their were 16S data (see ®gure 38) and 12S mtDNA Group 1. Nanorana GuÈnther, 1896, is the old- (331 bp). Alignment was done manually using Se- est available name for their ®rst group. quencher (Applied Biosystems). Gaps treated as Three nominal genera are de®nitely in- missing data. cluded in Paini: ``Chaparana'' (polyphyletic; see above); Nanorana; and ``Paa'' (paraphy- letic with respect to ``Chaparana'' and Nan- unpublished DNA sequence, suggested that orana20). We sampled Nanorana pleskei, Paini be transferred from Raninae to Dicog- Quasipaa exilispinosa and Q. verrucospino- lossinae. Jiang and Zhou (2001, 2005; ®g. sa but did not sample ``Chaparana'' or 41), Jiang et al. (2005; ®g. 42), Roelants et ``Paa'' (sensu stricto). al. (2004; ®g. 35), and Van der Meijden Jiang et al. (2005) did not mention or ad- (2005; ®g. 36) on the basis of published dress three supraspeci®c taxa usually asso- DNA sequence evidence, suggested that Di- ciated with Paini. The ®rst is Eripaa Dubois, croglossinae, with a forked omosternum, is 1992, whose type and only species is Rana paraphyletic with respect to Paini, with an fasciculispina Inger, 1970. Eripaa Dubois, unforked omosternum. For this reason Roe- 1992, was named and is currently treated as lants et al. (2004) and Jiang et al. (2005) a subgenus of Paa. Although Eripaa exhibits transferred Paini out of Raninae and into Di- spines on the entire chest and throat, such as croglossinae. Larvae in the group are exo- in group 2 of Jiang et al. (2005), they are trophic and aquatic (Altig and McDiarmid, uniquely distinct from all other ``Paa'', 1999). ``Chaparana'', and Nanorana species in that Jiang et al. (2005) recently provided a phy- these spines are clustered in groups of 5±10 logenetic study (®g. 42) of Paini on the basis on circular whitish tubercles. We cannot haz- of 12S and 16S rRNA fragments. Unfortu- ard a guess as to how Eripaa is related to nately, that study appeared too late to guide the rest of Paini. The second is Annandia our choice of terminals, but their results are important in helping us interpret our own re- 20 Without mentioning content, Dubois (2005) recog- sults. They found Paa to be paraphyletic with nized three genera: Chaparana, Nanorana, and Quasi- respect to Chaparana and Nanorana; Cha- paa. In light of the phylogenetic study by Jiang et al. parana to be polyphyletic with the parts im- (2005), it is not clear how Chaparana and Nanorana were intended to be delimited or what the content of bedded within ``Paa''; and Nanorana to be these taxa would be. We presume that Dubois' (2005) deeply imbedded within ``Paa''. Within Paini intention was to recognize a paraphyletic Chaparana they recognized two groups: (1) Group 1, within which a monophyletic Nanorana is imbedded. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 89

Fig. 40. Maximum-likelihood tree of ranoids of Delorme et al. (2004), based on sequences from 12S and 16S rRNA for a total of 1198 bp. Alignment was made using the program Se-Al (Rambaut, 1995; cost functions not provided) and by comparison with models of secondary structure. Gaps were treated as missing data. The maximum-likelihood nucleotide substitution model accepted was TrN ϩ I ϩ G.

Dubois, 1992, whose type and only species originally proposed as a subgenus of Cha- is Rana delacouri Angel, 1928. Annandia parana. This species also posseses spinules was originally named as a subgenus of Cha- only around the anus, prompting Dubois parana Bourret, 1939, but recently, Dubois (1987, ``1986'') to consider it evidence of a (2005), without discussion of evidence, treat- unique reproductive mode, and thus a close ed Annandia as a genus in Limnonectini. relative of Annandia delacouri. Unfortunate- Perhaps this was done because this species ly, we did not sample any of these three taxa, bears a smooth venter, with spinules only so their status will remain questionable. clustering around the anus (Dubois, 1987 LANKANECTINAE (1 GENUS,1SPECIES): This ``1986''). Regardless, this is a large taxo- subfamily was named for Lankanectes cor- nomic change (from Paini to Limnonectini) rugatus of Sri Lanka by Dubois and Ohler and because no evidence was produced or (2001). Its distinguishing features are (1) discussed to justify this change, we must forked omosternum (plesiomorphy); (2) vo- consider the status of this taxon questionable. merine teeth present (presumed plesiomor- The third is Ombrana Dubois, 1992, whose phy); (3) median lingual process absent (like- type and only species is Rana sikimensis Jer- ly plesiomorphy); (4) femoral glands absent don, 1870). Ombrana Dubois, 1992, was (likely plesiomorphy); (5) tips not en- 90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 41. Consensus of two parsimony trees of Chinese ranids from Jiang and Zhou (2005). Data were 1,005 bp of the mtDNA sequences of the 12S and 16S rRNA gene fragments (tree length ϭ 1485, ci ϭ 0.449). Sequences were aligned using ClustalX (Thompson et al., 1997), with manual modi®cations made subsequently. Gaps and ambiguously aligned sequences excluded from analysis. Generic names in parentheses re¯ect alternative usages. Generic taxonomy is updated to recognize Quasipaa (Jiang et al., 2005). larged (arguable polarity); (6) tarsal fold pre- nectes is far from Limnonectes, where it had sent (likely plesiomorphy at this level); and been placed by Dubois (1992). Roelants et (7) lateral line system present in adults (also al. (2004) placed it as the sister taxon of in Phrynoglossus and Euphlyctis, but pre- Nyctibatrachinae, and Delorme et al. (2004) sumably apomorphic). Roelants et al. (2004; placed it as the sister taxon of Nyctibatra- ®g. 35) and Delorme et al. (2004; ®g. 40) chinae ϩ Raninae. We sampled the sole spe- subsequently suggested on the basis of cies, . mtDNA and nuDNA evidence that Lanka- MICRIXALINAE (1 GENUS,11SPECIES): Trop- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 91

NYCTIBATRACHINAE (1 GENUS,12SPECIES): Nyctibatrachinae contains the Indian taxon and is characterized by hav- ing a forked omosternum (likely plesiom- orphic), vomerine teeth present, digital discs present, femoral glands present (shared with Ranixalinae and some Dicroglossinae) and an aquatic tadpole with a keratodont formula of 0/0 (likely apomorphic; Dubois et al., 2001). Of this taxon we sampled Nyctibatra- chus cf. aliciae and N. major. PETROPEDETINAE (2 GENERA,10SPECIES); PHRYNOBATRACHINAE (4 GENERA,72SPECIES) AND PYXICEPHALINAE (13 GENERA,57SPE- CIES): Until recently, members of Petrope- detinae and Phrynobatrachinae, as well as several genera now assigned to Pyxicephali- nae (e.g., , Arthroleptella, Cacosternum, Microbatrachella, Nataloba- trachus, Nothophryne, and ) were considered members of ``Petropedetidae'' (sensu lato), aggregated on the basis of over- Fig. 42. Consensus of four parsimony trees of all similarity, with no evidence for its mono- Paini by Jiang et al. (2005), based on 796 bp (of phyly ever suggested. Noble (1931) recog- which 174 were parsimony informative) of the nized his Petropedetinae ( and 12S and 16S rRNA framents of mtDNA. Se- Petropedetes), as united by the possession of quences were aligned using ClustalW (Thompson dermal scutes on the upper surface of each et al., 1994), cost functions not disclosed, with digit and otherwise corresponding osteolog- subsequent manual modi®cations. Gaps and am- ically and morphologically with Raninae. biguously aligned sequences were excluded from Noble (1931) also recognized Cacosterninae analysis (ci ϭ 0.584, ri ϭ 0.571). The trees were rooted on Hoplobatrachus chinensis and Fejer- for Cacosternum and Anhydrophryne, united varya fujianensis. A conclusion of Jiang et al. by lacking a clavicle and having palatal ridg- (2005) is that their Group 2 was recognized as es. He related the cacosternines to brevicip- Quasipaa. itines, and the remainder of the genera then named he allocated to Raninae. Laurent (1941 ``1940'') addressed the con- ical Asian Micrixalus (11 species) is the sole fusion between Arthroleptis and Phrynoba- member of this taxon, diagnosed by Dubois trachus and transferred Petropedetes, Anhy- (2001) as differing from Dicroglossinae in drophryne, Phrynobatrachus (including Na- lacking a forked omosternum (possibly apo- talobatrachus), Dimorphognathus, and Ar- morphic), lacking vomerine teeth, having throleptella into his Phrynobatrachinae. digital discs (present in some limnonectines Laurent (1941) subsequently provided an an- and otherwise widespread in Ranoidea) and atomical characterization of the group. having a larval keratodont formula in its Laurent (1951) transferred Cacosterninae aquatic tadpoles of 1/0 (likely apomorphic) into Ranidae and moved Microbatrachella (Dubois et al., 2001). On the basis of mtDNA into Cacosterninae. Poynton (1964a) sug- and nuDNA evidence, Roelants et al. (2004; gested that Phrynobatrachus is deeply para- ®g. 35) considered Micrixalinae to be the sis- phyletic with respect to Cacosterninae and ter taxon of Ranixalinae. We were able to therefore considered Laurent's Phrynobatra- sample and M. kottigehar- chinae (ϭ Petropedetinae) and Cacosterninae ensis. Although this provides only a minimal to be synonyms. Subsequent authors (e.g., test of the monophyly of Micrixalus, it al- Dubois, 1981; Frost, 1985) uncritically fol- lows us to place the taxon phylogenetically. lowed this unsupported suggestion, although 92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 there have been signi®cant instances of taxon was stated to be Anhydrophryne, Ar- workers continuing to recognize Cacosterni- throleptella, Cacosternum, Microbatrachel- nae and Petropedetinae as distinct (e.g., la, Nothophryne, Poyntonia (from Petrope- Liem, 1970; J.D. Lynch, 1973). detidae), and, possibly Strongylopus and To- Another morphologically compact African mopterna (from Ranidae). group was Pyxicephalinae (Dubois, 1992), Van der Meijden et al. (2005; ®g. 36) sug- composed of Pyxicephalus (2 species) and gested Phrynobatrachus to be the sister tax- (2 species). The taxon was diagnosed on of . On this basis Dubois by at least four synapomorphies (Clarke, (2005) recognized a ranid subfamily Phry- 1981): (1) cranial exostosis; (2) occipital ca- nobatrachinae, containing Phrynobatrachus, nal present in the frontoparietal; (3) zygo- but also allocated to this subfamily, without matic ramus being much shorter than otic ra- discussion, Dimorphognathus, Ericabatra- mus; and (4) sternal style a long bony ele- chus, and Phrynodon. Petropedetes and Con- ment tapering markedly from anterior to pos- raua formed successively more distant out- terior. Dubois' (1992) reasoning for groups of the southern African clade of Van excluding this taxon from Dicroglossinae is der Meijden et al. (2005), so Dubois (2005) not clear, but presumably had to do with the removed Conrauini (Conraua) from Dicog- distinctive appearances of Pyxicephalus and lossinae and placed it in its own subfamily, Aubria. Conrauinae, and recognized Petropedetinae Dubois (1992) also recognized a subfam- for Petropedetes, as well as the presumably ily Tomopterninae, for Tomopterna (sensu closely allied Arthroleptides. The southern lato, at the time including Sphaerotheca, now African clade of Van der Meijden et al. in Dicroglossinae, Limnonectini). The diag- (2005; ®g. 36) was composed of Cacoster- nosis provided by Clarke (1981) presumably num (formerly of Petropedetidae), Afrana applies inasmuch as he examined only Afri- and Strongylopus (formerly of Raninae), Na- can species (Tomopterna, sensu stricto), even talobatrachus (formerly of Petropedetidae), though the optimization of these characters Tomopterna (Tomopterninae), and Pyxice- on his cladogram may well be contingent on phalus (Pyxicephalinae), a group that Dubois being compared only with other African ra- (2005) allocated to an enlarged Pyxicephal- nids: (1) zygomatic ramus much shorter than inae. Aubria was asserted by Dubois (2005) otic ramus; (2) outline of anterior end of cul- to be in this group because it was grouped triform process pointed, with lateral borders by morphological evidence with Pyxicephal- tapering to a point; (3) distal end of the an- us. Amietia he transferred into the group terior pterygoid ramus overlapping the dorsal without discussion, but presumably because surface of the posterior lateral border of the they appeared to him to be related to Stron- palatine; (4) no overlap of the anterior border gylopus and Afrana. He transferred Arthro- of the parasphenoid ala by the medial ramus leptella, Microbatrachella, Nothophryne, and of the pterygoid in the anterior±posterior Poyntonia into Pyxicephalinae, presumably plane; (5) sternal style short, tapering poste- because he thought that they were more like- riorly; (6) dorsal protuberance of the ilium ly to be here than close to either Petropede- not or only slightly differentiated from the tinae or Phrynobatrachinae. spikelike dorsal prominence; and (7) terminal Of Dubois' (2005) Petropedetinae (which phalanges of the ®ngers and reduced, presumably is diagnosed as by Noble, 1931) almost conelike. we were able to sample both genera: Arthro- In 2003 this untidy, but familiar arrange- leptides sp. and Petropedetes cameronensis, ment began to unravel. Dubois (2003), re- P. newtoni, P. palmipes, and P. parkeri. moved Cacosterninae from ``Petropedetidae'' Of the newly constituted Phrynobatrachi- without discussion, apparently anticipating nae, we were also able to sample species evidence to be published elsewhere, although from three of four genera: Dimorphognathus Kosuch et al. (2001; ®g. 38) had suggested africanus, Phrynobatrachus auritus, P. cal- earlier that Cacosternum was more closely caratus, P. dendrobates, P. dispar, P. ma- related to Tomopterna and Strongylopus than babiensis, P. natalensis, and Phrynodon san- it was to Petropedetes. The content of this dersoni. We did not sample , 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 93 an unfortunate omission, inasmuch as we are The three nominal genera in the taxon are unaware of the evidence for Dubois' (2005) Ptychadena (47 species), (3 association of Ericabatrachus with Phryno- species), and Lanzarana (1 species) of which batrachinae, other than the statement that it we sampled only Ptychadena anchietae, P. is ``Phrynobatrachus-like'' (Largen, 1991). cooperi, and P. mascareniensis. Because we Phrynobatrachus, at least for the species did not sample Hildebrandtia and Lanzar- which it is known, have exotrophic larvae. ana, we did not adequately test the mono- Larvae are unknown in Dimorphognathus phyly of this group. Nevertheless, assuming and Ericabatrachus, and Phrynodon is en- the group to be monophyletic, our three spe- dotrophic (Amiet, 1981; Altig and Mc- cies of Ptychadena allow us to test the place- Diarmid, 1999). ment of Ptychadeninae within Ranoidea. For Of the reformulated Pyxicephalinae we his analysis Clarke (1981) assumed that Pty- were able to sample Aubria (Aubria subsi- chadeninae is imbedded within other African gillata [2 samples21]) and Pyxicephalus (Py- ranids, although a lack of comparison with xicephalus edulis) as well as several of the Asian members of the group makes this as- taxa recently transferred into this taxon in- sumption questionable. Van der Meijden et cluding Anhydrophryne rattrayi, Arthrolep- al. (2005; ®g. 36) suggested that Ptychadena tella bicolor, Cacosternum platys, and Na- is the sister taxon of Phrynobatrachus among talobatrachus bonebergi. We also sampled his exemplars, thereby implying that Pty- members of Afrana (A. angolensis and A. chadeninae is the sister taxon of Phrynoba- fuscigula), Tomopterna (T. delalandii), trachinae. Strongylopus (S. grayii), and Amietia (A. ver- ``RANINAE'' (CA.8GENERA, 309 SPECIES): tebralis), but for reasons having to do with ``Raninae'' is a catch-all largely Holarctic the evidentiary basis and history of taxono- and tropical Asian taxon united because the my in Raninae, considerable discussion of members do not ®t into the remaining sub- these genera is presented there. We did not families and have unforked omosterna. Until sample Microbatrachella, Nothophryne,or recently, ``Raninae'' included two tribes: Pai- Poyntonia. Pyxicephalines have exotrophic ni and Ranini (Dubois, 1992). However, Pai- larvae, with the exception of Anhydrophryne ni and Nanorana of Ranini were transferred and Arthroleptella, which are endotrophic; to Dicroglossinae on the basis of mtDNA and unknown in Nothophryne (Hewitt, 1919; nuDNA evidence (Roelants et al., 2004Юg. Procter, 1925; DeVilliers, 1929; Altig and 35; Jiang et al., 2005Юg. 42), so Raninae, McDiarmid, 1999). This selection should al- as we use it, is coextensive with Ranini of low us to test the phylogenetic results of Van Dubois (1992), itself dubiously monophylet- der Meijden et al. (2005). ic22. PTYCHADENINAE (3 GENERA,51SPECIES): ``Raninae'' is distributed on the planet co- Ptychadeninae is a morphologically compact extensively with the family and is united by group of sub-Saharan ranids diagnosed the lack of putative apomorphies, either in (Clarke, 1981; Dubois, 1987 ``1985'', 1992) the adult or in the larvae. There does not by having: (1) an otic plate of the squamosal appear to be any reason to suggest that this covering the crista parotica in dorsal view nominal taxon is monophyletic. and extending mesially to overlap the otoc- The starting point of any discussion of cipital; (2) palatines absent; (3) clavicles re- Ranini must be Dubois (1992), who provided duced; (4) sternal style a short compact ele- an extensive, and controversial, taxonomy. ment tapering anteriorly to posteriorly; (5) Because the distinction between ranks (sec- eighth presacral fused with sacral vertebra; and (6) the dorsal protuberance of 22 Van der Meijden et al. (2005; ®g. 36), provided ev- ilium smooth-surfaced and not prominent. idence from DNA sequences that suggests strongly that ``Raninae'' is polyphyletic, with at least Afrana and Strongylopus in a southern African clade (along with 21 We included two specimens of Pyxicephalus, Tomopterna, Natalobatrachus, and Ca- as separate terminals in the analysis because the identity costernum), far from other ranines, and in Pyxicephali- of one of the specimens was not determined conclusive- nae of Dubois (2005). We therefore treat ``Raninae'' in ly until after the analyses were complete. the following discussion as dubiously monophyletic. 94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 tion, subsection, genus, and subgenus) in Du- pole morphology (presence of a raised, bois' system appears to rest primarily on sub- sharply de®ned abdominal ). Like oth- jective perceptions of similarity and differ- er cascade-dwelling taxa, larvae of Amolops ence, the evidentiary basis of this taxonomy (sensu lato) all share high numbers of kera- is unclear, even though we accepted his sys- todont rows. Subsequently, Yang (1991b) tem as a set of bold phylogenetic hypotheses. recognized two other genera from within Nevertheless, most of these taxa are imper- Amolops: and Huia. Amolops fectly or incompletely diagnosed and to lay (sensu stricto) has one possible synapomor- the foundation for our results and concomi- phy (short ®rst metacarpal, also found in tant taxonomic remedies, we discuss this tax- Huia), and three synapomorphies joining onomy in greater depth than we do most of Huia and Meristogenys to the exclusion of the remainder of current amphibian taxono- Amolops (lateral glands present in larvae; my. Suf®ce it to say that we think that we four or more uninterrupted lower labial ker- sampled ``Rana'' diversity suf®ciently to atodont rows; and longer legs). provide at least a rudimentary phylogenetic Subsequently, Dubois (1992) treated Mer- understanding of the taxon as a starting point istogenys and Huia as subgenera of Amolops, for future, more densely sampled studies. and added a fourth subgenus, Amo (including Within his Ranini, Dubois (1992) recog- only ). Amo was diag- nized six genera: Amolops, Batrachylodes, nosed (Boulenger, 1918) as having a digital Nanorana, Micrixalus, Rana, and Staurois disc structure similar to species of Staurois (table 4). Of these, two continue to be placed (i.e., having a transverse groove or ridge on in this taxon (Amolops and Rana [sensu the posteroventral side of the disc continuous lato]) (Dubois, 2005). Staurois, Nanorana with a circummarginal groove to de®ne a and Micrixalus have subsequently been hemisphere; Boulenger, 1918) and as having transferred out of Ranini, Staurois to a new axillary glands (after Yang, 1991b) that are tribe, Stauroini (Dubois, 2005), Nanorana to otherwise unknown in Amolops. Dicroglossidae (Roelants et al., 2004; ®g. Although Dubois (1992) considered Amo- 35), and Micrixalus to a distant Micrixalinae lops (sensu stricto), Amo, Huia, and Meris- (Dubois et al., 2001). Batrachylodes was pro- togenys to be subgeneric parts of a mono- visionally transferred, without substantial phyletic genus Amolops, other authors (e.g., discussion, by Dubois (2005) to Ceratobatra- Yang, 1991b) considered at least Amolops, chinae. Huia, and Meristogenys as genera. For con- Within both Amolops and Rana, Dubois sistency we treat as genera Amo, Amolops, recognized several subgenera, that other au- Huia, and Meristogenys. Our samples were thors (e.g., Yang, 1991b) considered to be Amolops (A. chapaensis, A. hongkongensis), genera, as we do, although we arrange the Huia (H. nasica), and Meristogenys (M. or- discussions by Dubois' genera and subgen- phocnemis). We were unable to sample Amo era. Dubois (2003) arranged Raninae into larutensis. two tribes (Amolopini for the taxa with cas- Staurois: The de®nition of Staurois (digi- cade-adapted tadpoles, i.e., Amo, Amolops, tal discs broader than long; T-shaped termi- Huia, Meristogenys, , Eburana, nal phalanges in which the horizontal part of Odorrana) and Ranini (for everything else). the T is longer than the longitudinal part; This system represents typical nonevolution- outer metatarsals separated to base but joined ary A and not-A groupings, although Amo- by webbing; small nasals separated from lopini in this form is testable. Dubois (2005) each other and frontoparietal; omosternal subsequently did not embrace Amolopini, style not forked [Boulenger, 1918]) has also because it was too poorly understood, but he been used to de®ne Hylarana (Boulenger, did erect Stauroini for Staurois, because Roe- 1920; see below). Although some larval lants et al. (2004) placed Staurois as the pu- characters are shared among species of Stau- tative sister taxon of other ranines. rois (deep, cup-like oral disc in the tadpole, Amolops, Amo, Huia, and Meristogenys: no glands or abdominal disc in tadpole; In- Amolops has been recognized in some form ger, 1966), the diagnostic value of these char- since Inger (1966) noted the distinctive tad- acters is unknown due to the large number 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 95 of ranid species whose adults are morpho- (Inger, 1996) that are not based on any com- logically similar to those of Staurois, but prehensive comparative study of either inter- whose larvae remain undescribed. Our single nal or external morphology. For instance, lar- exemplar of Staurois, S. tuberilinguis, is not vae may have dorsal dermal glands, lateral suf®cient to test the monophyly of the genus. dermal glands, or ventral dermal glands in Although no one has suggested that Staurois various combinations (e.g., Yang, 1991b). is polyphyletic, or that it is paraphyletic with These characters have become larval dermal respect to any other group, both of these re- glands present or absent in Dubois' (1992) main untested possibilities. Roelants et al. diagnoses, thereby con¯ating the positional (2004; ®g. 35) provided evidence that Stau- homology of these features. Although we ad- rois is the sister taxon of remaining ranines. dress de®ciencies here and in the Taxonomy Rana (sensu Dubois, 1992)23: Rana of Du- section, for other critiques see Emerson and bois (1992) is diagnostically coextensive Berrigan (1993), Matsui (1994), Matsui et al. with his Ranini (our ``Raninae''), and no fea- (1995), Inger (1996), Bain et al. (2003), and tures provided in his paper exclude ``Rana'' Matsui et al. (2005). from being paraphyletic with respect to Stau- As noted earlier, several, if not most taxa rois, Amolops (sensu Dubois, 1992), or Ba- recognized by Dubois within his ``Rana'' are trachylodes. So, as we discuss the internal effectively undiagnosed in a utilitarian sense taxonomy of ``Rana'' as provided by Dubois, (i.e., they are diagnosed suf®ciently only to readers should bear in mind that Amolops make the names available under the Inter- (sensu lato), Batrachylodes, and Staurois,as national Code; ICZN, 1999). In addition, discussed by Dubois (1992), must be regard- several are demonstrably nonmonophyletic ed as potential members of all infrageneric (Matsui, 1994; Matsui et al., 1995; Inger, taxa that do not have characters that specif- 1996; Tanaka-Ueno et al., 1998a; Emerson et ically exclude them. (And, at least with re- al., 2000a; Marmayou et al., 2000; Vences et spect to Dubois', 1992, Rana subgenera, al., 2000a; B.J. Evans et al., 2003; Roelants Strongylopus and Afrana, DNA sequence et al., 2004; Jiang and Zhou, 2005). Unlike data have been published that suggest that the super®cially similar situation in Eleuth- they have little relationship with other rani- erodatylus (sensu lato) where it is straight- nes [Van der Meijden et al., 2005; ®g. 36].) forward to get speci®c information on indi- With respect to ``Rana'' speci®cally, Dubois vidual species and where the nominal sub- (1992) provided a system of sections, sub- genera and most related genera, even if they sections, and subgenera that has posed seri- do not rise to the level of synapomorphy ous challenges for us: Rather than a syna- schemes, have been diagnosed largely com- pomorphy scheme, or even a system of care- paratively, the subgeneric (and generic, in fully-evaluated characteristics, the various part) diagnoses of ranids are not comparable, taxa appear to represent postfacto character and the purported differentiating characters justi®cations of decidedly nonphylogenetic frequently do not bear up to specimen ex- and subjectively arrived-at groups. We found amination (e.g., Tschudi, 1838; Boulenger, Dubois' (1987 ``1985'', 1992) arrangement 1920; Yang, 1991b; Fei et al., 1991 ``1990''; to be inconsistent with the preponderance of Dubois, 1992). evidence in certain instances (see the discus- Historically, taxonomists approached sion of inclusion of Aquarana in his section Rana (sensu lato) as being composed of two Pelophylax, below) and the underlying di- very poorly de®ned similarity groupings: (1) agnostic basis of the system to contain over- those that have expanded toe tips (likely ple- ly-generalized statements from the literature siomorphic) that at one time or another have been covered by the name Hylarana; and (2) 23 Although Afrana, Amietia, and Strongylopus (now those that lack expanded toe tips, and that in Pyxicephalinae), Batrachylodes (now in Ceratoba- have more-or-less always been associated trachidae), Micrixalus (now in Micrixalinae), and Na- with the generic name Rana. Most authors norana (now in Dicroglossinae) have been transferred out of Raninae, we address them as part of the general since Boulenger (1920) recognized the lack discussion of ranine systematics prior to 2004. (See table of de®nitive ``breaks'' between the two 4) groups, and Dubois was the ®rst to attempt 96 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

TABLE 4 Generic and Subgeneric Taxonomy of Dubois' (1992) Ranini Nanorana transferred to Paiini by Roelants et al. (2004); Batrachylodes transferred to Ceratobatrachinae, without discussion of evidence by Dubois (2005); Micrixalus transferred to a new subfamily, Micrixalinae, by Dubois (2001); Afrana and Strongylopus transferred to Pyxicephalinae by Dubois (2005), based on evidence presented by Van der Meijden et al. (2005); and Staurois transferred to a new tribe, Stauroini, by Dubois (2005).

Number of Species sampled (re¯ecting Genus Section Subsection Subgenus species nomenclature used in this work) Amolops Amolops 22 Amolops chapaensis, A. hongkongensis Amolops Amo 1 Not sampled Amolops Huia 4 Huia nasica Amolops Meristogenys 8 Meristogenys orphnocnemis Batrachylodes 8 Batrachylodes vertebralis Micixalus 6 Micrixalus fuscus, M. kottigeharensis Nanorana Altirana 1 Not sampled Nanorana Nanorana 2 Nanorana pleskei Rana Amerana Amerana 2 Amerana muscosa Rana Amerana Aurorana 4 Aurorana aurora Rana Amietia Amietia 2 Rana Babina Babina 2 Not sampled Rana Babina Nidirana 6 Nidirana adenopleura, N. chapaensis Rana Hylarana Hydrophylax Amnirana 9 Amnirana albilabris Rana Hylarana Hydrophylax Humerana 3 Not sampled Rana Hylarana Hydrophylax Hydrophylax 2 Hydrophylax galamensis Rana Hylarana Hydrophylax Papurana 11 Papurana daemeli Rana Hylarana Hydrophylax 10 Not sampled Rana Hylarana Hydrophylax Sylvirana 21 Sylvirana guentheri, S. maosonensis, S. nigrovittata, S. temporalis Rana Hylarana Hylarana Chalcorana 9 Chalcorana chalconata Rana Hylarana Hylarana Clinotarsus 1 Clinotarsus curtipes Rana Hylarana Hylarana Eburana 5 Eburana chloronota Rana Hylarana Hylarana Glandirana 1 Glandirana minima Rana Hylarana Hylarana Hylarana 3 Hylarana erythraea, H. taipehensis Rana Hylarana Hylarana Nasirana 1 Not sampled Rana Hylarana Hylarana Odorrana 10 Odorrana grahami Rana Hylarana Hylarana Pterorana 1 Not sampled Rana Hylarana Hylarana Sanguirana 2 Not sampled Rana Hylarana Hylarana Tylerana 2 Tylerana arfaki Rana Lithobates Lithobates 3 Lithobates palmipes Rana Lithobates Sierrana 3 Sierrana maculata Rana Lithobates Trypheropsis 2 Trypheropsis warszewitschii Rana Lithobates Zweifelia 5 Not sampled Rana Pelophylax Aquarana 7 Aquarana catesbeiana, A. clamitans, A. grylio, A. heckscheri Rana Pelophylax Pantherana 22 Pantherana berlandieri, P. capito, P. chiricahuensis, P. forreri, P. pipiens, P. yavapaiensis Rana Pelophylax Pelophylax 17 Pelophylax nigromaculata, P. ridibunda Rana Pelophylax Rugosa 3 Not sampled Rana Pseudorana Pseudorana 3 Pseudorana johnsi Rana Rana Rana 27 Rana japonica, R. sylvatica, R. temporaria Rana Strongylopus Afrana 8 Afrana angolensis, Afrana fuscigula Rana Strongylopus Strongylopus 6 Strongylopus grayii Staurois 4 Staurois tuberlinguis 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 97 to summarize the relevant taxonomic litera- ture and to divide Rana (sensu lato) into enough groups to allow some illumination of the problem. Our issue with his system is that it is impossible to tell from the relevant pub- lication (Dubois, 1992) which species have actually been evaluated for characters and which have merely been aggregated on the basis of overall similarity or erected on the basis of specially-favored characters. Dubois' primary division of Rana was into eight sections of arguable phylogenetic pro- pinquity to each other or to other ranine gen- era (see table 4). We discuss these with ref- erence to his diagnoses and other literature relevant to their recognition: (1) Section Amerana. Dubois (1992) erect- ed his subgenera Amerana and Aurorana for parts of the Rana boylii group of Zweifel (1955), which he placed in their own section, Amerana. Most previous work (e.g., Case, 1978; Farris et al., 1979; Post and Uzzell, 1981; Farris et al., 1982b; Uzzell and Post, 1986) had placed these frogs from western North American close to, or within, the Eur- asian Rana temporaria group. Nevertheless, section Amerana was recognized by Dubois Fig. 43. Restriction-site tree of exemplars of (1992) on the basis of a combination of char- Holarctic Rana of Hillis and Davis (1986). Un- acters, none unique but corresponding to the derlying data were restriction sites of the nuclear Rana boylii group identi®ed by ribosomal rDNA gene; presence was considered to be evi- data by Hillis and Davis (1986; ®g. 43). This dence of relationship, absence was not. The tree group had been suggested by Hillis and Da- was rooted on Pyxicephalus and Pelophylax (as vis (1986) to be in a polytomy with what Rana ridibunda). The original ®gure treated all Dubois regarded as his section Rana (R. tem- species, save Pyxicephalus, as members of Rana. poraria and R. sylvatica were the exemplar We have noted on the right the nominal subgenera species in their analysis), a group composed of Dubois (1992; which we have treated as gen- of a part of Dubois' section Pelophyax era), to clarify discussion. (Aquarana), and his sections Lithobates and Pantherana. Moreover, Hillis and Davis' monophyletic group with Rana temporaria, (1986; ®g. 43) results suggested that neither to the exclusion of all other North American of the groups subsequently identi®ed by Du- Rana, inasmuch as this was an assumption of bois (1992) as the subgenera Aurorana and their analysis, based on earlier work (e.g., Amerana are monophyletic. Subsequent Case, 1978). work (Hillis and Wilcox, 2005; ®g. 44) has Dubois (1992) provided no unique mor- provided substantial amounts of evidence in phological features to diagnose section support of the nominal subgenus Aurorana Amerana, and because of his use of present- being polyphyletic, and the subgenus Amer- or-absent as a characteristic, the characters ana being paraphyletic. Hillis and Wilcox provided in his table 1 fail to rigorously dis- (2005) used the section Amerana ϩ Rana tinguish section Amerana from sections Hy- temporaria to root the remainder of their larana, Lithobates, Pelophylax, Rana,or tree, so their overall tree cannot be taken as Strongylopus (now in Pyxicephalinae on the additional evidence of evolutionary propin- basis of DNA sequence evidenceÐDubois, quity of the section Amerana being in a 2005; Van der Meijden et al., 2005). Within 98 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 44. Maximum-likelihood tree of Holarctic Rana of Hillis and Wilcox (2005). The underlying data are ca. 2kb of mtDNA of the 12S±16S region (spanning the tRNAVal gene). Sequence alignment was done initially using Clustal W (Thompson et al., 1994), costs not disclosed, and manually adjusted, guided by assumed secondary structure, ambiguously aligned sequences discarded. It was not stated whether gaps were treated as data, but we presume not. Substitution model GTR ϩ⌫ϩPINVAR was assumed for the maximum-likelihood analysis. On the basis of previous research, the root was assumed to be between the Rana temporaria ϩ Rana boylii group and the remainder of New World Rana. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 99

Amerana, Dubois recognized two subgenera, their taxonomy was presented for only the Amerana and Aurorana, differing in the ex- Chinese fauna, so the wider implication of pansion of toe tips (mildly expanded in this action is not known. Of this section we Amerana; not expanded in Aurorana), rows sampled no member of the subgenus Babina, of larval keratodonts (4±7/4±6 in Amerana; although we did sample Nidirana adenopleu- 2±3/3±4 in Aurorana) karyotype (derived in ra and N. chapaensis. Babina and Nidirana Amerana; primitive in Aurorana). This sub- have also been associated with ``Hylarana'' generic distinction is not phylogenetically (see below), so Dubois' (1992) reason for consistent with the results of Hillis and Davis recognizing this as a section distinct from (1986; ®g. 43), who presented evidence sug- section Hylarana is unclear. gesting that Dubois' Aurorana is paraphylet- (4) Section Lithobates. This section is not ic with respect to his Amerana (making one rigorously diagnosable by the features pre- wonder what the purpose was in naming two sented by Dubois' (1992: his table 1) from subgenera). Macey et al. (2001) subsequently sections Amerana, Hylarana, Rana,orStron- provided additional molecular evidence for gylopus. However, Lithobates is consistent paraphyly of Aurorana with respect to Amer- with the phylogenetic tree of American Rana ana. Examples of this section in our analysis provided by Hillis and Davis (1986; ®g. 43), are Amerana muscosa and Aurorana aurora presumably the source of the concept of this (see table 4). section. Hillis and Davis placed this taxon, (2) Section Amietia (including a single on the basis of DNA substitutions, as the sis- subgenus, Amietia, for two species in the Le- ter taxon of part of Dubois' section Pelophy- sotho Highlands of southern Africa). The lax, the subgenus Pantherana. Within section sole synapomorphy of Amietia is the umbra- Lithobates, Dubois recognized four subgen- culum over the eye in the larva. The diag- era: Lithobates (Rana palmipes group), Sier- nosis of section Amietia is otherwise phylo- rana (Rana maculata group), Trypheropsis genetically indistinguishable on the basis of (Rana warszewitschii group), and Zweifelia the table of characters provided by Dubois (Rana tarahumarae group). All of them are (1992), from Amerana, Hylarana, Lithoba- consistent with the tree provided by Hillis tes, Rana,orStrongylopus. We sampled and Davis (1986). Dubois (1992) offered the Amietia vertebralis. Amietia was transferred following morphological characters which into Pyxicephalinae by Dubois (2005) on the apparent but undiscussed assumption that it may be synapomorphies: Lithobates differs is closely related to Strongylopus, which was from other members of the section by having placed by Van der Meijden et al. (2005) in tympanum diameter larger or equal to the di- that group on the basis of DNA sequence ev- ameter of the eye; Sierrana without diagnos- idence. tic characters that differentiate it from the (3) Section Babina (for the Rana holsti section diagnosis; Trypheropsis by having an and Rana adenopleura groups). The unique outer metatarsal tubercle (unusual in Ameri- synapomorphy for this group is a large ``su- can ranids); and Zweifelia with sacrum not prabrachial'' gland (sensu Dubois, 1992) on fused with presacral vertebrae. Hillis and the sides of reproductive males (which can Wilcox (2005; ®g. 44) presented evidence be dif®cult to assess in nonreproductive an- that suggests that section Lithobates of Du- imals). The diagnosis of section Babina does bois (1992) is paraphyletic, with part of Du- not otherwise allow it to be practically sep- bois' subgenera Sierrana (R. maculata), and arated from the sections Amerana, Hylarana, all of his subgenera Trypheropsis, and Lith- Lithobates, Pelophylax, Rana,orStrongylo- obates falling within one monophyletic pus. Within section Babina, Dubois recog- group, but Zweifelia (the Rana tarahumarae nized two subgenera, Babina (with a large group) and another part of Sierrana (R. sier- ®ngerlike prepollical spine, an apomorphy) ramadrensis) forming the sister taxon of Du- and Nidirana (members of the Babina sec- bois' subgenus Pantherana, the Rana pipiens tion lacking the apomorphy of the subgenus group of Hillis and Wilcox (2001). Babina). Fei et al. (2005) considered Nidi- Our exemplars of this section are Lithob- rana to be a subgenus of their Hylarana, but ates palmipes, Sierrana maculata, and Try- 100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 pheropsis warszewitschii. We did not sample sidered Pelophylax and Rugosa to be a dis- Zweifelia. tinct genera, but these authors generalized (5) Section Pelophylax. The characters solely over the Chinese fauna rather than at- provided by Dubois for his section Pelophy- tempting to draw global distinctions. From lax will not rigorously diagnose it from Aquarana (Rana catesbeiana group) we Amerana, Hylarana, Rana,orStrongylopus. sampled Aquarana catesbeiana, A. clami- Further, the association of his subgenera tans, A. grylio, and A. heckscheri.OfPanth- Aquarana (former Rana catesbeiana group), erana (Rana pipiens group) we sampled Pantherana (former Rana pipiens group), Pantherana berlandieri, P. capito, P. chiri- Pelophylax (former Rana ``esculenta'' cahuensis, P. forreri, P. pipiens, and P. ya- group), and Rugosa (Rana rugosa group) is vapaiensis.OfPelophylax we sampled R. ni- curious inasmuch as we are unaware that gromaculata and P. ridibunda. We did not anyone had previously suggested such a re- sample Rugosa. lationship. All published evidence that was (6) Section Pseudorana. This section can- available to Dubois at the time of his writing not be rigorously diagnosed on the basis of (e.g., Case, 1978; Post and Uzzell, 1981; Hil- information given by Dubois (1992) from lis and Davis, 1986; Pytel, 1986; Uzzell and section Hylarana. Pseudorana was named by Post, 1986) suggested that this section is Fei et al. 1991 ``1990'') as a distinct genus polyphyletic, with Dubois' subgenus Panth- for , R. sangzhiensis, and R. erana (of his section Pelophylax) more weiningensis. Subsequently, Fei et al. (2000) closely related to his section Lithobates, than coined Pseudoamolops for Rana sauteri, to any other member of section Pelophylax. suggesting, on the basis of its having a large Indeed, the subgenera Aquarana and Panth- ventral sucker on the tadpole, that it is more erana of Pelophylax are both more closely closely related to Amolops (sensu lato) than related to both the sections Lithobates, Rana, to Pseudorana. Although the ventral sucker and Amerana, than they are to the Old World found in Pseudoamolops is associated with members of section Pelophylax according to the oral disc of the tadpole, in Amolops the the evidentiary literature (i.e, Case, 1978; ventral sucker sits posterior to the oral disc. Post and Uzzell, 1981; Hillis and Davis, Fei et al. (2000) suggested that Pseudoamo- 1986; Pytel, 1986; Uzzell and Post, 1986). lops is the sister taxon of the remainder of There never was any evidence for the mono- their Amolopinae (Amo, Amolops, Huia, and phyly of section Pelophylax sensu Dubois, Meristogenys) and derived with respect to a while there was considerable evidence paraphyletic Hylarana, although Tanaka- against it. Recently, Hillis and Wilcox (2005; Ueno et al. (1998a) had previous suggested ®g. 44) have provided molecular evidence on the basis of DNA sequence analysis that that Aquarana (their Rana catesbeiana Pseudorana sauteri is imbedded within the group) is the sister taxon of Rana sylvatica, brown frog clade (Rana temporaria group), and together the sister taxon of all other although that analysis had addressed no American Rana, with the exception of the member of nominal Amolopinae. We were section Amerana (their Rana boylii group). able to sample Pseudoamolops sauteri and The subgenera recognized by Dubois Pseudorana johnsi to test the placement of within section Pelophylax have more justi®- these species. cation for their monophyly. Aquarana is dis- (7) Section Rana. This section cannot be tinct on the basis of its large snout±vent diagnosed rigorously from sections Amer- length and its tympanum diameter, which is ana, Hylarana, Lithobates, Pelophylax,or greater than eye diameter in males. Rugosa Strongylopus on the basis of characters pre- is separated by its ``small'' adult snout±vent sented by Dubois (1992). The association of length. Pantherana and Pelophylax are sep- Rana sylvatica with the Rana temporaria arated from Aquarana and Rugosa by their group has been controversial, with Hillis and ``medium'' size and spots on the dorsum, but Davis (1986) providing weak evidence for its are otherwise undiagnosable from each other placement with Rana temporaria, and Case by features presented by Dubois (1992). Fei (1978) suggesting that Rana sylvatica is phy- et al. (1991 ``1990'', 2005) consistently con- logenetically within other North American 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 101

Rana (sensu lato). Hillis and Wilcox (2005; Hylarana to the end of this discussion be- ®g. 44) recently provided molecular evidence cause it represents the of the problem in support of Rana sylvatica being the sister of ``Rana'' systematics. The name Hylarana taxon of the Rana catesbeiana group (Aquar- has had an historically unstable application, ana of Dubois, 1992). In addition to noncon- alternatively being considered synonymous troversial members of the Rana temporaria with Rana, or treated as a distinct subgenus group (Rana japonica and R. temporaria)we or genus with an ill-de®ned content, and di- sampled Rana sylvatica to test whether it was agnosed in several different, even contradic- a member of the Rana temporaria group or, tory ways (e.g., Tschudi, 1838; GuÈnther, as suggested previously, imbedded within a 1859 ``1858''; Boulenger, 1882, 1920; Perret, North American clade. 1977; Poynton and Broadley, 1985; Laurent, (8) Section Strongylopus. This section also 1986; Fei et al., 1991 ``1990''; Dubois, is not phylogenetically diagnosable on the 1992), although it is almost always associ- basis of Dubois' (1992) suggested evidence ated with frogs that exhibit expanded toe from sections Amerana, Hylarana, Lithoba- tips. The original diagnostic character of the tes, Pelophylax,orRana. If the autapomor- genus Hylarana Tschudi, 1838 (type species: phies of Babina and Amietia are not consid- Rana erythraea Schlegel, 1827) is the pres- ered, there also is nothing in the diagnosis of ence of a dilated disc on the tips of the toes section Strongylopus that would prevent it (a character that can now be seen to encom- from being paraphyletic with respect to Ba- pass many of the species of Ranidae and its bina or Amietia. Nevertheless, DNA se- immediate outgroups). GuÈnther (1859 quence evidence of Van der Meijden et al. ``1858'') revised the diagnosis to include (2005; ®g. 36) places Strongylopus in Pyxi- ``males with an internal subgular vocal sac'' cephalinae, and Dubois (2005) presumed that (i.e., lacking gular pouches) as a character, Afrana and Amietia also should be so allo- and increased the composition to ®ve Asian cated. Section Strongylopus is seemingly a and African species (including Hylarana al- geographically determined unit, not a phy- bolabris and H. chalconota). logenetically determined one. Within section Because of the ambiguity of the diagnostic Strongylopus, Dubois recognized two sub- character of dilated toe disc, Boulenger genera that differ in size and color of larvae (1882, 1920) believed Hylarana to be a (long and dorsally black in Afrana; modest ``group of polyphyletic origin'', but suggest- length and entirely black in Strongylopus), ed that it was a subgenus of Rana, removing foot length (short in Afrana; long in Stron- vocal sac condition as a diagnostic character gylopus), and webbing (less webbing in Af- and expanding its de®nition: dilated digital rana than in Strongylopus). discs with circummarginal grooves, T-shaped Van der Meijden (2005; ®g. 36) provided terminal phalanges, and an unforked omos- a phylogenetic tree, based on mtDNA and ternal style (Boulenger, 1920: 123; as Hylor- nuDNA sequence data, that placed Strongy- ana). All of his putatively diagnostic char- lopus and Afrana in a heterogeneous clade acters have greater levels of generality than (which they termed the ``southern African ra- ``Hylarana''. He listed 62 species from Aus- nid clade'', and which Dubois, 2005, consid- tralasia, including Rana curtipes, R. guenth- ered as an expanded Pyxicephalinae), along eri, and R. taipehensis (the latter implicit, as with Tomopterna (Tomopterninae), Cacos- he synonomized it with R. erythraea; Bou- ternum and Natalobatrachus (``Petropedeti- lenger, 1920: 152±155). dae''), and Pyxicephalus (Pyxicephalinae). Perret (1977: 842) listed ten African spe- Because the evidence of Van der Meijden et cies of the genus Hylarana (including H. gal- al. (2005; ®g. 36) is the ®rst phylogenetic amensis), revising the diagnosis as follows: evidence that bears on this issue, we follow precoracoids ossi®ed, transverse, approach- that taxonomy, but note that nothing in mor- ing each other medially; metasternum ossi- phology so far supports this arrangement. ®ed, elongated; males with or without gular We sampled Afrana angolensis, A. fusci- pouches; males with brachial (humeral) gula, and Strongylopus grayii. glands. Poynton and Broadley (1985: 139) (9) Section Hylarana. We have left section revised the diagnosis in their account of Af- 102 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

group (O. andersoni, O. grahami, O. haina- nensis, and O. margaretae) have large chest spines, with small spines otherwise only in O. schmackeri. Chest spines were reported as absent in all other species of Odorrana that they studied: O. anlungensis, O. exiliversa- bilis, O. hejiangensis, O. kuangwuensis, O. livida, O. lungshengensis, O. nasuta, O. swinhoana, O. tiannanensis, O. versabilis, and O. wuchuanensis. Fei et al. (1991 ``1990'': 138±139) further divided Hylarana into two subgenera, Hylar- ana and Tenuirana based on the following characters (Tenuirana in parentheses): ante- rior process of hyoid long, curved outwards (long, straight); tips of digits with or without a horizontal groove (always present on toes); feet almost fully webbed (half webbed); Fig. 45. Tree of Chinese species of Odorrana body not long or slender (long, slender); of Ye and Fei (2001), based on 29 character trans- snout blunt and rounded (long, pointed); formations of morphology (ci ϭ 0.507). Tree root- limbs moderate (long, slender); dorsolateral ed on Rana japonica and Rana omeimontis. The folds distinct to extremely broad (narrow); subgenera Odorrana and Eburana of Fei et al. humeral gland or shoulder gland present in (2005) are noted on the right and the terminals males (absent); gular pouches present in noted with an asterisk (*) are members of Dubois' (1992) subgenus Eburana. male (absent); and tadpole vent tube dextral (medial). As part of the Chinese fauna, they included R. nigrovittata and R. guentheri (under the subgenus Hylarana) and R. tai- rican Hylarana: only some species with ex- pehensis (the type species of the subgenus panded digital discs; broad brown to golden Tenuirana)inHylarana. Although they did band from head to urostyle; upper lip white; not discuss R. erythraea (the type species of males with single or paired baggy gular pouches. Laurent (1986: 761) further revised Hylarana), its inclusion in the subgenus Hy- the diagnosis of Hylarana: without trans- larana was implied. verse grooves on ®nger discs. As noted earlier, Dubois (1992) partitioned Fei et al. (1991 ``1990'') moved some spe- species formerly associated with one or more cies from Hylarana into a new genus Odor- of the historical manifestations of Hylarana rana. They diagnosed their new genus Odor- into several sections, subsections, and sub- rana by having: omosternum extremely genera (see table 4) of which the sections small, colorless spines present on chest of Babina (subgenera Babina and Nidirana) male in breeding condition. Despite the ety- and Hylarana (subsections Hydrophylax and mology of the generic name, Fei et al. (1991 Hylarana) are particularly relevant to this ``1990''), did not include odoriferous secre- discussion of ``Hylarana''-like frogs (al- tions as one of the characters uniting the ge- though the section Hylarana, in Dubois' sys- nus. In addition, they included six species tem was not precluded by any evidence from (O. anlungensis, O. kwangwuensis, O. swin- being paraphyletic to any or all of the other hoana, O. tiannanensis, O. versabilis, and O. sections de®ned by him). Sections Babina wuchuanensis) known not to have colorless and Hylarana are distinguishable in Dubois' spinules on the chest of the male. Subse- system solely by the possession of a supra- quently, Ye and Fei (2001; ®g. 45), on the brachial gland (apomorphy) in section Ba- basis of a phylogenetic study of Chinese bina. This gland is not found in section Hy- Odorrana (including Eburana in their sense), larana which at least as portrayed by Dubois suggested that only the Odorrana andersoni (1992) and noted above, has no apomorphies. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 103

Fig. 46. Maximum-likelihood tree of Matsui et al. (2005) for East Asian ranids, based on mito- chondrial 12S and 16S rRNA sequences (total of 1,283 bp). Sequence alignment was done under ClustalX (Thompson et al., 1997) with cost functions not disclosed and subsequently adjusted manually, guided by secondary structure models as suggested by Kjer (1995). Modeltest 3.06 (Posada and Crandall, 1998) was used to select nucleotide evolutionary model (GTR) assumed for analysis. Fejervarya and were used to root the tree.

All other characters overlap or are identical chalconota and (subgenus) Hylarana (sub- between the two sections. section Hylarana) and that subsection Hylar- Dubois placed the collection of subgenera ana is polyphyletic with Hylarana (subge- that he aggregated under section Hylarana nus) and being in- into two subsections: a humeral gland-bear- dependently derived of the main group of ing group (subsection Hydrophylax) and a subsection Hylarana, which included all of group characterized by having indistinct or their exemplars of subgenera Eburana and absent humeral glands (subsection Hylar- Odorrana, as well as Chalcorana hosii. ana). The presence of a humeral gland is an Within the apomorphic subsection Hydro- apomorphy, so at least prior to analysis we phylax (well-developed humeral gland-bear- considered this single character as evidence ing group) Dubois (1992) recognized several of monophyly of Dubois' subsection Hydro- weakly or undiagnosed (except in the no- phylax, leaving the condition ``humeral menclatural sense) subgenera: Amnirana, glands indistinct or absent'' as plesiomorphic Humerana, Hydrophylax, Papurana, Pul- (although we would have liked to know the chrana, and Sylvirana. According to Dubois distribution of ``indistinct'' humeral glands (1992; his table II), Humerana is distin- within the groups where Dubois reported guished from other members of the subsec- them as indistinct or absent). During analy- tion by the absence of an outer metatarsal sis, however, Matsui et al. (2005; ®g. 46) tubercle; Amnirana and Pulchrana are not provided DNA sequence evidence suggesting rigorously diagnosable from each other; Pa- that that the subsection Hydrophylax is par- purana and Pulchrana are not rigorously di- aphyletic at least with respect to Chalcorana agnosable from each other; and Hydrophylax 104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 can be diagnosed from Sylvirana only on the and C. hosii close to members of Eburana. basis of the absence of an expanded disc and Matsui et al. (2005) suggested that this was lateral groove on ®nger III and toe IV. Mar- not surprising as Chalcorana chalconota lays mayou et al. (2000; ®g. 37) presented DNA pigmented eggs and has a larval keratodont sequence evidence that Sylvirana (a humeral formula of 4±5/3 (Inger, 1966), whereas gland-bearing taxon) is paraphyletic with re- Chalcorana hosii has pigmentless eggs and spect to Hylarana (subgenus) and Pelophy- larvae with a keratodont formula of 5±6/4. lax, both of which lack humeral glands, sug- Matsui et al. (2005) transferred Chalcorana gesting that his subsection Hydrophylax (of hosii into Odorrana (sensu lato, as including section Hylarana) is paraphyletic. We sam- Eburana), with the status of the remaining pled Amnirana albilabris, Hydrophylax gal- species of nominal Chalcorana left question- amensis, Papurana daemeli, Sylvirana able. guentheri, S. maosonensis, S. nigrovittata, Clinotarsus is a monotypic taxon (Clino- and S. temporalis. We were unable to sample tarsus curtipes) that is also poorly diagnosed, any member of Pulchrana, although Matsui with larvae attaining a large size and having et al. (2005; ®g. 46) provided evidence that a somewhat high (but not exclusively) larval it is related to a group of subsection Hydro- keratodont formula of 8/6±8 (Chari, 1962; phylax, including Sylvirana, as well as an im- Dubois, 1992), both characteristics found in bedded piece of subsection Hylarana, Chal- Nasirana as well. We sampled the single spe- corana chalconota. cies, Clinotarsus curtipes. The ``indistinct or absent'' humeral-gland Subgenera Eburana and Odorrana (sensu group (subsection Hylarana) is not rigorous- Dubois, 1992) are putatively distinguished ly diagnosable on the basis of apomorphies from each other by Eburana having (1) discs from any of the other sections of Rana (ex- with a circumlateral groove on ®nger III and cept for Amietia [now in Pyxicephalinae] and toe IV (present or absent in Odorrana); (2) Babina) or from other genera of Ranidae. external metatarsal tubercle present or absent We, therefore, must assume that it is a mix- (absent in Odorrana); (3) gular pouches (var- ture of groups with no necessary phyloge- iable, including the Eburana condition, in netic propinquity or to the exclusion of other Odorrana); (4) no unpigmented spines on the ranid groups. The subgenera coined and ag- chest in males (putatively present in Odor- gregated under subsection Hylarana by Du- rana, according to Dubois, 1992, but absent bois (1992) are variably diagnosable. Mar- in most species, being present in Odorrana mayou et al. (2000; ®g. 37) provided DNA only in the Odorrana andersoni group [see sequence evidence for the polyphyly of sub- above] and two species of the Odorrana section Hylarana (as well as for the para- schmackeri group [O. schmackeri and O. phyly of the other subsection, Hydrophylax; lungshuengensis]; see C.-C. Liu and Hu, see above), by placing Hylarana (subgenus) 1962; Hu et al., 1966, 1973; Yang and Li, and Chalcorana very distant from each other 1980; L. Wu et al., 1983; Fei, 1999; Fei and evolutionarily. Ye, 2001, Ye and Fei, 2001; see also Bain et Subgenus Chalcorana (Chalcorana chal- al., 2003; Bain and Nguyen, 2004); (5) ani- conota being our exemplar, and the type of mal pole of egg unpigmented (pigmented in the taxon) is a morphologically very poorly Odorrana, except O. anlungensis, O. exiliv- diagnosed subgenus within the subsection ersabilis, O. hejiangensis, O. kwangwuensis, Hylarana, with dermal glands present or not O. lungshengensis, O. nasuta, O. tiannanen- in the larvae, outer metatarsal tubercle pres- sis, O. versabilis [C.-C. Liu and Hu, 1962; ent or not, male with paired subgular vocal Hu et al., 1966; Yang and Li, 1980; Fei, pouches present or not, pole of egg 1999; Fei and Ye, 2001; Fei et al., 2001; Ye pigmented or not, and the only likely syna- and Fei, 2001; see also Bain et al., 2003; pomorphy is the relative size of the ®ngers Bain and Nguyen, 2004]). (I Ͻ II; Dubois, 1992). Matsui et al. (2005; Ye and Fei (2001; ®g 45) on the basis of ®g. 46) provided evidence that Chalcorana morphology, and Jiang and Zhou (2005; ®g. is broadly polyphyletic, with Chalcorana 41), on the basis of DNA sequence evidence chalconota close to subsection Hydrophylax have demonstrated that recognition of Ebur- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 105 ana renders Odorrana paraphyletic. With a into two subgenera: Bamburana and Odor- different sampling of species of Eburana and rana. Bamburana was distinguished from Odorrana, Matsui et al. (2005; ®g. 46) pro- subgenus Odorrana (sensu Fei et al., 2005) vided DNA sequence evidence that nominal by the following characters: dorsolateral Eburana is paraphyletic with respect to at folds present (absent in Odorrana), upper lip least one member of Odorrana (O. schmack- with sawtooth spinules (absent in Odorrana); eri) and one species of Chalcorana (C. ho- xiphisternum without notch (deeply notched sii). On this basis Matsui et al. (2005) con- in Odorrana); sternum widened posteriorly sidered Eburana to be part of Odorrana (sternum not widened posteriorly in Odor- (along with Chalcorana hosii). rana). Odorrana (Bamburana) versabilis As noted above, a number of characters (the type species) and O. (Bamburana) na- suggested by Dubois (1992) to diagnose var- suta do not have white spines on the chest ious taxa have taxonomic distributions to of the male, but the other species, O. (Bam- suggest more widespread occurrence. Col- burana) exiliversabilis does. According to orless chest spinules (a putative character of this diagnosis, Bamburana should also in- Odorrana) are also present in Huia nasica clude O. trankieni (Orlov et al., 2003). Nev- (B.L. Stuart and Chan-ard, 2005), Nidirana ertheless, Ye and Fei (2001; ®g. 45) provided adenopleura, and the holotype of N. cald- a cladogram based on 29 character transfor- welli (R. Bain, personal obs.). The one pu- mations of morphology that suggest strongly tative apomorphy of Eburana is character 5 that Bamburana renders the subgenus Odor- (lacking a pigmented animal pole on the egg) rana as paraphyletic. We did not sample any which is known from at least three other gen- species of nominal Bamburana, but on the era: Odorrana (see above), Amolops (e.g., A. basis of the study of Ye and Fei (2001) we chunganensis), and Chalcorana (e.g. C. ho- can reject its recognition. sii) (Bain et al., 2003; Bain and Nguyen, Glandirana was coined by Fei et al. (1991 2004). ``1990'') as a genus, a position they have Bain et al. (2003) transferred Rana chlo- maintained consistently (Fei et al., 2005). ronota (which they thought Dubois, 1992, Nevertheless, Glandirana was placed by Du- had in hand as his exemplar of ``Rana livi- bois (1992) within subsection Hylarana, da'') from Eburana to Odorrana on the fol- where it was diagnosed by Dubois as lacking lowing bases: it has odoriferous skin secre- digital and toe pads, although it retains a lat- tions (implied to be characteristic of Odor- eral groove on the toe tips as found in other rana by way of the formulation of the name groups that do have enlarged digital pads. by Fei et al., 1991 ``1990''); its chromo- With the exception of the lateral toe grooves somes have submetacentric pairs and posi- in Glandirana, we are unaware of any mor- tions of secondary constrictions more similar phological character that would prevent as- (in some cases almost identical) to other spe- signment of Glandirana to sections Amer- cies of Odorrana than to other species of ana, Pelophylax,orRana. Jiang and Zhou Eburana (Li and Wang, 1985; Wei et al., (2005), on the basis of DNA sequence evi- 1993; Matsui et al., 1995); and molecular dence, placed Glandirana as the sister taxon data (Murphy and Chen, unpublished), al- of Rugosa and together as the sister taxon of though it has unpigmented eggs and lacks a group composed of Amolops, Nidirana, Pe- pectoral spinules. The implication is that (1) lophylax, and Rana (®g. 41). We sampled odoriferous skin secretions may be unreport- Glandirana minima. ed for other Eburana species, or (2) odorif- Subgenus Hylarana is also weakly diag- erousness, presence of spinules, and egg col- nosed by comparative characters, with the or may be homoplastic. We sampled Ebur- only morphological apomorphies suggested ana chloronota and Odorrana grahami. Al- by Dubois (1992) being the low number of though this will not allow us to test the rows of labial keratodonts in larvae (shared monophyly of Eburana or Odorrana, it will with Glandirana and sections Amerana, Pe- help illuminate the extent of the problem. lophylax, and Rana; tadpoles unknown in Fei et al. (2005; ®g. 45) have since divided Pterorana and Tylerana). We sampled Hy- Odorrana (sensu Fei et al., 1991 ``1990'') larana erythraea and H. taipehensis. Matsui 106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 et al. (2005; ®g. 46) suggested, on the basis of Dicroglossinae on the basis of mtDNA of DNA sequence evidence that Hylarana (a and nuDNA sequence data. member of Dubois', 1992, subsection Hylar- We sampled two species of Indirana (In- ana) is imbedded within his subsection Hy- dirana sp. 1 and Indirana sp. 2). drophylax. RHACOPHORIDAE (10 GENERA, 267 SPECIES) Subgenus Tylerana is diagnosed from the AND MANTELLIDAE (5 GENERA, 157 SPECIES): remaining Hylarana-like taxa by having a Some authors consider Afro-Asian Rhaco- large oval gland on the inner side of the arm phoridae and Madagascan Mantellidae to be in males (Boulenger, 1920; Dubois, 1992). families (e.g., Vences and Glaw, 2001; Van We sampled Tylerana arfaki. der Meijden et al., 2005). Others consider Subgenera Sanguirana, Pterorana, and them subfamilies of Ranidae (e.g., J.D. Nasirana, which we did not study, were re- Lynch, 1973; Dubois, 1987 ``1985'', 1992; ported by Dubois (1992) to have dermal Roelants et al., 2004) or subfamilies of a glands on the larvae (unknown in Ptero- larger Rhacophoridae (e.g., J.A. Wilkinson rana), well-developed digital discs, and outer and Drewes, 2000; J.A. Wilkinson et al., metatarsal tubercles (unknown in Pterorana). 2002). Regardless, their taxonomic histories Two of the three subgenera, Nasirana and are deeply entwined and we treat them in our Pterorana, contain single species that have discussion as families. distinctive autapomorphies. Nasirana altico- Liem (1970) provided the ®rst character- la can be distinguished from other Hylarana- analysis-based study of phylogeny of the like frogs by the large size of its larvae group (including the mantellids in his sense) (shared with Clinotarsus), the ocellated color in which the mantellids were considered bas- pattern on the larval tail (larvae of Pterorana al to the remaining rhacophorids (®g. 47A). and Tylerana unknown), the ¯eshy promi- Channing (1989) followed with a more rig- nence on the of the adult, and the rel- orous analysis of Old World treefrogs and atively high 7±9/8±9 keratodont formula proposed that Buergeria is the sister taxon of (Dubois, 1992), which may suggest that it is the remaining rhacophorids (including the a member of one of the cascade-dwelling mantellines; ®g. 47B), which he called Buer- clades. Similarly, Pterorana khare is distin- geriinae and Rhacophorinae, respectively. In guished from other ranid frogs by the ¯eshy his arrangement the mantellids were included folds on the ¯anks of the adult. Matsui et al. as basal members of Rhacophorinae. Ford (2005) did not study Sanguirana or Pteror- and Cannatella (1993) noted at least four ana, but suggested that Nasirana is the sister synapomorphies that distinguish Rhacophor- taxon of a group composed of subsection Hy- idae ϩ Mantellidae from other ranoids: (1) drophylax and Chalcorana chalconota (nom- presence of intercalary elements (presuming inally part of subsection Hylarana). that hyperoliids are not the sister taxon); (2) RANIXALINAE (1 GENUS,10SPECIES): Ranix- one slip of the m. extensor digitorum com- alinae is another Indian endemic. It contains munis longus inserts on the distal portion of only Indirana, and is characterized by terres- the fourth metatarsal; (3) outermost slip of trial tadpoles with a keratodont formula of the m. palmaris longus inserts on the proxi- 3±5/3±4. Otherwise, it is diagnostically iden- molateral rim of the aponeurosis palmaris; tical to Nyctibatrachinae (Dubois et al., and (4) possession of a bifurcate terminal 2001). Dubois (1999a: 89) doubted that Nyc- phalanx. J.A. Wilkinson and Drewes (2000) tibatrachinae was distinguishable from Ra- discussed the analyses by Liem (1970) and nixalinae and suggested that Blommers- reanalysis of these data by Channing (1989) SchloÈsser's (1993) distinction between Ra- and suggested further analytical re®nements nixalinae (as Indiraninae), Nyctibatrachinae, but noted considerable instability in the mor- and Nannophrys (which Blommers-SchloÈsser phological evidence (®g. 47C). placed in the otherwise African Cacosterni- More recent work has suggested that man- nae and Dubois placed in Ranixalinae) might tellids are the sister taxon of rhacophorids be substantiated by additional evidence. (e.g., Emerson et al., 2000b; Richards et al., Van der Meijden (2005; ®g. 36), recently 2000; Roelants et al., 2004; Delorme et al., placed, weakly, Indirana as the sister taxon 2005), with this group imbedded within Ran- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 107

idae. Vences and Glaw (2001) suggested that Mantellidae is composed of three subfami- lies: Boophinae (), ( and ), and Man- tellinae ( and ``''). Vences et al. (2003d) arranged these subfam- ilies as Boophinae ϩ (Laliostominae ϩ Man- tellinae), with ``Mantidactylus'' deeply par- aphyletic with respect to Mantella, and sev- eral of the subgenera of ``Mantidactylus'' paraphyletic or polyphyletic. J.A. Wilkinson et al. (2002; ®g. 48) pro- posed a phylogeny of rhacophorines, based on mtDNA sequence data. They found man- tellines to be the sister taxon of rhacophori- nes, and that within rhacophorines, that Buergeria is the sister taxon of all others. They also found Chirixalus to be polyphy- letic, a problem that was addressed, in part, by the recognition of by Ye, Fei, and Dubois (In Fei, 1999), for ``Chirixalus'' eif®ngeri. Some other taxonomic problems were left open by J.A. Wilkinson et al. (2002): the recognition of ``Chirixalus'' pal- bebralis, which is isolated phylogenetically from the majority of rhacophorids; the mono- phyletic grouping of the type species of Chi- rixalus (Chirixalus doriae) with that of Chi- romantis ( xerampelina); and the weakly supported sister clade of Chirixalus- Chiromantis of Chirixalus vittatus, with the type species of , P. leucomystax. Delorme et al. (2005) have since proposed a taxonomy of Philautini (Rhacophoridae;

← Fig. 47. A, Rhacophorid and mantellid tree of Liem (1970) based on 36 direct to dendritic mor- phological transformation series, rooted on a hy- pothetical generalized ranid ancestor. This is one of six equally parsimonious trees constructed un- der the Combinatorial Method (Sharrock and Fel- senstein, 1975) that Liem considered to be the ``best''; B, Tree of Rhacophoridae (including Mantellidae) by Channing (1989) based on a re- interpretation and reanalysis of character transfor- mations from Liem (1970); C, Rhacophorid sec- tion of consensus tree of J.A. Wilkinson and Drewes (2000; their ®g. 14), based on reanalysis of Liem and Channing's data, as well as reinter- pretation of some characters on the basis of spec- imen study. Quotation marks denote nonmono- phyly. 108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 48. Consensus of weighted parsimony trees of Rhacophoridae suggested by J.A. Wilkinson et al. (2002), with their subfamily taxonomy on right. (This is Mantellidae and Rhacophoridae of other authors.) The tree was based on 2kb (of 12S and 16S mt rRNA as well as tRNAVal). Alignment was manual, guided by models of secondary structure with ambiguously aligned segments discarded. In analysis, transversions were weighted twice transitions. Whether reatment of gaps were treated as evi- dence of relationship or as missing data was not stated. Chirixalus eif®ngeri was placed in Kurixalus by Ye, Fei, and Dubois (In Fei, 1999), and Chirixalus idiootocus was transferred into an explicitly polyphyletic/paraphyletic Aquixalus by Delorme et al. (2005). The tree was rooted on Nidirana aden- opleura and Aquarana catesbeiana. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 109

Fig. 49. Delorme et al.'s (2005) dendrogram of rhacophorids, based on undisclosed molecular and morphological data (although characters were summarized for some genera and suprageneric groups), redrawn to illuminate the paraphyly of groupings.

®g. 49). Although a tree was provided, the (2002). Because none of the underlying data evidence (molecular or morphological) that were formally provided, methods of align- provided the tree structure was not provided, ment and analysis were also not provided. and inasmuch as phylogenetic propinquity Substantially less resolution is evident in the was not the organizing principle of their pro- Delorme et al. (2005) tree (®g. 49) than in posed taxonomy, their taxonomy is not con- the J.A. Wilkinson et al. (2002) tree (®g. 48), sistent with the phylogeny they proposed. although they agree that (1) mantellines are Although reported to be based largely on the the sister taxon of rhacophorines; (2) Buer- same data set as the rhacophorid study of geria is the sister taxon of all remaining rha- J.A. Wilkinson et al. (2002; 12S and 16S cophorids; (3) and rRNA), the tree proposed by Delorme et al. are sister taxa; (4) Chirixalus is paraphyletic (2005) also included data from rhodopsin with respect to Chiromantis and likely poly- and from morphology (number and content phyletic (see points 6 and 7); (5) Rhacopho- of transformations undisclosed), but Delorme rus may be paraphyletic with respect to a et al. (2005) did not include the tRNAValine possibly nonmonophyletic Polypedates; (6) a gene included by J.A. Wilkinson et al. monophyletic unit exists that is composed of 110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Kurixalus eif®ngeri and Aquixalus idiootocus us''), for which the predominance of their and A. verrucosus (the latter two were trans- own evidence, as demonstrated by their tree, ferred, respectively, by Delorme et al., 2005, does not reject paraphyly. In particular, it is from ``Chirixalus'' and ``'' into not clear why these authors transferred Chi- an explicitly paraphyletic or polyphyletic rixalus idiootocus into a paraphyletic Aquixalus, without disclosure of phylogenet- ``Aquixalus'', so for our overall discussion, ic evidence; see comment below); (7) ``Chi- we will not follow the transfer of ``Chirix- rixalus'' palpebralis is demonstrably not in alus'' idiootocus into a paraphyletic/poly- a monophyletic group with remaining Chi- phyletic ``Aquixalus'', because this taxonom- rixalus. ic change disagrees with the phylogenetic Delorme et al. (2005) recognized a para- tree (albeit, data free) proposed in the same phyletic/polyphyletic Aquixalus containing publication. two nominal subgenera: (1) Aquixalus (par- In our analysis we sampled Boophinae aphyletic/polyphyletic if Aquixalus idiooto- (Boophis albilabris, B. tephraeomystax); Lal- cus and A. verrucosus are included; if they iostominae (Aglyptodactylus madagascarien- are excluded from Aquixalus the monophyly sis, Laliostoma labrosum); of the remaining subgenus Aquixalus remains (Mantella aurantiaca, M. nigricans, Manti- arguable); (2) Gracixalus (type species: Phi- dactylus cf. femoralis, M. peraccae); Buer- lautus gracilipes Bourret, 1937) for the geriinae (Buergeria japonica); Rhacophori- ``Chirixalus'' gracilipes group, which they nae (``Aquixalus'' (Gracixalus) gracilipes treated as phylogenetically distant from ``C.'' [formerly in Chirixalus or Philautus], ``Chi- palpebralis, thereby suggesting that the pal- rixalus'' idiootocus, Chirixalus doriae, C. pebralis group of Fei (2001), composed, in vittatus, Chiromantis xerampelina, Kurixalus Fei's usage, of Philautus palpebralis, P. gra- eif®ngeri, , N. spinosus, cilipes, P. medogensis, P. ocellatus, and P. Philautus rhododiscus, , romeri, is nonmonophyletic. Nevertheless, P. leucomystax, Rhacophorus annamensis, R. because J.A. Wilkinson et al. (2002) and De- bipunctatus, R. calcaneus, R. orlovi, and lorme et al. (2005) presumably had so much ). underlying evidence in common, the fact of their substantial topological differences be- RESULTS tween their results is surprising, although SEQUENCE LENGTH VARIATION AND many of the internal branches of the J.A. NOTES ON ANALYSIS Wilkinson et al. (2002) tree are weakly sup- ported and possibly could be modi®ed by the Length variation among the four nuclear undisclosed rhodopsin and morphology data protein coding genes was minimal. Follow- of Delorme (2005). Nevertheless, a tree with- ing trimming of primers, all histone H3-com- out associated evidence (that of Delorme et plete products were 328 bp, and all SIA- al., 2005) cannot test a tree that has evidence complete products were 397 bp. All but one attached to it (the tree of J.A. Wilkinson et of the rhodopsin-complete products were 316 al., 2002). bp; the sequence for Alytes obstetricans Because Delorme et al. (2005; ®g. 49) do was 315 bp, as was the sequence of this not accept (apparently) phylogenetic propin- species deposited previously on GenBank quity as the organizing principle in taxono- (AY364385). Most tyrosinase products were my, they (1) created a new paraphyletic ge- 532 bp, exceptions being Xenophrys major nus, Aquixalus (including Chirixalus idioo- and Ophryophryne hansi, which were 538 tocus and Rhacophorus verrucosus, which bp. Tyrosinase was by far the most dif®cult they simultaneously ®gured to be closer evo- fragment to amplify (tyrosinase sequences lutionarily to Kurixalus eif®ngeri than to oth- were sampled for only 38% of the terminals), er members of their Aquixalus), (2) retained and this dif®culty impedes understanding of a nonmonophyletic Chirixalus (with respect the signi®cance of this length variation. The to Chiromantis and ``Chirixalus'' palpebral- ``closest'' taxa for which we were able to ob- is), and (3) recognized Philautini (Philautus tain sequences for this locus were Xenopus ϩ Theloderma ϩ Nyctixalus ϩ ``Aquixal- laevis (from GenBank AY341764) and Hem- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 111 isus marmoratus (both of which are 532 bp), (698 bp) which differ from close relatives by so it is unclear whether the greater length of Ͼ 50 bp and Ͼ 25 bp, respectively. Ascaphus this tyrosinase fragment is characteristic of truei, Leiopelma archeyi, and L. hochstetteri some megophryids or a more inclusive clade. are all 703 bp, as are the included species of The homologous tyrosinase sequence for Pe- Pelodytes and Spea. Similarly, Alytes and tropedetes parkeri downloaded from Gen- Discoglossus are the only sampled species Bank (AY341757) was 535 bp. As with the with a 28S fragment of 706 bp. megophryids, the generality of this length is Although these variations in length do not unclear. However, the length of Arthrolep- provide evidence of phylogeny independent tides sp. is 532, so it is likely that the in- of the underlying indel and nucleotide trans- creased length is restricted to some or all spe- formation events, their phylogenetic conser- cies of Petropedetes. vativeness makes them useful diagnostic Length variation was much more extensive tools, and we therefore note 28S sequence and taxonomically widespread in the ribo- length, where relevant, in the taxonomic sec- somal loci. Among complete H1 sequences, tions that follow. the shortest length of 2269 bp was found in Parsimony analysis by POY of the com- Afrana fuscigula. The longest sequence was bined data set resulted in a single most par- that of the outgroup terminal Latimeria chal- simonious solution of 127019 steps. Al- umnae (2530 bp), followed by Ptychadena though optimizing the implied alignment on mascareniensis (2494 bp) and Silurana tro- the topology found in POY veri®ed the picalis (2477 bp). Length variation was too length reported in POY, ratcheting of the im- extensive for clear phylogenetic patterns to plied alignment in NONA spawned from emerge. However, although extensive varia- Winclada resulted in four most parsimonious tion in the length of the 28S sequences oc- trees of length 127,017 steps, and these are curred even among closely related species our preferred hypotheses. The only differ- (e.g., 744 bp in Schoutedenella schubotzi and ences between the POY and NONA solutions 762 bp in S. xenodactyloides), numerous involve the placement of (1) Glandirana and clades may be characterized by their 28S (2) Brachytarsophrys feae. This con¯ict is length. For example, of the 20 salamander also seen among the four 127017-step trees, 28S fragments with no missing data, all had resulting in the polytomies seen in the strict a length of 694 bp, except Pseudoeurycea consensus (®g. 50 [provided as a multipage conanti and Desmognathus quadramacula- insert]). tus, which were 695 bp. The only other spe- cies of 694 bp in this study were the two TOPOLOGICAL RESULTS AND DISCUSSION turtles (Pelomedusa subrufa and Chelydra A consensus of the four equally most par- serpentina) and the pelodryadine frog, Nyc- simonious trees is shown in ®gure 50 (in- timystes dayi. Length variation in 28S is sert). Most clades are highly corroborated by greater among caecilians (683±727 bp), but molecular evidence (and in some places by it is still more restricted than in anurans morphological evidence). Although only an (685±830 bp). imperfect surrogate for a measure of support Among the sampled anurans, this 28S (something that so far eludes us), the Bremer fragment is Ͼ 700 bp in all but six species ϭ decay index) and jackknife values all (appendix 3). Mantella nigricans and M. au- ( speak to a highly corroborated tree. (See ap- rantiaca differ from all other taxa in that their 28S sequence is 685 bp (28S sequences were not generated for Mantidactylus, but they were for Laliostoma, Aglyptodactylus, and numerous rhacophorids, which have 28S sequences of 709±712 bp). As mentioned Figure 50 is the earlier, the 28S sequence of Nyctimystes dayi taxonomy tree of life, is 694 bp, and that of the related Litoria gen- inserted under the back cover. imaculata is 690. The remaining outliers are Bufo punctatus (700 bp) and Microhyla sp. 112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 pendix 4 for branch length, Bremer support, problematic amphibian groups and discuss and jackknife values.) Because this study aspects of our results that are relevant to the rests on the largest amount of data ever ap- systematics of that particular group, such as plied to the problem of the relationships monophyly of nominal genera and various among amphibians, we think that the ob- taxonomic remedies to problems that our re- tained tree is a step forward in the under- sults highlighted. standing of the evolutionary history of am- phibians. We do, of course, have reservations OUTGROUP RELATIONSHIPS about parts of the overall tree. But, upon re- In our results, Latimeria is outside of the ¯ection, we realized that most of the parts of tetrapod clade, and amniotes form the sister the tree that concerned us were those that (1) taxon of amphibians. This topology was con- we considered insuf®ciently sampled relative ventional, at least for paleontologists and to known species and morphological diver- morphologists (e.g., Gauthier et al., 1988a, sity (e.g., Bufonidae); or (2) are groups for 1988b; ®g. 2A). Within Amniota, we found which no other evidence-based suggestions turtles to be the sister taxon of diapsids (ar- of phylogeny had ever been provided (e.g., chosaurs ϩ lepidosaurs) and this inclusive parts of traditionally recognized Ranoidea). group to be the sister taxon of mammals. Our Nevertheless, familiarity has much to do with molecular data do not support the suggestion notions of plausibility, the root of the prob- by Rieppel and de Braga (1996), based on lem of social conservatism in amphibian sys- morphology, that turtles are more closely re- tematics. lated to lepidosaurs than to archosaurs. Our We discuss results under two headings and molecular results disagree with the results of with reference to several different ®gures. Mannen and Li (1999), Hedges and Poling The primary focus in this ®rst section, ``Re- (1999), and Iwabe et al. (2005), in which tur- sults'', is to address issues of relationship tles were found to be closely related to ar- among, and monophyly of, major groups chosaurs, with lepidosaurs, and mammals as (nominal families and subfamilies and no- successively more distant relations. An anal- menclaturally unregulated taxa). We also ysis of why our molecular results are con- make general taxonomic recommendations in gruent with the conventional tree of mor- this section. Under the second heading, phology (®g. 2A) and not with previous mo- ``Taxonomy'', we discuss further results and lecular results is largely outside the scope of various taxonomic issues under the appro- this paper. Nevertheless, our analysis was a priate taxonomic category. Bremer and jack- parsimony analysis, as were the studies of knife values are reported for each branch in Gauthier et al. (1988a; 1988b). The molec- ®gure 50 (insert; as well as in other ®gures, ular study of Hedges and Poling (1999) rest- where relevant) but are otherwise only oc- ed on a large amount of DNA evidence (ca. casionally mentioned in text. 5.2kb), but their alignment was made under The general tree shown in ®gure 50 (in- a different set of evolutionary assumptions sert), with 532 terminals, is obviously too from that used in their phylogenetic analysis. complex and detailed for easy discussion, so A stronger test of amniote relationships will we will refer to subtrees in different ®gures. be made by combining morphology and all Relevant taxa (branches) have the molecular available DNA evidence and analyzing these data summarized by name and/or number in data under a common set of assumptions. appendix 4. We ®rst discuss the results rel- ative to the Review of Current Taxonomy at AMPHIBIA (LISSAMPHIBIA) AND BATRACHIA or above the nominal family-group level, with reference to families that appear to be Our results (®gs. 50 [insert], 51) corrobo- monophyletic and those that are paraphyletic rate the monophyly of amphibians (Lissam- and polyphyletic. In the case of paraphyly phibia of Parsons and Williams, 1963; Am- and polyphyly we offer remedies in this sec- phibia of Cannatella and Hillis, 1993) with tion that are paralleled in more detail in the reference to other living taxa, although our Taxonomy section, where we propose a data obviously cannot shed any light on the monophyletic taxonomy for all but a few placement of the lissamphibians among fossil 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 113

and Hedges, 1998). Our data suggest strong- ly that the arrangement favored by morphol- ogists (e.g., Trueb and Cloutier, 1991; Ior- dansky, 1996; Zardoya and Meyer, 2000, 2001; Schoch and Milner, 2004) is also the arrangement favored by the preponderance of the molecular evidence (e.g., San Mauro et Fig. 51. Basal structure of our consensus tree al., 2005), that living amphibians form a (®g. 50 [insert]) with respect to outgroups and monophyletic group with respect to Amniota, major amphibian taxa. and that frogs and salamanders are more closely related to each other than either is to groups. We also found the three groups of the caecilians (contra Feller and Hedges, lissamphibians to be strongly supported (®g. 1998). The effect of including fossils and a 50 [insert], branches 7, 24, 74). Furthermore, much more complete morphological data set our DNA sequence data indicate that the cae- are not known, but we note that our molec- cilians are the sister taxon of the clade com- ular data are consistent with the preponder- posed of frogs plus salamanders (Batrachia; ance of morphological data so far published. ®g. 50 [insert], branch 23), the topology pre- Salamanders (Caudata) and frogs (Anura) ferred by Trueb and Cloutier (1991). Our are each also monophyletic, a result that will data reject (1) that living amphibians are par- surprise no one, even though the morpholog- aphyletic with respect to Amniota (Carroll ical evidence for monophyly of the salaman- and Currie, 1975; J.S. Anderson, 2001); (2) ders, in particular, is weak (Larson and Dim- that salamanders are paraphyletic with re- mick, 1993). spect to caecilians (Laurin, 1998a, 1998b, 1998c); and (3) the hypothesis, based on GYMNOPHIONA smaller amounts of evidence, that caecilians In general form our cladogram (®g. 50 [in- and salamanders are closest relatives (Feller sert], ®g. 52) agrees with the conventional

Fig. 52. Caecilian section of general tree (®g. 50 [insert]). 114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 view of caecilian relationships (®g. 3). Like Our results differ slightly from those pre- Nussbaum (1977, 1979) and later authors sented by M. Wilkinson et al. (2003), which (e.g., Duellman and Trueb, 1986; San Mauro were based on a smaller amount of sequence et al., 2004; San Mauro et al., 2005) we ®nd data (mt rRNA only). Like us, M. Wilkinson that Rhinatrematidae is the monophyletic sis- et al. (2003) found Scolecomorphidae and ter taxon of the remaining caecilians. This Typhlonectidae to be imbedded within ``Cae- placement appears well-corroborated on both ciliidae'', although in a different and less morphological and molecular grounds. strongly corroborated placement. Our place- Ichthyophiidae is paraphyletic with respect ment of Siphonops (South America) as the to Uraeotyphlidae (this being highly corrob- sister taxon of (Seychelles) and orated by our molecular data), and can be together the sister taxon of Gegeneophis (In- restated as Ichthyophis is paraphyletic with dia), is the only unanticipated result. In light respect to Uraeotyphlus. This outcome was of the strong support it received in our anal- arrived at previously by Gower et al. (2002). ysis, this conclusion deserves to be evaluated There is a single morphological character, carefully. angulate annuli anteriorly, that supports the monophyly of the ichthyophiids (sensu stric- CAUDATA to, excluding Uraeotyphlus), but the amount of molecular evidence in support of Uraeo- Among previously published cladograms typhlus being nested within Ichthyophis in- our results (®g. 53) most resemble the tree dicates that this character was either reversed of salamander families suggested by Gao and in Uraeotyphlus or independently derived in Shubin (2001; ®g. 5) and diverge slightly different lineages of ``Ichthyophis''. Under from the results presented by Larson and these circumstances, Uraeotyphlus must be Dimmick (1993; ®g. 4) and Wiens et al. transferred to Ichthyophiidae, and although (2005; ®g. 7) in placing sirenids (which lack treatment of ``Ichthyophis'' is beyond the spermatophore-producing organs) as the sis- scope of this study, we expect subsequent ter taxon of Proteidae (which, like other sal- work (denser sampling of ichthyophiids and amandroid salamanders has spermatophore- addition of new data) to delimit the nature of producing organs), rather than placing the this paraphyly and reformulate infrafamilial sirenids as the sister taxon of all other sala- taxonomy. The effect of this change is min- mander families. (The Bayesian analysis of imal, because Uraeotyphlidae contains a sin- Wiens et al., 2005, however, placed crypto- gle genus, and no hierarchical information is branchoids as the sister taxon of remaining lost by placing Uraeotyphlidae in the syn- salamanders, suggesting that there is internal onymy of Ichthyophiidae. con¯ict within their data set.) Other recent As expected from previously published results found, on the basis of RAG-1 DNA DNA sequence (M. Wilkinson et al., 2003) sequence evidence (Roelants and Bossuyt, and morphological evidence (M.H. Wake, 2005; San Mauro et al., 2005), and on the 1993; M. Wilkinson, 1997), we found Sco- basis of RAG-1, nuRNA, and morphology lecomorphidae to be imbedded within Cae- (Wiens et al., 2005), Sirenidae to be the sister ciliidae. The evidence for this is strong (ap- taxon of remaining salamanders, the tradi- pendix 4, branches 12, 14, 16), and we there- tional arrangement. Because our molecular fore consider Scolecomorphidae to be a sub- evidence did not overlap with theirs, and sidiary taxon (Scolecomorphinae) within with the arguable example of Wiens et al. Caeciliidae. Similarly, Typhlonectidae is (2005), their amount of evidence is smaller deeply imbedded within Caeciliidae, a result than ours, these results require additional previously noted (M.H. Wake, 1977; Nuss- testing. Our results do not reject the mono- baum, 1979; M. Wilkinson, 1991; Hedges et phyly of any of the nominal families of sal- al., 1993). Typhlonectidae is here regarded as amanders, a result that is consistent with pre- a subsidiary taxon (as Typhlonectinae) with- vious studies. Except as noted later, the re- in a monophyletic Caeciliidae, although the maining results are conventional. genera of the former ``Caeciliinae'' remain HYNOBIIDAE AND CRYPTOBRANCHIDAE: Un- incertae sedis within the Caeciliidae. like the results of Larson and Dimmick 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 115

Fig. 53. Salamander section of general tree (®g. 50 [insert]). See discussion in ``Taxonomy'' for subfamilies of Plethodontidae and Salamandridae. New taxonomy is on right. 116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(1993; ®g. 4), San Mauro et al. (2005; ®g. and Sirenidae). This taxon was diagnosed by 17), Roelants and Bossuyt (2005; ®g. 16), internal fertilization through the production and Wiens et al. (2005; ®g. 7) our results of spermatophores (produced by a complex place these taxa as the sister taxon of all oth- system of cloacal glands) and having angular er salamanders, and not as the sister taxon of and prearticular bones fused (also found in all salamanders excluding sirenids (the rela- Sirenidae). The hypothesis that sirenids and tionship recovered by Larson and Dimmick, proteids form a taxonomic group is quite old: 1993, San Mauro et al., 2005, and Roelants It was ®rst suggested by Ra®nesque (1815; and Bossuyt, 2005). The monophyly of hy- as Meantia; see the discussion in appendix nobiids plus cryptobranchids is not contro- 6). versial, nor is that of Cryptobranchidae. In RHYACOTRITONIDAE AND AMPHIUMIDAE:We the case of Hynobiidae, as noted in the tax- resolved the polytomy found in the tree of onomic review, our sampling is insuf®cient Gao and Shubin (2001) of Plethodontidae, to address any of the generic controversies Rhyacotritonidae, and Amphiumidae into (summarized by Larson et al., 2003: 43±45) Rhyacotritonidae ϩ (Amphiumidae ϩ Pleth- and is only a minimal test of the monophyly odontidae), a conclusion also of Wiens et al of Hynobiidae. (2005). Although we did not test the mono- SIRENIDAE AND PROTEIDAE: Unlike Larson phyly of either Rhyacotriton or Amphiuma, and Dimmick (1993) and more recent mor- in neither case is this seriously in question. phological and molecular studies (Roelants As noted earlier, the position of Amphiuma and Bossuyt, 2005; San Mauro et al., 2005; with respect to plethodontids is conventional Wiens et al., 2005), but like Gao and Shubin (Larson, 1991; Larson and Dimmick, 1993). (2001; ®g. 5), we recovered Sirenidae not as PLETHODONTIDAE: Our tree differs tren- the sister taxon of all other salamanders but chantly from those of authors prior to 2004 as the sister taxon of Proteidae. Our highly (e.g., D.B. Wake, 1966; Lombard and Wake, corroborated results and the results of Gao 1986), but is similar in general form to those and Shubin (2001) suggest that the perenni- of Mueller et al. (2004) on the basis of com- branch characteristics of Proteidae and Sir- plete mtDNA genomes, Macey's (2005) re- enidae are homologous. On this topology the analysis of those data, and the tree of Chip- cloacal apparatus for spermatophore forma- pindale et al. (2004), based on 123 characters tion is a synapomorphy at the level of all of morphology and about 2.9 kb of mtDNA salamanders, excluding Cryptobranchidae and nuDNA. In those studies and in ours and Hynobiidae, with a loss in Sirenidae. Al- Amphiumidae and Rhyacotritonidae were ternatively, it is a convergent development in obtained as successively more distant out- Proteidae and in the ancestor of Salamandri- groups of Plethodontidae. In the three pre- dae, Rhyacotritonidae, Dicamptodontidae, vious studies (Chippindale et al., 2004; Plethodontidae, Amphiumidae, and Ambys- Mueller et al., 2004; Macey, 2005) as well tomatidae. The effect of combining the mor- as in ours, the desmognathines are in a clade phological data presented by Wiens et al. with the plethodontines (Ensatina, and Pleth- (2005) with all of their and our molecular odon). Our data (as well as those of Mueller data remains an open question, although we et al., 2004, and Macey, 2005) also found note that their morphological-only data set Hydromantes and Speleomantes to be in this produced a result in which Sirenidae ϩ Pro- plethodontine clade, not with ``other'' boli- teidae form a monophyletic group. Thus, it toglossines. is not clear that this is a simple morphology- In our results, as well as those of Mueller versus-molecules issue. Rather than oversim- et al. (2004) and Chippindale et al. (2004), plify and misrepresent that paper, we leave all other plethodontids (the old Hemidacty- the question open as to what the result will liinae and Bolitoglossini) are placed in a be when all molecular and morphological group that forms the sister taxon of the ®rst data are combined. group. The evidence for these groupings is As noted earlier, our results reject a mono- strong (appendix 4; ®g. 53). The placement phyletic Salamandroidea (all salamanders, of Hydromantes and Speleomantes in the ®rst excluding Cryptobranchidae, Hynobiidae, group by our data is strongly corroborated, 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 117 being placed within the desmognathines (a also placed Hydromantes (including Speleo- result that runs counter to the morphological mantes) in Plethodontinae. These two genera evidence as presented by Schwenk and had previously been associated with Bolito- Wake, 1993). Mueller et al. (2004) obtained glossini (D.B. Wake, 1966; Elias and Wake, Hydromantes (including Speleomantes)in 1983). the same general group as we did, but placed Our results regarding placement of Hydro- as the sister taxon of Aneides. In the details mantes and Speleomantes imply either that of placement of Batrachoseps, Hemidacty- the morphological synapomorphies of the lium, and our few overlapping bolitoglossine Desmognathinae, mostly manifestations of genera, we differ mildly. Our differences the bizarre method of jaw opening in which from the tree of Macey (2005) are dif®cult the lower jaw is held in a ®xed position by to explain. The amount of evidence mar- ligaments extending to the atlas±axis com- shalled by Macey (the same aligned data set plex, are reversed in the hydromantine clade as Mueller et al., 2004), is on the order of or that this peculiar morphology is conver- 14kb of aligned mtDNA sequence. Our gent in Desmognathus and Phaeognathus. mtDNA set is a subset of that, but analyzed Previous to the study of Mueller et al. differently, particularly with respect to align- (2004), who found Plethodon to be mono- ment. Alignment of the data set of Mueller phyletic on the basis of analysis of mtDNA et al. (2004) was done with different trans- sequence data, all published evidence point- formation costs than used in analysis, and ed to paraphyly of Plethodon with respect to this alignment was accepted for reanalysis by Aneides (e.g., Larson et al., 1981; Mahoney, Macey (2005). Further, a number of our ex- 2001). Our analysis of a variety of DNA se- emplars (i.e., Plethodon dunni, P. jordani, quence data suggests also that the eastern and Desmognathus quadramaculatus, Phaeog- western components of Plethodon do not nathus, Hydromantes platycephalus, Eurycea have a close relationship, being united solely wilderae, Gyrinophilus porphyriticus, Thor- by symplesiomorphy. Had it not been for the ius sp., Bolitoglossa rufescens, and Pseu- appearance of the recent paper by Chippin- doeurycea conanti) are represented in our dale et al. (2004), we would have erected a analysis by sequences that are not part of the new generic name for western Plethodon (for mtDNA genome. Although we provisionally which no name is currently available). But, accept the results of Macey (2005; ®g. 10) the denser sampling of plethodons and dif- as based on a much larger amount of data ferent selection of genes in the Chippindale than our results, it may be that the single et al. (2004) paper suggests that a study in- biggest cause of different results between our cluding all of the available data and a denser analysis and his is the method of alignment. sampling is required before making any tax- One will know only when that data set is onomic novelties. analyzed using direct optimization. We recovered former Bolitoglossini as Chippindale et al. (2004; ®g. 11) suggest- polyphyletic, with the traditional three main ed a taxonomy, consistent with their tree, for components (supergenera Batrachoseps, Hy- Plethodontidae. Plethodontinae in their sense dromantes, and Bolitoglossa; D.B. Wake, corresponds to the group composed of the 1966) being found to have little in common former Desmognathinae and former Pletho- with each other. Our tree of bolitoglossines dontini. Within the second group composed (sensu stricto) is not strongly corroborated. of hemidactyliines and bolitoglossines they Nevertheless, that the three groups of boli- recognized Hemidactyliinae (Hemidacty- toglossines should be recovered as polyphy- lium), Spelerpinae Cope, 1859 (Eurycea letic is not shocking inasmuch as the amount [sensu lato], Gyrinophilus, Stereochilus, and of evidence that traditionally held them to- Pseudotriton), and Bolitoglossinae (for all of gether was small. the bolitoglossine genera studied). Macey SALAMANDRIDAE: Our results largely cor- (2005) came to the same taxonomy, but respond to those of Titus and Larson (1995) placed Hemidactyliinae as the sister taxon of and especially with those presented by Lar- remaining plethodontids, the relative position son et al. (2003). Our tree differs from the of the other groups remaining the same. He topology suggested by Larson et al. (2003), 118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

which was based on more extensive taxon ASCAPHIDAE AND LEIOPELMATIDAE: Asca- sampling but less DNA evidence, in that we phidae and Leiopelmatidae are recovered in get additional resolution of the group Neu- our analysis as parts of a monophyletic rergus ϩ (Triturus ϩ Euproctus), where in group, mirroring the results of Green et al. the tree provided by Larson et al. (2003) (1989), BaÂez and Basso (1996), and more re- these taxa are in a polytomy below the level cent authors (Roelants and Bossuyt, 2005; of Paramesotriton ϩ Pachytriton. San Mauro et al., 2005). The paraphyly of DICAMPTODONTIDAE AND AMBYSTOMATI- this grouping, as suggested by Ford and Can- DAE: Dicamptodon is recovered as the sister natella (1993), is rejected. If our results are taxon of Ambystomatidae, the same phylo- accurate, the ®ve morphological synapomor- genetic arrangement found by previous au- phies suggested by Ford and Cannatella thors (Sever, 1992; Larson and Dimmick, (1993) of Leiopelma plus all frogs excluding 1993; Wiens et al., 2005). The monophyly of Ascaphus must be convergences or synapo- Dicamptodon was only minimally tested, al- morphies of all living frogs that were lost in though Dicamptodon monophyly is not se- Ascaphus. Nevertheless, the hypothesis of riously in doubt (Good and Wake, 1992). In- Ford and Cannatella (1993) was based large- asmuch as Dicamptodontidae was recognized ly on the unpublished dissertation of Can- on the basis of its hypothesized phylogenetic natella (1985; cited by Ford and Cannatella, distance from Ambystomatidae (Edwards, 1993), who rooted his analysis of primitive 1976), a hypothesis now rejected, we pro- frogs on Ascaphus on the basis of two ple- pose the synonymy of Dicamptodontidae siomorphic characters found among frogs with Ambystomatidae, which removes the uniquely in Ascaphus: (1) facial nerve passes redundancy of having two family-group through the anterior acoustic foramen and names, each containing a single genus. The into the auditory capsule while still fused to reformulated Ambystomatidae contains two the auditory nerve; (2) salamander-type jaw sister genera, Dicamptodon and Ambystoma. articulation in which there is a true basal ar- Ambystomatidae was found to be mono- ticulation. All other characters placing Leio- phyletic, at least with reference to our ex- pelma as more closely related to all non-As- emplar taxa, and the sister taxon of former caphus frogs were optimized by this assump- Dicamptodontidae. Although we have not se- tion, requiring their polarity to be veri®ed. verely tested the monophyly of Ambystoma, Furthermore, the support for the Ascaphus ϩ others have done so (e.g., Shaffer et al., Leiopelma branch is very high (Bremer ϭ 1991; Larson et al., 2003), and its monophy- 41, jackknife ϭ 100%), so it is unlikely that ly is well corroborated. ®ve morphological characters (of which three have not been rigorously polarized) can re- ANURA verse this. Placing Ascaphus and Leiopelma as sister taxa allows some characters to be As mentioned earlier and in the taxonomic explained more ef®ciently. Thus, the absence review, the amount of morphological and of the columella in these two taxa can be DNA sequence evidence supporting the seen to be a synapomorphic loss. Ritland's monophyly of Anura is overwhelming. We (1955) suggestion that the m. caudalipubois- think that our data make a strong case for a chiotibialis in Leiopelma and Ascaphus may new understanding of frog phylogeny. Even not be homologous with the tail-wagging though most of our results are conventional muscles of salamanders, and the more tradi- with respect to understanding of frog phy- tional view of homology with these muscles logenetics, our purpose is not to conceal this are both consistent with our results. To re- understanding, but to bring the taxonomy of move the redundancy of the family-group frogs into line with their phylogenetic rela- names with the two genera (Ascaphus and tionships. For discussion we adopt the Ford Leiopelma), we assign Ascaphus to Leiopel- and Cannatella (1993) tree (®g. 14) as the matidae (as did San Mauro et al., 2005). Roe- traditional view of phylogeny (although not lants and Bossuyt (2005) retained Ascaphi- of nomenclature). We ®rst discuss the non- dae and Leiopelmatidae as separate families neobatrachian frogs (®g. 54). and resurrected the name Amphicoela Noble, 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 119

Fig. 54. Part 1 of anurans from the general tree (®g. 50 [insert]): non-neobatrachian frogs.

1931, for this taxon. Amphicoela is redun- of Pelobatoidea (Ford and Cannatella, 1993; dant with Leiopelmatidae (sensu lato) when their Mesobatrachia) or as the sister taxon of Ascaphidae and Leiopelmatidae are regarded all other frogs (Maglia et al., 2001; Pugener as synonymous, as we do. et al., 2003). All three of these arrangements PIPIDAE AND RHINOPHRYNIDAE: We found, are supported by morphological characters, as did Haas (2003) and San Mauro et al. although Haas' arrangement is more highly (2005), and as was suggested even earlier by corroborated. Haas (2003) suggested nine Orton (1953, 1957), Sokol (1975), and Mag- apomorphies that exclude Pipoidea and As- lia et al. (2001) that Rhinophrynidae ϩ Pip- caphidae from a clade composed of all other idae is the sister taxon of all non-leiopel- frogs. Pugener et al. (2003) suggested three matid frogs. This result is strongly supported synapomorphies for all frogs excluding pi- by our evidence (®g. 54; appendix 4, branch- poids. (This statement is based on examina- es 77, 78, 84). Recent suggestions had alter- tion of their ®gure 12; they provided no com- natively placed Pipoidea as the sister taxon prehensive list of synapomorphies.) Ford and 120 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 55. Trees of intergeneric relationships within Pipidae (from ®g. 19). A, Cannatella and Trueb (1988). B, BaÂez and Pugener (2003); C, Roelants and Bossuyt (2005); D, De Sa and Hillis (1990; results consistent with B, C, and E); E, This work. (Undirected network on lower right shows rooting points of each result, except for D.)

Cannatella (1993) suggested that four char- are the component families. A novel arrange- acters support Mesobatrachia: (1) closure of ment in our tree is Hymenochirus being the frontoparietal fontanelle by juxtaposition placed as the sister taxon of Pipa ϩ (Silur- of the frontoparietal bones (not in Pelodytes ana ϩ Xenopus). This result differs from the or Spea); (2) partial closure of the hyoglossal cladograms of Cannatella and Trueb (1988), sinus by the ceratohyals; (3) absence of the de Sa and Hillis (1990), BaÂez and Pugener taenia tecti medialis; and (4) absence of the (2003), and Roelants and Bossuyt (2005; ®g. taenia tecti transversum. However, on the ba- 16). Although our results are highly corrob- sis of Haas' (2003) morphological data orated by our data, a more complete test alone, these characters are rejected as syna- would involve the simultaneous analysis of pomorphies. However, the mtDNA molecular all of the sequence data with the morpholog- results presented by GarcõÂa-ParõÂs et al. ical data of all relevant living and fossil taxa. (2003) support the recognition of Mesobatra- As noted in ®gure 55, the rooting point of chia (Pelobatoidea ϩ Pipoidea). Neverthe- the pipid network appears to be more impor- less, these authors included only three non- tant to the estimates of phylogeny than dif- pipoid, non-pelobatoid genera (Ascaphus, ferences among networks. Discoglossus, and Rana) as outgroups, which The placement of Pipidae ϩ Rhinophryn- did not provide a strong test of mesobatra- idae as the sister taxon of all frogs, save chian monophyly. Placement of Pipoidea as Leiopelmatidae ϩ Ascaphidae, suggests the sister taxon of all other non-leiopelmatid strongly that the fusion of the facial and tri- frogs requires rejection of Discoglossanura, geminal ganglia (Sokol, 1977) found in pe- Bombinatanura, and Mesobatrachia of Ford lobatoids, pipoids, and neobatrachians, but and Cannatella (1993), a rejection that is not in Discoglossidae and Bombinatoridae is strongly supported by our study. homoplastic. Similarly, the absence of free In our analysis, as well as in all recent ribs in the adults of pelobatoids, neobatra- ones (Ford and Cannatella, 1993; BaÂez and chians, and pipoids, but their presence in Pugener, 2003; Haas, 2003), Pipoidea (Rhin- Leiopelma, Ascaphus, and Discoglossidae, ophrynidae ϩ Pipidae) is monophyletic, as requires either independent losses in pipoids 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 121 and pelobatoids ϩ neobatrachians, or an in- pole morphology. GarcõÂa-ParõÂs et al. (2003; dependent gain in discoglossids ϩ bombi- ®g. 18) also found Pelobatoidea to be mono- natorids. Roelants and Bossuyt (2005) noted phyletic, on the basis of their DNA evidence, fossil evidence that would support the inde- and suggested a topology of Scaphiopodidae pendent loss in pipoids and Acosmanura (Pe- ϩ (Pelodytidae ϩ (Megophryidae ϩ Pelo- lobatoidea ϩ Neobatrachia). batidae)), with relatively low Bremer values DISCOGLOSSIDAE AND BOMBINATORIDAE: on the branch tying Scaphiopodidae to the Ford and Cannatella (1993) partitioned the remaining taxa. In our results we recover all former Discoglossidae (sensu lato) into Dis- of these family-group units as monophyletic coglossidae (sensu stricto) and Bombinato- and highly supported. But, our data show ridae because their evidence suggested that strongly a relationship of (Pelodytidae ϩ former Discoglossidae was paraphyletic, Scaphiopodidae) ϩ (Pelobatidae ϩ Mego- with Bombinatoridae and Discoglossidae phryidae) (®g. 54). forming a graded series between the Asca- NEOBATRACHIA: As in all previous studies, phidae and Leiopelmatidae on one hand, and we found Neobatrachia to be highly corrob- all other frogs on the other hand. As noted orated by many transformations (®gs. 50 [in- in the taxonomic review, this partition was sert], 56, 58, 59, 60). What is particularly based on two characters shared by discog- notable in the broad structure of Neobatra- lossines and all higher frogs and absent in chia is the dismemberment of Leptodactyli- the bombinatorines. Haas (2003) rejected this dae and Hylidae as traditionally formulated, topology with six character transformations as well as the placement of Heleophrynidae supporting the monophyly of Bombinatori- outside of the two major monophyletic com- dae and Discoglossidae. In addition to Haas' ponents, for our purposes referred to here as characters, we have strong molecular evi- (1) Hyloidea, excluding Heleophrynidae and dence in support of the monophyly of this (2) Ranoidea. taxon (Discoglossidae ϩ Bombinatoridae), as HELEOPHRYNIDAE: Haas (2003) suggested well as the subsidiary families. that Heleophryne may be related to Peloba- Unlike Haas (2003), but like recent mo- toidea, a suggestion that is not borne out by lecular studies (Roelants and Bossuyt, 2005; our simultaneous analysis of Haas' data and San Mauro et al., 2005), we did not recover our molecular data. Earlier authors (e.g., J.D. Alytes as the sister taxon of the remaining Lynch, 1973) addressed the phylogenetic po- discoglossines and bombinatorines. We in- sition of Heleophryne and associated it with cluded Haas' six characters supporting that Limnodynastidae on the basis of overall sim- topology in our analysis, and the taxon sam- ilarity, or with Limnodynastidae ϩ Myoba- pling for this part of the tree is nearly iden- trachidae on the basis of DNA sequence data tical in the two studies, so it appears that mo- (Biju and Bossuyt, 2003). But recently San lecular evidence in support of a topology of Mauro et al. (2005) suggested, on the basis Alytes ϩ Discoglossus is decisive. The only of DNA sequence evidence, that Heleo- rationale for considering Discoglossidae and phrynidae is the sister taxon of remaining Bombinatoridae as separate families rested Neobatrachia. We obtained the same place- on the assertion of paraphyly of the group ment of Heleophrynidae as did San Mauro (Ford and Cannatella, 1993), a position now et al. (2005). rejected. Nevertheless, we retain the two- HYLOIDEA, EXCLUDING HELEOPHRYNIDAE: family arrangement because this re¯ects the Hyloidea, as traditionally composed, consists state of the literature and is consistent with of all arciferal groups of neobatrachians and recovered phylogeny. was expected (on the basis of absence of PELOBATOIDEA: Haas (2003) did not recov- morphological evidence) to be broadly par- er Pelobatoidea (Megophryidae, Pelobatidae, aphyletic with respect to Ranoidea, or ®r- Pelodytidae, Scaphiopodidae) as monophy- misternal frogs (Microhylidae, Ranidae, and letic. Although we included his morpholog- their satellites, Mantellidae, Rhacophoridae, ical data in our analysis, we ®nd Pelobato- Hyperoliidae, Arthroleptidae, Astylosterni- idea to be highly corroborated, which sug- dae, and Hemisotidae), or monophyletic on gests very interesting convergences in tad- the basis of molecular data (Ruvinsky and 122 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 56. Part 2 of anurans from the general tree (®g. 50 [insert]): Heleophrynidae and basal hyloids (Sooglossidae, Batrachophrynidae, Limnodynastidae, and Myobatrachidae).

Maxson, 1996; Feller and Hedges, 1998; Fai- ing literature: Leptodactylidae was found to vovich et al., 2005; San Mauro et al., 2005). be composed of several only distantly related In our results, Hyloidea is only narrowly par- groups, and Hylidae (in the sense of includ- aphyletic, with the bulk of the hyloids form- ing Hemphractinae) was con®rmed to be par- ing the sister taxon of ranoids and only He- aphyletic or polyphyletic (see below). leophrynidae outside of this large clade (a SOOGLOSSIDAE AND NASIKABATRACHIDAE: conclusion also reached by San Mauro et al., The South Indian Nasikabatrachus and the 2005). Within the restricted (non-heleo- Seychellean sooglossids form an ancient tax- phrynid) Hyloidea, a unit composed of Soog- on united by considerable amounts of molec- lossidae and the newly discovered Nasika- ular evidence (®g. 56). Biju and Bossuyt batrachidae forms the sister taxon of the re- (2003) placed Nasikabatrachus as the sister maining hyloids (cf. Biju and Bossuyt, 2003; taxon of the sooglossids and our results cor- San Mauro et al., 2005). For the most part, roborate this. We are unaware of any histor- the traditional family-group units within Hy- ical (in the sense of history of systematics) loidea were found to be monophyletic, the or other reason to regard Nasikabatrachus as exceptions being predictable from preexist- being in a family distinct from Sooglossidae, 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 123 and on the basis of the molecular evidence Limnodynastinae. Further discussion can be we consider Nasikabatrachus to be the sole found in the Taxonomy section. known mainland member of Sooglossidae. ``LEPTODACTYLIDAE'': The paraphyly and The antiquity of this united group is evident polyphyly of ``Leptodactylidae'' is starkly in its placement as the sister taxon of all oth- exposed by this analysis, being paraphyletic er non-heleophrynid hyloids. Its phylogenet- with respect to all hyloid taxa except Heleo- ic position as well as its presence both in phrynidae and Sooglossidae (®g. 57). Be- India and in the Seychelles suggests that the cause of the extensiveness of the paraphyly taxon existed before the ®nal breakup of Pan- and the complexity of the reassortment of the gaea in the late . subsidiary groupings, the various units of a MYOBATRACHIDAE,LIMNODYNASTIDAE, AND paraphyletic/polyphyletic ``Leptodactylidae'' RHEOBATRACHIDAE: Because of the absence must be dealt with before the remainder of of morphological synapomorphies uniting Hyloidea can be addressed. Speci®cally the the Australo-Papuan groups Myobatrachidae, following nominal families are imbedded Limnodynastidae, and Rheobatrachidae (in within ``Leptodactylidae'': Allophrynidae, our usage), and because of the suggestion of Brachycephalidae, Bufonidae, Centrolenidae, a special relationship between Myobatrachi- Dendrobatidae, Hylidae, Limnodynastidae, dae and Sooglossidae and between Limno- Myobatrachidae, and Rhinodermatidae. To dynastidae and Heleophrynidae (J.D. Lynch, provide the tools to allow us to discuss the 1973), we were surprised that the preponder- remainder of the hyloid families, we here ance of evidence corroborates a monophy- provide a new familial taxonomy with ref- letic Myobatrachidae ϩ Limnodynastidae ϩ erence to the old taxonomy provided in ®g- Rheobatrachidae (®g. 56). Nevertheless, ure 50 (insert). We start at the top of ®gure there is only one morphological character in- 56 and address the subfamilies of ``Lepto- volved in these alternatives (condition of the dactylidae'' as we come to them. cricoid ring: complete or incomplete), so, in ``TELMATOBIINAE'': ``Telmatobiinae'' is retrospect, our surprise was unwarranted. found to be polyphyletic (®gs. 56, 57, 58, With respect to Myobatrachidae (sensu 59), with the austral South American Calyp- stricto; Myobatrachinae of other authors), tocephalellini (Telmatobiinae-1: Telmatobufo our results are largely congruent with those ϩ Caudiverbera) forming the sister taxon of of Read et al. (2001). The positions of Me- the Australo-Papuan Myobatrachidae, Lim- tacrinia and Myobatrachus are reversed in nodynastidae, and Rheobatrachidae; Telma- the two studies. The trenchant difference be- tobiinae-2 being paraphyletic with respect to tween our results is in the placement of Par- Batrachyla (Telmatobiinae-3: Batrachylini); acrinia. Our results placed it strongly as the and Ceratophryini (Lepidobatrachus (Cera- sister taxon of Assa ϩ Geocrinia, whereas tophrys ϩ Chacophrys)); and Telmatobiinae- Read et al. (2001) placed it as the sister taxon 4(Hylorina, Alsodes, Eupsophus) being the of the myobatrachids that they studied, with sister taxon of a taxon composed of part of the exception of Taudactylus. Conclusive the polyphyletic Leptodactylinae (Limnome- resolution of this problem will require all dusa) and Odontophrynini (Proceratophrys available evidence to be analyzed simulta- and Odontophrynus; part of nominal Cera- neously. tophryinae). As noted in the taxonomic re- We include Mixophyes (formerly in Lim- view, Telmatobiinae was united by overall nodynastidae) and Rheobatrachus (sole plesiomorphic similarity (e.g., exotrophic member of former Rheobatrachidae) in tadpoles, non-bony sternum). That the mo- Myobatrachidae (sensu stricto); Read et al. lecular data show Telmatobiinae to be poly- (2001) did not include those taxa in their phyletic is neither surprising nor unconven- study. We obtain a sister-taxon relationship tional. between Mixophyes and Rheobatrachus (al- The Chilean and Peruvian telmatobiine though this is only weakly corroborated) and clade composed of Caudiverbera and Tel- association of Mixophyes (and Rheobatra- matobufo is monophyletic on both molecular chus) with Myobatrachinae, inasmuch as and morphological grounds; is highly corrob- Mixophyes has traditionally been assigned to orated as the sister taxon of the Australo- 124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 57. Fate of former Leptodactylidae (sensu lato) on our general tree (®g. 50 [insert]). Imbedded non-leptodactylid taxa are in bold.

Papuan Myobatrachidae ϩ Limnodynastidae ®g. 17). (The inclusion of Batrachophrynus ϩ Rheobatrachidae; and is phylogenetically is discussed under Batrachophrynidae in the distant from all other telmatobiine ``lepto- Taxonomy section.) This result is not unex- dactylids'' (see also San Mauro et al., 2005; pected as calyptocephallelines have long 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 125

Fig. 58. Part 3 of anurans from the general tree (®g. 50 [insert]): Hemiphractidae, Brachycephalidae, Cryptobatrachidae, Amphignathodontidae, and Hylidae. 126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 59. Part 4 of anurans from the general tree (®g. 50 [insert]): Centrolenidae, Leptodactylidae, Ceratophryidae, and Cycloramphidae. been suspected to be only distantly related to within some larger familial group, but to other telmatobiine leptodactylids (Cei, 1970; maintain familiar usage (and because we Burton, 1998a). Moreover, the region they have resolved Limnodynastidae, Myoba- inhabitat is also home to Dromiciops, a mar- trachidae, and Rheobatrachidae into rede- supial most closely related to some ®ned Limnodynastidae and Myobatrachidae) groups of Australian and not to we consider it as the family Batrachophryn- other South American marsupials (Aplin and idae (the oldest available name for calypto- Archer, 1987; Kirsch et al., 1991; Palma and cephallelines as currently understood; see Spotorno, 1999). The previous association of ``Taxonomy'' and appendix 6 for discussion Calyptocephalellini with the South American of application of this name). Telmatobiinae was based on overall similar- As suggested by Lynch (1978b), one part ity with geographically nearby groups. As of Telmatobiinae-2; (®g. 59), Telmatobiini, is the sister taxon of the Australian Myoba- paraphyletic with respect to Batrachylini trachidae ϩ Limnodynastidae, it would be (Batrachylus) as well as to Ceratophryinae-1 acceptable to place Calyptocephallelinae (Ceratophryini). The oldest name for the 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 127

clade Telmatobiinae-2 (Telmatobius, Batra- ELEUTHERODACTYLINAE AND BRACHYCE- chyla, Atelognathus) ϩ Ceratophryinae-1 PHALIDAE: Eleutherodactylinae is paraphylet- (Ceratophrys, Chacophrys, and Lepidobatra- ic with respect to Brachycephalidae (Brachy- chus) is Ceratophryidae. Within this family cephalus) (®g. 57, 58). There is nothing we recognize two subfamilies, Telmatobiinae about Brachycephalus being imbedded with- (Telmatobius) and Ceratophryinae (for all re- in Eleutherodactylus (sensu lato) that re- maining genera). Within Ceratophryinae we quires any signi®cant change in our under- recognize two tribes: Batrachylini (Batrachyla standing of morphological evolution, except ϩ Atelognathus) and Ceratophryini (for Cer- to note that this allows the large eggs and atophrys, Chacophrys, and Lepidobatrachus). direct development of Brachycephalus to be (See the Taxonomy section for further dis- homologous with those of eleutherodacty- cussion.) lines. This result was suggested previously As noted earlier, another former compo- (Izecksohn, 1971; Giaretta and Sawaya, nent of Telmatobiinae (Telmatobiinae-3; see 1998; Darst and Cannatella, 2004), and no ®gs. 57, 59) is recovered as the sister taxon evidence is available suggesting that we of one piece of ``Leptodactylinae'' (Limno- should doubt it. Further, to impose a mono- medusa) plus Odontophrynini (Ceratophryi- phyletic taxonomy, we follow Dubois (2005: nae-2, formerly part of Ceratophryinae). 4) in placing Eleutherodactylinae Lutz, 1954, (The polyphyly of ``Leptodactylinae'' will be into the synonymy of Brachycephalidae addressed under the discussion of that sub- GuÈnther, 1858. All ``eleutherodactyline'' familial taxon.) Because no documented genera are therefore assigned to Brachyce- morphological synapomorphies join the two phalidae. Previous authors (e.g., Heyer, 1975; groups of nominal Ceratophryinae (Odonto- J.D. Lynch and Duellman, 1997) have sug- phrynini and Ceratophrynini), and they had gested that Eleutherodactylus (and eleuthero- previously been shown to be distantly related dactylines) is an explosively radiating lineage. (Haas, 2003), this result does not challenge Our results, which places brachycephalids as credibility. (See further discussion in the the sister taxon of the majority of hyloid frogs Taxonomy section.) refocuses this issue. The questions now be- ``HEMIPHRACTINAE'': ``Hemiphractinae'', come (as suggested by Crawford, 2003): (1) which was transferred out of Hylidae and Why are the ancient brachycephalids mor- into Leptodactylidae by Faivovich et al. phologically and reproductively conservative (2005), is united by possessing bell-shaped as compared with their sister taxon (com- gills in developing and bearing eggs posed of Cryptobatrachidae, Amphignatho- on the dorsum in shallow depressions to ex- dontidae, Hylidae, Centrolenidae, Dendro- tensive cavities. The subfamily has not been batidae, and Bufonidae, as well as virtually found to be monophyletic by any recent au- all other ``leptodactylid'' species)? (2) Why thor (Darst and Cannatella, 2004; Faivovich are there so few species in the brachyce- et al., 2005). In our results (®gs. 57, 58) we phalid (eleutherodactyline) radiation relative found (1) Hemiphractus is the sister taxon of to their sister group (the former composed of hyloids, excluding Batrachophrynidae, some 700 species, mostly in nominal Eleuth- Myobatrachidae (including Rheobatrachi- erodactylus, and the latter consisting of more dae), Limnodynastidae, Sooglossidae (in- than twice as many species)? Additional cluding Nasikabatrachidae), and Heleo- comments on this taxon will be found under phrynidae; (2) Flectonotus ϩ Gastrotheca; Brachycephalidae in the Taxonomy section. and (3) Stefania ϩ Cryptobatrachus are suc- ``LEPTODACTYLINAE'': Although ``Lepto- cessively more distant from a clade [branch dactylinae'' has at least one line of evidence 371] bracketed by Hylidae and Bufonidae. in support of its monophyly (bony sternum), The evidence for this polyphyly is quite the molecular data unambiguously expose its strong, so we recognized three families to polyphyly, with its species falling into two remedy this: Hemiphractidae (Hemiphrac- units (®g. 57, 59). The ®rst of these (Lepto- tus), Cryptobatrachidae (Cryptobatrachus ϩ dactylinae 1±2), is paraphyletic with respect Stefania), and Amphignathodontidae (Flec- to the cycloramphine unit, called Cycloram- tonotus ϩ Gastrotheca). phinae-1 in ®gures 57 and 59, Paratelmato- 128 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 bius and Scythrophrys (an arrangement par- groups. Our molecular data overcome the tially consistent with the suggestion of J.D. few morphological characters that might be Lynch, 1971, that at least Paratelmatobius considered synapomorphies of the relevant belongs in Leptodactylinae). The second unit group. The ®rst of these groups, labeled Cy- (Leptodactylinae-3, Limnomedusa) is the sis- cloramphinae-1, is composed of Scythro- ter taxon of Odontophrynini (Ceratophryi- phrys and Paratelmatobius and is imbedded nae-2). Limnomedusa was previously united within Leptodactylidae (sensu stricto; as part with other leptodactylines solely by its pos- of Leptodactylinae, as discussed earlier.) The session of a bony sternum, but it lacks the second unit, which is labelled Cycloramphi- foam-nesting behavior found in most other nae-2, is Elosiinae (ϭ Hylodinae) of Lynch leptodactylines (exceptions being Pseudopa- (1971); although it is relatively weakly cor- ludicola, Paratelmatobius, and some species roborated by molecular evidence, it is united of Pleurodema). Regardless, the association by morphological evidence suggested by of Limnomedusa with Leptodactylinae has Lynch (1971, 1973). Cycloramphus (part of always been tentative (Heyer, 1975). So, our Cycloramphinae-2) is tightly linked to Rhi- discovery (corroborating the results of Fai- noderma (Rhinodermatidae), one of the vovich et al., 2005) that Limnomedusa is not points of paraphyly of former Leptodactyli- part of Leptodactylinae is not unexpected; dae. Cycloramphinae-2 forms a paraphyletic nor does it require extensive homoplasy in group with respect to Rhinodermatidae, Tel- the morphological data that are available. We matobiinae-2, Leptodactylinae-3, and Odon- recognize this unit (Leptodactylinae-1 ϩ Cy- tophrynini (Ceratophryinae-2). Because no cloramphinae-1 ϩ Leptodactylinae-2; ®gs. morphological characteristics that we are 57, 59, branch 430) as Leptodactylidae (sen- aware of would reject this larger grouping, su stricto), a taxon that is much diminished we place these ®ve units into a single family, compared with its previous namesake but for which the oldest available name is Cy- that is consistent with evolutionary history. cloramphidae. Within this, we recognize two Further discussion is found under Leptodac- subfamilies: Hylodinae (for Crossodactylus, tylidae in the Taxonomy section. Megaelosia, and Hylodes) and Cycloramphi- ``CERATOPHRYINAE'': ``Ceratophryinae'' nae for the remainder of this nominal family- (sensu lato) is polyphyletic, with its two con- group taxon. stituent tribes, Odontophrynini (Ceratophryi- Our DNA sequence evidence places Tho- nae-2) and Ceratophryninae (Ceratophryi- ropa (Cycloramphinae-3) as the sister taxon nae-1) (sensu Laurent, 1986) being only dis- of the monophyletic Dendrobatidae (®gs. 57, tantly related (®gs. 57, 59, branches 446, 60). We were surprised by this result, be- 458). As noted elsewhere in this section, cause none of the morphological characters there has never been any synapomorphic ev- that had been suggested to ally Hylodinae idence to associate these two groups. Thus, with Dendrobatidae are present in Thoropa their distant relationship is not surprising or (T. Grant, personal obs.), and Thoropa most even unconventional, inasmuch as Barrio recently has been associated with Batrachyla (1963; 1968) and Lynch (1971) suggested (J.D. Lynch, 1978b). Nevertheless, our mo- that these two units are distantly related. Cer- lecular data support this arrangement, and atophryini is imbedded in a taxon (®gs. 57, Thoropa has never been more than tentative- 59: Telmatobiinae-2; branch 441) that is ly associated with the grypiscines (ϭ cyclor- weakly corroborated, but is here recognized amphines; Heyer, 1975). Furthermore, man- as a family Ceratophryidae. Odonotophry- ual rearrangements of hylodines and Thoro- nini is resolved as the sister taxon of Lim- pa used as starting trees for further analysis nomedusa (formerly in Leptodactylinae), to- inevitably led to less parsimonious solutions gether residing in a group composed largely or returned to this solution as optimal (as im- of former cycloramphines. plied by the Bremer values). Our ®rst incli- ``CYCLORAMPHINAE'' AND RHINODERMATI- nation was to place Thoropa into Dendro- DAE: ``Cycloramphinae'' (sensu Laurent, batidae, so as not to erect a monotypic fam- 1986) was also found to be polyphyletic ily. However, Dendrobatidae, as traditionally (®gs. 57, 59) in three distantly related conceived, is monophyletic and has a large 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 129

Fig. 60. Part 5 of anurans from the general tree (®g. 50 [insert]): Thoropidae, Dendrobatidae, and Bufonidae. 130 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 literature associated with it that addresses a cal sac, we place Rhinodermatidae into the certain content and diagnosis that remained synonymy of Cycloramphidae. largely unchanged for nearly 80 years. For DENDROBATIDAE: We found Dendrobatidae this reason, we place Thoropa into a mono- to be monophyletic and the sister taxon of typic family, Thoropidae, to preserve the Thoropa. The former statement is conven- core diagnostic features of Dendrobatidae for tional, the latter, surprising. Nevertheless, the the large number of workers that are familiar highly corroborated nature of this placement with the taxon. (cladistically in the same neighborhood as CENTROLENIDAE AND ALLOPHRYNIDAE:As hylodines, with which it was considered suggested by Noble (1931), Austin et al. closely allied by some authors, e.g., Noble, (2002), and Faivovich et al. (2005), Allo- 1926, and Lynch, 1973) should close discus- phryne is closely related to Centrolenidae, to- sion of whether the ®rmisternal dendrobatids gether forming a monophyletic group that is are derived from some austral South Amer- the sister taxon of a group composed of most ican arciferal group (here strongly supported; of the former Leptodactylinae (®g. 59; for dendrobatid girdle architecture see Noble, branch 426). Our data reject a close relation- 1926; Kaplan, 1995) or related to some ran- ship of Centrolenidae to Hylidae, as well as oid or ranid group, a conclusion suggested the suggestion by Haas (2003), made on the by some lines of morphological evidence basis of larval morphology, that Centroleni- (Blommers-SchloÈsser, 1993; Ford, 1993; ϩ dae may not be a member of Neobatrachia. Grant et al., 1997). Thoropa Dendrobati- Allophryne shares with the centrolenids T- dae form the sister taxon of Bufonidae. This shaped terminal phalanges (J.D. Lynch and phylogenetic arrangement is highly corrobo- Freeman, 1966), which is synapomorphic at rated and suggests that Ameerega Bauer, this level. We regard Allophryne as a part of 1986 (a senior synonym of Centrolenidae, the sister taxon of a taxon Myers, 1987; see Walls, 1994) is polyphy- letic, a result that is consistent with previous composed of Centrolene ϩ Cochranella ϩ studies (e.g., Santos et al., 2003; Vences et Hyalinobatrachium (which has as a morpho- al., 2003b). Taxon sampling was limited in logical synapomorphy intercalary phalangeal all studies to date, however, and we leave it elements). to more exhaustive analyses to assess the de- BRACHYCEPHALIDAE: Our study found Bra- tails of the relationships within Dendrobati- chycephalus to be imbedded within Eleuth- dae. erodactyinae, indeed, within Eleutherodac- HYLIDAE: If hylids are considered to con- tylus (sensu lato; ®g. 57, 58). Previous au- tain Hemiphractinae (see above), then Hyli- thors (e.g., Izecksohn, 1971; Giaretta and Sa- dae would be catastrophically paraphyletic waya, 1998) suggested that Brachycephalus with respect to leptodactylids (excluding the is allied with Euparkerella (Eleutherodacty- former calyptocephalellines [Batrachophryn- linae) on the basis of sharing the character of idae]), dendrobatids, bufonids, Allophryne, digital reduction. We did not sample Eupar- and centrolenids (®gs. 57, 58, 59). This ar- kerella, which could be imbedded within a rangement suggests that the claw-shaped ter- paraphyletic Eleutherodactylus. This propo- minal phalanges and intercalary cartilages sition remains to be tested. As noted earlier, taken previously to be synapomorphies of Brachycephalidae and Eleutherodactylinae Hylidae (sensu lato) are homoplastic and not are synonyms, with Brachycephalidae being synapomorphic for Hylidae. Because Hylidae the older name. (sensu lato) is broadly para- or polyphyletic, RHINODERMATIDAE: We found Rhinoderma we adopt the concept of Hylidae adopted by to be imbedded within a clade composed Faivovich et al. (2005), that is Hylinae ϩ largely of South American cycloramphine Phyllomedusinae ϩ Pelodryadinae. leptodactylids (®gs. 57, 59), more speci®cal- Hylidae (sensu stricto, excluding ``Hemi- ly as the sister taxon of Cycloramphus. Be- phractinae'') is monophyletic and highly cor- cause the only reason to recognize Rhinod- roborated. Our results are largely congruent ermatidae has been its autapomorphic life with the results of Faivovich et al. (2005), history strategy of brooding larvae in the vo- which were based on more sequence evi- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 131 dence and denser sampling of hylids. Fai- of Bufo margaritifer with Rhamphophryne vovich et al. (2005) should be referenced for conforms with their morphological similari- the evidentiary aspects of hylid phylogenet- ty, but the nesting of this clade within a ics. The only signi®cant difference between group of Asian Bufo was unexpected. The our results and theirs is that our exemplars association of Bufo lemur (a species of for- of Hyla form a paraphyletic group with re- mer Peltophryne in the Antilles) with Schis- spect to Isthmohyla and Charadrahyla, and maderma (Africa) is novel, as is the place- Hypsiboas is paraphyletic with respect to ment of this group with Bufo viridis and Bufo Aplastodiscus, and the tribe Dendropsophini melanostictus, although Graybeal (1997), at is not monophyletic as delimited by Faivov- least in her parsimony analysis of molecular ich et al. (2005). However, because our den- data, suggested that Peltophryne was asso- sity of sampling and evidence is less than in ciated with Bufo melanostictus, an Asian tax- that study, our results do not constitute a test on. of those results, and we leave their taxonomy Obviously, denser sampling will be re- unchanged. quired to resolve bufonid relationships, but Hylinae has long been suspected of being the current topology provides an explicit hy- paraphyletic, but our results and those of Fai- pothesis for further investigation. Clearly, vovich et al. (2005) strongly corroborate the Bufo must be partitioned into several genera notion that Hylinae is monophyletic and the to remedy its polyphyly/paraphyly with re- sister taxon of Pelodyradinae ϩ Phyllome- spect to several other nominal genera and to dusinae, both of which are also strongly cor- provide a reasonable starting place from roborated as monophyletic. which to make progress. For more discussion The apparent polyphyly of Nyctimystes in and the beginnings of this partition, see Bu- our results may be real, although our paucity fonidae in the Taxonomy section. of sampling prevents us from delimiting the RANOIDEA: Monophyly of Ranoidea (in the problem precisely. Similarly, the long-rec- sense of excluding Dendrobatidae) was ognized (Tyler and Davies, 1978; King et al., strongly corroborated in our analysis, as well 1979; Tyler, 1979; Maxson et al., 1985; as by other recent analyses (Roelants and Hutchinson and Maxson, 1987; Haas, 2003; Bossuyt, 2005; San Mauro et al., 2005). Ran- Faivovich et al., 2005), pervasive paraphyly oidea in our analysis is divided into two ma- of Litoria in Pelodryadinae with respect to jor groups (see ®gs. 50 [insert], 56, 61, 62, both Cyclorana and Nyctimystes has obvi- 63, 65), which correspond to (1) a group ously been a major problem in understanding composed of a para- or polyphyletic Micro- relationships among pelodryadines. Ongoing hylidae, Hemisotidae, Hyperoliidae, para- research by S. Donnellan and collaborators phyletic Astylosternidae, and Arthroleptidae aims to rectify these issues in the near future. (®gs. 61, 62); and (2) a giant paraphyletic BUFONIDAE: That Bufonidae is a highly ``Ranidae'' and its derivative satellites, Man- corroborated monophyletic group is not sur- tellidae and Rhacophoridae (®g. 63, 65). This prising; that we have a reasonably well-cor- is summarized on the general tree (®g. 50 roborated phylogenetic structure within Bu- [insert]). fonidae is a surprise (®gs. 50 [insert], 60). MICROHYLIDAE AND HEMISOTIDAE: Our re- Like Graybeal (1997; ®g. 25), we found Me- sults (®gs. 50, 61, 62) do not support the tra- lanophryniscus (which lacks Bidder's or- ditional view of subfamilies and relationships gans) to form the sister taxon of the remain- suggested by Parker (1934) in the last revi- ing bufonids (which, excluding Truebella, sion of the family. The notion of polyphy- have Bidder's organs). Within this clade, Ate- letic Microhylidae falling into two monophy- lopus ϩ Osornophryne forms the sister taxon letic groupsÐ(1) Brevicipitinae (as the sister of the remaining taxa. taxon of Hemisotidae); and (2) the remaining The paraphyly of Bufo with respect to so microhylidsÐextends from the suggestion many other bufonid genera had previously by Blommers-SchloÈsser (1993) that Hemi- been detected (e.g., Graybeal, 1997; Cun- sotidae and Brevicipitinae are closely related. ningham and Cherry, 2004), but some asso- Because the Type II tadpole that was consid- ciations are unconventional. The relationship ered a synapomorphy in microhylids (Star- 132 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 61. Part 6 of anurans from the general tree (®g. 50 [insert]): Microhylidae. rett, 1973) is not present in brevicipitines family, Brevicipitidae. (We ®nd Dubois', (which have direct development) and hemi- 2005, proposal that Arthroleptidae, Astylos- sotids have a Type IV tadpole, there was nev- ternidae, Brevicipitidae, Hemisotidae, and er any particular evidence tying brevicipiti- Hyperoliiidae be considered subfamilies of nes to the remaining microhylids. Moreover, an enlarged Brevicipitidae, to be an unnec- only a single synapomorphy tied brevicipi- essary perturbation of familiar nomencla- tines to hemisotines (Channing, 1995), so the ture.) evidence for paraphyly/polyphyly of micro- Within the larger group of ``microhylids'', hylids also was not strong. As suggested by Microhylinae is broadly paraphyletic with re- Van der Meijden et al. (2004; and consistent spect to the remaining subfamilies, with with the results of Biju and Bossuyt, 2003, Phrynomantis (Phrynomerinae) being situat- and Loader et al., 2004, but contrary to the ed near the base of our sampled microhy- Scoptanura hypothesis of Ford and Canna- lines, Hoplophryne (Melanobatrachinae) tella, 1993), we ®nd Brevicipitinae and Hem- placed weakly next to Ramanella (Microhy- isotidae to form a monophyletic group, and linae), and Cophylinae (based on our exem- this taxon to be more closely related to Ar- plars of Anodonthyla, Platypelis, Plethodon- throleptidae, Astylosternidae, and Hyperoli- tohyla, and Stumpf®a) being found to be idae than to remaining Microhylidae. For this monophyletic and placed as the sister taxon reason we regard brevicipitines as a distinct of Ramanella (Microhylinae) ϩ Hoplophry- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 133

Fig. 62. Part 7 of anurans from the general tree (®g. 50 [insert]): Hemisotidae, Hyperoliidae, and Arthroleptidae. ne (Melanobatrachinae). Surprisingly, Sca- Microhylinae is nonmonophyletic, with phiophryne (Scaphiophryninae) is deeply im- (1) some taxa clustered around the base of bedded among the microhylids and the sister the Microhylidae and weakly placed (e.g., taxon of part of ``Microhylinae'' (branch Kalophrynus, Synapturanus, Micryletta); (2) 130, subtending Kaloula, Chaperina, Cal- a group of Asian taxa (e.g., Kaloula±Micro- luella, and Microhyla). Ford and Cannatella hyla) forming the sister taxon of Scaphio- (1993) and Haas (2003) had considered Sca- phryne; and (3) a New World clade (i.e., the phiophryne to form the sister taxon of the group composed of Ctenophryne, Nelsono- remaining microhylids on the basis of larval phryne, Dasypops, Hamptophryne, Elachis- features, but because we included Haas' tocleis, Dermatonotus, and Gastrophryne) (2003) morphological data in our analysis, placed as the sister taxon of Cophylinae ϩ we can see that these features must be ho- Melanobatrachinae ϩ Ramanella. moplastic. Our picture of ``Microhylinae'' runs coun- 134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 ter to the little phylogenetic work that has Laurent, 1984b; Dubois, 1987 ``1985'', been done so far, especially with respect to 1992). Within this group we found Hypero- the cladogram of New World taxa by Wild liidae (excluding Leptopelis)toforma (1995). Wild's (1995; ®g. 34) cladogram as- monophyletic group. sumed New World monophyly, was rooted on Phylogenetic structure within Hyperoliidae a composite outgroup, and is strongly incon- has been contentious, with various arrange- gruent with our topology. Our solution is to ments suggested by different authors. Our re- (1) recognize Gastrophryninae for the New sults differ signi®cantly from all previously World taxa that do form a demonstrably published hyperoliid trees (Drewes, 1984; monophyletic group (including Ctenophryne, Channing, 1989; Vences et al., 2003c). Like Nelsonophryne, Dasypops, Hamptophryne, Vences et al. (2003c), we found Leptopelis Elachistocleis, Dermatonotus and Gastro- (Hyperoliidae) to form a monophyletic group phryne); and (2) restrict Microhylinae to a that is separate from the remainder of Hy- monophyletic group including Calluella, peroliidae and placed with a group composed Chaperina, Kaloula, and Microhyla. The gen- of the Astylosternidae ϩ Arthroleptidae. The era that we have not assigned to either Gas- consideration of Leptopelinae as a subfamily trophryninae or Microhylinae (sensu stricto), of Hyperoliidae cannot be continued because or that are clearly outside of either group (e.g., it renders Hyperoliidae (sensu lato) paraphy- Synapturanus or Kalophrynus), we treat as in- letic. We restrict the name Hyperoliidae to certae sedis within Microhylidae. The ar- the former Hyperoliinae, which in addition rangement asserted without evidence by Du- to our molecular data, is supported by the bois (2005), of an Old World Microhylini and synapomorphic presence of a gular gland New World Gastrophrynini, within his Micro- (Drewes, 1984). hylinae, is speci®cally rejected by the basal We found Astylosternidae to be paraphy- position in our tree of Kalophrynus and Syn- letic with respect to Arthroleptidae, with Sco- apturanus, far from our Microhylinae and tobleps (Astylosternidae) being the sister tax- Gastrophryninae. on of Arthroleptidae (®g. 62). No previous As suggested by Savage (1973), Dysco- hypotheses of relationship within Astyloster- phinae is polyphyletic, with Calluella deeply nidae or Arthroleptidae have been rigorously imbedded within Asian microhylines and proposed (Vences et al., 2003c), so our re- Dyscophus placed as the sister taxon of a sults are the ®rst to appeal to synapomorphy. group composed of members of Asterophryi- Our ®nding that Schoutendenella is paraphy- nae (Cophixalus, Choerophryne, Genyophry- letic with respect to Arthroleptis is particu- ne, Sphenophryne, Copiula, Liophryne, larly noteworthy because recognition of Aphantophryne, Oreophryne) and Astero- Schoutedenella as distinct from Arthroleptis phryinae (Callulops). Genyophryninae is has been contentious (e.g., Laurent, 1954; clearly paraphyletic with respect to Astero- Loveridge, 1957; Schmidt and Inger, 1959; phryinae, as suggested by Savage (1973) and Laurent, 1961; Poynton, 1964b; Laurent, Sumida et al. (2000a). For this reason we re- 1973; Poynton, 1976; Poynton and Broadley, gard Asterophryinae and Genyophryninae as 1985; Poynton, 2003). Laurent and Fabrezi synonyms, with Asterophryinae being the (1986 ``1985'') suggested that Schoutedenel- older name for this taxon. This allows the la is more closely related to Cardioglossa optimization of direct development as a syn- than to Arthroleptis, an hypothesis rejected apomorphy for the combined taxon. here. ARTHROLEPTIDAE,ASTYLOSTERNIDAE AND RANIDAE,MANTELLIDAE, AND RHACOPHOR- HYPEROLIIDAE: We found an African group IDAE: Our results for this group are similar in composed of Hyperoliidae, Astylosternidae, some respects to those presented by Van der and Arthroleptidae to constitute a highly cor- Meijden et al. (2005; ®g. 36). Differences in roborated clade, the sister taxon of Hemiso- results may be due to our denser taxon sam- tidae ϩ Brevicipitidae (®g. 62). This exis- pling, to their greater number of analytical tence of this group was suggested previously assumptions, their inclusion of RAG-1 and but has not been substantiated by synapo- RAG-2, which we did not include, or their morphies (Laurent, 1951; Dubois, 1981; lack of 28S, seven in absentia, histone H3, 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 135 tyrosinase, and morphology, which we did bois (2005), the ``southern African clade'' of include. Final resolution will require analysis Van der Meijden et al. (2005): Tomopterna, of all of the data under a common assump- Arthroleptella, Natalobatrachus, Afrana, tion set. Amietia, Strongylopus, Cacosternum, and We found a taxon composed of a broadly Anhydrophryne. We place (1) Phrynobatra- paraphyletic ``Ranidae'', and monophyletic chus (and its satellites Phrynodon and Di- Mantellidae ϩ Rhacophoridae to form the morphognathus) in Phrynobatrachidae; (2) sister taxon of Microhylidae ϩ Hemisotidae Arthroleptides, Conraua, Indirana, and Pe- ϩ Hyperoliidae ϩ Arthroleptidae ϩ Astylos- tropedetes in Petropedetidae; (3) Afrana, ternidae (®g. 50 [insert], 61, 63). The results Amietia, Anhydrophryne, Arthroleptella, Au- are complex but are comparable to a group bria, Cacosternum, Natalobatrachus, Pyxi- of smaller studies that dealt overwhelmingly cephalus, Strongylopus, and Tomopterna in with Asian taxa (Tanaka-Ueno et al., 1998a, Pyxicephalidae, as had Dubois (2005). (See 1998b; Bossuyt and Milinkovitch, 2000; Em- ®g. 63 and further discussion of these groups erson et al., 2000a; Marmayou et al., 2000; in the Taxonomy section.) Bossuyt and Milinkovitch, 2001; Kosuch et Roelants et al. (2004), who did not include al., 2001; Grosjean et al., 2004; Roelants et any African taxa in their study, proposed In- al., 2004; Jiang and Zhou, 2005). This over- dirana to be the sister taxon of Micrixalinae, all result varies widely from Bossuyt and although their evidence did not provide res- Milinkovitch (2001), who found Mantellinae olution beyond a polytomy with (1) the Lan- ϩ Rhacophorinae as the sister taxon of Nyc- kanectes±Nyctibatrachus clade; and (2) the tibatrachinae ϩ Raninae; this clade sister to ranine-rhacophorine-mantelline clade. How- Dicroglossinae ϩ Micrixalinae, and Ranix- ever, we found Indirana to be deeply imbed- alinae sister to them all. ded in an African clade otherwise composed We ®nd Ptychadeninae (Ptychadena being of Conraua, Arthroleptides, and Petropede- our exemplar genus) to be the sister taxon of tes (a clade we consider a family, Petrope- the remaining ``Ranidae'', a highly corrobo- detidae). Dissimilarly, Van der Meijden et al. rated result (®g. 63). The sister taxon of Pty- (2005) found, albeit weakly, Indirana as the chadeninae is composed of Ceratobatrachi- sister taxon of Dicroglossinae. Nevertheless, nae (Ingerana, Discodeles, Ceratobatrachus, our result is highly corroborated, although it Batrachylodes, and Platymantis) and the re- is based on less overall evidence than that of maining ``ranids''. Here we differ signi®- Van der Meijden et al. (2005), although as cantly from Roelants et al. (2004), inasmuch noted previously, analyzed differently. Our as they considered Ingerana to be an occi- sequence evidence for Indirana is the same dozygine, whereas we ®nd Ingerana to be in 12S and 16S GenBank sequences produced/ Ceratobatrachinae, where it had originally used by Roelants et al. (2004), so contami- been placed by Dubois (1987 ``1985''). nation or misidenti®cation is not an issue. We ®nd a major African clade (®g. 63; Like Roelants et al. (2004), we ®nd occi- branch 192), similar to the results of Van der dozygines to form the sister taxon of Dicrog- Meijden et al. (2005). One clade (branch lossinae, with the latter containing Paini (our 193) is Phrynobatrachinae of Dubois (2005), exemplares being members of Nanorana and composed of a paraphyletic Phrynobatra- Quasipaa), which had been transferred from chus, within which Phrynodon and Dimor- Raninae into Dicroglossinae by Roelants et phognathus are imbedded. A second com- al. (2004). Unlike their data, ours place Na- ponent (branch 200) is composed of Con- norana not within Paa, but as the sister tax- rauinae (Conraua), Ranixalinae (Indirana), on of a clade composed of Fejervarya (which Petropedetinae, and Pyxicephalinae sensu we show to be paraphyletic), Sphaerotheca, Dubois (2005). Petropedetinae of Dubois Nannophrys, Euphlyctis, and Hoplobatra- (2005) (Petropedetes ϩ Arthroleptides, sub- chus. tended by branch 205), forms the sister taxon Our results are broadly consistent with of Indirana (Ranixalinae of Dubois, 2005). several other studies showing that Hoploba- Pyxicephalus ϩ Aubria (branch 210) form trachus (Limnonectini) is the sister taxon of the sister taxon of the Pyxicephalinae of Du- Euphlyctis (Dicroglossini) (Bossuyt and Mil- 136 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 63. Part 8 of anurans from the general tree (®g. 50 [insert]): Ptychadenidae, Ceratobatrachidae, Micrixalidae, Phrynobatrachidae, Petropedetidae, Pyxicephalidae, and Dicroglossidae. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 137 inkovitch, 2001; Kosuch et al., 2001; Gros- Paini to appear monophyletic. Examination jean et al., 2004; Roelants et al., 2004). Lim- of the trees and associated unrooted networks nonectini (sensu Dubois, 1992) is therefore (®g. 64) support this view. That Euphlyctis, rejected as nonmonophyletic. Limnonectes Hoplobatrachus, and Nannophrys lack (including Taylorana Dubois, 1987 ``1986'', spines on the forearms and belly as in Paini as a synonym; a result congruent with Em- is incongruent evidence. Nevertheless, it erson et al., 2000a) forms the sister taxon of does strengthen our view that Group 1 of a clade formed by Paini (Quasipaa), Nanor- Jiang et al. (2005) deserves generic recog- ana, Nannophrys, and the remaining mem- nition, and that Paini, as nonmonophyletic, bers of ``Limnonectini'' (Fejervarya, must be placed into the synonymy of Di- Sphaerotheca, and Hoplobatrachus) and Di- croglossinae. (See the account of Dicroglos- croglossini (Euphlyctis), a result congruent sinae in the Taxonomy section.) with Grosjean et al. (2004). Marmayou et al. A trenchant difference between our results (2000) found Fejervarya ϩ Sphaerotheca to and those of Roelants et al. (2004; but the form the sister taxon of a monophyletic Lim- same as found by Van der Meijden et al., nonectes ϩ Hoplobatrachus, but they did not 2005) is in the placement of Lankanectes ϩ include Euphlyctis in their study. Roelants et Nyctibatrachus. Roelants et al. (2004) placed al. (2004; ®g. 35), and Jiang et al. (2005; ®g. this taxon outside of most of ``Ranidae'' (ex- 42), and Jiang and Zhou (2005; ®g. 41) cepting Micrixalinae and Indiraninae, which found Paini to be imbedded within this group we also found to be placed elsewhere). We (Dicroglossinae), and our results con®rm ®nd Lankanectes ϩ Nyctibatrachus to be the their result. This suggests that a character sister taxon of Raninae, excluding Amietia, that has been treated as of particular impor- Afrana, and Strongylopus (and Batrachylo- tance to ranoid systematics, forked or entire des, transferred to Ceratobatrachidae, as dis- omosternum, is considerably more variable cussed earlier). than previously supposed (see Boulenger, Dubois' (1992) Amolops (containing the 1920: 4), regardless of the weight placed on subgenera Amo [which we did not study], this character by some taxonomists (e.g., Du- Amolops, Huia, and Meristogenys) is dem- bois, 1992). onstrated to be polyphyletic (a result congru- Our topology is not consistent with that of ent with Roelants et al., 2004; who did not Roelants et al. (2004), Jiang et al. (2005), study Huia; ®g. 65). At least with respect to and Van der Meijden et al. (2005) in that we our exemplars, the character of a ventral do not recover a monophyletic Paini, instead sucker on the larva is suggested by our re- ®nding our exemplars (2 species of Quasipaa sults to be convergent in Amolops (in the and1ofNanorana) to form a pectinate series sense of including Amo), Huia, and Meris- leading to ``Fejervarya'' ϩ Hoplobatrachus togenys (as well as in Pseudoamolops). (Euphlyctis and Nannophrys were pruned for As expected, the genus Rana (sensu Du- this discussion because they were not part of bois, 1992) is shown to be wildly nonmon- the study of Jiang et al., 2005; ®g. 42). Al- ophyletic, with Dubois' sections Strongylo- though our topological differences from the pus (Afrana and Strongylopus) and Amietia results of Roelants et al. (2004) apparently (Amietia) being far from other ``Rana'' in re¯ect differences in evidence and sampling, our results. (This result is consistent with that we have more of both. The difference be- of Van der Meijden et al., 2005, and was an- tween our results and those of Jiang et al. ticipated by Dubois, 2005.) In this position, (2005) seemingly do not re¯ect differences Section Strongylopus is paraphyletic with re- at the level of descriptive ef®ciency at the spect to Cacosternum ϩ Anhydrophyne (®g. level of unrooted network. We do have a bit 63). As noted earlier, we transfer Sections more resolution between their groups 1 and Strongylopus and Amietia out of Ranidae and 2 as a paraphyletic grade, rather than as a into a newly recognized family, Pyxicephal- polytomy. By treating Hoplobatrachus and idae, as was done by Dubois (2005). (See the Fejervarya as their outgroups on which to Taxonomy section for further discussion.) root a tree of Limnonectes ϩ Paini, the study As noted in the Review of Current Tax- by Jiang et al. (2005) inadvertantly forced onomy, understanding the phylogeny of Hy- 138 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 64. A, Original tree of Jiang et al. (2005; from ®g. 42) of Paini and (on right) its equivalent undirected network; B, Tree rerooted and with augmented resolution as implied by our general results, and, at right, its equivalent undirected network. We have applied the name Nanorana to Group 1 of Jiang et al. (2005); Quasipaa was applied by Jiang et al. (2005) for their Group 2. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 139

Fig. 65. Part 9 of anurans from the general tree (®g. 50 [insert]): Mantellidae, Rhacophoridae, Nyctibatrachidae, Ranidae. 140 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 larana-like frogs (Dubois' sections Babina is not surprising, as subsection Hylarana and Hylarana) is critical to understanding ra- never did have any suggested synapomor- nid systematics. Our results show that Bou- phies; again, this is consistent with the results lenger (1920) was correct that ``Hylarana'' of Matsui et al., 2005.) The component sub- (sensu lato) is polyphyletic, or at least wildly genus Hylarana is most closely related to paraphyletic. The plesiomorphic condition in Sylvirana guentheri (subsection Hydrophy- Ranidae is to have expanded toe digits, as in lax); subgenus Chalcorana (subsection Hy- Rhacophoridae ϩ Mantellidae and farther larana) is most closely related to Hydrophy- outgroups, so this discovery merely illumi- lax ϩ Amnirana (subsection Hydrophylax); nates that ``Hylarana'' was constructed on Tylerana (subsection Hylarana) is most the basis of plesiomorphy. Dubois' (1992) closely related to Papurana (subsection Hy- Section Hylarana is paraphyletic with re- drophylax); and Clinotarsus (subsection Hy- spect to Amolops, Meristogenys, and Huia, larana) forms the sister taxon of Meristogen- as well as most of sections Babina, Amerana, ys (subgenus of Amolops sensu Dubois, Rana, Pelophylax, and Lithobates. Further, 1992). Glandirana (subsection Hylarana)is the Hylarana subsection Hydrophylax (his the sister taxon of Pelophylax (section Pe- humeral-gland group) is polyphyletic (as lophylax). Eburana (subsection Hylarana)is hinted at by the results of Matsui et al., 2005; the sister taxon of Huia (subgenus of Amo- ®g. 46), in our results placing this group in lops sensu Dubois), and our exemplar of two places: (1) Sylvirana guentheri forms the Odorrana (subsection Hylarana) is the sister sister taxon of subgenus Hylarana (in the taxon of ``Amolops'' chapaensis, a result non-humeral-gland group) and (2) in a group similar to those of Jiang and Zhou (2005; ®g. containing Hydrophylax galamensis and Pa- 41), who found Eburana nested within Odor- purana daemeli. Our ®ndings are largely rana (see the Taxonomy section for further congruent with the results of Roelants et al. discussion). (2004), who suggested ``S.'' guentheri as sis- As suggested by Hillis and Davis (1986) ter to H. erythraea, but who also suggested and con®rmed by Hillis and Wilcox (2005), that this ``erythraea clade'' is sister to a clade Dubois' (1992) Section Pelophylax is poly- containing Sylvirana nigrovittata. Marmayou phyletic, with one part, Pelophylax (sensu et al. (2000; ®g. 37) found strong support for stricto), being found most closely related to H. erythraea and H. taipehensis as sisters, Glandirana (section Hylarana) and the part and weak support for ``S.'' guentheri to be composed of Aquarana and Pantherana be- part of that clade. They did show, weakly but ing paraphyletic with respect to Dubois' Sec- consistently, that Sylvirana is polyphyletic tion Lithobates, as well as one species in his with respect to Hylarana (Marmayou et al., Section Rana (R. sylvatica). Our results do 2000). Kosuch et al. (2001) found Amnirana not con¯ict with Roelants et al. (2004), who to be the sister taxon of Hydrophylax gala- found Pelophylax (P. lessonae, P. nigroma- mensis ϩ Sylvirana gracilis. Differences in culata) to be the sister taxon of Amolops cf. data size and taxon sampling may account ricketti (A. ricketti and P. lessonae not in- for differences in tree topology among these cluded in our study). Roelants et al. (2004) studies, but the substantial results are similar. also found that the Amolops±Pelophylax Roelants et al. (2004) included exemplars of clade is sister to a ``Sylvirana''±Hylarana± subgenus Hydrophylax sensu Dubois and Chalcorana±Hydrophylax±Pulchrana clade, subgenus Hylarana sensu Dubois, but not which is largely consistent with our ®ndings. Amnirana as in our study. Kosuch et al. (We did not study Pulchrana.) Jiang and (2001) included exemplars of Hydrophylax Zhou (2005) had results that were only partly and Amnirana, but not Hylarana, as was congruent with ours and with those of Roe- done for our study; but Roelants et al. lants et al. (2004). Jiang and Zhou (2005; ®g. (2004), Kosuch et al. (2001), and Marmayou 41) found Pelophylax to form a monophy- et al. (2000) did not include species of Pa- letic group with Nidirana and Rana, and this purana or Tylerana. group formed the sister taxon of Amolops. The subsection Hylarana (the non-humer- The next more inclusive group was found to al-gland group) is polyphyletic as well. (This include the Rugosa±Glandirana clade. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 141

Dubois' (1992) Section Amerana is recov- The basal dichotomy of Rhacophoridae is ered as monophyletic and the sister taxon of as suggested by Channing (1989), with Buer- Pseudorana ϩ Rana ϩ Pseudoamolops. Sec- geria forming the sister taxon of the remain- tion Rana (our exemplars being Rana japon- ing rhacophorids. But beyond that level, ica, R. temporaria, and R. sylvatica)isre- however, our results are quite different. This covered as polyphyletic, with one component is not surprising, given the inherent con¯ict (Rana japonica and R. temporaria) being and lack of resolution in the morphological paraphyletic with respect to Pseudoamolops, data gathered so far, as discussed by J.A. and another (R. sylvatica) forming the sister Wilkinson and Drewes (2000). We will not taxon of Pantherana (section Pelophylax) ϩ discuss in detail the minor differences be- Section Lithobates. tween our results and those of J.A. Wilkinson Excluding Dubois' (1992) section Amer- et al. (2002) because, although our taxon ana, we ®nd American Rana (i.e., Aquarana, sampling was somewhat different, we includ- Lithobates, Trypheropsis, Sierrana, Panth- ed all of the same genes used in that study, erana, and Rana sylvatica) to form a mono- as well as our own. phyletic group, a conclusion reached previ- Our tree suggests polyphyly of Chirixalus, ously by Hillis and Wilcox (2005; ®g. 44). a conclusion to which others had previously Section Amerana (subgenera Aurorana plus arrived (e.g., J.A. Wilkinson et al., 2002): (1) Amerana [former Rana aurora and R. boylii one relatively basal clade (our Kurixalus eif- groups]) is most closely related to the Rana ®ngeri and ``Chirixalus'' idiootocus) noted temporaria group (including Pseudorana and previously by J.A. Wilkinson et al.'s (2002) Pseudamolops), an arrangement that suggests study for which the name Kurixalus Ye, Fei, the results of Case (1978) and Post and Uz- and Dubois (In Fei, 1999) is available; (2) zell (1981). Further discussion and generic the group associated with the name Chirix- realignments are provided in the Taxonomy alus (Chirixalus doriae and C. vittatus) form- section. ing a paraphyletic grade with respect to Chi- MANTELLIDAE AND RHACOPHORIDAE:We romantis (also illustrated by Delorme et al., ®nd Mantellidae and Rhacophoridae to be 2005; ®g. 49); and (3) our ``Chirixalus'' gra- monophyletic sister taxa deeply imbedded cilipes, except for Buergeria, being the sister within the traditional ``Ranidae'', together taxon of all rhacophorids. We, unfortunately, placed as the sister taxon of Raninae ϩ Nyc- did not sample ``Chirixalus'' palpebralis, tibatrachinae (®g. 65). The monophyly of the which J.A. Wilkinson et al. (2002; ®g. 48) combined Mantellidae and Rhacophoridae is found in a similar, basal, position, although not controversial and was suggested by a as shown by the dendrogram published by number of authors on the basis of DNA se- Delorme et al. (2005; ®g. 49), ``Chirixalus'' quence data (e.g., Emerson et al., 2000b; palpebralis, which we did not study, will Richards et al., 2000; J.A. Wilkinson et al., likely be found to be quite distant from 2002; Roelants et al., 2004; Roelants and Aquixalus (Gracixalus) gracilipes, once Bossuyt, 2005; Van der Meijden et al., 2005) Aquixalus is adequately sampled for molec- as well as the morphological data of Liem ular analysis. (1970). For mantellids, the phylogenetic structure A TAXONOMY OF LIVING we obtained is identical to that obtained by AMPHIBIANS Vences et al. (2003d): Boophis ((Aglyptodac- tylus ϩ Laliostoma) ϩ (Mantidactylus ϩ The taxonomy that we propose is consis- Mantella)), but different from that of Van der tent with the International Code of Zoologi- Meijden (2005) ((Aglyptodactylus ϩ Lalio- cal Nomenclature (ICZN, 1999). It will ap- stoma) ϩ (Boophis ϩ (Mantella ϩ Manti- pear to some that we have adopted an un- dactylus)). Although Vences et al. (2003d) ranked taxonomy. This is partially true, but demonstrated that Mantidactylus is deeply only for above-family-group nomenclature paraphyletic with respect to Mantella, our unregulated by the Code. Regardless of limited taxon sampling did not allow us to widespread perception, the Code does not test that result rigorously. govern nomenclature above the family 142 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 group. In fact, it barely mentions the exis- ation in taxon content according to various tence of Linnaean nomenclature above the authors, controversies regarding diagnosis, rank of the family group, and it does not or, more subtly, the taxon sampling regime specify particular ranks above that category. (Delorme et al., 2004) and underlying data Our suggested taxonomy is predicated on the used to infer the existence of particular taxa. recognition that the community of taxono- In other words, taxonomies are constructions mists has largely discarded its concerns re- for verbal and written communication that garding ranks above the family-group level. are inherently limited because they represent For example, one no longer hears arguments sets of theories of relationship and do not regarding whether Aves is a , coordinate communicate information on underlying data with a Class Amphibia, or whether it is at or assumptions of analysis. Precision in com- most a family within Archosauria. The rea- munication is enhanced by background son for this withering of concerns about knowledge on the part of those using the sys- ranks is that the concerns do not constitute tem for communication or, even better, hav- an empirical issue. Notions of rank equiva- ing the relevant tree(s) and data set(s) avail- lency are always based on notions of levels able from which the taxonomy was derived. of divergence, age, content, or size that are For an example of how taxonomies always bound to fail for a number of theoretical or must be quali®ed, Ford and Cannatella empirical reasons24. But, because nominal (1993) explicitly de®ned Hylidae as the most families and the ranks below them have been recent common ancestral species of Hemi- regulated by a more or less universally ac- phractinae, Hylinae, Pseudinae, Pelodryadi- cepted rulebook for more than 160 years nae, and Phyllomedusinae and all of its de- (Stoll, 1961), we are not inclined to easily scendants. This de®nition was implicitly throw out that rulebook or the universal com- changed by Darst and Cannatella (2004) to munication that it has fostered. Even though be the ancestor of Pelodryadinae, Phyllo- several of the criticisms of Linnaean nomen- medusinae, and Hylinae, and all of its de- clature are accurate, the alternatives so far scendants, because Hemiphractinae was dis- suggested have their own drawbacks. The In- covered to be paraphyletic and phylogeneti- ternational Code can be changed, and we ex- cally distant from ``other'' hylids. A casual pect that changes will be made to meet the glance at our tree will show that an appli- needs of modern-day problems. cation of Ford and Cannatella's (1993) cla- All taxonomies are rough and ready in the dographic de®nition of Hylidae would render sense that, except for the most general level as hylids nearly all arciferal neobatrachians, of communication, they must be quali®ed with the exception of Batrachophrynidae, implicitly or explicitly with respect to vari- Heleophrynidae, Limnodynastidae, Myoba- trachidae, and SooglossidaeÐa far cry from any content familiar to any who have used 24 A major underlying reason for this failure is that these terms and certainly not promoting pre- there are no natural classes in evolution that correspond to taxonomic ranks such as genus (contra Van Gelder, cision in the discussion of synapomorphies 1977; Dubois, 1982, 1988b, 2005; see Fink, 1990), fam- or even casual notions of similarity25. Fur- ily, or phylum. A related logical error is the notion that thermore, the molecular evidence that opti- organismal characteristics are transitive to their inclusive mizes as synapomorphies for Hylidae (sensu clades, except in an operational sense that is dependent on simplifying analytical assumptions (Frost and Kluge, stricto) in the study of Darst and Cannatella 1994), rendering such mistaken ideas such that there are (2004) must differ from those proposed by ``generic'' or ``family'' characters (e.g., see recognition Faivovich et al. (2005) simply because the of Taylorana by Dubois, 2005). Further, inasmuch as no objective criteria can correspond to subjective and idi- osyncratic notions of organismal similarity and differ- 25 Note that this kind of instability of nomenclature ence (Ghiselin, 1966), the idea that ranks could be tied and diagnosis is, in part, what Phylogenetic Nomencla- to special characters or levels of organismal divergence ture is supposed to address. Compare this with the ex- is seen to be particularly futile. Ranks in the Linnaean ample of Linnaean nomenclatural instability provided by system are assigned to taxa as part of a formal nomen- de Queiroz and Gauthier (1992) to demonstrate that this clatural/mnemonic system, not through discovery of kind of instability is found in both systems but appar- Linnaean ranks. ently is more typical of Phylogenetic Nomenclature. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 143 ingroup and outgroup taxon sampling of the Kress and DePriest, 2001; Niklas, 2001; Pa- latter is so much denser than that of the for- pavero et al., 2001; Pennisi, 2001; Brummitt, mer. As taxa are sampled more and more 2002; Carpenter, 2003; Keller et al., 2003; densely, more and more nonhomology will Kojima, 2003; Nixon et al., 2003; Schuh, be detected, with concomitant improvements 2003; Kluge, 2005; Pickett, 2005). What we in estimates of phylogeny (W.C. Wheeler, do think is that the conversation will contin- 1992; Zwickl and Hillis, 2002). The contro- ue for some time and that changes will take versy as it exists today, regardless of slogan- place, all discussed fully and not driven by eering, is about how to portray in words hy- the overheated sloganeering that, unfortu- potheses of monophyly, and revolves not nately, characterizes so much of the rhetoric about precision of communicating tree struc- at this timeÐon all sidesÐinasmuch as this ture or underlying data, but about how to is a political, not a scienti®c controversy (see maintain consistency of communication Pickett, 2005, for discussion). With respect among authors and across studies with a min- to our approach to taxonomy, we, in effect, imum of quali®cation. All systems so far take the easy way out, we follow the Inter- suggested have limitations; like all maps they national Code of Zoological Nomenclature must have limitations to be useful. Linnaean (ICZN, 1999) for regulated taxa (family taxonomy does promote useless rank contro- group and down) and apply an unranked tax- versies, but, as noted above and discussed onomy for unregulated taxa (above family more fully below, rigid application of cla- group), the hypotheses for these taxa being dographic de®nitions of taxonomic names derived from their included content and di- (such as the method proposed by de Queiroz agnostic synapomorphies. and Gauthier, 1992) brings other kinds of no- We expect that regulated nomenclature menclatural instability as well. will increasingly be pushed toward the ter- It is beyond the scope of this work to dis- minal taxa and that unregulated taxa will in- cuss at length the theory and practice of tax- creasingly be rankless. The reason for this is onomy and nomenclature. The ranked and that there really is a practical limit to the rankless alternatives to expressing phyloge- number of ranks that workers are willing to netic relationships in words theoretically are use. Systematists seemingly are not enam- endless but most recently and most clearly ored of new ranks such as grandorders, hy- discussed by Kluge (2005). To oversimplify perfamilies, epifamilies, and infratribes (e.g., his paper, currently competing systems for Lescure et al., 1986) or of the redundancies expressing phylogenetic relationships in and controversies over rank that are part and words are (1) Linnaean system (Linnaeus, parcel of ranked nomenclature (e.g., see Du- 1758); (2) Annotated Linnaean system (Wi- bois, 2005). So, our observation is that so- ley, 1981); (3) what Kluge termed ``Descent ciological pressures will push workers to- Classi®cation'' and proponents call ``Phylo- wards ever smaller families, especially be- genetic Taxonomy'' (de Queiroz and Gau- cause there is no scienti®c or sociological thier, 1992); (4) the ``Set Theory Classi®ca- pressure to construct larger families. Regard- tion'' system of Papavero et al. (2001), as less, we think that this process will corre- termed by Kluge; and (5) Kluge's (2005) spond with enormous progress in phyloge- ``Phylogenetic System''. netic understanding. We have taken a sixth approach, one that We suggest that the content of an above- we think is based on common sense, espe- family taxon as originally formed by an au- cially with respect to how systematists use thor renders an implied hypothesis of de- taxonomies and with respect to the state of scent, even if the concept of that taxon pre- the discussion, which is still very preliminary dates any particular theory of descent with and re¯ecting a deep ambivalence on the part modi®cation. We spent considerable time de- of taxonomists (for all sides of the contro- termining the original intent of various tax- versy see: Wiley, 1981; de Queiroz, 1988; de onomic names. Unfortunately, an examina- Queiroz and Gauthier, 1994; Cantino et al., tion of the original content of the groups de- 1997; Cantino et al., 1999; Benton, 2000; noted by these taxonomic names obviated the Nixon and Carpenter, 2000; Withgott, 2000; need to use many of them because they de- 144 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 viated so widely from all but a few of our types, or diagnoses but to tree topology26. phylogenetic hypotheses (e.g., in Avoiding this instability requires great cau- the original sense of Laurenti, 1768, not only tion in the application of that naming con- includes all frogs, but shares Proteus with his vention. Nevertheless, in our judgment it is Gradientia, a novel phylogenetic hypothe- unlikely that a fourth ``order'' of living am- sis!). phibians will be discovered, so application of In some cases (e.g., Caudata), we set aside the cladographic rules suggested by de Quei- the intent of the original author in favor of roz and Gauthier (1992) governing the ap- widespread current usage as suggested by plication of the names Anura, Caudata, and subsequent authors. The wisdom of this kind Gymnophiona could be salutary for purposes of action is open for discussion (see Dubois, of discussing fossil relatives of these crown 2004b, 2005), but increasingly the Interna- groups. tional Commission of Zoological Nomencla- Our strategy in designing a taxonomy for ture appears to be moving toward usage rath- unregulated taxa is to preserve, as nearly as er than priority as an important criterion to practical, the originally implied phylogenetic decide issues, so we take this to be the ap- content of named above-family-group taxa. propriate strategy. We also attempted to apply older names for As noted above, we are unconvinced that above family-group taxa, but because of the cladographic rules governing name assign- constant rede®nition of many of these taxa, ment (sensu de Queiroz and Gauthier, 1992) we could solve these only on an ad hoc basis, necessarily engender enhanced stability or depending on use, original intent, and recen- precision of discussion (except in the special cy of coining of the name(s). case of the crown-group approach to delim- In several cases, we changed the ranks of itation). However, we do think that associ- some regulated taxa from subfamilies to fam- ating names of extant taxa with content-spe- ilies to provide ¯exibility and help workers in the future with the problems inherent in ci®c, ostensively derived concepts (cf. Pat- ranked hierarchies. Because all names above terson and Rosen, 1977) will go a long way the regulated family group are unaddressed toward reducing the ``wobble'' of diagnoses by the International Code of Zoological No- associated with extant taxa as membership menclature (ICZN, 1999) we have regarded changes. One need only look at the history all of these names as unranked, but within of the use of ``Amphibia'' to see how the the zone normally associated with class and lack of an overarching concept of the taxon order (whatever that might mean to the read- has resulted in considerable drift of content er). We have not been constrained by rec- and diagnosis. As noted by Laurin (1998a: ommendations regarding name formations 10), until Huxley (1863), the term Amphibia and endings for ranks above the level of fam- applied only to Recent taxa. Haeckel (1866) ily group simply because we believe that and Cope (1880) rendered Amphibia para- these are unworkable and that they merely phyletic by the addition of some fossil taxa, exacerbate the previously recognized prob- with other authors (e.g., Romer, 1933) con- lems of taxonomic ranks (de Queiroz and tinuing the trend until all fossil tetrapods that Gauthier, 1992). were not ``'' were considered to be Although we argue that taxonomy should members of ``Amphibia''. Amphibia was re- re¯ect knowledge of phylogeny as closely as turned to monophyly only by Gauthier et al. possible, by eliminating all paraphyly and (1989) and subsequently restricted back to the groups of original intent by de Queiroz 26 ϩ and Gauthier (1992). If the application of a name for a taxon A (B C) is governed by the cladographic rule ``the ancestor of A Although the discussion is generating con- and B and all of its descendants'', and if new data show siderable self-examination by systematists, that the phylogenetic structure of this taxon has to we think that cladographically assigned tax- change to C (A ϩ B), the cladographically assigned ϩ onomic names (de Queiroz and Gauthier, name has to apply to A B and exclude C, even though the content of the taxon A ϩ B ϩ C has not changed. 1992) introduce a new kind of nomenclatural Linnaean nomenclature would be unaffected by this to- instability by tying names, not to content, pological change. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 145 recognizing all clades, we focused our atten- morphology presented by Hass (2003), the tion primarily on the taxonomy of clades only comparable data set across all frogs. above the ``genus level'' for three reasons. Despite the fact that this is, so far, the most First, for the most part our taxon sampling data-heavy analysis of amphibians, we ex- was inadequate to test prior hypotheses of pect to be criticized for presenting this tax- intrageneric relationships for most genera. onomy for four reasons: The practical implication of this inadequacy (1) This taxonomy will be criticized both is that we lack evidence to refer the majority as premature and as not conservative. How- of species in a more re®ned generic taxono- ever, the underlying cladogram re¯ects the my, which would require those species to be best overall estimate of phylogeny on the placed as incertae sedis, a cumbersome so- most thorough dataset applied to the issue. lution with little payoff. The other alterna- The alternativeÐto stick for sociological rea- tiveÐexpanding the content of genera to en- sons to an old taxonomy that is clearly mis- force monophyly is equally unsatisfactory in leading and based on relatively little evi- these cases, as it overlooks the ®ner-level denceÐcertainly will not ef®ciently promote knowledge of phylogeny that exists but, for additional research. Some will attempt to de- practical reasons was not brought to bear in fend as conservative the old arrangements, this analysis. Secondly, the bulk of phylo- especially favored paraphyletic groups, but genetic research since the mid-1970s has fo- mostly this will mean socially conservative, cused primarily on ``genus-level'' diversity, not scienti®cally conservative, something which means that a considerable amount of detrimental to scienti®c progress. As re- evidence, both molecular and morphological, vealed in the ``Review of the Current Tax- has been generated for those groups, most of onomy'', much of the existing taxonomy of which was not included in the present study. amphibians stands on remarkably little evi- Third, we see the value of the present con- dence and has simply been made plausible tribution to be in framing ®ner level prob- through decades of repetition and rei®cation. lems that are better addressed by regional A similar argument is that we should re- specialists who can achieve more exhaustive tain the status quo with respect to taxonomy taxon and character sampling. until we are ``more sure'' of a number of Our consensus tree is shown in ®gure 50 weakly recovered relationships. This position (insert), which also displays the current and ignores how little evidence underlies the ex- recommended family-group taxonomy. We isting classi®cations. Indeed, our taxonomy modify the current generic taxonomy in plac- explains more of the evolution of amphibian es in this section, but those changes are not characteristics than the existing classi®ca- re¯ected in the ®gure for purposes of clarity tion(s) and has the distinction of attempting in ``Results''. With minor exceptions, all to be explicitly monophyletic over all of the clades are highly corroborated by molecular evidence analyzed. We are surely mistaken evidence (and morphological evidence on in several places, but this is better than con- many branches as well) as estimated by Bre- tinuing to recognize taxonomic groups that mer values and parsimony jackknife frequen- are known to be inconsistent with evolution- cies (see below and appendix 4 for these val- ary history, regardless of social convention. ues by branch). Because this study rests on We do go beyond our data in several places the largest amount of data applied to the (e.g., Brachycephalidae, Bufonidae) and rec- problem of the relationships among living ognize some groups whose monophyly we amphibians, we provide a new taxonomy that have not rigorously tested. The reason for we think will provide a better reference for this is to attempt to delimit new hypotheses additional progress. and not sit idly by while major problems are This taxonomy of living amphibians is concealed by convention. Critics may charge based on a phylogenetic analysis of 532 ter- that this is no different from post facto ``di- minals, on the basis of a total of 1.8 million agnosis'' of subjective similarity groupings bp of nuDNA and mtDNA sequence data (xÅ (e.g., Dubois, 1987 ``1985'', 1992). Howev- ϭ 3.7 kb/terminal) in addition to the mor- er, in each case we think there is good reason phological data from predominantly larval to expect our taxa to obtain as monophylet- 146 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 icÐand that leaving the taxonomy as it exists bined analysis this will constitute a test of does nothing to promote improved under- our results and taxonomy. This problem calls standing of evolutionary history. for careful evaluation of all morphological (2) Some will be critical of the fact that characters across all taxonomic groups con- we have not included all of the morpholog- comitant with the evaluation of relevant fos- ical data that have been presented by other sil groups. This is a big task, but one worth authors. Early in the development of this doing well. Unfortunately, this kind of in- work, we made an attempt to marshal the frastructural science is not ¯ashy and there- disparate but extensive number of characters fore will not attract funding from already presented by such authors as J.D. Lynch oversubscribed and under®nanced granting (1973), Estes (1981), Duellman and Trueb agencies. (See Maienschein, 1994, for an es- (1986), Milner (1988), Nussbaum and Wil- say on the dangers to science from the pre- kinson (1989), Trueb and Cloutier (1991), occupation by administrators and funding Ford and Cannatella (1993), Larson and agencies with the ``cutting edge''.) Dimmick (1993), Milner (1993 (1994), (3) Some will criticize our analytical McGowan and Evans (1995), Shubin and methods. We have been conservative with re- Jenkins (1995), M. Wilkinson and Nussbaum spect to analytical assumptions. Beyond at- (1996), Laurin and Reisz (1997), Laurin tempting to maximize explanatory ef®ciency, (1998a), Maglia (1998), Carroll et al. (1999), some workers prefer to incorporate assump- M. Wilkinson and Nussbaum (1999), Carroll tions about the evolutionary process by the (2000a), Laurin et al. (2000), Milner (2000), addition of particular evolutionary models. J.S. Anderson (2001), Gardner (2001), Kap- This is obviously a discussion that we think lan (2001), Zardoya and Meyer (2001), will continue for a long time because of the Gardner (2002), Gower and Wilkinson serious philosophical and evidentiary issues (2002), Laurin (2002), Scheltinga et al. involved. (2002), and BaÂez and Pugener (2003). What Some will be uncomfortable that such a we found, not surprisingly, is that different large proportion of our data are molecular studies tended to generalize across different (even though most of our results are gener- exemplars, even if they were working on the ally conventional). We believe that it is better same groups, and that in some cases putative to present a taxonomy that represents explic- synapomorphies had been so rei®ed through repetition in the literature that it was dif®cult, it, evidence-based hypotheses of relation- if not impossible, to ascertain which taxa ships than to retain a taxonomy solely be- (much less which specimens) had actually cause we are used to it. Some will want to been evaluated for which characters. We also exclude all sequence data that require align- found that many of the new characters re- ment. Unfortunately, this assumes that same- main in unpublished dissertations (e.g., Can- length sequences lack evidence of having natella, 1985; Ford, 1990; S.-H. Wu, 1994; had length variation, an assumption not sup- Graybeal, 1995; da Silva, 1998; Scott, 2002), ported by evidence (Grant, unpubl.). Others where ethics dictates they not be mined for will want to ``correct'' alignments manually information if they are new, and prudence (although this is likely to increase the num- dictates that the information in them not be ber of transformations required to explain se- taken at face value if they are old and still quence variation). Although such methodo- unpublished. logical choices are crucial and should contin- Further, most of the paleontological liter- ue to be debated (indeed, we urge authors ature re¯ects such incomplete sampling of and editors of empirical papers to be more living taxa as to oversimplify living diversi- explicit about both their methods of align- ty. (One does not read evolution from the ment and analysis and their reasons for em- rocks, but the rocks certainly are an under- ploying them), the issue at hand is that it is sampled component of our study.) Reconcil- time to move away from a taxonomy known ing all morphological descriptions of char- to be fatally ¯awed and that promotes mis- acters in comparable form, obviously, is the understanding and into a scienti®c dialogue next big step, for someone else, and in com- that will promote a much improved under- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 147 standing of the evolution of amphibian taxic, changes, or incorrect subsequent spellings. life history, and morphological diversity. More extensive discussion of speci®c no- (4) We will be trivially criticized for for- menclatural issues are dealt with in appendix mulating new taxonomic names with 19 au- 6. A summary of generic name changes is thors. Times change and collaborations on presented in appendix 7. We do not address this scale are necessary to answer global fossil taxa, although they can be placed with- questions. That a new name can have 19 au- in this framework with relatively little effort. thors may be cumbersome, but, authorship is Dubois (2005) recently provided a taxonomy not part of the scienti®c name. And, regard- of living amphibians and their fossil relatives less of recommendations made in the Code (Neobatrachii in his sense). Because his tax- (ICZN, 1999) this authorship re¯ects accu- onomy appeals to a taxonomic philosophy rately the extensive effort in collecting sam- deeply steeped in the importance of ranks ples, sequencing, data analysis, and writing and personal authority and the unimportance that work on this scale requires. of evidence and logical consistency with Although our results will undoubtedly al- evolutionary history, we comment on it only low considerable progress to be made, by where necessary. nearly doubling the number of amphibian For taxa above the family group, which species for which DNA sequences are avail- are not regulated by the Code, homonymy able in GenBank, projects such as this one remains an unresolved issue in amphibian generate questions as well as answers. Our nomenclature because, even if the original results therefore will provide a reasonably author intended one content (i.e., one hy- well-tested departure point for future studies pothesis of relationship), subsequent authors by identifying outstanding problems that are saw (and may see) little problem in rede®n- especially worthy of investigation. ing these names to ®t revised hypotheses of relationship. For these taxa we do not pro- TAXONOMIC ACCOUNTS vide a synonymy because in the absence of any regulatory tradition of above-family- Below we present ancillary information group nomenclature, we have tried to opti- and discussion to accompany the taxonomy mize on the hypothesis of relationship in- presented in ®gures 50 (insert) and 66 (a re- tended by the author (or rede®ner) of that duced tree of family-group taxonomy). (Ta- taxon. Although we do not provide a ``syn- ble 5 provides names of taxa/branches on the onymy'' in the accounts of unregulated taxa, interior of the tree shown in ®gure 66, and we variably note in appendix 6 (``Nomencla- ®gure 67 provides the taxonomy of amphib- ture'') synonyms, near-synonyms, and prob- ians in condensed form.) Most morphologi- lematic nomenclatural issues. cal evidence is addressed in accounts, but The structure of the taxonomic accounts is molecular synapomorphies are provided straight-forward with several categories of where relevant in appendix 5, with branch information: (1) the name and author of the numbers corresponding with those noted in taxon (and where appropriate and to enhance the various ®gures. We are conservative in navigation among records, bracketed num- the scienti®c sense in that we stick close to bers are associated that correspond to the the preponderance of evidence and not to tra- numbered branches in our various ®gures dition. Genera in bold listed under Content and tables in ``Results''); (2) a list of avail- represent those from which one or more spe- able names if application of the name is reg- cies were included in our analysis (as DNA ulated by the International Code of Zoolog- sequences either generated or by us or others ical Nomenclature; (3) an etymology if the and available via GenBank). A justi®cation name of a taxon is used for the ®rst time; (4) is provided for inclusion of taxa that were the name and branch number of the imme- not sampled. Synonymies provided in the diately more inclusive taxon; (5) the name family group and below conform to the In- and branch number of the sister taxon; (6) a ternational Code of Zoological Nomenclature statement of the geographic distribution of (ICZN, 1999). We include citations only to the taxon; (7) the concept of the taxon in original uses and not to emendations, rank terms of content; and (8) a characterization 148 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

TABLE 5 Branch Numbers and Taxon Names Corresponding to Internal Branches on Figure 50 Left side, sorted by branch number; right side, sorted by taxon name.

Branch Branch number Taxon name number Taxon name 6 Amphibia 91 Acosmanura 7 Gymnophiona 192 Africanura 8 Rhinatrematidae 143 Afrobatrachia 9 Stegokrotaphia 460 Agastrorophrynia 23 Batrachia 244 Aglaioanura 24 Caudata 109 Allodapanura 25 Cryptobranchoidei 191 Ametrobatrachia 29 Diadectosalamandroidei 6 Amphibia 30 Hydatinosalamandroidei 92 Anomocoela 31 Perennibranchia 74 Anura 35 Treptobranchia 371 Athesphatanura 49 Plethosalamandroidei 319 Australobatrachia 50 Xenosalamandroidei 23 Batrachia 74 Anura 24 Caudata 77 Lalagobatrachia 440 Chthonobatrachia 78 Xenoanura 366 Cladophrynia 84 Sokolanura 85 Costata 85 Costata 25 Cryptobranchoidei 91 Acosmanura 461 Dendrobatoidea 92 Anomocoela 29 Diadectosalamandroidei 93 Pelodytoidea 425 Diphyabatrachia 96 Pelobatoidea 7 Gymnophiona 105 Neobatrachia 448 Hesticobatrachia 107 Phthanobatrachia 30 Hydatinosalamandroidei 108 Ranoides 314 Hyloides 109 Allodapanura 77 Lalagobatrachia 143 Afrobatrachia 148 Laurentobatrachia 144 Xenosyneunitanura 424 Leptodactyliformes 148 Laurentobatrachia 349 Meridianura 180 Natatanura 321 Myobatrachoidea 183 Victoranura 180 Natatanura 189 Telmatobatrachia 105 Neobatrachia 191 Ametrobatrachia 348 Nobleobatrachia 192 Africanura 318 Notogaeanura 200 Pyxicephaloidea 96 Pelobatoidea 220 Saukrobatrachia 93 Pelodytoidea 244 Aglaioanura 31 Perennibranchia 245 Rhacophoroidea 107 Phthanobatrachia 269 Ranoidea 49 Plethosalamandroidei 314 Hyloides 200 Pyxicephaloidea 318 Notogaeanura 269 Ranoidea 319 Australobatrachia 108 Ranoides 321 Myobatrachoidea 245 Rhacophoroidea 348 Nobleobatrachia 8 Rhinatrematidae 349 Meridianura 220 Saukrobatrachia 366 Cladophrynia 84 Sokolanura 368 Tinctanura 9 Stegokrotaphia 371 Athesphatanura 189 Telmatobatrachia 424 Leptodactyliformes 368 Tinctanura 425 Diphyabatrachia 35 Treptobranchia 440 Chthonobatrachia 183 Victoranura 448 Hesticobatrachia 78 Xenoanura 460 Agastorophrynia 50 Xenosalamandroidei 461 Dendrobatoidea 144 Xenosyneunitanura 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 149

Fig. 66. A simplied tree of our results (®g. 50) tree showing families. Numbers on branches allow branch lengths, Bremer, and jackknife values, as well as molecular synapomorphies to be identi®ed in appendices 4 and 5. See table 5 for taxon names associated with internal numbered branches and ®gure 67 for a complete summary of taxonomy. 150 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 67. Summary taxonomy of living amphibians. Quotation marks around names denote nonmon- ophyly. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 151

Fig. 67. Continued. 152 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 67. Continued. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 153

Fig. 67. Continued. 154 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 67. Continued. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 155

Fig. 67. Continued. 156 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 67. Continued. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 157

Fig. 67. Continued. 158 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 67. Continued. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 159

Fig. 67. Continued. 160 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 67. Continued. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 161

Fig. 67. Continued. 162 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 67. Continued. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 163

Fig. 67. Continued. 164 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 and diagnosis, which is merely a general scienti®c community will be to let sleeping summary of the salient features of the ani- dogs lie. mals that are included in the taxon under dis- In a few places in the taxonomy, we do cussion, and characters (either synapomorph- not render taxonomic changes suggested by ic or not) that differentiate this taxon from our tree. In the cases of ``Eleutherodactylus'' others. Where a character is thought to be a and ``Centrolene'', our sampling density is synapomorphy, this is stated. If the explicit so low compared to the species diversity that statement is not made, then the character our results could not be practically translated should be assumed to be of unknown polar- into an informative taxonomy. In two other ity. Because we included Haas' (2003) char- cases, the reason is that we do not consider acters in the analysis, for each group we list our results to constitute a suf®cient test of a all unambiguously optimized synapmorphies published cladogram, based on a data set that for that data set, reported using Haas' origi- includes as a subset the data over which we nal numbering scheme (e.g., Haas 34.1). Oth- generalized. The ®rst of these is in Hylinae, erwise, we have not attempted to be exhaus- where our data represent a subset of the data tive nor to make these differentia explicitly (and concomitant results) of Faivovich et al. comparable for the simple reason that the (2005), meaning that our analysis does not challenge of sorting out the published record constitute an adequate test of their results. regarding the morphological characteristics The second is plethodontid salamanders, of amphibians will be enormous and, clearly, where the placement of certain taxa (i.e., is outside of the scope of this work27. Re- Hemidactylium and Batrachoseps) in our tree gardless, that next step is an important one is based on a subset of data in a published in elucidating the morphological evolution of tree (Macey, 2005), which came to different amphibians. The characterization and diag- conclusions regarding those critical taxa, nosis is followed by (9) various systematic based, at least with respect to those taxa, on comments and discussion. Considerable tax- a more inclusive data set (although the as- onomic ``sausage making'' is evident in these sumptions of analysis were subtly different). sections, particularly with respect to the larg- In these two cases we do not reject the con- er and more chaotic genera, which we have clusions of these authors, pending even more not been shy about partitioning because con- inclusive analyses. siderable redistribution of taxonomic names needs to happen if we are going to progress [6] AMPHIBIA GRAY, 1825 towards a taxonomy that re¯ects evolution- Amphibia Gray, 1825: 213. (See appendix 6 for ary history. In some places our changes have further nomenclatural discussion.) not been successful in producing a taxonomy that is entirely monophyletic. Our rationale RANGE: Worldwide on all continents ex- for failing to propose a more precise taxon- cept Antarctica and most oceanic islands, in omy was given earlier, and we are con®dent cold-temperate to tropical habitats. that future work will correct this shortcoming CONCEPT AND CONTENT: Amphibia is a in our proposal. To that end, we emphasize monophyletic taxon composed of [7] Gym- and discuss the speci®c problems and inad- nophiona J. MuÈller, 1832, and [23] Batrachia equacies for each of these cases. Some work- Latreille, 1800, constituting the crown group ers will not appreciate the loose-ends that re- (i.e., living) amphibians (sensu Amphibia main untied and will prefer the old approach Gray, 1825; not Amphibia of Linnaeus, of concealing these questions. Our position, 1758; cf. de Queiroz and Gauthier, 1992). however, is that unless these problems are CHARACTERIZATION AND DIAGNOSIS: Be- advertised, the sociological response of the yond our molecular data, Amphibia is diag- nosed by many morphological characters. 27 For some clades, diagnosis by nongenetic characters Amphibians, like mammals, retain plesiom- is not currently possible. To make molecular diagnosis orphically the glandular skin of ancestral tet- more tangible and descriptively simple, we also report salient charateristics, such as length variation in 28S se- rapods. They do not have the apomorphy of quences (appendix 3), as well as unambiguous molecular epidermal scales found in sauropsids (turtles transformations (appendix 5), where needed. and diapsids). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 165

Trueb and Cloutier (1991) and Ruta et al. lians are a bizarre group of legless amphib- (2003) provide extensive discussions of the ians, primitively oviparous with aquatic lar- synapomorphies of Amphibia (as Lissamphi- vae (Rhinatrematidae, Ichthyophiidae), al- bia) in the context of fossil groups. Syna- though some species are ovoviparous (with pomorphies of Amphibia include (Trueb and or without direct development) and - Cloutier, 1991): (1) loss of the ing, as re¯ected by considerable numbers of bones; (2) loss of the supratemporal bone; (3) osteological modi®cations. loss of the tabular bone; (4) loss of the post- Beyond our molecular data, the following orbital bones; (5) loss of the jugal bone; (6) morphological characters have been suggest- loss of the interclavicle; (7) loss of the cleith- ed to be synapomorphies (Nussbaum and rum; (8) papilla amphibiorum present in ear; Wilkinson, 1989; Trueb and Cloutier, 1991): (9) opercular element associated with the (1) lacking limbs and girdles (except for one columella; (10) fat bodies present that orig- antecedent fossil taxon not included in the inate from the germinal ridge associated with crown group; Carroll, 2000b); (2) presence the gonads; and (11) pedicellate and bicuspid of a dual jaw-closing mechanism; (3) pres- teeth that are replaced mediolaterally (re- ence of an eversible phallodeum in males versed in some taxa). formed by a portion of the cloacal wall; (4) SYSTEMATIC COMMENTS: Amphibia is high- annuli encircling the body; (5) paired sen- ly corroborated as a taxon, but this only im- sory tentacles on the snout. plies that all living amphibians are more SYSTEMATIC COMMENTS: M. Wilkinson has closely related to each other than to any other an extensive morphological matrix of more living species and does not address the place- than 180 character transformations (see also ment of amphibian groups within the larger Nussbaum and Wilkinson, 1995; M. Wilkin- structure of relevant fossil tetrapods. All son and Nussbaum, 1996, 1999), which will work so far on the overall placement of am- appear elsewhere, analyzed in conjunction phibians (lissamphibians) among fossil with this and additional evidence. groups has depended on inadequate sampling of living taxa and, with the exception of Gao [8] FAMILY: RHINATREMATIDAE NUSSBAUM, and Shubin (2001), has ignored available 1977 molecular data. We hope that additional work Rhinatrematidae Nussbaum, 1977: 3. Type genus: on fossil groups, combined with the data pre- Rhinatrema DumeÂril and Bibron, 1841. sented here, and a better account of living diversity, will further elucidate those rela- IMMEDIATELY MORE INCLUSIVE TAXON: [7] tionships. Gymnophiona J. MuÈller, 1832. SISTER TAXON: [9] Stegokrotaphia Canna- [7] GYMNOPHIONA J. MUÈ LLER, 1832 tella and Hillis, 1993. RANGE: Tropical northern South America Gymnophiona J. MuÈller, 1832: 198. (See appendix 6 for nomenclatural discussion.) from Amazonian Peru and Brazil, through eastern Ecuador, Colombia, , and IMMEDIATELY MORE INCLUSIVE TAXON: [6] the Guianas. Amphibia Gray, 1825. CONTENT: Epicrionops Boulenger, 1883; SISTER TAXON: [23] Batrachia Latreille, Rhinatrema DumeÂril and Bibron, 1841. 1800. CHARACTERIZATION AND DIAGNOSIS: Rhina- RANGE: Pantropical, except for Madagas- trematids are oviparous with aquatic larvae. car and southeast of Wallace's Line; not yet They are strongly annulated with numerous reported from central equatorial Africa. secondary and tertiary grooves. Like ichth- CONCEPT AND CONTENT: Gymnophiona is a yophiids, rhinatrematids have a short tail and monophyletic taxon containing the living the are visible, although they lie beneath caecilians (cf. J. MuÈller, 1832; Cannatella the skin in bony sockets. The tentacle arises and Hillis, 1993): [8] Rhinatrematidae Nuss- near the anterior edge of each eye, and the baum, 1977, and [9] Stegokrotaphia Canna- contains a (Nussbaum, tella and Hillis, 1993. 1977). CHARACTERIZATION AND DIAGNOSIS: Caeci- Beyond the molecular evidence, the fol- 166 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 lowing morphological characters have been eye; (3) frontal and squamosal articulate; (4) suggested to be synapomorphies (Duellman stegokrotaphic skull; (5) vomers in contact and Trueb, 1986; M. Wilkinson and Nuss- throughout their entire length; (6) sides of the baum, 1996): (1) dorsolateral process of the parasphenoid converge anteriorly; (7) quad- os basale present; (2) loss or fusion of the rate and maxillopalatine lack articulation; (8) prefrontal with the maxillopalatine; (3) sec- squamosal and frontal in contact; (9) ptery- ondary annulus/primary annulus greater than goid reduced; (10) basipterygoid present; one; and (4) fourth ceratobranchial absent. In (11) retroarticular process long and usually addition, the prefrontals are fused with the curved dorsally; (12) third and fourth cera- maxillopalatine as in caeciliids, but not in tobranchial fused; (13) anterior ®bers of the ichthyophiids and outgroups, rendering the m. interhyoideus do not insert on ceratohyal; optimization of this character arguable. (14) m. interhyoideus posterior in two bun- dles; (15) orientation of m. interhyoideus [9] STEGOKROTAPHIA CANNATELLA AND posterior is longitudinal rather than oblique; HILLIS, 1993 and (16) m. depressor mandibulae longitu- Stegokrotaphia Cannatella and Hillis, 1993: 2. dinally oriented rather than vertically orient- ed. IMMEDIATELY MORE INCLUSIVE TAXON: [7] Gymnophiona J. MuÈller, 1832. [10] FAMILY: ICHTHYOPHIIDAE TAYLOR, 1968 SISTER TAXON: [8] Rhinatrematidae Nuss- Epicria Fitzinger, 1843: 34. Type genus: Epicrium baum 1977. Wagler, 1828. Suppressed for purposes of pri- RANGE: Tropics of southern North Ameri- ority but not homonymy in favor of Ichthy- ca, South America, equatorial East and West ophiidae by Opinion 1604 (Anonymous, 1990: Africa, islands in the Gulf of Guinea, Sey- 166). chelles, and India; Philippines and India to Ichthyophiidae Taylor, 1968: 46. Type genus: southern China, Thailand, Indochina and the Ichthyophis Fitzinger, 1826. Placed on Of®cial Malayan archipelago. List of Family-Group Names in Zoology by Opinion 1604 (Anonymous, 1990: 166±167). CONCEPT AND CONTENT: Stegokrotaphia is a monophyletic group containing [10] Ich- Uraeotyphlinae Nussbaum, 1979: 14. Type genus: Uraeotyphlus Peters, 1880 ``1879''. thyophiidae Taylor, 1968, and [12] Caecili- idae Ra®nesque, 1814 (cf. Cannatella and IMMEDIATELY MORE INCLUSIVE TAXON: [9] Hillis, 1993). Stegokrotaphia Cannatella and Hillis, 1993. CHARACTERIZATION AND DIAGNOSIS: Ste- SISTER TAXON: [12] Caeciliidae Ra®n- gokrotaphian caecilians show variation in re- esque, 1814. productive mode (from aquatic larvae to RANGE: India to southern China, Thailand, ) and morphology, with some and through the Malayan archipelago to the retaining tails (Ichthyophiidae) and others Greater Sunda Islands and Philippines. (Caeciliidae) having lost them (even though CONTENT: Caudacaecilia Taylor, 1968; a pseudotail may be present). The eyes may ``Ichthyophis'' Fitzinger, 1826 (see System- be visible (e.g., Ichthyophis), completely hid- atic Comments); Uraeotyphlus Peters, 1880 den beneath bone (e.g., Scolecomorphus), or ``1879''. completely absent (Boulengerula). Unlike in CHARACTERIZATION AND DIAGNOSIS: Ichth- rhinatrematids, the tentacle originates in front yophiids are oviparous with aquatic larvae, of the eye and may be nearly as far forward both features being plesiomorphies. Like as the nostril. A stapes is generally present rhinatrematids, ichthyophiids plesiomorphi- but is lost in some taxa (Nussbaum, 1977). cally retain a true tail. Eyes are externally Beyond the molecular evidence, the fol- visible beneath the skin and are in bony lowing morphological characters have been sockets. The tentacle arises between the nos- suggested to be synapomorphies (Duellman tril and the eye, generally closer to the eye and Trueb, 1986; M. Wilkinson and Nuss- in Ichthyophis and Caudacaecilia and ante- baum, 1996): (1) mouth subterminal or re- rior near the nostril in Uraeotyphlus. A sta- cessed rather than terminal; (2) tentacular pes is present (Nussbaum, 1977). opening anterior to the anterior edge of the Beyond the molecular evidence supporting 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 167 the monophyly of this group the following Caeciliadae Gray, 1825: 217. Type genus: Cae- morphological characters have been suggest- cilia Linnaeus, 1758. ed to be synapomorphies (M. Wilkinson and Siphonopina Bonaparte, 1850: 1 p. Type genus: Nussbaum, 1996): (1) vomers in contact an- Siphonops Wagler, 1828. teriorly (convergent in Siphonops, Scoleco- Typhlonectidae Taylor, 1968: xi, 231. Type genus: Typhlonectes Peters, 1880 ``1879''. morphus, and Gegeneophis); (2) atria divided Scolecomorphidae Taylor, 1969a: 297. Type ge- externally; (3) anterior pericardial sac long nus: Scolecomorphus Boulenger, 1883. and extensive; (4) posterior internal ¯exures Dermophiinae Taylor, 1969b: 610. Type genus: in the m. rectus lateralis II; (5) tracheal lung Dermophis Peters, 1880 ``1879''. present (also in Typhlonectes). Herpelinae Laurent, 1984a: 199±200. Type genus: SYSTEMATIC COMMENTS: As noted in ``Re- Herpele Peters, 1875. sults'', the preponderance of evidence sug- Geotrypetoidae Lescure et al., 1986: 162. Type gests that ``Ichthyophis'' is paraphyletic with genus: Geotrypetes Peters, 1880. respect to Uraeotyphlus. Unfortunately, the Grandisoniilae Lescure et al., 1986: 164. Type ge- number of species currently assigned to nus: Taylor, 1968. Indotyphlini Lescure et al., 1986: 164. Type ge- ``Ichthyophis'' is large and mostly unsam- nus: Taylor, 1960. pled, and the relationships among them (and Afrocaeciliiti Lescure et al., 1986: 164. Type ge- Caudacaecilia [unsampled by us] and nus: Afrocaecilia Taylor, 1968. Uraeotyphlus) are unclear. Nussbaum and Brasilotyphlili Lescure et al., 1986: 166. Type ge- Wilkinson (1989: 31) suggested that Cau- nus: Taylor, 1968. dacaecilia and Ichthyophis might both be Pseudosiphonopiti Lescure et al., 1986: 166. Type polyphyletic inasmuch as they are diagnosed genus: Pseudosiphonops Taylor, 1968. solely on single characters of known vari- Oscaecilioidae Lescure et al., 1986: 167. Type ge- ability. We do not place Caudacaecilia and nus: Taylor, 1968. Uraeotyphlus into the synonymy of Ichthy- Gymnopiilae Lescure et al., 1986: 168. Type ge- nus: Peters, 1874. ophis, although to do so would certainly ren- Potamotyphloidea Lescure et al., 1986: 169. Type der a monophyletic taxonomy. Ongoing genus: Potamotyphlus Taylor, 1968. work by M. Wilkinson, Nussbaum, and col- Pseudotyphlonectini Lescure et al., 1986: 170. laborators should provide a monophyletic Type genus: Pseudotyphlonectes Lescure, Ren- taxonomy without resorting to that minimal- ous, and Gasc, 1986. ly informative one. In the interim we place quotation marks around ``Ichthyophis'' (the IMMEDIATELY MORE INCLUSIVE TAXON: [9] only ichthyophiid genus for which we have Stegokrotaphia Cannatella and Hillis, 1993. evidence of paraphyly). In the face of strong SISTER TAXON: [10] Ichthyophiidae Taylor, evidence of paraphyly of ``Ichthyophis'', 1968. maintaining a family-group name for Uraeo- RANGE: Tropics of Mexico, Central Amer- typhlus is unnecessary, and we therefore ica, and South America; equatorial East and place Uraeotyphlinae in the synonymy of West Africa and islands in the Gulf of Guin- Ichthyophiidae. Other than assuming that the ea, Seychelles, and India. morphological synapomorphies are suf®- CONTENT: Nussbaum and cient, stronger evidence of monophyly of Wilkinson, 1995; Boulengerula Tornier, Ichthyophiidae will require sampling of Cau- 1896; Brasilotyphlus Taylor, 1968; Caecilia dacaecilia and more ``Ichthyophis''. Never- Linnaeus, 1758; Peters, 1880; theless, we make the hypothesis that Ichth- Crotaphatrema Nussbaum, 1985; Dermo- yophiidae is a monophyletic taxon and trust phis Peters, 1880; Gegeneophis Peters, 1880; that others will elucidate this further. Geotrypetes Peters, 1880; Grandisonia Tay- lor, 1968; Gymnopis Peters, 1874; Herpele [12] FAMILY: CAECILIIDAE RAFINESQUE, 1814 Peters, 1880; Hypogeophis Peters, 1880; Idi- Cecilinia Ra®nesque, 1814: 104. Type genus: ocranium Parker, 1936; Indotyphlus Taylor, Caecilia Linnaeus, 1758. See Dubois (1985: 1960; Taylor, 1968; Micro- 70). Authorship but not spelling to be con- caecilia Taylor, 1968; Mimosiphonops Tay- served following Opinion 1830 (Anonymous, lor, 1968; Taylor, 1968; Oscae- 1996: 68±69). cilia Taylor, 1968; Parvicaecilia Taylor, 168 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

1968; Potomotyphlus Taylor, 1968; ly coordinate with Scolecomorphinae and Boulenger, 1909; Parker, Typhlonectinae would merely push the par- 1941; Scolecomorphus Boulenger, 1883; Si- aphyly to the subfamily level, as was done phonops Wagler, 1828; Wake, by Hedges et al., 1993.) Although molecular 1987; Typhlonectes Peters, 1880. evidence corroborates the monophyly of CHARACTERIZATION AND DIAGNOSIS: Caeci- Scolecomorphinae, the following morpho- liids represent the bulk of caecilian diversity logical characters also diagnose that taxon and, not surprisingly, show considerable (Nussbaum and Wilkinson, 1989; M. Wilkin- morphological and reproductive variation. son and Nussbaum, 1996): (1) temporal fossa Some taxa are oviparous with aquatic larvae secondarily large (also in Typhlonectinae, (e.g., Praslinia), whereas others are ovipa- though not homologously); (2) premaxillae rous with direct development in the egg (e.g., separate; (3) septomaxilla present; (4) pre- Hypogeophis, Idiocranium, and Boulengeru- frontals present; (5) basipterygoid process la), and others are viviparous (e.g., Schisto- small; and (6) no stapes. Similarly, for Ty- metopum, Dermophis, and typhlonectines). phlonectinae, the following apomorphic Unlike Ichthyophiidae and Rhinatrematidae, characters diagnose that taxon (Nussbaum no caeciliid possesses a true tail, although and Wilkinson, 1989; M. Wilkinson and some (e.g., typhlonectines) have a pseudotail. Nussbaum, 1996): (1) temporal fossa second- Most species are terrestrial and burrowing, arily large (also in Scolecomorphinae, although some (e.g., typhlonectines) are sec- though not homologously); and (2) choanae ondarily aquatic. At least one species (Atre- large, with well-developed valves. tochoana eiselti: Typhlonectinae) is totally lungless (Nussbaum and Wilkinson, 1995). [23] BATRACHIA LATREILLE, 1800 Most taxa have stapes, but all scolecomor- phines lack them (Nussbaum, 1977). Batrachii Latreille, 1800: xxxvii. A Latinization of Batraciens Brongniart, 1800b, emended to Beyond the molecular evidence, the fol- Batrachia by Ra®nesque, 1814: 103. (See ap- lowing morphological characters have been pendix 6 for nomenclatural discussion.) suggested to be synapomorphies of this group (M. Wilkinson and Nussbaum, 1996): IMMEDIATELY MORE INCLUSIVE TAXON: [6] (1) tail absent; (2) premaxillae and nasal Amphibia Gray, 1825. bones fused; (3) septomaxillae reduced or SISTER TAXON: [7] Gymnophiona J. MuÈller, absent (reversed in Scolecomorphus); (4) 1832. pterygoid absent; (5) basiptergygoid process RANGE: Cosmopolitan in cold-temperate to large (small in Scolecomorphus); (6) fused tropical habitats, except for extreme northern third and fourth ceratobranchials greatly ex- latitudes, Antarctica, and most oceanic is- panded; and (7) vent circular or transverse, lands. not longitudinally oriented (reversed in Sco- CONCEPT AND CONTENT: Batrachia is a lecomorphus). monophyletic taxon containing [24] Caudata SYSTEMATIC COMMENTS: Recognition of the Fischer von Waldheim, 1813, and [74] Anura nominal families Typhlonectidae (Atreto- Fischer von Waldheim, 1831 (cf. Cannatella choana, Chthonerpeton, Nectocaecilia, Po- and Hillis, 1993; cf. Latreille, 1800). tomotyphlus, Typhlonectes) and Scolecomor- CHARACTERIZATION AND DIAGNOSIS: Batra- phidae (Crotaphatrema and Scolecomor- chia is a taxon whose living members of the phus) renders Caeciliidae paraphyletic. Al- two component groups (salamanders and though we expect that ongoing work by M. frogs) are so different (and mutually apo- Wilkinson, Nussbaum, and collaborators will morphic) that their synapomorphies are not provide a more re®ned taxonomy, these cur- obviously re¯ected in external appearance. rently recognized taxa can be retained as sub- The annectant members of the taxon are all families (Scolecomorphinae Taylor, 1969, fossil and not well known. For practical pur- and Typhlonectinae Taylor, 1968) with no poses, Batrachia is composed of living am- paraphyly implied as long as the remaining phibians that are not members of Gymno- caeciliids are not placed within a subfamily. phiona. (A Caeciliinae recognized as nomenclatural- Beyond our molecular evidence, the fol- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 169 lowing morphological characters have been spelling, Caudata (see Stejneger, 1907). Not suggested to be synapomorphies of this Caudata Scopoli (1777), as attributed incorrect- group (Trueb and Cloutier, 1991): (1) loss of ly by Stejneger, 1907: 215. (See appendix 6 for a postfrontal bone; (2) loss of the surangular nomenclatural discussion.) bone; (3) loss of splenial bone; (4) loss of IMMEDIATELY MORE INCLUSIVE TAXON: [23] dermal scales; (5) absence of an articulation Batrachia Latreille, 1800. of the anterior ptergygoid ramus with the pal- SISTER TAXON: [74] Anura Fischer von atine; (6) absence of an ectopterygoid; (7) Waldheim, 1831. absence of a stapedial foramen; (8) presence RANGE: Temperate Eurasia, northwestern of a papilla neglecta; (9) presence of a ca- Africa, and North America, and in disjunct rotid labyrinth; (10) choanal tube opens into populations throughout tropical America. the archenteron during development; and CONCEPT AND CONTENT: Caudata is a (11) pronephros modi®ed for trans- monophyletic group composed of all living port. salamanders (cf. Cannatella and Hillis, SYSTEMATIC COMMENTS: Feller and Hedges 1993), the subsidiary taxa being [25] Cryp- (1998) coined the name (for which tobranchoidei Noble, 1931, and [29] Diadec- Homomorpha Fitzinger [1835] is an avail- tosalamandroidei new taxon. able older name) for a clade composed of CHARACTERIZATION AND DIAGNOSIS: Sala- salamanders and caecilians that they believed manders are immediately recognizable be- to be monophyletic. Procera was supported by analysis of 2.7 kb of sequence from four cause they are the only living amphibians to mtDNA genes. We have not attempted to re- have both forelimbs and tails. Their primitive analyze the data of Feller and Hedges (1998), aspect is restricted only to general body plan. but we note that we also used 12S and 16S Salamanders show many osteological losses fragments of the mt rRNA genes and t- and morphological simpli®cations from their RNAValine. They also used sequences from a non-caudatan ancestors. Unlike the other two portion of the tRNALeucine gene, which we did major clades of living amphibians, whole not. Unlike Feller and Hedges (1998), we in- groups of salamanders are known for pae- cluded substantial evidence from nuDNA se- domorphic lineages with varying degrees of quences (see ``Materials''), with the result retention of larval characteristics in the that we have employed almost half again as aquatic adults (e.g., Cryptobranchidae, Sir- much sequence as they did and more than 43 enidae, Proteidae, and various members of times as many terminals. Our results strongly the Ambystomatidae [e.g., Amybystoma du- support the relationship corroborated by merilii] and Plethodontidae [e.g., Eurycea morphological evidence (Trueb and Cloutier, tridentifera]). Most salamanders transfer 1991), which is caecilians ϩ (frogs ϩ sala- sperm via the production of spermatophores, manders). This arrangement, in turn, is con- but like frogs and caecilians, salamanders sistent with the recognition of Batrachia La- primitively have external fertilization with treille (1800) and as intended by Trueb and free-living aquatic larvae. Cloutier (1991). Furthermore, for our data al- Beyond our molecular evidence, Caudata ternative topologies required considerably is diagnosed by the following morphological more steps: (1) frogs ϩ (caecilians ϩ sala- characters, judged to be synapomorphies manders) required 84 additional steps; and (modi®ed from Trueb and Cloutier, 1991; (2) salamanders ϩ (caecilians ϩ frogs) so- Larson and Dimmick, 1993; Larson et al., lution required an additional 85 steps. 2003): (1) incomplete maxillary arcade; (2) presence of a tuberculum interglenoideum; [24] CAUDATA FISCHER VON WALDHEIM, 1813 (3) scapulocoracoid and scapula fused (re- versed in sirenids); (4) no operculum and Caudati Fischer von Waldheim, 1813: 58, an ap- columella detached (modi®ed in some hy- parent latinization and reranking of Caudati A.M.C. DumeÂril, 1806: 95 (which was coined nobiids, plethodontids, salamandrids, and as a family-group taxon and is therefore un- ambystomatids); and (5) male anterior ven- available for above-family-group taxonomy). tral glands present (reversed in sirenids). In Emended here to conform to the traditional addition, Trueb and Cloutier (1991) dis- 170 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 cussed a number of other features that may Salamandropes Fitzinger, 1843: 34. Type genus: be synapomorphic but are highly contingent Salamandrops Wagler, 1830. on cladogram topology. Megalobatrachi Fitzinger, 1843: 34. Type genus: Megalobatrachus Tschudi, 1837. Sieboldiidae Bonaparte, 1850: 1 p. Type genus: [25] CRYPTOBRANCHOIDEI NOBLE, 1931 Sieboldia Gray, 1838. Cryptobranchoidea Noble, 1931: 473. Explicit or- Protonopsidae Gray, 1850a: 52. Type genus: der emended to Cryptobranchoidei by Tamaru- ``Protonopsis Barton, 1824'' (ϭ Protonopsis nov, 1964b: 159. (See appendix 6 for nomen- LeConte, 1824). clatural note.) IMMEDIATELY MORE INCLUSIVE TAXON: [25] IMMEDIATELY MORE INCLUSIVE TAXON: [24] Cryptobranchoidei Noble, 1931. Caudata Fischer von Waldheim, 1813. SISTER TAXON: [26] Hynobiidae Cope, SISTER TAXON: [29] Diadectosalamandro- 1859. idei new taxon. RANGE: Central China; Japan; eastern tem- RANGE: Eastern United States and south- perate North America. eastern Canada in North America; in Eurasia CONTENT: Andrias Tschudi, 1837; Cryp- from Kamchatka west through Siberia to tobranchus Leuckart, 1821. eastern European to Turkmenistan, CHARACTERIZATION AND DIAGNOSIS: Cryp- Afghanistan, and Iran and eastward through tobranchidae is a taxon composed of three central China to Korea and Japan. species of giant, obligately aquatic paedo- CONCEPT AND CONTENT: Cryptobranchoidei morphs. Like other cryptobranchoids, they is a monophyletic taxon composed of [27] lack internal fertilization and share a suite of Cryptobranchidae Fitzinger, 1826, and [26] internal characters primitive for Caudata. Hynobiidae Cope, 1859. Adults lack gills and the lungs are nonfunc- CHARACTERIZATION AND DIAGNOSIS: Cryp- tional, so nearly all respiration is across the tobranchoidei exhibits external fertilization extensively folded and wrinkled skin (Noble, (one genus showing a unique kind of sper- 1931; Bishop, 1943). matophore formation) and other features Beyond the molecular evidence, the fol- primitive for Caudata. Although one group lowing morphological characters have been (Cryptobranchidae) consists of paedomor- suggested to be synapomorphies (Larson and phic giants with distinctive apomorphies Dimmick, 1993; Larson et al., 2003): (1) dor- such as lateral folds of skin, the bulk of spe- soventrally ¯attened bodies; (2) presence of cies (Hynobiidae) are generalized forms that folds of skin forming ¯aps along the lateral are similar in many ways to the ancestral sal- margins of the body; and (3) septomaxilla amander. absent (also in some salamandrids, Am- Beyond the molecular evidence, the fol- phiumidae, and Perennibranchia). lowing morphological characters are likely SYSTEMATIC COMMENT: The monophyly of synapomorphies (Noble, 1931; Larson and Cryptobranchidae was never seriously in Dimmick, 1993; Larson et al., 2003): (1) fu- doubt, but our results (appendix 5) and those sion of the m. pubotibialis and m. pubois- of Larson et al. (2003) demonstrate that chiotibialis; and (2) ribs unicapitate (also in Cryptobranchus is the sister taxon of An- Anura). drias, an arrangement suggested, but not sub- stantiated, by Estes (1981). [27] FAMILY: CRYPTOBRANCHIDAE FITZINGER, 1826 [26] FAMILY: HYNOBIIDAE COPE, 1859 (1856) Cryptobranchoidea Fitzinger, 1826: 42. Type ge- nus: Cryptobranchus Leuckart, 1821. Ellipsoglossidae Hallowell, 1856: 11. Type genus: Menopomatidae Hogg, 1838: 152. Type genus: Ellipsoglossa DumeÂril, Bibron, and DumeÂril, Menopoma Harlan, 1825. 1854. Andriadini Bonaparte, 1839: 131. Type genus: Hynobiidae Cope, 1859: 125. Type genus: Hy- Andrias Tschudi, 1837. nobius Tschudi, 1838. Protonopsina Bonaparte, 1840: 101 (p. 11 of off- Protohynobiinae Fei and Ye, 2000: 64. Type ge- print). Type genus: Protonopsis LeConte, 1824. nus: Protohynobius Fei and Ye, 2000. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 171

IMMEDIATELY MORE INCLUSIVE TAXON: [25] RANGE: Temperate and tropical regions of Cryptobranchoidei Noble, 1931. North America, tropical South America, and SISTER TAXON: [27] Cryptobranchidae Fit- Palearctic Eurasia and North Africa. zinger, 1826. CONCEPT AND CONTENT: Diadectosalaman- RANGE: Japan, Korea, and Kamchatka droidei is a monophyletic group of salaman- west through Siberia and China to eastern ders containing [30] Hydatinosalamandroidei European Russia to Turkmenistan, Afghani- new taxon and [49] Plethosalamandroidei stan, and Iran. new taxon. CONTENT: Boulenger, CHARACTERIZATION AND DIAGNOSIS: Dia- 1878; Hynobius Tschudi, 1838; Onychodac- dectosalamandroids represent the bulk of liv- tylus Tschudi, 1838; Fei, Qu, ing salamander diversity. All are character- and Wu, 1983; Protohynobius Fei and Ye, ized by internal fertilization through the use 2000; Ranodon Kessler, 1866; Salamandrel- of a spermatophore. The exception is Siren- la Dybowski, 1870. idae, which in our analysis appears to have CHARACTERIZATION AND DIAGNOSIS: Hyno- lost this complex reproductive feature (inas- biids are unremarkable salamanders, predom- much as the secretory structures are absent), inantly exhibiting a biphasic life history with although this is optimization-dependent, the external fertilization and females lacking alternative being that Proteidae gained the spermathecae. Lungs are usually developed, characteristic independently of other sala- except in Onychodactylus. mander families that have spermatophore Beyond the molecular evidence (which is production. Morphological diversity is enor- of limited value in testing the monophyly of mous, from the large and obligately aquatic this group; see ``Review of Current Taxon- amphiumas to arboreal web-footed tropical omy'' and ``Results''), the following mor- bolitoglossine plethodontids to various pae- phological characters are likely synapomor- domorphic perennibranch lineages such as in phies (Larson and Dimmick, 1993; Larson et Ambystoma. All families within Diadectosa- al., 2003): (1) ®rst hypobranchial and ®rst lamandroidei primitively show a biphasic life ceratobranchial fused (also in amphiumids); history. However, because of the enormous and (2) vomerine dentition replacement from species diversity of direct-developing pleth- posterior (also in Rhyacotriton and Ambys- odontids, most species within this taxon lack tomatidae). a free-living larval stage. SYSTEMATIC COMMENTS: Monophyly of Beyond the molecular evidence (appendix Hynobiidae requires additional testing, es- 5), the following are likely synapomorphies pecially with respect to Cryptobranchidae. (modi®ed from Larson and Dimmick, 1993; Larson et al. (2003) suggested that Batrachu- Larson et al., 2003): (1) maxilla with a single perus is polyphyletic. Unfortunately, al- center of ossi®cation (maxilla lost in Nectu- though the resultant tree was published, the rus); (2) angular bone absent; (3) spinal underlying data were not, leaving the prob- nerve foramina present in at least some ver- lem unaddressable at this time. The status of tebrae; (4) spermathecae present (lost in Sir- Protohynobiinae also requires phylogenetic enidae); (5) posterior ventral glands present corroboration to determine the placement of (lost in amphiumids and sirenids); (6) Kings- Protohynobius within the remaining hyno- bury's glands present (lost in sirenids); and biids. (7) dorsal pelvic glands present in females (lost in sirenids). [29] DIADECTOSALAMANDROIDEI NEW TAXON [30] HYDATINOSALAMANDROIDEI ETYMOLOGY: Diadectos (Greek: transmit- NEW TAXON ter) ϩ salamandroidei- (Greek: of the form of a salamander). (See appendix 6 for no- ETYMOLOGY: Hydatino- (Greek: of the wa- menclatural note.) ter) ϩ salamandroidei (Greek: of salamander IMMEDIATELY MORE INCLUSIVE TAXON: [24] form), denoting that these salamanders gen- Caudata. erally spend at least part of their in wa- SISTER TAXON: [25] Cryptobranchoidei. ter. 172 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

IMMEDIATELY MORE INCLUSIVE TAXON: [29] synapomorphies (from Larson and Dimmick, Diadectosalamandroidei new taxon. 1993):(1) metamorphosis absent so adults re- SISTER TAXON: [49] Plethosalamandroidei tain numerous paedomorphic characteristics, new taxon. such as large bushy (also in RANGE: Coextensive with Caudata, exclud- various paedomorphic lineages in Ambysto- ing the Americas south of the Mexican Pla- matidae and Plethodontidae); and (2) ypsi- teau. loid cartilage absent (also lacking in Am- CONCEPT AND CONTENT: Hydatinosalaman- phiumidae ϩ Plethodontidae, and the hyno- droidei is a monophyletic group composed of biid Onychodactylus); (3) second ceratobran- [31] Perennibranchia Latreille, 1825, and chial in three or four elements; and (4) [35] Treptobranchia new taxon. maxilla reduced or absent (also reduced in CHARACTERIZATION AND DIAGNOSIS: Hyda- Batrachuperus). tinosalamandroidei is the predominant group SYSTEMATIC COMMENTS: The monophyly of of transforming salamanders, with a few pae- Perennibranchia requires additional testing domorphic lineages in the major families. although the preponderane of our evidence There are no morphological characters of un- supports strongly its recognition. Wiens et al. ambiguous placement (i.e., morphological (2005) did not support the monophyly of synapomorphies) of this clade. Molecular Perennibranchia in their parsimony analysis, synapomorphies are summarized in appendix instead placing Sirenidae as the sister taxon 5. of all other salamanders. Their evidence in- cluded morphological and molecular evi- [31] PERENNIBRANCHIA LATREILLE, 1825 dence (from RAG-1) that we did not have, Perennibranchia Latreille, 1825: 105. (See appen- although our total amount of molecular evi- dix 6 for nomenclatural note.) dence is greater. These authors treated in- ferred gaps as unknown characters, while we IMMEDIATELY MORE INCLUSIVE TAXON: [30] treated inferred gaps as evidence. (As noted Hydatinosalamandroidei new taxon. in ``Methods'', we see gaps as a logical con- SISTER TAXON: [35] Treptobranchia new sequence of indels and like other characters taxon. that are consequences of deductive reason- RANGE: Extreme northeastern Mexico ing, such as morphological reversals, we are north through the eastern United States to inclined to include them as evidence.) A southeastern Canada; Adriatic seaboard as strong test of Perennibranchia will involve far north as Istrian region and as far south as analyzing all of the data of Wiens et al. Montenegro; isolated population in north- (2005) along with our evidence, under a sin- eastern Italy. gle analytical assumption-set (e.g., the same CONCEPT AND CONTENT: Perennibranchia assumption set for alignment and analysis, Latreille, 1825, is a monophyletic group as inclusion as evidence of gaps and morpho- implied by its original content, containing logical reversals, and nonexclusion of mor- [32] Proteidae Gray, 1825, and [33] Sireni- phological characters deemed paedomor- dae Gray, 1825. phic). CHARACTERIZATION AND DIAGNOSIS: Peren- nibranchia is a clade composed of moderate [32] FAMILY: PROTEIDAE GRAY, 1825 to large obligately aquatic paedomorphic species, with permanent bushy external gills, Proteina Gray, 1825: 215. Type genus: Proteus no eyelids, and laterally compressed tails. All Laurenti, 1768. species have well-developed forelimbs and Phanerobranchoidea Fitzinger, 1826: 43. Type ge- one group has lost hindlimbs. Lungs are pre- nus: Phanerobranchus Leuckart, 1821. Necturi Fitzinger, 1843: 35. Type genus: Necturus sent, and although internal fertilization ap- Ra®nesque, 1819. pears to be the plesiomorphic condition in Hypochthonina Bonaparte, 1840: 101 (p. 11 of this group, it may have been lost in sirenids, offprint). Type genus: Hypochthon Merrem, although this is optimization-dependent. 1820. Beyond the molecular evidence, the fol- Necturina Bonaparte, 1845: 6. Type genus: Nec- lowing morphological characters are likely turus Ra®nesque, 1819. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 173

Hylaeobatrachidae Abel, 1919: 329±330. Type lack of eyelids, bushy external gills, and lat- Genus: Hylaeobatrachus Dollo, 1884. (Whether erally compressed tailÐbut also lack pre- this fossil taxon is inside the crown group is maxillary teeth and have keratinized jaw unknown and it is placed here provisionally.) pads (a synapomorphy). Unlike all other sal- Menobranchida Knauer, 1883: 96. Type genus: amanders, sirens lack hind limbs; unlike near Menobranchus Harlan, 1825. relatives they appear to have lost internal fer- IMMEDIATELY MORE INCLUSIVE TAXON: [31] tilization (Noble, 1931). They typically live Perennibranchia Latreille, 1825. in heavily vegetated lakes, ponds, and SISTER TAXON: [33] Sirenidae Gray, 1825. swamps (Bishop, 1943). RANGE: Eastern United States and adjacent Beyond our molecular evidence, the fol- southeastern Canada; Adriatic seaboard as lowing morphological characters have been far north as Istrian region and as far south as suggested to be synapomorphies (Larson et Montenegro; isolated population in north- al., 2003) or are synapomorphies in our to- eastern Italy. pology: (1) hindlimbs lost; (2) scapulocora- CONTENT: Necturus Ra®nesque, 1819; coid and scapula separate elements (a rever- Proteus Laurenti, 1768. sal); (3) teeth absent (present in some fossil CHARACTERIZATION AND DIAGNOSIS: Protei- forms, outside of the crown group), replaced dae is a group of obligately aquatic paedo- by keratinized beaklike pads; (5) all spinal morphic salamanders characterized by hav- nerves exit through foramina except for ®rst ing bushy external gills throughout life, lack- two vertebrae (also in salamandrids); and (7) ing eyelids, having laterally compressed tails. all glands and spermathecae lost that were Unlike their sister taxon, Sirenidae, they ex- associated with spermatophore production. hibit internal fertilization and have hind legs ANATOMICAL COMMENT: Sirenid nasal (Noble, 1931). All of these characteristics are bones have been suggested to be nonhomol- either synapomorphic with their sister taxon ogous with those in spermatophore-produc- or plesiomorphic with respect to Perenni- ing taxa (Salamandroidea sensu Duellman branchia. and Trueb, 1986) because they ossify from Beyond our molecular evidence, the fol- anlagen positioned medially to the dorsal lowing morphological characters are likely process of the premaxillae (laterally to the synapomorphic (modi®ed from Larson and paired premaxillary processes in ``salaman- Dimmick, 1993; Larson et al., 2003): (1) re- droids''; Larson et al., 2003). Our placement cessus amphibiorum with vertical orientation of sirenids within Diadectosalamandroidei (also in Plethodontidae); (2) basilaris com- suggests that the ossi®cation center has plex absent (also in plethodontids and some moved from lateral to medial in sirenids, salamandrids); and (3) maxilla absent. with the nasal bones themselves remaining homologous as nasal bones. [33] FAMILY: SIRENIDAE GRAY, 1825 [35] TREPTOBRANCHIA NEW TAXON Sirenina Gray, 1825: 215. Type genus: Siren Lin- naeus, 1767 (ϭ Siren OÈ sterdam, 1766). ETYMOLOGY: Greek: Treptos (Greek: Sirenes Fitzinger, 1843: 35. Type genus: Siren turned) ϩ branchia (Greek: gill), noting that Linnaeus, 1767 (ϭ Siren OÈ sterdam, 1766). the bulk of the salamanders in this group are IMMEDIATELY MORE INCLUSIVE TAXON: [13] transforming (or a few further derived in Perennibranchia Latreille, 1825. having direct development). SISTER TAXON: [32] Proteidae Gray, 1825. IMMEDIATELY MORE INCLUSIVE TAXON: [30] RANGE: Southeastern United States and ex- Hydatinosalamandroidei new taxon. treme northeastern Mexico. SISTER TAXON: [31] Perennibranchia La- CONTENT: Pseudobranchus Gray, 1825; treille, 1825. Siren OÈ sterdam, 1766. RANGE: British Isles and Scandinavia east- CHARACTERIZATION AND DIAGNOSIS: Sirens ward to the Ural Mountains, southward into are a group of slender, obligately aquatic pae- the Iberian Peninsula and Asia Minor; north- domorphic salamanders that exhibit the stan- central India and China to northern Indochi- dard suite of paedomorphic characteristicsÐ na; extreme northwestern Africa; southern 174 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Canada and southern Alaska south to the nus Bathysiredon being distinguished from southern edge of the Mexican Plateau. Ambystoma on the basis of its cat®sh-like CONCEPT AND CONTENT: Treptobranchia is habitus (Dunn, 1939). a monophyletic group containing Ambysto- Beyond our molecular evidence, the fol- matidae Gray, 1850, and Salamandridae lowing morphological characters have been Goldfuss, 1820. suggested to be synapomorphies: (1) vomer- CHARACTERIZATION AND DIAGNOSIS: Am- ine dentition replacement from posterior bystomatids and salamandrids are commonly (also in Hynobiidae and Rhyacotritonidae; encountered salamanders in North America Larson and Dimmick, 1993; Larson et al., and temperate Eurasia. Their life history is 2003); (2) presence of conspicuous folds in biphasic and they have internal fertilization. cloacal tube in males (also in Rhyacotriton- With the exception of a few paedomorphic idae; Larson and Dimmick, 1993; Larson et lineages, they transform into adults that lack al., 2003); and (3) ring-shaped otoglossal gills and have eyelids. Their body forms run cartilage (also in Rhyacotriton; Cope, 1887; from moderately slender to robust; the limbs Tihen, 1958). are well-developed and robust. SYSTEMATIC COMMENTS: We place Dicamp- No morphological synapomorphies have todon in Amybstomatidae, because doing so been suggested for this taxon, and although renders a more ef®cient taxonomy and be- this group is uniformly characterized by sev- cause the reason for removing Dicamptodon eral of the included morphological charac- originally from Ambystomatidae (that it was ters, none of them optmizes unambiguously thought to be distantly related to Ambystoma to this taxon. Unambiguously optimized mo- [Edwards, 1976]) has now been rejected lecular synapomorphies of Treptobranchia (Larson et al., 2003). are listed in appendix 5. [40] FAMILY: SALAMANDRIDAE [36] FAMILY: AMBYSTOMATIDAE GRAY, 1850 GOLDFUSS, 1820 Ambystomina Gray, 1850a: 32. Type genus: Am- Salamandrae Goldfuss, 1820: 129. Type genus: bystoma Tschudi, 1838. Salamandra Laurenti, 1768. Siredontina Bonaparte, 1850: 1 p. Type genus: Si- Tritonidae Boie, 1828: 363. Type genus: Triton redon Wagler, 1830. Laurenti, 1768. Dicamptodontinae Tihen, 1958: 3. Type genus: Pleurodeles Tschudi, 1838: 91. Type genus: Pleu- Dicamptodon Strauch, 1870. New synonymy. rodeles Michahelles, 1830. Salamandrinae Fitzinger, 1843: 33. Type genus: IMMEDIATELY MORE INCLUSIVE TAXON: [35] Fitzinger, 1826. Treptobranchia new taxon. Molgidae Gray, 1850a: 14. Type genus: Molge SISTER TAXON: [40] Salamandridae - Merrem, 1820. fuss, 1820. Seiranotina Gray, 1850a: 29. Type genus: Seir- RANGE: Alaska and southern Canada south anota Barnes, 1826. to the southern edge of the Mexican Plateau. Bradybatina Bonaparte, 1850: 1 p. Type genus: Bradybates Tschudi, 1838. CONTENT: Ambystoma Tschudi, 1838; Di- Geotritonidae Bonaparte, 1850: 1 p. Type genus: camptodon Strauch, 1870. Geotriton Bonaparte, 1832 (ϭ Triturus Ra®n- CHARACTERIZATION AND DIAGNOSIS: Am- esque, 1815). bystomatids are thick-bodied salamanders with well-developed limbs. They inhabit a IMMEDIATELY MORE INCLUSIVE TAXON: [35] wide variety of habitats from semidesert Treptobranchia new taxon. grassland to boreal conifer and decid- SISTER TAXON: [36] Ambystomatidae Gray, uous forest, generally returning to water only 1850. for . Nevertheless, the most fa- RANGE: British Isles and Scandinavia east- mous paedomorphic lineage, Ambystoma ward to the Ural Mountains, southward into mexicanum () of central Mexico, is in the Iberian Peninsula and Asia Minor; north- this family. Some of the paedomorphic lake- central India and China to northern Indochi- form species have assumed extreme and na; extreme northwestern Africa; northeast- large forms, with the formerly recognized ge- ern and extreme northwestern Mexico 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 175 through western and eastern United States the ``true'' salamanders (Chioglossa, Lycias- north to Alaska and southeastern Canada. alamandra, Mertensiella, and Salamandra) CONTENT: Chioglossa Bocage, 1864; Cy- and Pleurodelinae Tschudi, 1838 (for nops Tschudi, 1838; Nussbaum ``'', the remaining genera). Salamandra and Brodie, 1982; Euproctus GeneÂ, 1838 is our sole exemplar of Salamandrinae and (see Systematic Comments); Lissotriton Bell, likely some of the molecular characters for 1838; Lyciasalamandra Veith and Steinfartz, this genus (appendix 5) are synapomorphies 2004; Mertensiella Wolterstorff, 1925; Me- of the subfamily. Branch 41 in appendix 5 is sotriton Bolkay, 1927; Neurergus Cope, equivalent to Pleurodelinae as we hypothe- 1862; Notophthalmus Ra®nesque, 1820; size it. Pachytriton Boulenger, 1878; Paramesotri- GarcõÂa-ParõÂs et al. (2004a: 602) suggested ton Chang, 1935; Pleurodeles Michahelles, in brief comment that the date of publication 1830; Salamandra Laurenti, 1768; Salaman- of Euproctus Gene is not 1838, but 1839, drina Fitzinger, 1826; Gray, 1850; rendering it a junior synonym of Megapterna Triturus Ra®nesque, 1815 (see Systematic Savi, 1838. They also suggested that ongoing Comments); Tylototriton Anderson, 1871. molecular work will show Euproctus to be CHARACTERIZATION AND DIAGNOSIS: The paraphyletic and render Euproctus asper as salamandrid body plans range from moder- asper (DugeÁs, 1852) as well as ately slender to robust with four well-devel- show that Triturus vittatus should not be in- oped limbs. Most species are periodically cluded within Triturus, the oldest available (e.g., Taricha, Notophthalmus) or completely name for this monotypic taxon being Om- (e.g., Cynops, Pleurodeles, and Pachytriton) matotriton Gray, 1850. Pending publication aquatic and typically have biphasic life his- of the relevant evidence we retain the status tories, except for Mertensiella, which has di- quo. rect-development from terrestrial eggs, and some populations of Salamandra that have live birth. Notophthalmus exhibits three dis- [49] PLETHOSALAMANDROIDEI NEW TAXON tinct life-history stages, an aquatic larva, ter- ETYMOLOGY: Pletho- (Greek: great num- restrial subadult (eft), and aquatic adult ber) ϩ salamandrodei (Greek: of salamander (Bishop, 1943). There are a few paedomor- form), to denote the large number of species phic populations of Notophthalmus and Tri- in this taxon, and with passing reference to turus, that, although they retain external gills, do develop eyelids (Duellman and Trueb, the largest contributor to this enormity, 1986; Zug et al., 2001). Plethodontidae. Beyond our molecular evidence, the fol- IMMEDIATELY MORE INCLUSIVE TAXON: [29] lowing morphological characters have been Diadectosalamandroidei new taxon. suggested to be synapomorphies (Larson and SISTER TAXON: [30] Hydatinosalamandro- Dimmick, 1993; Larson et al., 2003): (1) idei new taxon. periotic present (also in RANGE: Temperate and tropical North and plethodontids); (2) periotic cistern small tropical South America; Korea; and Mediter- (also in plethodontids); and (3) vomerine ranean Europe. dentition medially replaced. CONCEPT AND CONTENT: Plethosalamandro- SYSTEMATIC COMMENTS: Our results, al- idei is a monophyletic group containing Rhy- though based on less dense sampling, are acotritonidae Tihen, 1958, and [50] Xenosa- broadly similar to those of Titus and Larson lamandroidei new taxon. (1995; see ``Results''). Various authors (e.g., CHARACTERIZATION AND DIAGNOSIS: Ple- Risch, 1985) have recognized subfamilies, thosalamandroidei contains the vast majority although none so far suggested has been con- of species of salamanders, being dominated sistent with the phylogeny of the group. Cur- by the very large family Plethodontidae. rent understanding of relationships among Morphological and life-history variation is salamandrids (e.g., Larson et al., 2003) is extensive, from the obligately aquatic am- consistent with the recognition of two sub- phiumas to the arboreal species of Bolito- families: Salamandrinae Goldfuss, 1820, for glossa. Although primitively exhibiting a bi- 176 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

phasic life history, the bulk of the plethosa- IMMEDIATELY MORE INCLUSIVE TAXON: [49] lamandroids are direct-developers. Plethosalamandroidei new taxon. No unambiguous evidence for this taxon SISTER TAXON: Rhyacotritonidae Tihen, extends from morphology, and only molec- 1958. ular evidence documents the existence of this RANGE: Extreme southern Alaska and clade, summarized in appendix 5. Nova Scotia (Canada) south to Amazonian Brazil and central ; southern Europe FAMILY: RHYACOTRITONIDAE TIHEN, 1958 and the Korean Peninsula. Rhyacotritoninae Tihen, 1958: 3. Type genus: CONCEPT AND CONTENT: Xenosalamandro- Rhyacotriton Dunn, 1920. idei is a monophyletic group containing Am- phiumidae Gray, 1825, and [51] Plethodon- IMMEDIATELY MORE INCLUSIVE TAXON: [49] tidae Gray, 1850. Plethosalamandroidei. CHARACTERIZATION AND DIAGNOSIS: Xenos- SISTER TAXON: [50] Xenosalamandroidei alamandroids share no externally obvious new taxon. synapomorphies and have widely divergent RANGE: Extreme northwestern United life histories and morphologies (e.g., troglob- States. itic paedomorphs; large -like obligately CONTENT: Rhyacotriton Dunn, 1920. aquatic predators; burrowers; and arboreal CHARACTERIZATION AND DIAGNOSIS: Ani- salamanders). The two nominal families are mals in this taxon are relatively small, semi- also dissimilar in most aspects of their biol- aquatic transforming salamanders of stout ogy. body and limbs, resembling the ambysto- Beyond our molecular evidence (appendix matids in general aspect, a group with which 5), the following characters have been sug- they were once considered to be allied. Rhy- gested to be synapomorphies (Larson and acotritonids exhibit a biphasic life history Dimmick, 1993): (1) maxillae fused (also in and have internal fertilization. Notophthalmus and some Hynobius); and (2) Beyond our molecular evidence, the fol- ypsiloid cartilage absent (also absent in sir- lowing characters have been suggested to be enids and Onychodactylus). synapomorphies (modi®ed from Larson and Dimmick, 1993; Larson et al., 2003): (1) vo- FAMILY: AMPHIUMIDAE GRAY, 1825 merine dentition replacement from posterior (also in Hynobiidae and Ambystomatidae); Amphiumidae Gray, 1825: 216. Type genus: Am- (2) conspicuous folds present in the male clo- phiuma Garden, 1821. acal tube (also in Ambystomatidae); (3) male IMMEDIATELY MORE INCLUSIVE TAXON: [50] vent gland extremely enlarged and secretes Xenosalamandroidei new taxon. through pores lateral to the cloacal ori®ce SISTER TAXON: [51] Plethodontidae Gray, rather than into the cloacal ori®ce as in other 1850. spermatophore-producing groups; and (4) no RANGE: Southeastern United States. dorsal ossi®cations of the maxilla. In addi- CONTENT: Amphiuma Garden, 1821. tion, the otoglossal cartilage is ring-shaped CHARACTERIZATION AND DIAGNOSIS: Am- as in Ambystomatidae (Dunn, 1920; Tihen, phiumas are large, obligately aquatic sala- 1958) manders with cylindrical bodies up to 1.16 meters in length, tiny legs, and unpleasant [50] XENOSALAMANDROIDEI NEW TAXON dispositions. Transformation is partial, the ETYMOLOGY: Xenos (Greek: strange) ϩ gills being lost but eyelids never developing. salamandroidei (Greek: of salamander form), Beyond our molecular evidence, the fol- to denote the fact that some of the more ex- lowing morphological characters have been otic salamanders (e.g., Nyctanolis, Thorius, suggested to be synapomorphies (modi®ed and Amphiuma) are in this clade and that from Larson and Dimmick, 1993): (1) sep- some of the stranger biogeographical distri- tomaxilla absent (also absent in some sala- butions of on the planet are attri- mandrids, Sirenidae, and Cryptobranchidae); buted to members of this group (e.g., Hydro- (3) ®rst hypobranchial and ®rst ceratobran- mantes ϩ Speleomantes). chial fused (also in hynobiids); (4) second 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 177 ceratobranchial in four elements; and (5) idactylium Tschudi, 1838; Hydromantes posterior ventral glands absent. Beyond this, Gistel, 1848; Karsenia Min, Yang, Bonett, their elongate body and tiny limbs are clearly Vieites, Brandon, and Wake, 2005; Nototri- synapomorphies. ton Wake and Elias, 1983; Nyctanolis Elias SYSTEMATIC COMMENT: Because Amphiuma and Wake, 1983; Oedipina Keferstein, 1868; is clearly the sister taxon of Plethodontidae, Parvimolge Taylor, 1944; see Systematic under normal circumstances it would be de- Comment; Phaeognathus Highton, 1961; sirable to place them in a single family to Plethodon Tschudi, 1838; Pseudoeurycea avoid having a monotypic Amphiumidae. Taylor, 1944 (including Ixalotriton Wake But, because Amphiumidae and Plethodon- and Johnson, 1989, and Lineatriton Tanner, tidae have never been considered to consti- 1950; see Systematic Comments and new tute one nominal family and there is no par- combinations in appendix 7); Pseudotriton aphyly to be eliminated, little is to be gained Taylor, 1944; Speleomantes Dubois, 1984; by a nomenclatural change, so we stay with Stereochilus Cope, 1869; Thorius Cope, traditional usage. 1869. CHARACTERIZATION AND DIAGNOSIS: Pleth- [51] FAMILY: PLETHODONTIDAE GRAY, 1850 odontids demonstrate a spectacular radiation Plethodontidae Gray, 1850a: 31. Type genus: in the Americas, with representatives also Plethodon Tschudi, 1838. found in Mediterranean Europe and one spe- Desmognathina Gray, 1850a: 40. Type genus: cies on the Korean Peninsula. Plethodontids Desmognathus Baird, 1850. are all lungless and uniquely exhibit distinc- Oedipina Gray, 1850a: 42. Type genus: Oedipus tive nasolabial grooves in transformed adults. Tschudi, 1838. Most species show direct development, Ensatinina Gray, 1850a: 48. Type genus: Ensatina which has arisen within the clade several Gray, 1850. times (Chippindale et al., 2004). A few lin- Bolitoglossidae Hallowell, 1856: 11. Type Genus: Bolitoglossa DumeÂril, Bibron, and DumeÂril, eages are perennibranch paedomorphs, but 1854. they are all contained within genera that oth- Hemidactylidae Hallowell, 1856: 11. Type Genus: erwise are composed of salamanders with Hemidactylium Tschudi, 1838. terrestrial adults. Spelerpinae Cope, 1859: 123. Type Genus: Spe- Beyond our molecular evidence, the fol- lerpes Ra®nesque, 1832. lowing morphological characters are likely Thoriidae Cope, 1869: 110. Type Genus: Thorius synapomorphies (modi®ed from Larson and Cope, 1869. Dimmick, 1993; Larson et al., 2003): (1) loss Typhlomolgidae Stejneger and Barbour, 1917: 2. of stylus from opercular apparatus; (2) peri- Type Genus: Typhlomolge Stejneger, 1896. otic connective tissue present (also in sala- IMMEDIATELY MORE INCLUSIVE TAXON: [50] mandrids); (3) periotic cistern small (also in Xenosalamandroidei new taxon. salamandrids); (4) basilaris complex absent SISTER TAXON: Amphiumidae Gray, 1825. (also absent in proteids and some salaman- RANGE: Extreme southeastern Alaska and drids); (5) recessus amphibiorum with verti- Nova Scotia (Canada) south to eastern Brazil cal orientation (also in Proteidae); (6) palatal and central Bolivia; Mediterreanean Europe; dentition replacement both laterally and pos- southwestern Korea. teriorly; and (7) loss of lungs. CONTENT: Aneides Baird, 1851; Batracho- SYSTEMATIC COMMENTS: Chippindale et al. seps Bonaparte, 1839; Bolitoglossa DumeÂril, (2004) suggested on the basis of their study Bibron, and DumeÂril, 1854; of DNA and morphology that two major Wake and Elias, 1983; Tay- groups could be discerned within Plethodon- lor, 1944; Cryptotriton GarcõÂa-ParõÂs and tidae: (1) Plethodontinae, including former Wake, 2000; Wake and Elias, Desmognathinae and Plethodontini (Aneides, 1983; Desmognathus Baird, 1850; Ensatina Desmognathus, Ensatina, Plethodon, and Gray, 1850; Eurycea Ra®nesque, 1822 (in- Phaeognathus); and (2) an unnamed taxon cluding Haideotriton Carr, 1939; see System- composed of (a) Hemidactyliinae (Hemidac- atic Comments and new combination in ap- tylium); (b) Spelerpinae (Eurycea, Gyrino- pendix 7); Gyrinophilus Cope, 1869; Hem- philus, Stereochilus, and Pseudotriton); and 178 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(c) Bolitoglossinae (Batrachoseps, Bolito- which would require a new name). Inasmuch glossa, Nyctanolis, and Pseudoeurycea). as a partition of Pseudoeurycea does not ap- Chippindale et al. (2004) assumed the fol- pear to be forthcoming in the near future, we lowing (which they had not included in their prefer to recognize a monophyletic Pseu- analysis) to be in Bolitoglossinae: Chiropter- doeurycea, which requires the synonymy of otriton, Cryptotriton, Dendrotriton, Hydro- Lineatriton and Ixalotriton. (See appendix 7 mantes, Nototriton, Oedipina, Speleomantes, for name changes caused by these generic and Thorius). However, our results and those changes.) Although our results suggest that of Mueller et al. (2004) and Macey (2005) Ixalotriton and Parvimolge are outside of suggest that Hydromantes and Speleomantes Pseudoeurycea, in the ®rst case this conclu- are not bolitoglossines, but fall inside Pleth- sion is likely an illusion due to sparse taxon odontinae. Beyond this, the placement of sampling. In the case of Parvimolge, our data Hemidactylium is problematic. Chippindale place it outside of this clade and as the sister et al. (2004) on the basis of mtDNA and taxon to Bolitoglossa, a taxon not in Parra- nuDNA and morphology, placed it as the sis- Olea's (2002) analysis. We therefore retain ter taxon of Bolitoglossinae; Mueller et al. Parvimolge and regard the clade subtended (2004), on the basis of a Bayesian analysis by our branch 72 to be Pseudoeurycea.A of mtDNA, placed it as the sister taxon of densely sampled study including all bolito- Batrachoseps; Macey (2005), on the basis of glossine taxa (especially Bolitoglossa), and a parsimony analysis of mtDNA, placed it as all available evidence, should be the next the sister taxon of all other plethodontids; step. and we place it as imbedded in a group com- We have been unable to discern any char- posed of the traditional Bolitoglossinae and acters (see D.B. Wake, 1966) other than Hemidactyliinae. But, our placement of sev- those related to paedomorphy (such as those eral of the terminals in this group (notably that formerly distinguished Typhlomolge and Batrachoseps and Hemidactylium) is based Typhlotriton from Eurycea) to distinguish the solely on a fraction of the mtDNA of Macey monotypic Haideotriton Carr, 1939, from (2005) and barring differences due to align- Eurycea Ra®nesque, 1822. We, like Dubois ment, our placement of these taxa does not (2005), regard the former to be a synonym constitute a strong test of Macey's (2005) of the latter. Bonett and Chippindale (2004) placement of these taxa or, concomitantly, of recently placed Typhlotriton Stejneger, 1892, the taxonomy that he adopted. A strong test, into the synonymy of Eurycea as well. (See of course, would be the analysis, using direct appendix 7 for new combinations produced optimization, of all of the data presented by by these generic changes.) As noted in ``Re- us, Mueller et al. (2004), and by Chippindale sults'', the status of Plethodon is equivocal et al. (2004) to see what the preponderance inasmuch as our evidence suggests its para- of evidence actually is. Regardless, the ear- phyly, but more densely sampled studies lier taxonomy (e.g., D.B. Wake, 1966) has based on more and different assortments of been speci®cally rejected. evidence (Chippindale et al., 2004; Macey, We consider Lineatriton to be a junior syn- 2005) suggest its monophyly. onym of Pseudoeurycea. Parra-Olea (2002) presented DNA sequence evidence for the [74] ANURA FISCHER VON WALDHEIM, 1813 polyphyly of Lineatriton and that both ``Li- Anuri Fischer von Waldheim, 1813: 58. Latini- neatriton'' lineages rendered Pseudoeurycea zation and reranking of Anoures of A.M.C. Du- paraphyletic. She also provided DNA se- meÂril, 1806 (which was coined explicitly as a quence evidence that Parvimolge and Ixalo- family and therefore unavailable for regulated triton extended from within a paraphyletic nomenclature). Emended to Anura by Hogg, Pseudoeurycea. She recommended, but did 1839a: 270. (See appendix 6 for nomenclatural not execute, a partition of Pseudoeurycea to note.) maintain ``Lineatriton'', Parvimolge, and IMMEDIATELY MORE INCLUSIVE TAXON: [23] Ixalotrition, that presumably would require Batrachia Latreille, 1825. the recognition of several new genera to pre- SISTER TAXON: [24] Caudata Fischer von serve the two Lineatriton clades (one of Waldheim, 1813. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 179

RANGE: Worldwide in tropical to cold-tem- phology, several of which appear to be re- perate habitats, excluding Antarctica and lated to the major evolutionary step in anuran most oceanic islands. larvaeÐsuspension feedingÐwhereas others CONCEPT AND CONTENT: Anura Fischer von have no apparent relation to feeding ecology: Waldheim, 1813, is a monophyletic group (1) operculum fused to abdominal wall (Haas containing all living frogs (i.e., [75] Leio- 16.1); (2) m. geniohyoideus origin from cer- pelmatidae Mivart, 1869, and [77] Lalago- atobranchials I/II (Haas 19.1); (3) m. inter- batrachia new taxon). hyoideus posterior absent (Haas 23.0); (4) CHARACTERIZATION AND DIAGNOSIS: Frogs larval jaw depressors originate from palato- are so distinctive among tetrapods that little quadrate (Haas 42.1); (5) ramus maxillaris introduction is required. Among frogs, how- (cranial nerve V2) medial to the muscle (m. ever, there is enormous variation in mor- levator manidbulae longus; Haas 63.1); (6) phology, behavior, reproductive mode, and ramus mandibularis (cranial nerve V3) ante- life-history. Plesiomorphically, frogs exhibit rior (dorsal) to the m. levator mandibulae the textbook amphibian biphasic life history, longus (Haas 64.2); (7) ramus mandibularis with an aquatic tadpole transforming to an (cranial nerve V3) anterior (dorsal) to the ex- air-breathing adult. However, many species ternus group (Haas 65.2); (8) cartilago labi- of frogs exhibit mild to extreme variations alis superior (suprarostral cartilage) present on this theme, with many exhibiting direct (Haas 84.1; (9) two perilymphatic foramina development within the egg capsule (e.g., (Haas 97.1); (10) hypobrachial skeletal parts Eleutherodactylus) and others bearing the de- as planum hypobranchiale (Haas 104.1); (11) veloping young in dermal vacuities on the processus urobranchialis short, not reaching dorsum (Pipa), in vocal sacs (Rhinoderma), beyond the hypobranchial plates (Haas or even in the stomach (Rheobatrachus). 108.1); (12) commisura proximalis present Beyond our molecular data, the following (Haas 109.1); (13) commisura proximalis II morphological characteristics have been sug- present (Haas 110.1); (14) commisura prox- imalis III present (Haas 111.1); (15) cerato- gested to be synapomorphies of this group hyal with diarthrotic articulation present, me- (Trueb and Cloutier, 1991; Ford and Canna- dial part broad (Haas 115.1); (16) cleft be- tella, 1993): (1) loss of ; (2) tween hyal arch and branchial arch I closed loss of prearticular bone; (3) loss of a pala- (Haas 123.0); (17) ligamentum cornuquad- tine (reversed in Acosmanura); (4) reduction ratum present (Haas 125.1); (18) ventral val- of vertebrae to nine or fewer; (5) atlas with vular velum present (Haas 128.1); and (19) a single centrum; (6) ®rst spinal nerve exits branchial food traps present (Haas 134.1). from spinal nerve canal via intervertebral fo- Haas (2003) also suggested that the fol- ramen; (7) fusion of caudal vertebral seg- lowing are nonlarval synapomorphies not ments into a urostyle; (8) hindlimbs signi®- mentioned as such by Ford and Cannatella cantly longer than forelimbs (with excep- (1993): (1) amplexus present (Haas 138.1); tions), including elongation of ankle bones; (2) vertical pupil shape (Haas 143.0); (3) (9) fusions of radius and ulna and tibia and clavicle overlapping scapula anteriorly (Haas ®bula; (6) fusion of hyobranchial elements 145.1); (4) cricoid as a closed ring (Haas into a hyoid plate; (10) presence of kerati- 148.1); and (5) tibiale and ®bulare elongate nous jaw sheaths and keratodonts on larval and fused at ends (Haas 150.1). mouthparts (lost in some lineages); (11) a In addition, the following optimize as syn- single median spiracle in the larva (a char- apomorphies (Trueb and Cloutier, 1991) of acteristic of Type III tadpolesÐthis being Salientia (ϭ Proanura [fossil taxon] ϩ Anu- highly contingent on phylogenetic structure); ra): (1) loss of ; (2) presence of and (12) skin with large subcutaneous lymph a frontoparietal bone; (3) long and slender spaces; (13) two m. protractor lentis attached ilium; and (4) ribs unicapitate (also in Cryp- to lens (based on very narrow taxon sam- tobranchoidei). pling; Saint-Aubain, 1981; Ford and Canna- tella, 1993). [75] FAMILY: LEIOPELMATIDAE MIVART, 1869 In addition, Haas (2003) reported 19 un- Liopelmatina Mivart, 1869: 291. Type genus: Lio- ambiguous synapomorphies from larval mor- pelma GuÈnther, 1869. Emended to Leiopelma- 180 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

tidae by N.G. Stephenson (1951: 18±28); Lio- without a fossa (Haas 94.3); (6) processus pelmatina considered an incorrect original postcondylaris of ceratohyals present (Haas spelling and Leiopelmatidae placed on the Of- 118.1, shared with Alytes and Discoglossus); ®cial List of Family-Group Names in Zoology (7) intracranial endolymphatic system with by Opinion 1071 (Anonymous, 1977: 167). anterior recessus ascendens present (Haas Ascaphidae FejeÂrvaÂry, 1923: 178. Type genus: Ascaphus Stejneger, 1899. 122.1, also present in Alytes, and Acosman- ura); and (8) larval lungs present and func- IMMEDIATELY MORE INCLUSIVE TAXON: [74] tional (Haas 133.1). Anura Fischer von Waldheim, 1813. SYSTEMATIC COMMENTS: Ascaphus and SISTER TAXON: [77] Lalagobatrachia new Leiopelma had long been associated with taxon. each other in Leiopelmatidae (e.g., Noble, RANGE: New Zealand; Paci®c northwest- 1931), but were placed in different families ern United States and adjacent Canada. by Savage (1973) on biogeographic grounds CONTENT: Ascaphus Stejneger, 1899; and by subsequent authors (Ford and Can- Leiopelma Fitzinger, 1861. natella, 1993; Green and Cannatella, 1993) CHARACTERIZATION AND DIAGNOSIS: Leio- on the basis of suggested paraphyly with re- pelmatidae is a group of pervasively ple- spect to all other frogs. We return them to siomorphic frogs, in many aspects of their the same family-group taxon (as had Roe- anatomy, including vertical pupils (likely a lants and Bossuyt, 2005; but against the synapomorphy of frogs), retention of short judgment of D.M. Green) to avoid having ribs in adults, and amphicoelous vertebrae. monotypic families and to recognize that the Nevertheless, they are apomorphic in many only reason for treating Ascaphus and Leio- ways, including highly derived, high gradi- pelma as representing separate familiesÐthat ent-adapted tadpoles in Ascaphus; and nidic- they are not each other's closest living rela- olous endotrophy to direct development in tivesÐhas not survived testing. Leiopelma. Ascaphus is unique among frogs Characters suggested by Ford and Canna- in having an . tella (1993) to unite Leiopelma with all frogs, In addition to our molecular evidence, a excluding Ascaphus, must be considered ei- likely synapomorphy of Leiopelmatidae is ther convergences between Leiopelma and all loss of columella. Presence of an accessory other non-Ascaphus frogs, or characters that coccygeal head of the m. semimembranosus are apomorphies of Anura that have been (Ritland, 1955; observationally equivalent to secondarily lost in Ascaphus: (1) elongate the m. caudalipuboischiotibialis of other au- arms on the sternum; (2) loss of the ascend- thors, although not homologous if so named) ing process of the patalatoquadrate; (3) sphe- may be synapomorphic, but plesiomorphic nethmoid ossifying in the anterior position; retention of the m. caudalopuboischiotibialis (4) root of the facial nerve exits the braincase is also consistent with our cladogram. through the facial foramen, anterior to the Larval characters in our analysis (from auditory capsule, rather than via the anterior Haas, 2003) that optimize on the Ascaphus acoustic foramen into the auditory capsule branch may be characters solely of Ascaphus (Slabbert and Maree, 1945; N.G. Stephenson, or may be synapomorphies of Leiopelmati- 1951); and (5) palatoquadrate articulation dae (although possibly further modi®ed with- with the braincase via a pseudobasal process, in the endotrophy of Leiopelma). These char- rather than a basal process (Pusey, 1943). acters are (1) larval subdermal serous glands present (Haas 2.1); (2) three heads of the m. [77] LALAGOBATRACHIA NEW TAXON subarcualis obliquus originates from cerato- branchialia II, III, and IV (Haas 31.2); (3) ETYMOLOGY: Lalago (Greek: calling) ϩ larval m. levator mandibulae externus inserts batrachos (Greek: frog), in reference to the on soft tissue (Haas 55.2); (4) larval m. le- fact that the frogs of the sister taxon of Lal- vator mandibular internus inserts broadly agobatrachia, Leiopelmatidae, do not call, across the jaw articulation (Haas 59.2); (5) whereas the vast majority of the Lalagoba- distal end of cartilago meckeli broad and ¯at trachia have a wide variety of calls. Although with processus dorsomedialis absent and it may be that vocal behavior is not a syna- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 181 pomorphy of this taxon, it certainly is char- Ford and Cannatella (1993) had also includ- acteristic. ed the Type IV tadpole as a synapomorphy IMMEDIATELY MORE INCLUSIVE TAXON: [74] of Pipanura; however, besides the fact that Anura Fischer von Waldheim, 1831. Orton's larval types are not characters by SISTER TAXON: [75] Leiopelmatidae Mi- themselves but a complex mosaic of multi- vart, 1869. ple, independent character transformations, RANGE: Coextensive with the range of An- the Type I tadpole of Xenoanura is most par- ura, excluding New Zealand. simoniously derived from the Type III tad- CONCEPT AND CONTENT: Lalagobatrachia pole of Costata and Ascaphus, with the Type Fischer von Waldheim, 1813, is a monophy- IV tadpole a synapomorphy of Acosmanura. letic group containing [78] Xenoanura Sav- In addition, Abourachid and Green (1999) age, 1973, and [84] Sokolanura new taxon. noted that members of this taxon swim with CHARACTERIZATION AND DIAGNOSIS: Mem- coordinated thrusts of the hind legs rather bers of Lalagobatrachia are the familiar frogs than alternating sweeps, as in Leiopelmati- of pools and streams, and meadows, dae. We think that this character may well be , and canyons throughout the world. As a unique and unreversed synapomorphy of with Anura, morphological and life-history Lalagobatrachia. diversity is so great that it de®es detailed de- scription. [78] XENOANURA SAVAGE, 1973 Larval characters (Haas, 2003) that opti- Xenoanura Savage, 1973: 352. (See appendix 6 mize to this branch are (1) m. transversus for nomenclatural comment.) ventralis IV absent (Haas 22.1, reversed else- where but also absent in Heleophryne, Hem- IMMEDIATELY MORE INCLUSIVE TAXON: [77] isus, and hyperoliids among the taxa that Lalagobatrachia new taxon. Haas studied); (2) single m. subarcualis ob- SISTER TAXON: [84] Sokolanura new tax- liquus originates from ceratobranchial II on. (Haas 31.0); (3) insertion of the m. rectus RANGE: Tropical Africa and South Amer- cervicis at the processus branchiales II or III ica, north to southern North America. (Haas 39.1); (4) m. hyoangularis present CONCEPT AND CONTENT: Xenoanura Sav- (Haas 43.1); (5) m. levator mandibulae in- age, 1973, is a monophyletic crown taxon ternus anterior (Haas 58.1); (6) m. levator containing [79] Pipidae Gray, 1825, and mandibulae longus originates from posterior Rhinophrynidae GuÈnther, 1859 ``1858'' (and palatoquadrate (Haas 60.1); and (7) and pal- presumably a number of fossil taxa internal atoquadrate connection to trabecula cranii to this clade, including palaeobatrachids; rostral (Haas 69.1). Savage, 1973; cf. Ford and Cannatella, The synapomorphies associated with Dis- 1993). coglossanura of Ford and Cannatella (1993) CHARACTERIZATION AND DIAGNOSIS: The optimize on this branch as well (with reversal highly aquatic bizarre pipids and equally in Bombinatoridae): (1) bicondylar sacrococ- strange, but terrestrial, Rhinophrynus share cygeal articulation; and (2) episternum pre- the distinctive Type I tadpole (Orton, 1953; sent. Starrett, 1973). Several characters suggested by Ford and Several characters of larval morphology Cannatella (1993) as synapomorphies of their (originally from Haas, 2003) used in our Pipanura would optimize on our tree alter- analysis optimize on this branch: (1) eye po- natively as synapomorphies of Lalagobatra- sition lateral (Haas 11.1); (2) opercular canal chia and reversed in Costata (Alytidae ϩ absent and spiracle paired (Haas 17.0); (3) Bombinatoridae), or independently derived m. constrictor branchialis I absent (Haas in Xenoanura (Pipidae ϩ Rhinophrynidae) 27.0); (4) m. levator mandibulae internus an- and Acosmanura (Anomocoela ϩ Neobatra- terior (Haas 58.2); (5) m. levator mandibulae chia). These characters are (1) torsion of car- longus originates exclusively from the arcus pal elements; (2) absence of free ribs in suborcularis (Haas 60.2); (6) posterolateral adults; (3) presence of vocal sacs; and (4) projections of the crista parotica are expan- fusion of the trigeminal and facial ganglia. sive ¯at chondri®cations (Haas 67.2); (7) ar- 182 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 cus subocularis with a distinct processus la- Xenopoda Fitzinger, 1843: 33. Type genus: Xen- teralis posterior projecting laterally from the opus Wagler, 1827. posterior palatoquadrate (Haas 81.3); (8) ar- Hymenochiridae Bolkay, 1919: 348. Type genus: ticulation of cartilago labialis superior with Hymenochirus Boulenger, 1896. cornu trabeculae fused into rostral plate Siluraninae Cannatella and Trueb, 1988: 32. Type genus: Silurana Gray, 1864. (Haas 85.2); and (9) forearm erupts out of limb pouch outside peribranchial space (Haas IMMEDIATELY MORE INCLUSIVE TAXON: [78] 132.0). Xenoanura Savage, 1973. Characters suggested by Ford and Canna- SISTER TAXON: Rhinophrynidae GuÈnther, tella (1993) in support of their Mesobatrachia 1859 ``1858''. (Xenoanura ϩ Anomocoela) are on our to- RANGE: South American and Panamanian pology required to be convergent in their Pi- tropics; sub-Saharan Africa. poidea (our Xenoanura) and their Pelobato- CONTENT: Hymenochirus Boulenger, idea (our Anomocoela), and they are there- 1896; Pipa Laurenti, 1768; Pseudhymeno- fore independent apomorphies for each chirus Chabanaud, 1920; Silurana Gray, group: (1) closure of the frontoparietal fon- 1864; Xenopus Wagler, 1827. tanelle by juxtaposition of the frontoparietal CHARACTERIZATION AND DIAGNOSIS: Pipids bones; (2) partial closure of the hyoglossal are highly aquatic frogs that have inguinal sinus by the ceratohyals; (3) absence of the amplexus and that vocalize using the hyoid taenia tecti medialis; and (4) absence of the apparatus to make clicks (Rabb, 1960), a taenia tecti transversum (Sokol, 1981). characteristic that is likely synapomorphic. Characters that Ford and Cannatella Pipids share with Rhinophrynidae the dis- (1993) listed as apomorphies of their Pipo- tinctive Type I tadpole (Orton, 1953, 1957; idea also optimize on this branch: (1) lack of Starrett, 1973) but are highly apomorphic mentomeckelian bones; (2) absence of lateral with respect to that group. Morphological alae of the parasphenoid; (3) fusion of the characters (from Haas, 2003) addressed in frontoparietals into an azygous element; (4) our analysis provided a large number of lar- greatly enlarged otic capsule; (5) tadpole val and adult synapomorphies: (1) interbran- with paired spiracles and lacking keratinized chial septum invaded by lateral ®bers of the jaw sheaths and keratodonts (Type I tadpole). m. subarcualis rectus I±IV (Haas 29.2); (2) J.D. Lynch (1973: 169) reported Rhino- anterior insertion of the m. subarcualis rectus phrynidae to have opisthocoelous vertebrae, II±IV on ceratobranchial III (Haas 37.2); (3) in which case opisthocoely may be a syna- commissurae craniobranchiales present pomorphy of Xenoanura (and independently (Haas 75.1); (4) one perilymphatic foramen of Costata), or alternatively opisthocoely on the otic capsule (Haas 97.0); (5) ventral may be a character of Lalagobatrachia and centra formation perichordal (Haas 99.1; but subsequently modi®ed at the level of Acos- see Systematic Comment under Xenoanura); manura. (6) free basihyal present (Haas 105.0); (7) Xenoanura in our sense is coextensive processus urobranchialis absent (Haas with the Recent content of the redundant 108.0); (8) ventral valvular velum absent ranks Pipoidia Gray, 1825 (epifamily) and (Haas 128.0); (9) advertisement call without Pipoidea Gray, 1825 (superfamily) of Dubois air¯ow (Haas 140.3); (10) pupil shape round (2005). (Haas 143.3); (11) with epi- coracoids abutting and functionally ®xed [79] FAMILY: PIPIDAE GRAY, 1825 (Haas 144.2); (12) tongue absent (Haas 149.0). Piprina Gray, 1825: 214. Type genus: ``Pipra Ford and Canntella (1993) provided 11 Laurent'' (ϭ Pipa Laurenti, 1768). Incorrect original spelling. characters in support of the monophyly of Dactylethridae Hogg, 1838: 152. Type genus: this group, although we are not sure of the Dactylethra Cuvier, 1829. character optimization of all of them because Astrodactylidae Hogg, 1838: 152. Type genus: these authors did not provide a character ma- Astrodactylus Hogg, 1838 (ϭ Asterodactylus trix and our different placement of this taxon Wagler, 1827). within Anura may have resulted in some 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 183

reoptimization: (1) lack of a quadratojugal; CONTENT: Rhinophrynus DumeÂril and Bi- (2) presence of an epipubic cartilage; (3) un- bron, 1841. paired epipubic muscle; (4) free ribs in lar- CHARACTERIZATION AND DIAGNOSIS: Rhino- vae; (5) fused articulation between coccyx phrynidae contains a single species, Rhino- and sacrum; (6) short stocky scapula; (7) phrynus dorsalis, which is of medium-size, elongate septomaxillary bones; (8) ossi®ed with a cone-shaped head and globular body, pubis; (9) a single median palatal opening of re¯ecting its burrowing life history. Like the the eustachian tube; (10) lateral line organs pipids, it has inguinal amplexus and a Type persisting in adults; and (11) loss of tongue. I tadpole (Orton, 1953; J.D. Lynch, 1973). BaÂez and Trueb (1997) added to this list Several larval characters in our analysis (fossil taxa pruned for purposes of this dis- optimize as synapomorphies of this group: cussion): (1) the possession of an optic fo- (1) m. geniohyoideus lost (Haas 19.5); (2) m. ramen with a complete bony margin formed levator mandibulae externus present in two by the sphenethmoid; (2) anterior ramus of portions (profundus and super®cialis; Haas the pterygoid arises near the anteromedial 54.1); (3) ramus mandibularis (of cranial corner of the otic capsule; (3) parasphenoid nerve V3; Haas 65.0); (4) larval ribs absent, fused at least partially with the overlying the feature convergent with the condition in braincase; (4) vomer lacks an anterior pro- Lalagobatrachia (Haas 102.0); (5) processus cess, if the bone is present; (5) mandible urobranchialis reaching far beyond hyobran- bears a broad-based, bladelike coronoid pro- chial plates (Haas 108.2); (6) endolymphatic cess along its posteromedial margin; (6) ster- spaces extend into more than half of verte- nal end of the coracoid not widely expanded; bral canal (presacral vertebrae IV or beyond; (7) anterior ramus of pterygoids dorsal with Haas 121.1); (7) branchial food traps divided respect to the maxilla; and (8) premaxillary and crescentic (Haas 135.1); and (8) cartilag- alary processes expanded dorsolaterally. inous cricoid ring with a dorsal gap (Haas Finally, Burton (1998a) suggested that the 148.3). In addition, Rhinophrynus has lost dorsal origin of the mm. ¯exores teretes III ribs in the adults. and IV relative to the corresponding mm. Ford and Cannatella (1993, following transversi metacarpum I and II is a synapo- Henrici, 1991) suggested the following as morphy. synapomorphies of the group: (1) division of SYSTEMATIC COMMENT: Our data do not the distal condyle of the femur into lateral support the recognition of sister-subfamilies and medial condyles; (2) modi®cation of the Pipinae GuÈnther, 1859 ``1858'' (Hymenochi- prehallux and distal phalanx of the ®rst digit rus, Pseudhymenochirus, and Pipa) and Dac- into a spade for digging; (3) tibiale and ®- tylethrinae Hogg, 1839 (Silurana and Xeno- bulare short and stocky, with distal ends pus), as found by de Sa and Hillis (1990) and fused; (4) an elongate atlantal neural arch; BaÂez and Pugener (2003). Instead, our data and (5) sternum absent. Although these char- indicate that Hymenochirus (a member of acters are not available in matrix form, pre- nominal Pipinae) is the sister taxon of the cluding careful evaluation of level of univer- remainder of Pipinae ϩ Dactylethrinae. sality, we have no reason to doubt these sug- gestions. FAMILY: RHINOPHRYNIDAE GUÈ NTHER, 1859 ``1858'' [84] SOKOLANURA NEW TAXON Rhinophrynina GuÈnther, 1859 ``1858'': xiv. Type ϩ genus: Rhinophrynus DumeÂril and Bibron, ETYMOLOGY: Sokol (Otto Sokol) 1841. (Greek: tailless, i.e., frog). We commemorate Otto Sokol with this name. Dr. Sokol was an IMMEDIATELY MORE INCLUSIVE TAXON: [78] anatomist of great talent who would have Xenoanura Savage, 1971. continued to make important contributions to SISTER TAXON: [79] Pipidae GuÈnther, 1859 comparative larval anatomy had his life not ``1858''. been cut short by a tragic automobile acci- RANGE: Tropical and subtropical lowland dent. (See appendix 6 for nomenclatural North and Central America. note.) 184 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

IMMEDIATELY MORE INCLUSIVE TAXON: [77] on this branch are (1) origin of m. interman- Lalagobatrachia new taxon. dibularis restricted to the medial side of the SISTER TAXON: [78] Xenoanura Savage, cartilago meckeli corpus (Haas 52.1); (2) lar- 1973. val m. levator mandibulae externus in two RANGE: Coextensive with Anura, exclud- parts (profundus and super®cialis; Haas ing New Zealand. 54.1); (3) posterior processes of the pars alar- CONCEPT AND CONTENT: Sokolanura is a is double (Haas 88.0); (4) vertebral central monophyletic taxon composed of [85] Cos- formation epichordal (Haas 99.1); and (5) tata Lataste, 1879, and [91] Acosmanura processus urobranchialis absent (Haas Savage, 1973. 108.0). CHARACTERIZATION AND DIAGNOSIS: Larval Costata is also characterized by opistho- morphological synapomorphies (from Haas, coelous vertebrae, which is found in Xe- 2003) optimized by our analysis on this noanura (J.D. Lynch, 1973), making it either branch are (1) m. mandibulolabialis present a synapomorphy of Lalagobatrachia and sub- (Haas 48.1); upper jaw cartilages powered by sequently modi®ed at the level of Acosman- jaw muscles (Haas 53.1); (2) main part of ura, or nonhomologous synapomorphies. larval m. levator mandibulae externus inserts SYSTEMATIC COMMENTS: We could have on on upper jaw cartilages (Haas 55.1); (3) combined Alytidae and Bombinatoridae as a insertion of the larval m. levator mandibulae single family with two subfamilies, but rather internus is lateral to jaw articulation (Haas than continue an arbitrary rank controversy, 59.1); (4) m. levator mandibulae longus in we retain Alytidae and Bombinatoridae as two portions (profundus and super®cialis; families for the sake of continuity of dis- Haas 61.1); (5) processus muscularis on the course (but see comments by Dubois, 2005). lateral margin of the palatoquadrate present Costata in our sense is identical in Recent (Haas 79.1); and (6) ligamentum mandibu- content to the redundant taxa Bombinatoro- losuprarostrale present (Haas 127.1). idia Gray, 1825 (epifamily), Bombinatoro- idea Gray, 1825 (superfamily, and Bombi- [85] COSTATA LATASTE, 1879 natoridae (family) of Dubois (2005).

Costati Lataste, 1879: 339. Emended to Costata [86] FAMILY: ALYTIDAE FITZINGER, 1843 by Stejneger, 1907: 49. (See appendix 6 for no- menclatural note.) Alytae Fitzinger, 1843: 32. Type genus: Alytes Wagler, 1829. IMMEDIATELY MORE INCLUSIVE TAXON: [84] Colodactyli Tschudi, 1845: 167. Type genus: Col- Sokolanura new taxon. odactylus Tschudi, 1845 (ϭ Discoglossus Otth, SISTER TAXON: [91] Acosmanura Savage, 1837). 1973. Discoglossidae GuÈnther, 1858b: 346. Type genus: RANGE: Western Europe, North Africa, and Discoglossus Otth, 1837. (See appendix 6 for Israel, possibly into Syria; east to eastern nomenclatural comment.) Russia and Turkey, China, Korea, and north- IMMEDIATELY MORE INCLUSIVE TAXON: [85] ern Indochina; Borneo (western Kalimantan, Costata Lataste, 1879. Indonesia), and the Philippines. SISTER TAXON: [88] Bombinatoridae Gray, CONCEPT AND CONTENT: Costata Lataste, 1825. 1879, is a monophyletic group containing RANGE: Western Europe, North Africa, and [86] Alytidae Fitzinger, 1843, and [88] Bom- Israel, possibly into Syria. binatoridae Gray, 1825. CONTENT: Alytes Wagler, 1830; Discoglos- CHARACTERIZATION AND DIAGNOSIS: Mem- sus Otth, 1837. bers of Costata are relatively small, unre- CHARACTERIZATION AND DIAGNOSIS: Alyti- markable frogs in external appearance, which dae represents small frogs that reproduce via exhibit the typically biphasic life history via typical Type III pond-type larvae that post- a Type III larva with postmetamorphs retain- metamorphically retain ribs and are generally ing ribs (Noble, 1931; J.D. Lynch, 1973; Zug found around water, with Alytes being more et al., 2001). Larval characters (Haas, 2003) terrestrial than Discoglossus (Noble, 1931; that optimize unambiguously in our analysis J.D. Lynch, 1973). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 185

Haas (2003) did not consider our Alytidae ic frogs that are generally brightly colored to have synapomorphies, because his shortest ventrally (less so in Barbourula) and exhibit tree placed Alytes as the sister taxon of Dis- a typically biphasic life history (unknown in coglossus ϩ Bombina. Nevertheless, com- Barbourula). Like Alytidae, they have Type bined with our molecular data, the larval III larvae. Postmetamorphs retain ribs (No- characters from Haas study that optimize on ble, 1931; J.D. Lynch, 1973). The only mor- this branch are (1) admandibular cartilage phological synapomorphy from our analysis present (Haas 95.1, also found in Heleophry- (originally from Haas, 2003) that optimizes ne); and (2) processus postcondylaris of cer- on this branch is m. mandibulolabialis su- atohyal present (Haas 118.1). As noted under perior present (Haas 50.1). Lalagobatrachia, characters suggested by The implication of our topology is that the Ford and Cannatella (1993) to be synapo- two characters suggested by Ford and Can- morphies of Discoglossanura are here con- natella (1993) as synapomorphies of Discog- sidered to be synapomorphies of Lalagoba- lossanura (bicondylar sacrococcygeal articu- trachia, with reversal of these in Bombina- lation and episternum present) optimize to toridae: (1) monocondylar sacrococcygeal ar- Lalagobatrachia, with a loss in Bombinato- ticulation; and (2) episternum absent. Ford ridae (Bombina ϩ Barbourula). Bombinato- and Cannatella (1993) also suggested that V- ridae was suggested (Ford and Cannatella, shaped parahyoid bones (convergent in Pe- 1993) to have as synapomorphies (1) an ex- lodytes) and a narrow epipubic cartilage plate panded ¯ange of the quadratojugal; and (2) are synapomorphies of this taxon. presence of enchochondral ossi®cations in SYSTEMATIC COMMENTS: Haas (2003) sug- the hyoid plate. gested that Alytes is the sister taxon of Dis- coglossus ϩ Bombina on the basis of three [91] ACOSMANURA SAVAGE, 1973 characters (epidermal melanocytes irregular Acosmanura Savage, 1973: 354. (See appendix 6 in shape and not forming reticulaton [Haas for nomenclatural comment.) 1.1], inspiratory advertisement call [Haas 140.1]; and pupil shape triangular [Haas IMMEDIATELY MORE INCLUSIVE TAXON: [84] 143.2]) considered to be synapomorphies of Sokolanura new taxon. Discoglossus ϩ Bombina. Nevertheless, SISTER TAXON: [85] Costata Lataste, 1879. placing Alytes as the sister taxon of Discog- RANGE: Coextensive with Anura, exclud- lossus requires only two additional steps in ing New Zealand. his data set. CONCEPT AND CONTENT: Acosmanura Sav- age, 1973, is, as originally conceived, a [88] FAMILY: BOMBINATORIDAE GRAY, 1825 monophyletic group containing [92] Anom- ocoela Nicholls, 1916, and [105] Neobatra- Bombinatorina Gray, 1825: 214. Type genus: chia Reig, 1958. Bombinator Merrem, 1820. Bombitatores Fitzinger, 1843: 32. Type genus: CHARACTERIZATION AND DIAGNOSIS: As no- Bombitator Wagler, 1830. ted by Starrett (1973), Acosmanura is char- Bombininae FejeÂrvaÂry, 1921: 25. Type genus: acterized by a Type IV tadpole, differing Bombina Oken, 1816. from the ancestral Type III tadpole (of Leio- pelmatidae and Costata) in having a sinistral IMMEDIATELY MORE INCLUSIVE TAXON: [85] spiracle in the larvae, although there are oth- Costata Latste, 1879. er character differences (summarized below). SISTER TAXON: [86] Alytidae Fitzinger, Beyond the molecular synapomorphies of 1843. the group, several larval morphological char- RANGE: France and Italy east to western acters in our analysis (from Haas, 2003) op- Russia and Turkey; China, Korea, and north- timized on this branch: (1) labial ridge with ern Indochina; Borneo and the Philippines. a single row of keratodonts (Haas 4.0); (2) CONTENT: Barbourula Taylor and Noble, paired venae caudales laterales long (Haas 1924; Bombina Oken, 1816. 15.1); (3) spiracle position sinistral (Haas CHARACTERIZATION AND DIAGNOSIS: Mem- 18.1, also in Scaphiophryne); (4) m. subar- bers of Bombinatoridae are distinctive aquat- cualis rectus I portion with origin from cer- 186 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 atobranchial II present (Haas 34.1); (5) in- tidae is notochordal/amphicoelous; that Xe- sertion site of the m. subarcualis rectus I on noanura and Costata exhibit opisthocoelous the ventral muscle portion lateral (Haas vertebrae; and that Anomocoela and more 35.1); (6) anterior insertion of m. subarcualis ``basal'' groups within Hyloides have inter- rectus II±IV on ceratobranchial II (Haas vertebral bodies unfused to the centra, at 37.1); (7) m. suspensoriohyoideus present least in subadults. (Sooglossidae most likely (Haas 45.1); (8) m. interhyoideus and m. in- has amphicoelous vertebrae as an apomorphy termandibularis well separated by a gap at that level of universality.) Much work (Haas 47.1); (9) functional larval m. levator needs to be accomplished, but currently it ap- mandibulae lateralis present (Haas 56.1; lost pears that the fusion of intervertebral bodies in Gastrotheca); (10) articulation of cartilago has taken place in Hyloides and Ranoides in- labialis superior with cornua trabeculae by dependently. pars alaris (Haas 85.1); (11) larval ribs ab- sent (Haas 102.0; also in Rhinophrynus); (12) [92] ANOMOCOELA NICHOLLS, 1916 commissura proximalis I absent (Haas 109.0; Anomocoela Nicholls, 1916: 86. 109.1 in microhylids); (13) spicula present and long (Haas 112.1; lost in Ceratophrys ϩ IMMEDIATELY MORE INCLUSIVE TAXON: [91] Lepidobatrachus and independently gained Acosmanura Savage, 1973. in Alytes); (14) anterior processus ascendens SISTER TAXON: [105] Neobatrachia Reig, of intracranial endolymphatic system present 1958. (Haas 122.1; also in Ascaphus, Alytes); (15) RANGE: Southern Canada and United ligamentum cornuquadratum inserting on States south to south-central Mexico; Europe cornu trabeculae (Haas 126.1; reversed in and northwestern Africa; western Asia to Ceratophrys); (16) clavicula in adult not tropical southeastern Asia southeast to the overlapping (Haas 145.2; see also J.D. Greater Sunda Islands. Lynch, 1973: 147); (17) palatine bones pres- CONCEPT AND CONTENT: Anomocoela Nich- ent (Haas 146.1; independently lost in sev- olls, 1916, is here conceived as originally eral groups, including microhylids, and den- formed, a monophyletic group containing drobatids). [96] Pelobatoidea Bonaparte, 1850, and [93] SYSTEMATIC COMMENTS: Presence of a Pelodytoidea Bonaparte, 1850. (neo) as a synapomorphy of CHARACTERIZATION AND DIAGNOSIS: Only Acosmanura could be seen as controversial. one of the morphological characters in our It is not controversial that a palatine is char- analysis optimized on this branch: partes cor- acteristic of Neobatrachia, but its presence in pores medially separate (Haas 87.0). Anomocoela is. Some authors favored the Characters suggested by Ford and Canna- view that the palatine develops in pelobatids tella (1993) in support of their Pelobatoidea (sensu lato) but later fuses to the maxilla (our Anomocoela) are (1) sternum ossi®ed (Zweifel, 1956; Kluge, 1966; Estes, 1970); into a bony style, and (2) pupil vertical (ple- others have asserted that the palatine is fused siomorphic for Anura and possibly here; con- with either the vomer or maxilla (Jurgens, vergent with phyllomedusine and some pe- 1971; RocÏek, 1981 ``1980''). Wiens (1989) lodryadine hylids and Africanura, except suggested that the palatine never forms, at Brevicipitidae and Hyperoliidae). least not in Spea. Hall and Larsen (1998) dis- Characters suggested by Ford and Canna- cussed the issue and provided evidence that tella (1993) in support of their Mesobatrachia palatine centers of ossi®cation do exist in we found to be convergent in their Pipoidea Spea and in other anomocoelans. Without ev- (our Xenoanura) and their Pelobatoidea (our idence that the ``palatine'' center of ossi®- Anomocoela), and therefore independent cation in anomocoelans is anything other apomorphies for each group: (1) closure of than the palatine, Hennig's auxiliary princi- the frontoparietal fontanelle by juxtaposition ple (Hennig, 1966) suggests that we accept of the frontoparietal bones; (2) partial closure it as homologous with the palatine of neo- of the hyoglossal sinus by the ceratohyals; batrachians. (3) absence of the taenia tecti medialis; and J.D. Lynch (1973) noted that Leiopelma- (4) absence of the taenia tecti transversum 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 187

(Sokol, 1981). We have some reservations, ytids are small terrestrial frogs that live in however, because the characters were not moist habitats and have a typically biphasic presented in matrix form so we are not sure life history. A number of morphological of the distribution of any of these characters characters (Haas, 2003) in our analysis op- away from their Mesobatrachia. J.D. Lynch timize on this branch (although some of these (1973: 148) provided a character distribution may actually apply to some subset of Pelod- that suggests a dorsally incomplete cricoid ytes): (1) epidermal melanocytes of irregular ring as a synapomorphy at this level (con- shape not forming reticulation (Haas 1.1, vergent in Rhinophrynus). also in Discoglossus and Bombina); (2) up- SYSTEMATIC COMMENT: The monophyly of per labial papillae continuous (Haas 8.0); (3) this group seems assured and the reason for interbranchial septum IV invaded by ®bers maintaining four families within it, rather of m. subarcualis rectus II±IV (Haas 29.1); than having a single larger Pelobatidae, is (4) m. subarcualis rectus I portion with origin that no clarity is gained by changing the cur- from ceratobranchial I absent (Haas 33.0); rent taxonomy (contra Dubois [2005: 3], who (5) larval m. levator mandibulae externus in aggregated the content as four subfamilies two portions (profundus and super®cialis; within a larger Pelobatidae). Haas 54.1); (6) dorsal connection from pro- cessus muscularis to commissura quadrato- [93] SUPERFAMILY: PELODYTOIDEA orbitalis (Haas 78.2); (7) articulation of car- BONAPARTE, 1850 tilago labialis superior with cornua trabecu- IMMEDIATELY MORE INCLUSIVE TAXON: [92] lae by pars corporis (Haas 85.0); (8) verte- Anomocoela Nicholls, 1916. bral centra formation epichordal (Haas 99.1); SISTER TAXON: [96] Pelobatoidea Bonapar- (9) larval ribs present (Haas 102.1); (10) te, 1850. commissura proximalis II absent (Haas RANGE: Southwestern Europe and the Cau- 110.0); (11) eggs laid in strings (Haas 141.1; casus; southern Canada and United States (also in Pelobates and elsewhere in Acos- south to south-central Mexico. manura); (12) parahyoid ossi®cation present CONTENT: Pelodytidae Bonaparte, 1850, (Haas 147.1); and (13) tibiale and ®bulare and [94] Scaphiopodidae Cope, 1865. elongate and fully fused (Haas 150.2; con- CHARACTERIZATION AND DIAGNOSIS: Only vergent in Neobatrachia). one of the morphological characters in our analysis (originally from Haas, 2003) opti- [94] FAMILY: SCAPHIOPODIDAE COPE, 1865 mizes on this branch: basibranchial long Scaphiopodidae Cope, 1865: 104. Type genus: (Haas 105.0). Nevertheless, the molecular Scaphiopus Holbrook, 1836. data are decisive (appendix 5). The length of the 28S fragment is diagnostic for this taxon, IMMEDIATELY MORE INCLUSIVE TAXON: [93] being 703 bp (appendix 3; as in Leiopelma- Pelodytoidea Bonaparte, 1850. tidae), but differing from that taxon in all of SISTER TAXON: Pelodytidae Bonaparte, the morphological characters of the interven- 1850. ing taxa. RANGE: Southern Canada and United States south to south-central part of the Pla- FAMILY: PELODYTIDAE BONAPARTE, 1850 teau of Mexico. CONTENT: Scaphiopus Holbrook, 1836; Pelodytina Bonaparte, 1850: 1 p. Type genus: Pe- Spea Cope, 1866. lodytes Bonaparte, 1838. CHARACTERIZATION AND DIAGNOSIS: Sca- IMMEDIATELY MORE INCLUSIVE TAXON: [93] phiopodids are -like frogs characterized Pelodytoidea Bonaparte, 1850. by the possession of large metatarsal spades, SISTER TAXON: [94] Scaphiopodidae Cope, as found in Pelobatidae, with which they bur- 1865. row. Their life-history is typically biphasic RANGE: Southwestern Europe and the Cau- with a Type IV tadpole and inguinal amplex- casus. us. Morphological characters in our analysis CONTENT: Pelodytes Bonaparte, 1838. (from Haas, 2003) that optimize on this CHARACTERIZATION AND DIAGNOSIS: Pelod- branch are (1) paired venae caudales laterales 188 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 short (Haas 15.0); (2) m. subarcualis rectus I [97] FAMILY: PELOBATIDAE BONAPARTE, 1850 portion with origin from ceratobranchial III Pelobatidae Bonaparte, 1850: 1 p. Type genus: absent (Haas 35.0); and (3) m. mandibulo- Pelobates Wagler, 1830. labialis superior absent (Haas 50.0). Because Haas' (2003) study included only Spea with- IMMEDIATELY MORE INCLUSIVE TAXON: [92] in Scaphiopodidae, these characters may ac- Pelobatoidea Bonaparte, 1850. tually be synapomorphies of Scaphiopus ϩ SISTER TAXON: [98] Megophryidae Bona- Spea or some subset of Spea. Additional tax- parte, 1850. on sampling is needed to elucidate the ap- RANGE: Europe, western Asia, and north- propriate level of universality of these char- western Africa. acters. CONTENT: Pelobates Wagler, 1830. Other possible synapomorphies at this lev- CHARACTERIZATION AND DIAGNOSIS: Pelo- el are (1) fusion of the sacrum and coccyx batids are toad-like frogs, that have a dis- (although J.D. Lynch, 1973: 141, disagreed tinctive metatarsal spade (as does Scaphio- with this); (2) exostosed frontoparietals; and podidae) and their life history is typically bi- (3) presence of a metatarsal spade supported phasic with inguinal amplexus and a Type IV by a well-ossi®ed prehallux (Ford and Can- tadpole. Morphological characters (from natella, 1993). These appear convergently in Haas, 2003) that optimize on this branch are Pelobatidae, possibly relating to their bur- (1) larval eye positioned dorsolaterally (Haas rowing habits. 11.1); (2) posterolateral projections of the crista parotica present (Haas 67.1); (3) arcus [96] SUPERFAMILY: PELOBATOIDEA subocularis with an irregular margin (Haas BONAPARTE, 1850 81.1); (4) vertebral centra epichordal (Haas 99.1); (5) processus branchialis closed (Haas IMMEDIATELY MORE INCLUSIVE TAXON: [92] 114.1); (6) endolymphatic spaces extend into Anomocoela Nicholls, 1916. more than half of vertebral canal (presacral SISTER TAXON: [93] Pelodytoidea Bona- vertebra IV or beyond; Haas 121.1); (7) eggs parte, 1850. laid in strings (Haas 141.1; convergent else- RANGE: Europe, western Asia, and north- where within Acosmanura). western Africa; Pakistan and western China, Because this diagnosis is based solely on Indochinese peninsula, east to the Philippines Pelobates fuscus, some of these characters and the Greater Sunda Islands. may optimize on some subset of the species CONTENT: [97] Pelobatidae Bonaparte, of Pelobates and not on the Pelobatidae 1850, and [98] Megophryidae Bonaparte, branch. Increased density of sampling is 1850. needed. CHARACTERIZATION AND DIAGNOSIS: Mor- Other possible synapomorphies at this lev- phological characters in our analysis (from el are (1) fusion of the sacrum and coccyx; Haas, 2003) that optimized on this branch are (2) exostosed frontoparietals; and (3) pres- (1) m. interhyoideus posterior present (Haas ence of a metatarsal spade supported by a 23.1); (2) m. diaphragmatopraecordialis pres- well-ossi®ed prehallux (Ford and Cannatella, ent (Haas 25.1); (3) m. constrictor branchial- 1993). These appear convergently in Sca- is I absent (Haas 27.0); (4) m. mandibulola- phiopodidae, possibly relating to their bur- bialis superior present (Haas 50.1); and (5) rowing habits. adrostral cartilage very large and elongate (Haas 90.2). Because this generalization is [98] FAMILY: MEGOPHRYIDAE BONAPARTE, based solely on Pelobates, Megophrys, and 1850 Leptobrachium, taxon sampling needs to be expanded for further elucidation of the dis- Megalophreidina Bonaparte, 1850: 1 p. Type ge- nus: Megalophrys Wagler, 1830 (ϭ Megophrys tribution of these characters. J.D. Lynch Kuhl and Van Hasselt, 1822). (1973) noted that Pelobates and mego- Leptobrachiini Dubois, 1980: 471. Type genus: phryids have a monocondylar sacrococcy- Leptobrachium Tschudi, 1838. geal articulation, which is likely a synapo- Oreolalaxinae Tian and Hu, 1985: 221. Type ge- morphy at this level. nus: Oreolalax Myers and Leviton, 1962. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 189

IMMEDIATELY MORE INCLUSIVE TAXON: [96] and a very large tubercle starting at the base Pelobatoidea Bonaparte, 1850. and extending out and onto the ®rst ®nger SISTER TAXON: [97] Pelobatidae Bonaparte, [Lathrop, 2003]), ``Leptobrachiinae'' is rec- 1850. ognized solely on the basis of plesiomorphies RANGE: Montane Pakistan and western (lacking the umbelliform mouth and the large China, Indochinese peninsula, east to the tubercle extending out on the ®nger), so there Philippines and the Greater Sunda Islands. is little point in recognizing these taxa. CONTENT: Atympanophrys Tian and Hu, Moreover, Delorme and Dubois' (2001; ®g. 1983; Brachytarsophrys Tian and Hu, 1983; 20) own analysis rejects leptobrachiine Leptobrachella Smith, 1925; Leptobrachium monophyly. Beyond rejecting the monophyly Tschudi, 1838; Leptolalax Dubois, 1980; of Leptobrachiinae, Delorme and Dubois' Megophrys Kuhl and Hasselt, 1822; Ophry- (2001; ®g. 20) cladogram suggests that the ophryne Boulenger, 1903; Oreolalax Myers currently recognized nominate subgenus of and Leviton, 1962; Scutiger Theobald, 1868; the genus Scutiger, Scutiger (paraphyletic Vibrissaphora Liu, 1945; Xenophrys GuÈn- with respect to Aelurophryne), must be re- ther, 1864. jected, as must the monotypic subgenus Ae- CHARACTERIZATION AND DIAGNOSIS: Mego- lurolalax of genus Oreolalax that renders the phryids are small to large frogs that are gen- subgenus Oreolalax paraphyletic. erally found hopping in leaf litter along streams, although some species extend up- [105] NEOBATRACHIA REIG, 1958 wards in elevation to 5,100 meters on the Neobatrachia Reig, 1958: 115. (See nomenclatur- southern slopes of the Himalayas (Lathrop, al comment in appendix 6.) 2003). Reproduction is typically biphasic with inguinal amplexus and a Type IV tad- IMMEDIATELY MORE INCLUSIVE TAXON: [91] pole. Although our morphological data for Acosmanura Savage, 1973. this group were restricted to Megophrys SISTER TAXON: [92] Anomocoela Nicholls, montana and , our 1916. preferred tree (®gs. 50, 54) suggests that syn- RANGE: Coextensive with Anura, exclud- apomorphies subtending these two species ing New Zealand. are likely synapomorphies of Megophryidae. CONCEPT AND CONTENT: Neobatrachia Characters of morphology (from Haas, 2003) Reig, 1958, is conceived here as it was orig- that optimize on this branch are (1) m. su- inally intended by Reig (1958), a monophy- barcualis rectus accessorius present (Haas letic group of all frogs excluding his Ar- 32.1); and (2) suspensorium low (Haas 71.2). chaeobatrachia (Leiopelmatidae, Discoglos- In addition, Ford and Cannatella (1993) sidae [sensu lato], Pipidae, Rhinophrynidae, suggested that the following are synapomor- and Pelobatidae [sensu lato]). In other words, phies for Megophryidae: (1) complete or al- it is a monophyletic group composed of our most complete absence of ceratohyals in [106] Heleophrynidae Noble, 1931, and adults; (2) intervertebral cartilages with an [107] Phthanobatrachia new taxon. ossi®ed center; and (3) paddle-shaped CHARACTERIZATION AND DIAGNOSIS: Neo- tongue. batrachia includes approximately 96% of the SYSTEMATIC COMMENTS: The recognition of diversity of frogs, most of which have com- Xenophrys as a genus distinct from Mego- pletely ossi®ed vertebrae. Only one character phrys (e.g., Ohler et al., 2002) appears jus- in our analysis (from Haas, 2003) optimizes ti®ed, inasmuch as Megophrys and Xeno- on this branch: m. sartorius discrete from the phrys do not appear to be each other's closest m. semitendinosus (Haas 153.1). Although relatives, with Megophrys most closely re- many authors (e.g., Ford and Cannatella, lated to Ophryophryne. The subfamilies 1993; Trueb, 1993) have reported the pres- [101] Megophryinae Bonaparte, 1850, and ence of palatine bones as a synapomorphy of Leptobrachiinae Dubois, 1980, were not re- Neobatrachia, it is reasonably clear (Haas, jected by our molecular data (®g. 54). Nev- 2003; see also Acosmanura account) that this ertheless, although Megophryinae has apo- characteristic is a synapomorphy of Acos- morphies (an umbelliform oral disc in larvae manura. Nevertheless, one could argue that 190 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 the developmentally distinct palatine of neo- cularis absent (Haas 79.0); (6) partes cor- batrachians is a synapomorphy of this group, pores forming medial body (Haas 87.2); (7) although polarization between the anomo- adrostral cartilage very large and elongate coelan condition and the neobatrachian con- (Haas 90.2); (8) admandibular cartilage pre- dition has to be made on the basis of consid- sent (Haas 95.1); (9) free basihyal absent erations other than outgroup comparison. (Haas 105.0); and (10) processus branchialis Ford and Cannatella (1993) suggested closed (Haas 114.1). In addition, Ford and these additional characters as synapomor- Cannatella (1993) suggested that the loss of phies of the Neobatrachia: (1) third distal keratinous jaw sheaths in the larvae is a syn- carpal fused to other carpals (convergent in apomorphy of this group. Channing (2003) Pelodytes); (2) accessory head of m. adduc- corrected this, noting that larvae lack kera- tor longus present; and (3) parahyoid ossi®- tinized jaw sheaths, except for Heleophryne cation present. In addition, there are substan- rosei, which retains the lower jaw sheath. tial numbers of molecular synapomorphies Channing also noted that during the repro- (appendix 5) that support recognition of this ductive aquatic period, males develop folds taxon. of loose skin that increase the respiratory COMMENT: Neobatrachia in our sense is surface. Both of these characteristics are like- equivalent to the Recent content of epifamily ly apomorphic. Ranoidia Ra®nesque, 1814, of Dubois (2005). [107] PHTHANOBATRACHIA NEW TAXON

[106] FAMILY: HELEOPHRYNIDAE NOBLE, 1931 ETYMOLOGY: Phthano- (Greek: anticipate, do ®rst) ϩ batrachos (Greek: frog). We pro- Heleophryninae Noble, 1931: 498. Type genus: pose this name to honor Arnold G. Kluge Heleophryne Sclater, 1898. and James S. Farris's contribution to phylo- Heleophrynidae Hoffman, 1935: 2. Type genus: genetics generally and to amphibian system- Heleophryne Sclater, 1898. Coined as new fam- ily apparently in ignorance of Noble, 1931. atics speci®cally, especially with reference to the paper that started modern quantitative IMMEDIATELY MORE INCLUSIVE TAXON: phyleticsÐKluge and Farris, 1969). [105] Neobatrachia Reig, 1958. IMMEDIATELY MORE INCLUSIVE TAXON: SISTER TAXON: [107] Phthanobatrachia [105] Neobatrachia Reig, 1958. new taxon. SISTER TAXON: [106] Heleophrynidae No- RANGE: Mountainous areas of the Cape ble, 1931. and Transvaal regions of South Africa, from RANGE: Coextensive with Anura, exclud- sea level to about 3,000 meters elevation. ing New Zealand. CONTENT: Heleophryne Sclater, 1898. CONCEPT AND CONTENT: Phthanobatrachia CHARACTERIZATION AND DIAGNOSIS: Heleo- is a monophyletic group composed of all phrynidae is composed of moderately small neobatrachians, excluding Heleophrynidae treefrog-like anurans with expanded trian- Noble, 1931. In other words, it is composed gular digit tips, a typically biphasic life his- of our [314] Hyloides new taxon and [108] tory, prolonged larval development, Type IV Ranoides new taxon. larvae, and inguinal amplexus, that live in CHARACTERIZATION AND DIAGNOSIS: Mor- rocky, high-gradient habitats (J.D. Lynch, phological characters (from Haas, 2003) that 1973; Passmore and Carruthers, 1979). Con- optimize on this branch are (1) upper mar- siderable numbers of morphological charac- ginal papillae with a broad diastema (Haas ters (from Haas, 2003) in our analysis opti- 8.1; reversed in several subsidiary lineages); mized on this branch: (1) m. transversus ven- (2) m. interhyoideus posterior present (Haas tralis IV present (Haas 22.1); (2) interbran- 23.1); (3) m. diaphragmatopraecordialis pre- chial septum IV musculature invaded by sent (Haas 25.1); (4) m. constrictor bran- lateral ®bers of m. subarcualis rectus II±IV chialis I absent (Haas 27.0); and (5) secretory (Haas 29.1); (3) m. subarcualis rectus acces- ridges present (Haas 136.1). sorius present (Haas 32.1); (4) processus as- COMMENTS ON CHARACTER DISTRIBUTIONS: cendens thin (Haas 72.1); (5) processus mus- Another likely synapomorphy at this level is 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 191 widely-separated atlantal cotyles (ϭ Type I posed of arciferal (at least plesiomorphically of J.D. Lynch, 1971, 1973), although appar- within any of the groups) phthanobatrachian ently reversed in some taxa (e.g., Limnodyn- frogs. In other words, it is composed of [315] astidae, Bufonidae, part of Cycloramphidae Sooglossidae Noble, 1931, and [318] Noto- [Rhinoderma, Hylorina, Alsodes, Eupsophus, gaeanura new taxon. Proceratophrys, Odontophrynus], and part of CHARACTERIZATION AND DIAGNOSIS: Only Ceratophryidae [Ceratophrys, Lepidobatra- one morphological character in our analysis chus]). optimizes on this branch: m. diaphragmato- Presence of an outer metatarsal tubercle praecordialis meeting m. interhyoideus pos- (J.D. Lynch, 1971, 1973) is coherent on our terior in a smooth arch (Haas 26.0). Procoely tree and also may be a synapomorphy at the may also be a synapomorphy, but with re- level of Phthanobatrachia, because outer versals to an anomocoelous condition in at metatarsal tubercles are never found outside least some taxa (e.g., Myobatrachidae). Nev- of this clade, although clearly this character ertheless, substantial numbers of molecular has been lost and regained several times synapomorphies support this clade (appendix within Phthanobatrachia. Optimization of 5). this character requires more work, but we COMMENT: Hyloides in our sense is not co- suggest that it will provide additional evi- extensive with the Recent content of Hylo- dence of relationship. Our current under- idea of Dubois (1983), which included He- standing is that it is absent in Batrachophryn- leophrynidae; of Dubois (2005), which ex- idae, except for Batrachophrynus; absent in cluded Heleophrynidae and Sooglossidae; or Limnodynastidae, except for Limnodynastes of Hyloidea (sensu stricto) of Biju and Bos- tasmaniensis and Adelotus; and present in suyt (2003) and Darst and Cannatella (2004), Myobatrachidae, except for six species of which excluded Batrachophrynidae (by im- Crinia and Taudactylus, and, presumably, plication), Limnodynastidae, Myobatrachi- Mixophyes and Rheobatrachus. Within Mer- dae, and Sooglossidae, rendering Hyloidea idianura they are present, with the exclusion (sensu stricto) of these authors coextensive of some Hylidae, Centrolenidae, Rhinoderma with our [348] Nobleobatrachia. (Cycloramphidae), and Lepidobatrachus (Ceratophryidae). Interestingly, within Ran- [315] FAMILY: SOOGLOSSIDAE NOBLE, 1931 oides the trends are much less clear and much less well documented, the tubercle be- Sooglossinae Noble, 1931: 494. Type genus: Sooglossus Boulenger, 1906. ing absent in most Arthroleptidae (including Nasikabatrachidae Biju and Bossuyt, 2003: 711. Astylosternidae), most Hyperoliidae, Hemi- Type genus: Nasikabatrachus Biju and Bossuyt, sotidae, and Rhacophoridae, some Microhy- 2003. New synonym. linae, Cophylinae, Phrynomerus, and some Ranidae (J.D. Lynch, 1973). IMMEDIATELY MORE INCLUSIVE TAXON: [314] Hyloides new taxon. [314] HYLOIDES NEW TAXON SISTER TAXON: [318] Notogaeanura new taxon. ETYMOLOGY: Hyloides is, for the most part, RANGE: Granitic islands of the Seychelles Hyloidea of traditional usage, excluding He- and the Western Ghats of South India. leophrynidae (as suggested by Haas, 2003), CONTENT: Nasikabatrachus Biju and Bos- removed from regulated nomenclature, and suyt, 2003; Sooglossus Boulenger, 1906 (in- with the ending changed so as to not imply cluding Nesomantis Boulenger, 1909; see family-group regulation. Systematic Comments and appendix 7). IMMEDIATELY MORE INCLUSIVE TAXON: CHARACTERIZATION AND DIAGNOSIS: Soog- [107] Phthanobatrachia new taxon. lossids are tiny to moderate-size frogs with SISTER TAXON: [108] Ranoides new taxon. weakly expanded digits in Sooglossus and RANGE: Coextensive with Anura, exclud- unexpanded digits in Nasikabatrachus. The ing New Zealand. species of Sooglossus and Nesomantis, that CONCEPT AND CONTENT: Hyloides as con- are known, have inguinal amplexus and have ceived here is the monophyletic group com- either endotrophic larvae or direct develop- 192 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 ment (Nussbaum, 1980; Thibaudeau and Al- evidence supporting this view rests on the tig, 1999), whereas Nasikabatrachus has assumption that a complex call (shared by free-living exotrophic larvae (Dutta et al., Sooglossus sechellensis and Nesomantis tho- 2004). Sooglossus sechellensis is biphasic masseti) is apomorphic (Nussbaum et al., with presumably endotrophic larvae; Neso- 1982), as well as on the basis of allozymic mantis life history is unknown, but presum- distance measures (Green et al., 1988). Nev- ably has direct development as no larvae ertheless, there has never been any evidence have ever been found; and Sooglossus gar- suggested to support the monophyly of the dineri has direct development (Nussbaum, three species of Sooglossus with respect to 1980). Nesomantis, so the current taxonomy sug- This taxon was not studied by Haas gests a level of knowledge that is not war- (2003), so none of our morphological char- ranted. For this reason we place Nesomantis acters could optimize on this branch. Nev- into the synonymy of Sooglossus. We could ertheless, substantial numbers of molecular have placed quotation marks around ``Soog- synapomorphies exist (appendix 5). Ford and lossus'' to note the lack of phylogenetic ev- Cannatella (1993) suggested the following to idence, but this seems to us to be an extreme be synapomorphies of Sooglossidae, but be- step to preserve a monotypic genus (i.e., Ne- cause these characters have not been reported somantis). (This synonymy affects only one for Nasikabatrachus, they may only be syn- species name, Nesomantis thomasseti Bou- ϩ apomorphies of Sooglossus Nesomantis lenger, 1909, which becomes Sooglossus tho- and should be veri®ed for Nasikabatrachus, masseti [Boulenger, 1909].) as well: (1) tarsal sesamoid bones present Because preservation of Nasikabatrachi- (see Nussbaum, 1982, for description and dae as a family would require us to have two discussion of differences among sesamoids sister families, each composed of monotypic among several taxa); (2) ventral gap in cri- genera, we consider it bene®cial for taxo- coid ring present (the universality of this nomic ef®ciency to place Nasikabatrachidae characteristic is highly speculative); (3) m. into the synonymy of Sooglossidae. Our en- semitendinosus passing dorsal to m. gracilis larged Sooglossidae is identical to Sooglos- (level of universality highly speculative); (4) soidea of Dubois (2005). alary (ϭ anterolateral) process of hyoid winglike (the level of universality specula- tive); and (5) sphenethmoid divided. In ad- [318] NOTOGAEANURA NEW TAXON dition, J.D. Lynch (1973) reported the colu- ETYMOLOGY: Noto- (Greek: southern) ϩ mella as absent in sooglossids, although he ϩ did not examine Nasikabatrachus (not dis- Gaea (Greek: earth) anoura (Greek: tail- covered for another 30 years). Biju and Bos- less, i.e., frog), denoting the Gondwanaland suyt (2003) reported the tympanum in Nasi- origin of this taxon. kabatrachus sahyadrensis as ``inconspicu- IMMEDIATELY MORE INCLUSIVE TAXON: ous'', and Dutta et al. (2004) reported it to [314] Hyloides new taxon. be absent in their unnamed species of Nasi- SISTER TAXON: [315] Sooglossidae Noble, kabatrachus. The condition of the columella 1931. in Nasikabatrachus remains unreported. J.D. RANGE: Coextensive with Anura, exclud- Lynch (1973) also reported Sooglossidae as ing New Zealand and the Seychelles. exhibiting an ossi®ed omosternum, which CONCEPT AND CONTENT: Notogaeanura is a would be a synapomorphy at this level of monophyletic taxon composed of all hyloid universality. taxa except Sooglossidae Noble, 1931. In SYSTEMATIC COMMENTS: Nussbaum et al. other words, it is composed of our [319] (1982) and Green et al. (1989) discussed the Australobatrachia new taxon and [348] Nob- phylogeny of the Seychellean taxa (i.e., not leobatrachia new taxon. including Nasikabatrachus) and suggested CHARACTERIZATION AND DIAGNOSIS: None that Sooglossus is paraphyletic with respect of our morphological characters optimize at to Nesomantis, although outgroup compari- this level of universality, so its diagnosis is son for this suggestion was lacking and the based completely on molecular data, which 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 193 are decisive. Unambiguous molecular trans- Calyptocephalellinae Reig, 1960: 113. Type ge- formations are listed in appendix 5. nus: Strand, 1928. New syn- onym. (See nomenclatural comment in appen- [319] AUSTRALOBATRACHIA NEW TAXON dix 6.) ETYMOLOGY: Australo- (Greek: southern) IMMEDIATELY MORE INCLUSIVE TAXON: ϩ batrachos (Greek: frog), denoting the southern continental distribution of these [319] Australobatrachia new taxon. frogs, primarily in Australia and New Guin- SISTER TAXON: [321] Myobatrachoidea ea, with outliers in South America, in Chile Schlegel, 1850. and Peru. RANGE: Southern Chile and north into southern Andean Peru and Bolivia. IMMEDIATELY MORE INCLUSIVE TAXON: [318] Notogaeanura new taxon. CONTENT: Batrachophrynus Peters, 1873; SISTER TAXON: [348] Nobleobatrachia new Caudiverbera Laurenti, 1768; Telmatobufo taxon. Schmidt, 1952. RANGE: New Guinea and Australia; south- CHARACTERIZATION AND DIAGNOSIS: None ern Chile and north into southern Andean of the morphological characters in our anal- Peru and Bolivia. ysis optimize on this branch because no part CONCEPT AND CONTENT: Australobatrachia of it was studied by Haas (2003). Neverthe- new taxon is a monophyletic taxon com- less, Burton (1998a) suggested that the pres- posed of [320] Batrachophrynidae Cope, ence of the m. lumbricalis longus digiti III is 1875, and [321] Myobatrachoidea Schlegel, a synapomorphy (although convergently 1850. found in Heleophryne and Petropedetes). In CHARACTERIZATION AND DIAGNOSIS: None addition, Batrachophrynus, Caudiverbera, of the morphological characters in our anal- and Telmatobufo share the presence of two ysis optimize unambiguously to this branch origins of the m. lumbricalis brevis digiti III, because the only exemplar of this group for otherwise unknown outside of Ranoides which we have morphological data is Lim- (e.g., Cardioglossa, Discodeles, most of Hy- nodynastes peronii. Therefore, any of the fol- peroliidae, Mantellidae [excluding Aglypto- lowing characters could be synapomorphies dactylus], Petropedetes, Phrynomantis, Pla- of Australobatrachia, Myobatrachoidea, Lim- typelis, Plethodonthyla, Rhacophoridae, Sco- nodynastidae, Limnodynastes, or some subset tobleps, and Trichobatrachus). See appendix of Limnodynastes: (1) m. transversus ven- 5 for molecular synapomorphies of Telma- tralis IV present (Haas 22.1); (2) lateral ®bers tobufo ϩ Caudiverbera, which we hypothe- of m. subarcualis rectus II±IV invade inter- size are synapomorphies of Batrachophryni- branchial septum IV (Haas 29.1); (3) two dae. clearly separate heads of m. subarcualis ob- SYSTEMATIC COMMENTS: The association of liquus originate from ceratobranchialia II and Batrachophrynus with Calyptocephalellini is III (Haas 32.1); (4) processus ascendens thin arguable, Batrachophrynus more traditional- (Haas 72.1); (5) processus muscularis present ly having been allied with Telmatobius (Lau- (Haas 79.0); (6) partes corpores forming me- rent, 1983; Sinsch and Juraske, 1995; Sinsch dial body (Haas 87.2); (7) adrostral cartilage et al., 1995), although this association seems very large and elongate (Haas 90.2); (8) ad- to have been assumed because of overall mandibular cartilage present (Haas 95.1); (9) similarity. (Of course, both Caudiverbera free basihyal absent (Haas 105.0); (10) com- and Telmatobufo had also been associated missura proximalis II absent (Haas 110.0); (11) commissura proximalis III absent (Haas with Telmatobius [J.D. Lynch, 1971].) J.D. 111.0); and (12) processus branchialis closed Lynch (1971: 26) noted that Caudiverbera (Haas 114.1). Unambiguous molecular trans- and Batrachophrynus (as well as Odonto- formations are listed in appendix 5. phrynus and Telmatobius) have dextral larval vent tubes, possibly an apomorphy. Another [320] FAMILY: BATRACHOPHRYNIDAE COPE, character is pupil shape, vertical in Caudiv- 1875 erbera and Telmatobufo (presumably the Batrachophrynidae Cope, 1875: 9. Type genus: apomorphic condition at this level of univer- Batrachophrynus Peters, 1873. sality) and horizontal in Batrachophrynus. 194 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Telmatobufo and Caudiverbera exhibit the lecular synapomorphies are summarized in condition of the trigeminal nerve passing appendix 5. Further study is needed. medial to the m. adductor mandibulae (the ``E'' condition); the condition is unknown in [322] FAMILY: LIMNODYNASTIDAE LYNCH, 1971 Batrachophrynus. This characteristic is oth- erwise known sporadically in Ceratophrys Limnodynastini J.D. Lynch, 1971: 83. Type ge- and Lepidobatrachus, some bufonids, Crau- nus: Limnodynastes Fitzinger, 1843. gastor, Mixophyes, some hyperoliids, most microhylids, a few ranids, rhacophorids, IMMEDIATELY MORE INCLUSIVE TAXON: Rhinophrynus, and in Sooglossus thomasseti [321] Myobatrachoidea Schlegel, 1850. (J.D. Lynch, 1986). We regard this condition SISTER TAXON: [334] Myobatrachidae as a likely synapomorphy of Batrachophryn- Schlegel, 1850. idae (or minimally Caudiverbera ϩ Telma- RANGE: Australia and New Guinea, includ- tobufo), although the distribution of this fea- ing the Aru Islands. ture across the anuran tree requires further CONTENT: Adelotus Ogilby, 1907; Heleio- study. porus Gray, 1841; Lechriodus Boulenger, 1882; Limnodynastes Fitzinger, 1843; Neo- [321] SUPERFAMILY: MYOBATRACHOIDEA batrachus Peters, 1863; Notaden GuÈnther, SCHLEGEL, 1850 1873; Opisthodon Steindachner, 1867 (see IMMEDIATELY MORE INCLUSIVE TAXON: Systematic Comments and appendix 7); Phi- [319] Australobatrachia new taxon. loria Spencer, 1901 (including Kyarranus SISTER TAXON: [320] Batrachophrynidae Moore, 1958). Cope, 1875. CHARACTERIZATION AND DIAGNOSIS: Lim- RANGE: Australia and New Guinea. nodynastids are predominantly small to mod- CONTENT: [322] Limnodynastidae Lynch, erately-sized toad-like terrestrial frogs. Am- 1971, and [334] Myobatrachidae Schlegel, plexus is inguinal, and with the exception of 1850. Neobatrachus and Notaden, all species are CHARACTERIZATION AND DIAGNOSIS: See the foam-nesters (Martin, 1967). characterization and diagnosis of Australo- See ``Characterization and diagnosis'' of batrachia for morphological characters that Australobatrachia for morphological charac- may optimize on this branch with further ters that may optimize on this branch. Ford study. At present, there are no morphological and Cannatella (1993) suggested the follow- characters that can be documented to opti- ing to be a morphological synapomorphy of mize on this branch so justi®cation for rec- Limnodynastidae: connection between the m. ognizing this taxon is based entirely on mo- submentalis and m. intermandibularis. But, lecular evidence (listed in appendix 5). with the transfer of Mixophyes to Myoba- SYSTEMATIC COMMENTS: We recognize two trachidae on the basis of molecular evidence, families within Myobatrachoidea, corre- this morphological character requires veri®- sponding substantially to Limnodynastidae cation. Davies (2003a) noted that Limnodyn- and Myobatrachidae of previous usage (Zug astidae are united by the character of fusion et al., 2001; Davies, 2003a, 2003b), differing of the ®rst two vertebrae. Molecular evidence mildly only in the transfer of Mixophyes is decisive in support of this taxon; see ap- from Limnodynastidae to Myobatrachidae pendix 5 for diagnostic transformations. and the ®rm attachment of Rheobatrachus SYSTEMATIC COMMENTS: Our results sug- (formerly Rheobatrachidae) to Myobatrachi- gest strongly that Limnodynastes as currently dae. See those accounts for further discus- formulated is polyphyletic. SchaÈuble et al. sion. Haas (2003) suggested a number of (2000) provided a tree of species of Limno- morphological characters that optimize on dynastes which corresponds in some ways his terminal taxon, Limnodynastes peronii. with our results, but which differs in others. Because this was the only myobatrachoid in Their maximum-likelihood results, based on that study, all of these characters might be 450 bp of 16S mtDNA and 370 bp of ND4, synapomorphies of various monophyletic suggest that Adelotus sits within the Limno- groups within this taxon. Hypothesized mo- dynastes ornatus group (L. ornatus and L. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 195 spenceri), and that this overall group forms Uperoliidae GuÈnther, 1858b: 346. Type genus: the sister taxon of the remaing Limnodynas- Uperoleia Gray, 1841. tes in the arrangement Limnodynastes dor- Criniae Cope, 1866: 89. Type genus: Crinia salis group ϩ (L. peronii group ϩ L. salmini Tschudi, 1838. Rheobatrachinae Heyer and Liem, 1976: 11. Type group). Our results, based on denser taxon genus: Rheobatrachus Liem, 1973. sampling and substantially more data, place Adelotus outside of Limnodynastes (sensu IMMEDIATELY MORE INCLUSIVE TAXON: lato), but place Neobatrachus, Notaden, [321] Myobatrachoidea Schlegel, 1850. Lechriodus, and Heleioporus within a para- SISTER TAXON: [322] Limnodynastidae phyletic Limnodynastes, or, alternatively, Lynch, 1971. place Limnodynastes ornatus as the sister RANGE: Australia and New Guinea. taxon of Lechriodus ¯etcheri, and far away CONTENT: Arenophryne Tyler, 1976; Assa from Limnodynastes (including Megistolotis Tyler, 1972; Crinia Tschudi, 1838; Geocri- as a synonym as suggested by SchaÈuble et nia Blake, 1973; Metacrinia Parker, 1940; al., 2000), which is the sister taxon of Hel- Mixophyes GuÈnther, 1864; Myobatrachus eioporus. In order to alleviate the polyphyly Schlegel, 1850; Paracrinia Heyer and Liem, of Limnodynastes, we resurrect the name Op- 1976; Pseudophryne Fitzinger, 1843; Rheo- isthodon Steindachner, 1867 (type species: batrachus Liem, 1973; Spicospina Roberts, Opisthodon frauenfeldi Steindachner, 1867, Horwitz, Wardell-Johnson, Maxson, and Ma- by monotypy [ϭ Discoglossus ornatus Gray, hony, 1997; Taudactylus Straughan and Lee, 1842]) for the former Limnodynastes ornatus 1966; Uperoleia Gray, 1841. group (i.e., Opisthodon ornatus [Gray, 1842] CHARACTERIZATION AND DIAGNOSIS: Myob- and O. spenceri [Parker, 1940]). This renders atrachids are predominantly small frogs of Opisthodon as the sister taxon of Lechriodus, heterogeneous appearance. All are assumed and Limnodynastes as the sister taxon of Hel- to have inguinal amplexus, except Mixophyes eioporus, assuming that both Opisthodon and (which has axillary amplexus). Davies Lechriodus are monophyletic. We suggest (2003b) noted that although Mixophyes had that the molecular characters that optimize on traditionally been associated with Limnodyn- the branch labeled Limnodynastes ornatus astidae, its placement there was always prob- are synapomorphies of Opisthodon (appen- lematic due to its lack of most limnodynas- dix 5). See appendix 7 for new combinations tinae characteristics. All myobatrachids are produced by this generic change. assumed to have a typical biphasic life his- J.D. Lynch (1971: 76) distinguished two tory (Martin, 1967). None of our morpholog- tribes within his Cycloraninae (equivalent to ical characters optimize on this branch be- our Limnodynastinae with the removal of cause no member of this taxon was studied Cyclorana to Pelodryadinae): Cycloranini by Haas (2003), although Ford and Canna- (Cyclorana, Heleioporus, Mixophyes, Neo- tella (1993) suggested the following to be a likely synapomorphy: (1) broad alary process batrachus, and Notaden), characterized by of premaxilla (absent in Mixophyes, but also laying eggs in dry in a foam nest, present in the leptodactylids Adenomera, and Limnodynastini (Adelotus, Lechriodus, Pseudopaludicola, and Physalaemus [in the Limnodynastes, and Philoria), which lay sense of including Engystomops and Eupem- their eggs in water or in moist terrestrial phix]). In our topology, this character could sites. When these characteristics are opti- be reversed in Mixophyes or convergent in mized on our cladogram, they provide a rath- Rheobatrachus and the clade bracketed by er confusing picture of life history evolution Taudactylus and Arenophryne. Regardless, in limnodynastine frogs, and our data do not the molecular evidence appears to be deci- support recognition of these taxa. sive (see appendix 5 for molecular synapo- morphies for branch 334). [334] FAMILY: MYOBATRACHIDAE SCHLEGEL, SYSTEMATIC COMMENT: We expected 1850 Myobatrachus and Arenophryne to obtain as Myobatrachinae Schlegel In Gray, 1850b: 10. sister taxa because they both are head-®rst Type genus: Myobatrachus Schlegel, 1850. burrowers in sandy soil (Tyler, 1989), with 196 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 all of the concomitant morphological features per to the Amazonian slopes that are associated with this behavior. of the . CONTENT: Hemiphractus Wagler, 1828. [348] NOBLEOBATRACHIA NEW TAXON CHARACTERIZATION AND DIAGNOSIS: None ETYMOLOGY: Noble (Gladwyn K. Noble) of the morphological characters in our anal- ϩ batrachos (Greek: frog), to note one of the ysis optimize on this branch because, as most in¯uential herpetologists of the twenti- these species are direct-developers and there- eth century and the father of modern inte- fore were not studied by Haas (2003). Nev- grative herpetology. Although Noble died ertheless, Hemiphractus/Hemiphractidae is relatively young, at age 47 (Adler, 1989), his easily diagnosed by its wild appearance and contributions to amphibian systematics, life triangular skull. Like most basal species of history, comparative anatomy, and experi- Meridianura, Hemiphractus exhibits direct mental biology remain important milestones. development and bears the developing em- IMMEDIATELY MORE INCLUSIVE TAXON: bryos on the back until hatching, but unlike [318] Notogaeanura new taxon. species of Amphignathodontidae and Cryp- SISTER TAXON: [319] Australobatrachia tobatrachidae, it does not have a dorsal new taxon. pouch in which to carry developing embryos RANGE: Coextensive with Anura, exclud- (Noble, 1931). ing New Zealand and the Seychelles. SYSTEMATIC COMMENT: Hemiphractus has CONCEPT AND CONTENT: Nobleobatrachia is two pairs of bell-shaped gills in embryos, de- a monophyletic group containing Hemi- rived from branchial arches I and II (del Pino phractidae Peters, 1862, and [349] Meridi- and Escobar, 1981; Mendelson et al., 2000), anura new taxon. as do members of Cryptobatrachidae and CHARACTERIZATION AND DIAGNOSIS: Opti- Amphignathodontidae (except for Flectono- mization of claw-shaped terminal phalanges tus pygmaeus). This suggests that the char- and intercalary elements is ambiguous, acter of bell-shaped gills optimizes on Mer- placed on this branch only under accelerated idianura, with a reversal in Athesphatanura. optimization. Under delayed optimization, however, the characters appear convergently [349] MERIDIANURA NEW TAXON in Hemiphractidae and in Cladophrynia. The ETYMOLOGY: Meridianus (Greek: southern) character of bell-shaped gills optimizes on ϩ anoura (Greek: tailless, i.e., frog), refer- Meridianura, with a reversal at Athesphatan- encing the South American center of distri- ura. Nevertheless, the bulk of evidence for bution of this worldwide group. the existence of this clade is molecular; see IMMEDIATELY MORE INCLUSIVE TAXON: appendix 5 for molecular synapomorphies. [348] Nobleobatrachia new taxon. The length of the 28S fragment likely be- comes much longer (greater than 740 bp) at SISTER TAXON: Hemiphractidae Peters, this branch than found below this point (see 1862. appendix 3), although this must be con®rmed RANGE: Coextensive with Anura, exclud- by examining the 28S fragment in Hemi- ing New Zealand and the Seychelles. phractus. CONCEPT AND CONTENT: Meridianura is a monophyletic group containing [350] Bra- FAMILY: HEMIPHRACTIDAE PETERS, 1862 chycephalidae GuÈnther, 1858, and [366] Cla- dophrynia new taxon. Hemiphractidae Peters, 1862: 146. Type genus: CHARACTERIZATION AND DIAGNOSIS:No Hemiphractus Wagler, 1828. morphological characters in our analysis op- IMMEDIATELY MORE INCLUSIVE TAXON: timize on this branch, and no authors have [348] Nobleobatrachia new taxon. suggested morphological characters that SISTER TAXON: [349] Meridianura new tax- would optimize here. Nevertheless, it is well- on. corroborated by molecular characters (see RANGE: Panama; Paci®c slopes of Colom- appendix 5 for molecular synapomorphies). bia and northwestern Ecuador; Brazil, Co- Meridianura is characterized by a length of lombia, Ecuador, Peru, and Bolivia in the up- the 28S DNA fragment in excess of 740 bp 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 197

(appendix 3). This may be plesiomorphic, exhibits the further derived character of ovo- shared with Sooglossidae and reversed in (Drewery and Jones, 1976). In ad- Australobatrachia, but long 28S molecules dition, embryonic egg teeth have been re- are characteristic nonetheless. ported for Brachycephalus and Eleuthero- dactylus (Duellman and Trueb, 1986; Pom- [350] FAMILY: BRACHYCEPHALIDAE È bal, 1999). Additional survey may ®nd that GUNTHER, 1858 this characteristic is synapomorphic for some Brachycephalina GuÈnther, 1858a: 321. Type ge- larger group; is likely coextensive with direct nus: Brachycephalus Fitzinger, 1826. development in this group; and therefore is Cornuferinae Noble, 1931: 521. Type genus: possibly synapomorphic for the entire Bra- Cornufer Tschudi, 1838. chycephalidae. For molecular transformations Eleutherodactylinae Lutz, 1954: 157. Type genus: associated with this taxon see appendix 5. Eleutherodactylus DumeÂril and Bibron, 1841. SYSTEMATIC COMMENTS: We ®nd nominal IMMEDIATELY MORE INCLUSIVE TAXON: Eleutherodactylus (sensu latoÐsubtended by [349] Meridianura new taxon. branch 350) to be in the same situation as SISTER TAXON: [366] Cladophrynia new nominal ``Hyla'' prior to its partition by Fai- taxon. vovich et al. (2005) into several tribes and RANGE: Tropical North and South Ameri- many new generaÐthat of a gigantic and ill- ca; Antilles. de®ned group where the enormity of the tax- CONTENT: Adelophryne Hoogmoed and on and lack of understanding of its species Lescure, 1984; Atopophrynus Lynch and diversity has largely restricted taxonomic Ruiz-Carranza, 1982; Barycholos Heyer, work for the past 45 years to two individuals 1969; Brachycephalus Fitzinger, 1826; Dis- (John D. Lynch and Jay M. Savage)28. Nev- chidodactylus Lynch, 1979; Craugastor ertheless, the current taxonomy of Eleuther- Cope, 1862 (see Systematic Comments and odactylus (sensu lato, subdivided into the appendix 7); ``Eleutherodactylus'' DumeÂril taxa Craugastor, Euhyas, Eleutherodactylus and Bibron, 1841 (see Systematic Comments [sensu stricto], Pelorius, and Syrrhophus) ex- and appendix 7); ``Euhyas'' Fitzinger, 1843 tends from the work of Hedges (1989) in (see Systematic Comments and appendix 7); which he named Pelorius for the Eleuthero- Euparkerella Grif®ths, 1959; Geobatrachus dactylus inoptatus group and placed Tomo- Ruthven, 1915; Holoaden Miranda-Ribeiro, dactylus as a synonym of Syrrhophus and his 1920; Ischnocnema Reinhardt and LuÈtken, enlarged Syrrhophus as a subgenus of 1862 ``1861''; ``Pelorius'' Hedges, 1989 (see Eleutherodactylus. Systematic Comments and appendix 7); Hedges' (1989) systematic arrangement Phrynopus Peters, 1873; Phyllonastes Heyer, was based on an allozymic study of six loci 1977; Heyer, 1977; Syrrho- (223 alleles) focused on West Indian species, phus Cope, 1878 (including Tomodactylus with a narrative discussion of evidence sup- GuÈnther, 1900; see Systematic Comments porting recognition of non-West Indian taxa. and appendix 7). In his UPGMA tree, Hedges' Group I (native CHARACTERIZATION AND DIAGNOSIS: Bra- Jamaican species, except E. nubicola) ap- chycephalids are predominantly leaf-litter pears monophyletic. His Group II (E. ricordii frogs with axillary amplexus and direct de- group) obtained as polyphyletic, with two velopment (J.D. Lynch, 1971, 1973). None groups placed far from each other, one group of the morphological characters in our anal- (paraphyletic to group I) and another group ysis optimized on this branch due to incom- much more basal. Group III (E. auriculatus plete taxon sampling in our morphological group) obtained as polyphyletic, with both a data set (from Haas, 2003, who restricted his basal and a ``central'' monophyletic group. study to groups with larvae). Nevertheless, Group IV (E. inoptatus group) obtained as a Brachycephalidae is characterized by pos- sessing very large terrestrial eggs and exhib- 28 G.A. Boulenger (1882), in his extraordinarily in¯u- iting direct development in all species so far ential ``Catalogue of the Batrachia Salientia'', had to examined (J.D. Lynch, 1971), with the ex- deal with only about 50 species of what is now Eleuth- ception of Eleutherodactylus jasperi, which erodactylus (sensu lato). Life was much simpler then. 198 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 monophyletic group. In the same paper a son to believe this will not worsen as sam- Distance Wagner tree also obtained Group I pling density increases. In addition to the as monophyletic, Group II as polyphyletic, paraphyly of ``Eleutherodactylus'' (sensu Group III as polyphyletic, and IV as mono- lato) with respect to Brachycephalus, dis- phyletic. cussed in ``Results'', we found Ischnocnema, After performing these analyses, Hedges Barycholos, and Phrynopus to be imbedded rejected the idea that any signi®cant evolu- within it, as was anticipated by Ardila-Ro- tionary meaning attached to these results, bayo (1979). and suggested that Groups I±IV are each As regards Ischnocnema, J.D. Lynch monophyletic on the basis of possessing (1972b: 9) noted its extreme resemblance to unique alleles (no overall analysis of pres- species of the E. binotatus group and could ence±absence provided): Group I (Icdq2), not eliminate the possibility that Ischnocne- Group II (PgmjB), Group III (Icdf1), Group IV ma represents ``a single morphological di- (Icdp5, Lgla1, Pgm0). (This survey of loci was vergence of the binotatus group of Eleuth- based solely on Antillean taxa, without any erodactylus''. Our placement of E. binotatus sampling of the nominal subgenera Craugas- and Ischnocnema quixensis as sister taxa tor or Syrrhophus, and with very limited supports that hypothesis (see below). sampling of mainland species of subgenus The sole basis for recognizing Phrynopus Eleutherodactylus.) Hedges then considered as distinct from ``Eleutherodactylus'' (sensu Group I and Group II together to form his lato or sensu stricto) is the absence of ex- subgenus Euhyas; the rationale for this uni- panded digital discs (J.D. Lynch, 1975). Ex- ®cation was not provided. His Group III he panded discs are also absent in several spe- regarded as the E. auriculatus section of a cies of ``Eleutherodactylus'' (sensu stricto), presumed paraphyletic subgenus Eleuthero- which J.D. Lynch (1994: 201) considered to dactylus (referred to later as Eleutherodac- be evidence that ``Phrynopus are simply tylus [sensu stricto]), and Group IV he con- Eleutherodactylus having greatly reduced sidered to be his monophyletic subgenus Pe- digital tips''. Our taxon sampling was inad- lorius. In subsequent discussion, he noted equate to address the relationships among all that J.D. Lynch (1986) had provided a mor- brachycephalids (i.e., eleutherodactylines) phological synapomorphy for a group that with unexpanded discs and provided only a Hedges had not examined, Craugastor (the mandibular ramus of the trigeminal nerve ly- minimal test of Phrynopus monophyly, but ing medial [deep] to the m. adductor man- our results leave little doubt that Phrynopus dibulae externus super®cialis, the ``E'' con- is nested within ``Eleutherodactylus'' (sensu dition of Starrett in J.D. Lynch, 1986), which lato). Hedges also accepted as a subgenus. Hedges J.D. Lynch (1980) considered Barycholos brie¯y discussed why he rejected Savage's to be most closely related to Eleutherodac- (1987) contention that Tomodactylus and tylus nigrovittatus (then placed in the E. dis- Syrrhophus are distantly related, and then re- coidalis group but subsequently transferred garded them as synonymous (as Syrrhophus) to the new E. nigrovittatus group by J.D. and considered Syrrhophus to be a subgenus Lynch, 1989). We did not sample any species of Eleutherodactylus. As with other authors of the E. nigrovittatus group in this study and before and since, Hedges provided no evi- therefore did not test the assertion of a Bar- dence for the monophyly of Eleutherodac- ycholos±E. nigrovittatus relationship direct- tylus (sensu lato) with respect to other eleuth- ly. However, our ®nding (following Cara- erodactyline genera. J.D. Lynch and Duell- maschi and Pombal, 2001) that Barycholos man (1997) disputed some assignments to ternetzi is nested within ``Eleutherodactylus'' Euhyas, but otherwise accepted Hedges' ar- (sensu lato) is consistent with J.D. Lynch's rangement. hypothesis. We also did not test the mono- Our results showed Eleutherodactylus phyly of Barycholos, which is characterized (sensu lato) to be rampantly nonmonophy- by sternal architecture (primarily the occur- letic (indicated below by quotation marks rence of a calci®ed sternal style; J.D. Lynch, surrounding the name), and there is no rea- 1980), but the 3,200 km separation between 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 199 the only known species is strongly sugges- subgenera is meager or lacking, several less tive that it may not be monophyletic. inclusive species groups are delimited by Adelophryne, Brachycephalus, and Eupar- synapomorphy, and we anticipate that several kerella share the characteristic of reduction of these will be recognized formally as of phalanges in the fourth ®nger, a presumed knowledge increases. synapomorphy, but this does not prevent this As a preliminary step in this direction, we group from being imbedded within ``Eleuth- take the action of treating all of the nominal erodactylus'' (sensu lato or sensu stricto)29, subgenera of ``Eleutherodactylus'' (sensu nor are we aware of any other characters that lato) as genera. (As noted in the ``Review of would exclude any of the other nominal gen- Current Taxonomy'', Crawford and Smith, era of Brachycephalidae (including former 2005, on the basis of molecular data, recently Eleutherodactylinae) from ``Eleutherodacty- considered Craugaster a genus.) As dis- lus'' (sensu lato or sensu stricto). cussed later, this is only partially successful Given the extent of the demonstrated non- inasmuch as it leaves ``Eleutherodactylus'', monophyly and lack of any evidence to dis- ``Euhyas'', and ``Pelorius'' of dubious tinguish even a phenetic ``Eleutherodacty- monophyly or even demonstrated polyphyly lus'' assemblage from other brachycephalid (denoted by the quotation marks). Neverthe- genera, the only immediately available rem- less, this illuminates the attendant problems edy, and the most scienti®cally conservative of brachycephalid relationships and leaves us action in that it enforces monophyly as the in an operationally healthier place than organizing principle of taxonomy, would be where we had been. That is, the extent of our to place all species of the former Eleuthero- knowledge of monophyly is represented by dactylinae in a single genus (coextensive the recognition of Brachycephalidae and the with our Brachycephalidae), for which the demonstrably monophyletic units within it, oldest available name is Brachycephalus Fit- and other genera are merely provisional units zinger, 1826. But, as much as this appeals to of convenience. (See appendix 7 for new us in principle, we believe that, in this par- combinations produced by these generic ticular caseÐwhere knowledge is so limited changes.) and species diversity is so great, and where Among the previous subgenera of we have sampled so few of the nominal gen- ``Eleutherodactylus'' (sensu lato) we found era of Brachycephalidae (i.e., we have not [358] Syrrhophus to be monophyletic (tested sampled Adelophryne, Atopophrynus, Dis- by inclusion of S. marnocki of the S. mar- chidodactylus, Euparkerella, Geobatrachus, nocki group of J.D. Lynch and Duellman, Holoaden, Phyllonastes,orPhyzelaphry- 1997, and S. nitidus of the S. nitidus group ne)Ðthe scienti®c payoff from enforcing of J.D. Lynch and Duellman, 1997; see ap- monophyly is not worth the practical cost of pendix 5 for molecular synapomorphies). obscuring so much diversity under a single Our sole representative of the Antillean Eu- generic name and thereby concealing a con- hyas (represented by E. planirostris of the E. siderable number of phylogenetic hypotheses ricordii group of J.D. Lynch and Duellman, that we would rather advertise in order to 1997) did not allow us to test the monophyly attract more work. Moreover, we strongly be- of this taxon. lieve that progress in the scienti®c under- In an admittedly weak test of monophyly, standing of these frogs will be achieved by we included two species of Eleutherodacty- partitioning ``Eleutherodactylus'' into multi- lus (sensu stricto), both of the E. binotatus ple monophyletic genera. Indeed, although group: E. binotatus and E. juipoca. Never- evidence for the monophyly of the nominal theless, we refuted the monophyly of ``Eleutherodactylus'' (sensu stricto) (and the 29 Complicating this, Adelophryne and Phyzelaphryne E. binotatus group), showing E. binotatus (Hoogmoed and Lescure, 1984) and at least some mem- and E. juipoca to be more closely related to bers of the Eleutherodactylus diastema group (T. Grant, Ischnocnema and Brachycephalus, respec- personal obs.) possess conspicuously pointed tips on the toe discs. This suggests that, beyond reformulation of tively. Although this ®nding was unantici- genera within former ``Eleutherodactylus'' (sensu lato), pated (but see above regarding Ischnocne- some of the other genera will have to be redemarcated. ma), no synapomorphy has yet been identi- 200 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

®ed to unite the species of ``Eleutherodac- oventrad between that muscle and the deeper tylus'' (sensu stricto) (J.D. Lynch and m. adductor mandibulae posterior (``E'' mus-

Duellman, 1997; but see below), and the E. culature), either. Instead, V3 lies entirely pos- binotatus group in particular was delimited terior to both muscles and runs ventrolaterad only by overall similarity and biogeographic toward the jawÐthat is, it does not run proximity (J.D. Lynch, 1976). around the anterior face of the m. adductor It should be noted that, although no syn- mandibulae posterior. J.D. Lynch (1986) re- apomorphy is known for ``Eleutherodacty- ported a similar pattern for one of three spec- lus'' (sensu stricto), J.D. Lynch and Duell- imens of ``E.'' angelicus and one of two man (1997) argued that a large number of its specimens of ``E.'' maussi (now ``E.'' bipor- species form a clade delimited by the ®fth catusÐSavage and Myers, 2002; the other toe being much longer than the third. Insofar specimens exhibited the ``E'' condition), and as neither of the species of ``Eleutherodac- further sampling could show the present ob- tylus'' (sensu stricto) in our sample exhibits servations to be individual anomalies as well. this state, this hypothesis remains to be tested It should also be noted that we have not ex- critically. amined the m. adductor mandibulae of the Our results also indicate that Craugastor, other species of the ``E.'' fraudator group. the so-called Middle American clade delim- Nevertheless, these observations are reason ited by the synapomorphic ``E'' pattern of enough to question the placement of this Bo- the m. adductor mandibulae (J.D. Lynch, livian group in Craugastor, which is further 1986), was polyphyletic. However, the Mid- validated by the strongly supported place- dle American species we sampled, repre- ment of ``E.'' pluvicanorus well outside of senting 5 of the 11 Middle American groups the Craugastor clade. Consequently, we re- recognized by J.D. Lynch (2000)ÐC. bufon- move the ``E.'' fraudator group from Crau- iformis, C. bufoniformis group; C. alfredi, C. gastor and return it to the already demon- alfredi group; C. augusti, C. augusti group; strably polyphyletic ``Eleutherodactylus'', C. punctariolus and C. cf. ranoides, C. ru- where J.D. Lynch and McDiarmid (1987) gulosus group; and C. rhodopis, C. rhodopis placed ``E.'' fraudator originally. Another groupÐwere monophyletic, and the sole out- option would be to name the ``E.'' fraudator lier was the Bolivian species ``Eleuthorodac- group as a new genus. However, the rela- tylus'' pluvicanorus. De la Riva and Lynch tionship of this group to ``E.'' mercedesae (1997) placed this species and ``E.'' frauda- (which shares with this group the occurrence tor (grouped subsequently with ``E.'' ash- of a frontoparietal fontanelle; J.D. Lynch and kapara as the ``E.'' fraudator group by KoÈh- McDiarmid, 1987) and the hundreds of other ler, 2000) in Craugastor on the basis of its unsampled brachycephalids is unknown, and jaw musculature, although they noted that no given that its placement in ``Eleutherodac- other species of Craugastor is known to ex- tylus'' (sensu stricto) simply in¯icts addition- tend farther south than northwestern Colom- al damage on an already polyphyletic genus, bia (e.g., C. bufoniformis; J.D. Lynch, 1986; we consider it to be premature to name this J.D. Lynch and Duellman, 1997), a possible group at present. but certainly unexpected biogeographic sce- With the exclusion from nominal Crau- nario. gastor of the ``E.'' fraudator group, which Dissection of the jaw muscles of two spec- J.D. Lynch (2000) considered to be outside imens of ``E.'' pluvicanorus (both sides of the scope of his paper, Craugastor corre- AMNH A165194, right side of AMNH sponds to the clade subtended (appendix 5) A165211) showed it to differ from the ``E'' our topology to branch 351, and generally pattern of other species (T. Grant, personal corroborates the topology of Craugastor sug- obs.). A single muscle (the m. adductor man- gested by J.D. Lynch (2000). Within Crau- dibulae externus) originates on the zygomatic gastor, J.D. Lynch (2000: 151, his ®g. 9) ramus of the squamosal, and the mandibular proposed a clade delimited by extreme sex- ramus of the trigeminal nerve (V3) does not ual dimorphism in tympanum size. In our lie lateral (super®cial) to it (so it is not the tree C. alfredi, C. augusti, and C. bufonifor- ``S'' pattern), but it does not extend poster- mis, all with nondimorphic tympana, form a 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 201 basal grade, while C. punctariolus, C. rho- Cryptobatrachidae new family and [368] dopis, and C. cf. ranoides, with strongly sex- Tinctanura new taxon. ually dimorphic tympana, are monophyletic. CHARACTERIZATION AND DIAGNOSIS: None ``Pelorius'' has had four allozymic fea- of the morphological characters in our anal- tures suggested to be synapomorphies ysis (from Haas, 2003) optimize on this (Hedges, 1989), but as noted earlier, this is branch, so its recognition depends entirely on based on sparse taxon sampling. J.D. Lynch molecular evidence, which is decisive. (See (1996) suggested not only that are there no appendix 5 for molecular synapomorphies morphological synapomorphies of this associated with this taxon.) group, but that there is a lot less than meets the eye in Hedges' (1989) study, particularly [367] FAMILY: CRYPTOBATRACHIDAE NEW with respect to how the allozymic data were FAMILY interpreted. Alternatively, J.D. Lynch (1996: IMMEDIATELY MORE INCLUSIVE TAXON: 153) suggested that ``Pelorius'' is united [366] Cladophrynia new taxon. with at least some ``Euhyas'' by the posses- SISTER TAXON: [368] Tinctanura new tax- sion of an epiotic ¯ange. So, the evidence for on. ``Pelorius'' and ``Euhyas'' monophyly seems RANGE: Northern Andes and Sierra Santa to be equivocal as well. Marta of Colombia; moderate to high ele- In summary, we recognize 16 genera with- vations of the Guayana Shield in Guyana, in Brachycephalidae. Based on our limited Venezuela, and adjacent Brazil. sampling, we recognize as monophyletic CONTENT: Cryptobatrachus Ruthven, 1916 Craugastor, Syrrhophus, Phrynopus, as du- (type genus of the family); Stefania Rivero, biously monophyletic ``Euhyas'' and ``Pelor- 1968 ``1966''. ius''; and as demonstrably nonmonophyletic CHARACTERIZATION AND DIAGNOSIS: Cryp- ``Eleutherodactylus'' (sensu stricto). We in- tobatrachidae is characterized by claw- cluded in our analysis, but did not test the shaped terminal phalanges and intercalary el- monophyly of Barycholos, Brachycephalus, ements (like Hylidae, Hemiphractidae, and and Ischnocnema, all of which fall within Amphignathodontidae) and endotrophic lar- ``Eleutherodactylus'' (sensu stricto). We did vae that develop on the back of the adult not include any representative of Adelophry- (like Hemiphractidae and Amphignathodon- ne, Atopophrynus, Dischidodactylus, Eupar- tidae). Unlike Amphignathodontidae, but like kerella, Geobatrachus, Phyllonastes, and Hemiphractidae, Cryptobatrachidae does not Phyzelaphryne. (See appendix 7 for new develop a dorsal pouch but differs from combinations produced by these generic Hemiphractus in lacking fang-like teeth. changes.) None of the morphological characters in our analysis (from Haas, 2003) optimize on this [366] CLADOPHRYNIA NEW TAXON branch, because no member of Cryptoba- trachidae was studied by Haas (2003). (Mo- ETYMOLOGY: Clados (Greek: branch) ϩ lecular transformations associated with this phrynos (Greek: toad), referring to the ob- taxon are listed in appendix 5.) servation that this taxon is a clade but not obviously united by any morphological syn- [368] TINCTANURA NEW TAXON apomorphies. ETYMOLOGY: Tincta (Greek: colored, tint- IMMEDIATELY MORE INCLUSIVE TAXON: ed) ϩ anoura (Greek: frog), denoting the fact [349] Meridianura new taxon. that many of the frogs in this clade are spec- SISTER TAXON: [350] Brachycephalidae tacularly colored (although some groups GuÈnther, 1858. within itÐnotably, most species in Bufoni- RANGE: Coextensive with Anura, exclud- daeÐcertainly lack this characteristic). ing New Zealand, Madagascar, and the Sey- IMMEDIATELY MORE INCLUSIVE TAXON: chelles. [366] Cladophrynia new taxon. CONCEPT AND CONTENT: Cladophrynia is a SISTER TAXON: [367] Cryptobatrachidae monophyletic taxon composed of [367] new family. 202 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

RANGE: Cosmopolitan in temperate and the group. Amphignathodontidae can be dif- tropical areas of the continents, Madagascar, ferentiated from other frog taxa by its pos- Seychelles, and New Zealand. session of a dorsal pouch for brooding eggs, CONCEPT AND CONTENT: Tinctanura is a a likely synapomorphy. Molecular synapo- monophyletic taxon containing [369] Am- morphies are presented in appendix 5. phignathodontidae Boulenger, 1882, and [371] Athesphatanura new taxon. [371] ATHESPHATANURA NEW TAXON CHARACTERIZATION AND DIAGNOSIS: None ETYMOLOGY: Athesphatos (Greek: inex- of the morphological characters in our anal- pressible, marvelous) ϩ anoura (Greek: tail- ysis optimized on this branch, so its recog- less, i.e., frog), denoting the fact that even nition depends entirely on molecular data. though much research has been done on (See appendix 5 for molecular synapomor- these frogs, they continue to surprise. phies associated with this taxon.) IMMEDIATELY MORE INCLUSIVE TAXON: [369] FAMILY: AMPHIGNATHODONTIDAE [368] Tinctanura new taxon. BOULENGER, 1882 SISTER TAXON: [369] Amphignathodonti- dae Boulenger, 1882. Amphignathodontidae Boulenger, 1882: 449. RANGE: Coextensive with Anura, exclud- Type genus: Amphignathodon Boulenger, 1882. Gastrothecinae Noble, 1927: 93. Type genus: ing Madagascar, Seychelles, and New Zea- Gastrotheca Fitzinger, 1843. land. Opisthodelphyinae Lutz, 1968: 13. Type genus CONCEPT AND CONTENT: As here conceived, Opisthodelphys GuÈnther, 1859 ``1858''. Athesphatanura is a monophyletic group composed of [372] Hylidae Ra®nesque, IMMEDIATELY MORE INCLUSIVE TAXON: 1815, and [424] Leptodactyliformes new [368] Tinctanura new taxon. taxon. SISTER TAXON: [371] Athesphatanura new CHARACTERIZATION AND DIAGNOSIS: Athes- taxon. phatanura is a monophyletic group composed RANGE: Costa Rica and Panama, northern of Hylidae and the bulk of the former non- and western South America southward to brachycephalid, non-batrachophrynid lepto- northwestern Argentina; eastern and south- dactylids. The following characters suggest- eastern Brazil; Trinidad and Tobago. ed by Haas (2003) on the basis of a relatively CONTENT: Flectonotus Miranda-Ribeiro, small number of exemplars are potential syn- 1920; Gastrotheca Fitzinger, 1843. apomorphies of this group: (1) pars alaris and CHARACTERIZATION AND DIAGNOSIS: Haas pars corporis separated by deep distal notch (2003) suggested the following characters (Haas 86.1); (2) commissura proximalis II that optimize on his exemplar Gastrotheca absent (Haas 110.0); and (3) commissura riobambae of amphignathodontids and may proximalis III absent (Haas 111.0). In addi- be synapomorphies of Amphignathodonti- tion, molecular synapomorphies are summa- dae: (1) m. subarcualis rectus I portion with rized in appendix 5. origin from ceratobranchial III absent (Haas 35.0); (2) functional larval m. levator man- [372] FAMILY: HYLIDAE RAFINESQUE, 1815 dibulae lateralis present (Haas 56.0); (3) ra- mus mandibularis (cranial nerve V3) poste- IMMEDIATELY MORE INCLUSIVE TAXON: rior runs through the m. levator mandibulae [371] Athesphatanura new taxon. externus group (Haas 65.1); (4) posterior pal- SISTER TAXON: [424] Leptodactyliformes atoquadrate clearly concave with bulging and new taxon. pronounced margin (Haas 68.1); (5) proces- RANGE: North and South America, the sus pseudopterygoideus long (Haas 77.2); West Indies, and the Australo-Papuan Re- and (6) dorsal connection from processus gion; temperate Eurasia, including extreme muscularis to commissura quadrato-orbitalis northern Africa and the Japanese Archipela- (Haas 78.2). All of these have the potential go. to be synapomorphies of Amphignathodon- CONTENT: [386] Hylinae Ra®nesque, 1815 tidae, although some or all may be located ϩ [373] ([374] Phyllomedusinae GuÈnther, as less inclusive levels of universality within 1858 ϩ [377] Pelodryadinae GuÈnther, 1858). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 203

CHARACTERIZATION AND DIAGNOSIS: Mor- don Miranda-Ribeiro, 1920; Aplastodiscus phological characters in our analysis (from Lutz In Lutz, 1950; Argenteohyla Trueb, Haas, 2003) that optimize on this branch are 1970; Faivovich et al., (1) m. mandibulolabialis superior present 2005; Bromeliohyla Faivovich et al., 2005; (Haas 50.1); and (2) shape of arcus subocu- Charadrahyla Faivovich et al., 2005; Cory- laris in cross-section with margin sloping thomantis Boulenger, 1896; Dendropsophus ventrally and laterally (Haas 82.1). As noted Fitzinger, 1843; Duellmanohyla Campbell by Haas (2003), traditional characters of Hy- and Smith, 1992; Ecnomiohyla Faivovich et lidae, such as claw-shaped terminal phalan- al., 2005; Exerodonta Brocchi, 1879; Hyla ges and intercalary phalangeal elements, do Laurenti, 1768; Hyloscirtus Peters, 1882; not optimize as synapomorphies of this Hypsiboas Wagler, 1830; Isthmohyla Fai- group because they are shared with Crypto- vovich, et al., 2005; Itapotihyla Faivovich et batrachidae, Hemiphractidae, and Amphig- al., 2005; Lysapsus Cope, 1862; Megasto- nathodontidae. matohyla Faivovich et al., 2005; COMMENT: Because Hylidae is so large, we Faivovich et al., 2005; Nyctimantis Boulen- deviate from our practice of providing ac- ger, 1882; Osteocephalus Steindachner, counts only for families and higher taxa and 1862; Osteopilus Fitzinger, 1843; Phyllody- here provide accounts for the three nominal tes Wagler, 1830; Plectrohyla Brocchi, 1877; subfamilies of Hylidae, which have been Pseudacris Fitzinger, 1843; Pseudis Wagler, very recently revised (Faivovich et al., 1830; Taylor, 1944; Scarthyla 2005). Duellman and de SaÂ, 1988; Scinax Wagler, 1830; Smilisca Cope, 1865 (including Pter- [386] SUBFAMILY: HYLINAE RAFINESQUE, 1815 nohyla Boulenger, 1882); Sphaenorhynchus Hylarinia Ra®nesque, 1815: 78. Type genus: Hy- Tschudi, 1838; Tepuihyla AyarzaguÈena, SenÄ- laria Ra®nesque, 1814. aris, and Gorzula, 1993 ``1992''; Tlalocohyla Hylina Gray, 1825: 213. Type genus: Hyla Lau- Faivovich et al., 2005; renti, 1768. Tschudi, 1838 (including Phrynohyas Fitzin- Dryophytae Fitzinger, 1843: 31. Type genus: Dry- ger, 1843); Triprion Cope, 1866; ophytes Fitzinger, 1843. Izecksohn, 1998 ``1996''. Dendropsophi Fitzinger, 1843: 31. Type genus: CHARACTERIZATION AND DIAGNOSIS: Only Dendropsophus Fitzinger, 1843. one morphological character in our analysis Pseudae Fitzinger, 1843: 33. Type genus: Pseudis optimizes on this taxon: two clearly separate Wagler, 1830. Acridina Mivart, 1869: 292. Type genus: Acris heads of m. subarcualis obliquus originate DumeÂril and Bibron, 1841. from ceratobranchialia II and III (character Cophomantina Hoffmann, 1878: 614. Type genus: 31.1 of Haas, 2003). Faivovich et al. (2005) Cophomantis Peters, 1870. noted another morphological synapomorphy: Lophiohylinae Miranda-Ribeiro, 1926: 64. Type tendo super®cialis digiti V (manus) with an genus: Lophyohyla Miranda-Ribeiro, 1926. additional tendon that arises ventrally from Triprioninae Miranda-Ribeiro, 1926: 64. Type ge- the m. palmaris longus (da Silva In Duell- nus: Triprion Cope, 1866. man, 2001) and, likely, the 24 chromosome Trachycephalinae Lutz, 1969: 275. Type genus: condition. Nevertheless, substantial numbers Trachycephalus Tschudi, 1838. of molecular synapomorphies exist (appen- IMMEDIATELY MORE INCLUSIVE TAXON: dix 5). [372] Hylidae Ra®nesque, 1815. SYSTEMATIC COMMENTS: The latest revision SISTER TAXON: [373] unnamed taxon of this taxon was by Faivovich et al. (2005), ([374] Phyllomedusinae GuÈnther, 1858 ϩ who provided a much larger analysis, denser [377] Pelodryadinae GuÈnther, 1858). taxonomic sampling, and more molecular RANGE: North and South America, the data per terminal than we do. Our results, West Indies; temperate Eurasia, including ex- therefore, do not constitute a suf®cient test treme northern Africa and the Japanese Ar- of those results. Nevertheless, differences chipelago. were noted. We did not ®nd either Hypsiboas CONTENT: Acris DumeÂril and Bibron, or Hyla to be monophyletic. We presume that 1841; Anotheca Smith, 1939; Aparaspheno- these differences are due to our less-dense 204 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 taxon sampling and application of fewer data monophyly of Pelodryadinae remains unset- than in the earlier study (Faivovich et al., tled. One character suggested by Haas (2003) 2005). that may optimize on this branch is larval Faivovich et al. (2005) recognized four upper labial papillation complete (Haas 8.0), tribes within Hylinae: (1) Cophomantini which is a reversal from the Phthanobatra- Hoffmann, 1878 (Aplastodiscus, Bokerman- chian condition. However, the number of pe- nohyla, Hyloscirtus, Hypsiboas, and Myer- lodryadines with complete papillation is siohyla); (2) Dendropsophini Fitzinger, 1843 small, and because Cruziohyla and Phryno- (Dendropsophus, Lysapsus, Pseudis, Scar- medusa (basal taxa in Phyllomedusinae) also thyla, Scinax, Sphaenorhynchus, and Xeno- have complete papillation it may be that this hyla); (3) Hylini Ra®nesque, 1815 (Acris, characteristic is a synapomorphy of Phyllo- Anotheca, Bromeliohyla, Charadrahyla, medusinae ϩ Pelodryadinae. Alternatively, Duellmanohyla, Ecnomiohyla, Exerodonta, more dense sampling may show convergence Hyla, Isthmohyla, , Plec- between the phyllomedusinae condition and trohyla, Pseudacris, Ptychohyla, Smilisca, that found in pelodryadines, with this con- Tlalocohyla, Triprion); and (4) Lophiohylini dition in pelodryadines, a character of some Miranda-Ribeiro, 1926 (Aparasphenodon, subset of ``Litoria'' ϩ Nyctimystes ϩ Cy- Argenteohyla, Corythomantis, Itapotihyla, clorana. Nyctimantis, Osteocephalus, Osteopilus, Haas (2003) recovered the subfamily as Phyllodytes, Tepuihyla, and Trachycephal- paraphyletic with respect to phyllomedusines us). We refer the reader to that revision for on the basis of six exemplars. Tyler (1971c) a detailed discussion of the phylogenetics of noted the presence of supplementary ele- the group. ments of the m. intermandibularis in both Pe- lodryadinae (apical) and Phyllomedusinae [377] SUBFAMILY: PELODRYADINAE (posterolateral). These characters were inter- È GUNTHER, 1858 preted by Duellman (2001) as nonhomolo- GuÈnther, 1858b: 346. Type genus: gous and therefore synapomorphies of their Pelodryas GuÈnther, 1858. respective groups. If these conditions are ho- Chiroleptina Mivart, 1869: 294. Type genus: Chi- mologues, however, the polarity between the roleptes GuÈnther, 1858. two states is ambiguous because either, the Cycloraninae Parker, 1940: 12. Type genus: Cy- pelodryadine or the phyllomedusinae condi- clorana Steindachner, 1867. tion, might be ancestral at the Phyllomedu- Nyctimystinae Laurent, 1975: 183. Type genus: sinae ϩ Pelodryadinae level of generality Nyctimystes Stejneger, 1916. (Faivovich et al., 2005). IMMEDIATELY MORE INCLUSIVE TAXON: One character in our analysis (originally [373] unnamed taxon composed of [374] from Haas, 2003) optimizes on an [373] un- Phyllomedusinae GuÈnther, 1858 ϩ [377] Pe- named taxon joining Pelodryadinae and lodryadinae GuÈnther, 1858). Phyllomedusinae: ramus mandibularis (cra-

SISTER TAXON: [374] Phyllomedusinae nial nerve V3) posterior runs through m. le- GuÈnther, 1858. vator mandibulae externus group (Haas RANGE: Australia and New Guinea; intro- 65.1). As noted by Faivovich (2005), how- duced into New Zealand. ever, another morphological synapomorphy CONTENT: Litoria Tschudi, 1838 (including of Phyllomedusinae ϩ Pelodryadinae is the Cyclorana Steindachner, 1867, and Nycti- presence of a tendon of the m. ¯exor ossis mystes Stejneger, 1916; see Systematic Com- metatarsi II arising only from distal tarsi 2± ments and appendix 7). 3. See also appendix 5 for molecular syna- CHARACTERIZATION AND DIAGNOSIS:No pomorphies of Phyllomedusinae ϩ Pelodry- morphological character optimizes unambig- adinae. uously as a synapomorphy of Pelodryadinae. The extensive paraphyly of ``Litoria'' with The molecular data, however, are decisive respect to Cyclorana and ``Nyctimystes'' re- (see Systematic Comments below and appen- mains the elephant in the room for Australian dix 5). herpetology, and for reasons that escape us SYSTEMATIC COMMENTS: Evidence of this spectacular problem has largely been ig- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 205 nored until recently; S. Donnellan and col- present (Haas 96.2); (5) cleft between hyal laborators are currently addressing pelodry- arch and branchial arch I closed (Haas adine relationships. A further dimension of 123.0); and (6) pupil shape vertically ellip- this problem is that our results not only reject tical (Haas 143.0). Additionally, Faivovich Litoria monophyly; they also show Nycti- (2005) noted that ventrolateral position of the mystes nonmonophyly, even though morpho- spiracle is a likely synapomorphy. logical evidence would suggest that Nycti- mystes is monophyletic. Our resolution at [424] LEPTODACTYLIFORMES NEW TAXON this time is to consider Nyctimystes as a syn- ETYMOLOGY: Leptodactyli- (with reference onym of Litoria and Cyclorana as a subge- to the former leptodactylids [Greek: leptos ϭ nus within Litoria. It is unfortunate to have narrow ϩ dactylos ϭ toe]) ϩ formes [Greek: to embrace such an uninformative taxonomy, shaped]). but the generic taxonomy as it exists is se- IMMEDIATELY MORE INCLUSIVE TAXON: riously misleading and no good alternatives [371] Athesphatanura new taxon. present themselves pending the resolution of SISTER TAXON: [372] Hylidae Ra®nesque, this problem by S. Donnellan and collabo- 1815. rators. (See appendix 7 for new combinations RANGE: Coextensive with Anura, exclud- produced by these generic changes.) ing Australo-Papuan region, Madagascar, Seychelles, and New Zealand. [374] SUBFAMILY: PHYLLOMEDUSINAE ONCEPT AND CONTENT GUÈ NTHER, 1858 C : Leptodactylifor- mes is a monophyletic taxon composed of GuÈnther, 1858b: 346. Type ge- [425] Diphyabatrachia new taxon and [440] nus: Phyllomedusa Wagler, 1830. Chthonobatrachia new taxon. Pithecopinae Lutz, 1969: 274. Type genus: Pithe- CHARACTERIZATION AND DIAGNOSIS: Be- copus Cope, 1866. yond our molecular data, no characters in our IMMEDIATELY MORE INCLUSIVE TAXON: analysis (originally from Haas, 2003) opti- [373] unnamed taxon composed of [374] mize on this branch. (See appendix 5 for mo- Phyllomedusinae GuÈnther, 1858 and [377] lecular synapomorphies of this taxon.) J.D. Pelodryadinae GuÈnther, 1858. Lynch (1973) reported most of the taxa out- SISTER TAXON: [377] Pelodryadinae GuÈn- side of Leptodactyliformes to have moder- ther, 1858. ately to broadly dilated sacral diapophyses, RANGE: Tropical Mexico to Argentina. so round sacral diapophyses may be a syna- CONTENT: Agalychnis Cope, 1864; Cru- pomorphy of this taxon, although reversed in ziohyla Faivovich et al., 2005; Hylomantis Centrolenidae and Bufonidae. Peters, 1873 ``1872''; Pachymedusa Duell- man, 1968; Phasmahyla Cruz, 1991 [425] DIPHYABATRACHIA NEW TAXON ``1990''; Phrynomedusa Miranda-Ribeiro, ETYMOLOGY: Diphya- (Greek: two-nature) 1923; Phyllomedusa Wagler, 1830. ϩ batrachos (Greek: frog), referencing the CHARACTERIZATION AND DIAGNOSIS: Phyl- fact that the two components of this taxon lomedusinae is a group of bizarre hylids (Centrolenidae and Leptodactylidae) have characterized by vertical pupils and a loris- very different morphologies and life-histo- like movement. Haas (2003) suggested sev- ries. eral larval characters that are good candi- IMMEDIATELY MORE INCLUSIVE TAXON: dates for being synapomorphies of Phyllo- [424] Leptodactyliformes new taxon. medusinae, although they could also be syn- SISTER TAXON: [440] Chthonobatrachia apomorphies of less inclusive groups: (1) new taxon. suspensorium ultralow (Haas 71.3); (2) pro- RANGE: Neotropics of North, Central, and cessus pseudopterygoideus short (Haas South America. 77.1); (3) arcus subocularis with three dis- CONCEPT AND CONTENT: Diphyabatrachia is tinct processes (Haas 81.2); (4) cartilaginous a monophyletic taxon containing [426] Cen- roo®ng of the cavum cranii with taeniae trolenidae Taylor, 1951, and [430] Leptodac- transversalis et medialis (fenestrae parietales) tylidae Werner, 1896 (1838). 206 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

CHARACTERIZATION AND DIAGNOSIS:No on the basis of anatomical and external sim- morphological characters suggested by Haas ilarity (Noble, 1931), but subsequent molec- (2003) optimize on this branch, although ular work (Austin et al., 2002; Faivovich et substantial amounts of molecular evidence al., 2005) has substantiated this relationship. are synapomorphic (see appendix 5). We recognize within Centrolenidae the sub- families Allophryninae for Allophryne, and [426] FAMILY: CENTROLENIDAE TAYLOR, 1951 [427] , for Centrolene, Coch- ranella, and Hyalinobatrachium. Centrolen- Centrolenidae Taylor, 1951: 36. Type genus: Cen- inae is united by the possession of intercalary trolene JimeÂnez de la Espada, 1872. Allophrynidae Goin et al., 1978: 240. Type genus: elements between the ultimate and penulti- Allophryne Gaige, 1926. New synonym. mate phalanges, fusion of the ®bula and tibia (Taylor, 1951; but see SanchõÂz and de la IMMEDIATELY MORE INCLUSIVE TAXON: Riva, 1993), and the presence of a medial [425] Diphyabatrachia new taxon. projection on the third metacarpal (Hayes SISTER TAXON: [430] Leptodactylidae Wer- and Starrett, 1981 ``1980''). ner, 1896 (1838). Our results showed ``Centrolene'' to be RANGE: Tropical southern Mexico to Bo- paraphyletic with respect to ``Cochranella''. livia, northeastern Argentina, and southeast- Morphological evidence for the monophyly ern Brazil. of Centrolene consists of a single synapo- CONTENT: Allophryne Gaige, 1926; ``Cen- morphy, the presence of a humeral spine in trolene'' JimeÂnez de la Espada, 1872 (see adult males (Ruiz-Carranza and Lynch, Systematic Comments); ``Cochranella'' Tay- 1991a), which is conspicuously present (al- lor, 1951; Hyalinobatrachium Ruiz-Carran- beit morphologically different) in both ``Cen- za and Lynch, 1991. trolene'' geckoideum and ``Centrolene'' pro- CHARACTERIZATION AND DIAGNOSIS: Haas soblepon (Ruiz-Carranza and Lynch, 1991b: (2003) suggested several characters for his 3, their ®g. 1). Nevertheless, the of exemplar, , that are some species of Cochranella exhibits a con- candidates for being synapomorphies of Cen- spicuously developed ventral crest (e.g., C. trolenidae (including Allophryne, which Haas armata, C. balionota, and C. grif®thsi; J.D. did not examine): (1) anterior insertion of m. Lynch and Ruiz-Carranza, 1997 ``1996''; see subarcualis rectus II±IV on ceratobranchial also Ruiz-Carranza and Lynch, 1991a), III (Haas 37.2); (2) larval m. levator man- which at least suggests that coding this char- dibulae externus present as one muscle body acter as the presence or absence of a humeral (Haas 54.0); (3) processus anterolateralis of spine may be simplistic. Indeed, the basis is crista parotica absent (Haas 66.0); (4) partes unclear for coding the bladelike ``spine'' of corpores medially separate (Haas 87.0); (5) Centrolene grandisonae as the same condi- cleft between hyal arch and branchial arch I tion as the smooth, rounded, and protruding closed (Haas 123.0); and (6) terminal pha- spine of C. geckoideum and as distinct from langes T-shaped (Haas 156.2). Cochranella the strongly developed bladelike crest of C. granulosa, Haas' examplar species, lacks lar- armata. We urge centrolenid workers to ex- val labial keratodonts (Haas 3.0) but this is amine the different ``spines'' in greater detail unlikely to be a synapomorphy of the group and to evaluate hypothesized homologies (Altig and McDiarmid, 1999). Burton carefully. (Note that our ®ndings do not rule (1998a) provided evidence suggesting that out homology of the humeral spines. It is the ventral origin of the m. ¯exor teretes III equally parsimonious for it to have been relative to the corresponding m. transversi gained independently in the two lineages of metacarporum I is a synapomorphy. Regard- ``Centrolene'' or gained once and lost in less of whether all of these only apply to Cochranella.) Centroleninae, there are substantial numbers We did not test the monophyly of Coch- of molecular synapomorphies (see appendix ranella or Hyalinobatrachium. No synapo- 5). morphy has been identi®ed for Cochranella SYSTEMATIC COMMENTS: The association of (Ruiz-Carranza and Lynch, 1991a) and Darst Allophryne with centrolenids was ®rst made and Cannatella (2004; ®g. 22) have presented 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 207 molecular evidence for its nonmonophyly, 1863; Hydrolaetare Gallardo, 1963; Lepto- whereas Hyalinobatrachium is delimited by dactylus Fitzinger, 1826 (see Systematic the occurrence of a bulbous liver (Ruiz-Car- Comments and appendix 7 for treatment of ranza and Lynch, 1991a, 1998). subsidiary taxa and synonyms Adenomera Given our topology, the questions sur- Steindachner, 1867, Lithodytes Fitzinger, rounding the homology of the humeral 1843, and Vanzolinius Heyer, 1974); Para- spines, and the lack of evidence for the telmatobius Lutz and Carvalho, 1958; Phys- monophyly of Cochranella, we were tempted alaemus Fitzinger, 1826; Pleurodema Tschu- to place Cochranella in the synonymy of di, 1838; Pseudopaludicola Miranda-Ribei- Centrolene. A behavioral synapomorphy for ro, 1926; Scythrophrys Lynch, 1971; Somun- the inclusive clade is male±male physical curia Lynch, 1978. combat undertaken by hanging upside down CHARACTERIZATION AND DIAGNOSIS: This by the feet and grappling venter-to-venter taxon corresponds reasonably closely to the (BolõÂvar-G. et al., 1999), a behavior other- former Leptodactylinae, excluding Limno- wise known only in phyllomedusines (Py- medusa (to Cycloramphidae) and adding burn, 1970; Lescure et al., 1995; Wogel et Paratelmatobius and Scythrophrys (from the al., 2004) and some species of Hypsiboas and former Cycloramphinae). Most species are Dendropsophus (Hylidae; J. Faivovich and found on the forest ¯oor, although a diversity C.F.B. Haddad, personal obs.). However, the of tropical biomes are inhabited. Many spe- resulting genus, though monophyletic, would cies are foam-nest builders (excluding Par- be unwieldy (with 100 species). In light of atelmatobius, some Pleurodema, Pseudopa- the poverty of our taxon sampling, and our ludicola, Scythrophrys, and Somuncuria; anticipation of more thorough phylogenetic Barrio, 1977; Pombal and Haddad, 1999; C. studies of this charismatic group, we retain Haddad, personal obs.), and this may be syn- the current taxonomy and place quotation apomorphic of the group. Several of the marks around ``Centrolene'' to denote its ap- characters in our analysis (from Haas, 2003) parent paraphyly and around ``Cochranella'' optimize on our topology on the [431] to denote its nonmonophyly as well. branch subtending Physalaemus and Pleu- rodema. Because Haas did not study other [430] FAMILY: LEPTODACTYLIDAE WERNER, members of our Leptodactylidae, these char- 1896 (1838) acters are candidates for being synapomor- Cystignathi Tschudi, 1838: 26, 78. Type genus: phies of our Leptodactylidae: (1) m. subar- Cystignathus Wagler, 1830. cualis rectus I portion with origin from cer- Leiuperina Bonaparte, 1850: 1 p. Type genus: atobranchial III absent (Haas 35.0); and (2) Leiuperus DumeÂril and Bibron, 1841. dorsal connection from processus muscularis Plectromantidae Mivart, 1869: 291. Type genus: to neurocranium and pointed (Haas 78.1). Plectromantis Peters, 1862. We also suggest that the bony sternum of Adenomeridae Hoffmann, 1878: 613. Type genus: the former Leptodactylinae (J.D. Lynch, Adenomera Steindachner, 1867. 1971) is a synapomorphy of this taxon, but Leptodactylidae Werner, 1896: 357. Type genus: reversed to the cartilaginous condition in the Leptodactylus Fitzinger, 1826. ϩ Pseudopaludicolinae Gallardo, 1965: 84. Type ge- [435] branch subtending Paratelmatobius nus: Pseudopaludicola Miranda-Ribeiro, 1926. Scythrophrys. The bony sternum occurs in- dependently in Limnomedusa (J.D. Lynch, IMMEDIATELY MORE INCLUSIVE TAXON: 1971) in Cycloramphidae, and a calci®ed [425] Diphyabatrachia new taxon. sternum occurs in Barycholos (Brachyce- SISTER TAXON: [426] Centrolenidae Taylor, phalidae; J.D. Lynch, 1980). 1951. SYSTEMATIC COMMENTS: Hydrolaetare Gal- RANGE: Extreme southern USA and trop- lardo, 1963, is associated with this group be- ical Mexico throughout Central America and cause of its presumed association with Lep- South America. todactylus (Heyer, 1970), although we sug- CONTENT: Edalorhina JimeÂnez de la Es- gest that this proposition needs to be evalu- pada, 1871 ``1870''; Engystomops JimeÂnez ated carefully. We place Somuncuria de la Espada, 1872; Eupemphix Steindachner, provisionally in this group on the basis of the 208 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 evidence (though not the conclusions) sug- Ceratophryidae Tschudi, 1838, and [448] gested by J.D. Lynch (1978b), who placed Hesticobatrachia new taxon. Somuncuria as the sister taxon of Pleurode- CHARACTERIZATION AND DIAGNOSIS: None ma. On the basis of our evidence, Leptodac- of the larval characters studied by Haas tylus is paraphyletic with respect to Vanzo- (2003) optimize on this branch, so the diag- linius; this agrees with the results of Heyer nosis of this taxon rests entirely on molecular (1998) who presented evidence to place Van- evidence. See appendix 5 for molecular syn- zolinius deeply within a paraphyletic Lepto- apomorphies. dactylus, and likely the sister taxon of Lep- todactylus diedrus.DeSa et al. (2005) also [441] FAMILY: CERATOPHRYIDAE TSCHUDI, 1838 came to this conclusion and placed Vanzoli- nus in the synonymy of Leptodactylus.We Ceratophrydes Tschudi, 1838: 26. Type genus: regard Vanzolinius as a subjective junior syn- Ceratophrys Wied-Neuwied, 1824. onym of Leptodactylus. Although our data Telmatobii Fitzinger, 1843: 31. Type genus: Tel- are agnostic on the subject, Heyer (1998) and matobius Wiegmann, 1834. New synonym. Stombinae Gallardo, 1965: 82. Type genus: Stom- Kokubum and Giaretta (2005) also presented bus Gravenhorst, 1825. evidence that recognizing Adenomera ren- Batrachylinae Gallardo, 1965: 83. Type genus: ders Leptodactylus paraphyletic and that Batrachylus Bell, 1843. New synonym. Lithodytes is the sister taxon of Adenomera. On the basis of this evidence, as well as Hey- IMMEDIATELY MORE INCLUSIVE TAXON: er's (1998) and Kokubum and Giaretta's [440] Chthonobatrachia new taxon. (2005) evidence, we place Adenomera Stein- SISTER TAXON: [448] Hesticobatrachia new dachner, 1867, as a synonym of Lithodytes taxon. Fitzinger, 1843, and Lithodytes as a subgenus RANGE: Southern Andean and tropical of Leptodactylus, without delimiting any oth- lowland South America from Colombia and er subgenera so as not to construct or imply Venezuela south to extreme southern Argen- tina and Chile. any paraphyletic groups (see appendix 7 for CONTENT: Atelognathus Lynch, 1978; Ba- new combinations). Leptodactylus, therefore trachyla Bell, 1843; Ceratophrys Wied-Neu- is equivalent to the taxon subtended by wied, 1824; Chacophrys Reig and Limeses, branch 436 in our tree. 1963; Insuetophrynus Barrio, 1970 (see Sys- J.D. Lynch (1971: 26) noted that Pseudo- tematic Comments); Lepidobatrachus Budg- paludicola and Physalaemus (including En- ett, 1899; Telmatobius Wiegmann, 1834. gystomops and Eupemphix in his sense) share CHARACTERIZATION AND DIAGNOSIS: Mor- the feature of dextral vents in larvae, as do phological characters for this group were de- Edalorhina and some Paratelmatobius (Altig rived solely from Ceratophrys and Lepidob- and McDiarmid, 1999). atrachus. Inasmuch as these are very derived taxa, the characters in our analysis that op- [440] CHTHONOBATRACHIA NEW TAXON timize on them are likely characters of Lep- idobatrachus ϩ Ceratophrys, although some ETYMOLOGY: Chthonos- (Greek: ground) of them may optimize at other hierarchic lev- ϩ batrachos (Greek: frog), referencing the els (including Ceratophryidae in our sense) fact that most of the included species are once relevant specimens have been exam- ground-dwelling. ined. Relevant morphological characters IMMEDIATELY MORE INCLUSIVE TAXON: (from Haas, 2003) are (1) m. diaphragmato- [424] Leptodactyliformes new taxon. praecordialis absent (Haas 25.0; a reversal SISTER TAXON: [425] Diphyabatrachia new from the phthanobatrachian condition, also in taxon. bufonids, microhylids, and some ranoids); RANGE: Coextensive with Anura, exclud- (3) mm. levatores arcuum branchialium I and ing Australo-Papuan region, Madagascar, II narrow with a wide gap between them Seychelles, and New Zealand. (Haas 40.0; a reversal from the phthanoba- CONCEPT AND CONTENT: Chthonobatrachia trachian condition, also in some Litoria and is a monophyletic group composed of [441] Atelopus); (4) m. suspensoriohyoideus absent 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 209

(Haas 45.0; a reversal of the acosmanuran We did not study Insuetophrynus and it is condition, also in some hylines and Atelo- therefore only provisionally allocated to this pus); (5) ramus mandibularis (cranial nerve family. Lynch (1978bb), on the basis of a V3) runs through the m. levator mandibulae phylogenetic analysis of morphology, consis- externus group (Haas 65.1; one of many re- tently recovered Insuetophrynus as the sister versals on the overall tree); (6) anterolateral taxon of Atelognathus, while Diaz et al. base of processus muscularis without con- (1983) considered the relationships of Insue- spicuous projection (Haas 86.0); (7) tectum tophrynus to lie with Alsodes (Cycloramphi- of cavum cranii almost completely chondri- dae) or Telmatobius (Ceratophryidae). The ®ed (Haas 96.4); (8) spicula short or absent characters suggested by Diaz et al. (1983) in (Haas 112.0); and (9) branchial food traps support of their arrangement all are likely absent (Haas 134.0). Molecular synapomor- plesiomorphies, however, so we retain the phies for Ceratophryidae appear in appendix hypothesis of Lynch (1978bb) pending ad- 5. ditional evidence. SYSTEMATIC COMMENTS: Within Cerato- phryidae, the association of the genera is rel- atively weak, with the exception of Lepidob- [448] HESTICOBATRACHIA NEW TAXON atrachus ϩ Ceratophrys ϩ Chacophrys and Batrachyla ϩ Atelognathus. ETYMOLOGY: Hestico- (Greek: agreeable, We recognize two subfamilies within Cer- pleasing) ϩ batrachos (Greek: frog), denot- atophryidae: [442] Telmatobiinae (for Tel- ing the agreeable nature of these frogs, par- matobius) and [444] Ceratophryinae. Within ticularly with respect to the nature of the type Ceratophryinae we recognize two tribes: genus of their sister taxon, Ceratophryidae. [445] Batrachylini (for Atelognathus, Batra- IMMEDIATELY MORE INCLUSIVE TAXON: chyla, and, presumably Insuetophrynus), and [440] Chthonobatrachia new taxon. [446] Ceratophryini (for Lepidobatrachus, SISTER TAXON: [441] Ceratophryidae Ceratophrys, and Chacophrys). Ceratophryi- Tschudi, 1838. nae has a continuous row of papilla on the RANGE: Coextensive with Anura, exclud- upper lip in larvae (Haas 8.0), a synapomor- ing Australo-Papuan region, Madagascar, phy. Batrachylini is also diagnosed on the ba- Seychelles, and New Zealand. sis of molecular evidence (appendix 5), and CONCEPT AND CONTENT: Hesticobatrachia is Ceratophryini is diagnosed on the basis of a monophyletic group composed of [449] molecular evidence as well as on traditional Cycloramphidae Bonaparte, 1858, and [460] morphological characters associated with this Agastorophrynia new taxon. cluster of genera (J.D. Lynch, 1971, 1982b): CHARACTERIZATION AND DIAGNOSIS: Char- (1) transverse processes of anterior presacral acters proposed by Haas (2003) that optimize vertebrae widely expanded; (2) cranial bones on this branch are (1) posterior palatoquad- dermosed; (3) teeth fang-like, nonpedicellate; rate curvature clearly concave with bulging and (4) absence of pars palatina of maxilla and pronounced margin (Haas 68.1); and (2) and premaxilla. Another character, presence of a vertebral shield, may be a synapomor- presence of a dorsal connection from proces- phy of Ceratophryini although the optimi- sus muscularis to neurocranium ligament zation of this feature is ambiguous, requires (Haas 78.1). Molecular synapomorphies are detailed study, and was not considered a syn- summarized in appendix 5. apomorphy by Lynch (1982b). The shield is SYSTEMATIC COMMENTS: We did not study present in Ceratophrys aurita, C. cranwelli, Rupirana Heyer, 1999, and cannot allocate it, C. ornata, C. joazeirensis, and in Lepidoba- even provisionally, although Heyer (1999) trachus asper and L. llanensis (in these two thought that it might have some kind of re- the morphology of the shield is quite differ- lationship, not close, with either Batrachyla ent from that of Ceratophrys; J. Faivovich, (Cycloramphidae) or Thoropa (Thoropidae). personal obs.), and absent in C. calcarata, C. The data to support either contention are am- cornuta, C. testudo, C. stolzmanni, Chaco- biguous at best. The position of Rupirana re- phrys pierottii, and Lepidobatrachus laevis. mains to be elucidated. 210 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

[449] FAMILY: CYCLORAMPHIDAE BONAPARTE, molecular synapomorphies (see appendix 5), 1850 Hylodinae is diagnosed by the synapomor- Cyclorhamphina Bonaparte, 1850: 1 p. Type ge- phy of having a lateral vector to the alary nus: Cycloramphus Tschudi, 1838. processes (J.D. Lynch, 1971: 39), T-shaped Rhinodermina Bonaparte, 1850: 1 p. Type genus: terminal phalanges, and dermal scutes on the Rhinoderma DumeÂril and Bibron, 1841. New top of the digital discs (J.D. Lynch, 1971, synonym, considered a junior synonym of Cy- 1973). This latter character is also found in clorhamphina Bonaparte, 1850, under Article Petropedetidae (Ranoides) and Dendrobati- 24.2.1 (Rule of First Revisor) of the Interna- tional Code of Zoological Nomenclature dae. (ICZN, 1999). Cycloramphinae is not readily diagnosed Hylodinae GuÈnther, 1858b: 346. Type genus: Hy- on the basis of morphology, but it is com- lodes Fitzinger, 1826. New synonym. posed of two tribes. The ®rst of these is [453] Alsodina Mivart, 1869: 290. Type genus: Alsodes Cycloramphini Bonaparte, 1850 (Cycloram- Bell, 1843. New synonym. phus, Crossodactylodes, and ), Grypiscina Mivart, 1869: 295. Type genus: Gry- corresponding to Grypiscini Mivart, 1869, of piscus Cope, 1867 ``1866''. J.D. Lynch (1971) with the addition of Rhi- Elosiidae Miranda-Ribeiro, 1923: 827. Type ge- nus: Elosia Tschudi, 1838. noderma. The second is [454] Alsodini Mi- Odontophrynini J.D. Lynch, 1969: 3. Type genus: vart, 1869 (composed of the remaining gen- Odontophrynus Reinhardt and LuÈtken, 1862 era). Alsodini is diagnosed by its possession ``1861''. (Odontophrynini subsequently named of Type II cotylar arrangement (cervical co- more formally by J.D. Lynch, 1971: 142.) tyles narrowly separated with two distinct ar- ticular surfaces; J.D. Lynch, 1971). This oc- IMMEDIATELY MORE INCLUSIVE TAXON: curs otherwise in Hyloides only in Batrach- [448] Hesticobatrachia new taxon. ophrynidae, Limnodynastidae, Megaelosia, SISTER TAXON: [460] Agastorophrynia new and Telmatobius (J.D. Lynch, 1971), so is taxon. likely synapomorphic at this level. (See ap- RANGE: Southern tropical and temperate pendix 5 for relevant molecular synapomor- South America. phies.) CONTENT: Alsodes Bell, 1843; Crossodac- tylodes Cochran, 1938; Crossodactylus Du- meÂril and Bibron, 1841; Cycloramphus [460] AGASTOROPHRYNIA NEW TAXON Tschudi, 1838; Eupsophus Fitzinger, 1843; ETYMOLOGY: Agastoro- (Greek: near kins- Hylodes Fitzinger, 1826; Hylorina Bell, ϩ 1843; Limnomedusa Fitzinger, 1843; Macro- man) phrynia (Greek: having the nature of genioglottus Carvalho, 1946; Megaelosia a toad), noting the surprisingly close rela- Miranda-Ribeiro, 1923; Odontophrynus tionship of Dendrobatoidea and Bufonidae. Reinhardt and LuÈtken, 1862 ``1861''; Pro- IMMEDIATELY MORE INCLUSIVE TAXON: ceratophrys Miranda-Ribeiro, 1920; Rhinod- [448] Hesticobatrachia new taxon. erma DumeÂril and Bibron, 1841; Zachaenus SISTER TAXON: [449] Cycloramphidae Bon- Cope, 1866. aparte, 1850. CHARACTERIZATION AND DIAGNOSIS: One of RANGE: Coextensive with Anura, exclud- the morphological characters suggested by ing Australo-Papuan region, Madagascar, Haas (2003) optimizes as a synapomorphy of Seychelles, and New Zealand. this group: anterior insertion of m. subarcu- CONCEPT: Agastorophrynia is a monophy- alis rectus II±IV on ceratobranchial I (Haas letic taxon composed of [461] Dendrobato- 37.0). All decisive evidence for the existence idea Cope, 1865, and [469] Bufonidae Gray, of this clade is molecular (see appendix 5). 1825. SYSTEMATIC COMMENTS: Within Cyclor- CHARACTERIZATION AND DIAGNOSIS: None amphidae we recognize two sister subfami- of the morphological characters suggested by lies, [450] Hylodinae GuÈnther, 1858 (con- Haas (2003) optimize in a way that would taining Crossodactylus, Megaelosia, and Hy- suggest their possible candidacy as synapo- lodes) and [452] Cycloramphinae Bonaparte, morphies of Agastorophrynia. All decisive 1850 (for the remaining genera). Other than evidence for the recognition of this taxa is 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 211 molecular. These molecular synapomorphies of paired dermal scutes atop the digits has are summarized in appendix 5. been claimed as a synapomorphy of Dendro- batidae ϩ Hylodinae (e.g, Noble, 1926), but [461] SUPERFAMILY: DENDROBATOIDEA COPE, its optimization on our optimal topology re- 1865 quires only a single extra step, versus the 39 IMMEDIATELY MORE INCLUSIVE TAXON: steps required to disrupt the relationship be- [460] Agastorophrynia new taxon. tween Thoropa and Dendrobatidae. Insofar SISTER TAXON: [469] Bufonidae Gray, as there is no compelling evidence against 1825. our optimal solution, and despite our aston- RANGE: Central America (Nicaragua to ishment at the result, we recognize these sis- Panama) and South America (Guianas, Am- ter taxa as Dendrobatoidea and leave it to azon drainage, south to Bolivia and south- future tests based on greater character (in- eastern Brazil). cluding morphology) and taxon sampling to CONTENT: Thoropidae new family and assess the reality of this clade. Alternatively, [462] Dendrobatidae Cope, 1865. we could have left Thoropa insertae sedisÐ CHARACTERIZATION AND DIAGNOSIS: Evi- an obviously de®cient solutionÐor have dence for Dendrobatoidea is derived entirely placed it inside Dendrobatidae. from DNA sequence data, as summarized in appendix 5. FAMILY: THOROPIDAE NEW FAMILY SYSTEMATIC COMMENT: The sister group re- IMMEDIATELY MORE INCLUSIVE TAXON: lationship of Dendrobatidae and Thoropa,to [461] Dendrobatoidea. our knowledge, has never been proposed, SISTER TAXON: [462] Dendrobatidae Cope, and this is one of the most heterodox of our 1865. results. No morphological synapomorphies RANGE: Eastern, southeastern, and south- are apparent, and a large number of charac- ern Brazil. ters differ between the two taxa. Neverthe- CONTENT: Thoropa Cope, 1865. less, evidence for alternative placement of CHARACTERIZATION AND DIAGNOSIS: Be- Thoropa appears to be lacking (although the cause this taxon was not studied by Haas larvae of Thoropa and Cycloramphus are (2003) none of the morphological characters very similar and semiterrestrial; Haddad and in our analysis could optimize on this branch. Prado, 2005), and most of the characters that All evidence for the phylogenetic placement differ between Dendrobatidae and Thoropa of this taxon as distinct from Cycloramphi- are either of unclear polarity or unique to nae is molecular, although Thoropa larvae Dendrobatidae among hyloids (e.g., thigh can be distinguished from all near relatives musculature, epicoracoid fusion and nonov- by being very attenuate and ¯attened (J.D. erlap). Furthermore, it does not appear that Lynch, 1971: 124). For additional differentia this result is due to inadequate algorithmic see J.D. Lynch (1972a). searching. At numerous points in the analysis SYSTEMATIC COMMENTS: See comment un- we placed Dendrobatidae, Thoropa, the hy- der Hesticobatrachia regarding Rupirana. lodine genera, and various other cycloram- J.D. Lynch (1971) considered Thoropa to be phines and bufonids in alternative arrange- closely related to Batrachyla, sharing a Type ments and submitted those topologies either I cotylar arrangement, although the polarity as starting points or as constraint ®les for fur- of the character was unclear in his study, and ther searching, but our analysis invariably led dendrobatids have the Type I condition as away from those solutions. The Bremer val- well, rendering this character uninformative. ues and jackknife frequencies are both strong for this clade (39% and 100%, respectively). [462] FAMILY: DENDROBATIDAE COPE, 1865 The arrangement Thoropa (Hylodinae ϩ (1850) Dendrobatidae) requires 56 extra steps, and Phyllobatae Fitzinger, 1843: 32. Type genus: placing Thoropa in the more conventional ar- DumeÂril and Bibron, 1841. rangement Thoropa ϩ (Hylorina ϩ (Alsodes Eubaphidae Bonaparte, 1850: 1 p. Type genus: ϩ Eupsophus) and (Hylodinae ϩ Dendroba- Eubaphus Bonaparte, 1831. tidae) requires 87 extra steps. The occurrence Hylaplesidae GuÈnther, 1858b: 345. Type genus: 212 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Hylaplesia Boie, 1827 (ϭ Hysaplesia Boie, rently in a state of ¯ux. Dendrobatid mono- 1826). phyly has been upheld consistently (e.g., Dendrobatidae Cope, 1865: 100. Type genus: Myers and Ford, 1986; Ford, 1993; Haas, Dendrobates Wagler, 1830. 2003; Vences et al., 2003b) since ®rst pro- Colostethidae Cope, 1867: 191. Type genus: Co- posed by Noble (1926; see Grant et al., lostethus Cope, 1866. Calostethina Mivart, 1869: 293. Type genus: Ca- 1997), but the relationships among dendro- lostethus Mivart, 1869. batids remain largely unresolved. The most generally accepted view of den- IMMEDIATELY MORE INCLUSIVE TAXON: drobatid systematics, as summarized by My- [461] Dendrobatoidea Cope, 1865. ers et al. (1991; see also Kaplan, 1997), al- SISTER TAXON: Thoropidae new family. locates approximately two-thirds of the spe- RANGE: Central America (Nicaragua to cies to a ``basal'' grade of usually dully col- Panama) and South America (Guianas, Am- ored, nontoxic frogs (including Aromobates, azon drainage, south to Bolivia and central, Colostethus, Mannophryne, and Nepheloba- southern, and southeastern Brazil). tes), while the remaining one-third is hypoth- CONTENT: Zimmermann and esized to form a clade of putatively apose- Zimmermann, 1988; Ameerega Bauer, 1986 matic frogs (including Allobates, Ameere- (including Epipedobates Myers, 1987); Aro- ga30, Dendrobates, Minyobates, Phobobates, mobates Myers, Paolillo O., and Daly, 1991; and Phyllobates). Colostethus Cope, 1866; Cryptophyllobates Compelling evidence for the monophyly LoÈtters, Jungfer, and Widmer, 2000; Dendro- of most genera is lacking. This is especially bates Wagler, 1830 (including the case for the ``basal'' taxa. The nonmon- Bauer, 1988, and Ranitomeya Bauer, 1986); ophyly of Colostethus has been recognized Mannophryne La Marca, 1992; Minyobates for decades (J.D. Lynch, 1982a; J.D. Lynch Myers, 1987; Nephelobates La Marca, 1994; and Ruiz-Carranza, 1982), and the naming of Phobobates Zimmermann and Zimmermann, Aromobates, Epipedobates, Mannophryne, 1988; Phyllobates DumeÂril and Bibron, and Nephelobates has merely exacerbated the 1841. problem (Kaiser et al., 1994; Coloma, 1995; CHARACTERIZATION AND DIAGNOSIS: Den- Meinhardt and Parmalee, 1996; Grant et al., drobatids are well-known, mostly diurnal, 1997; Grant, 1998; Grant and Castro-Herre- terrestrial, and frequently brightly colored ra, 1998). Molecular evidence for the mono- frogs that have the exotic parental behavior phyly of Mannophryne and Nephelobates of carrying tadpoles on their back to water. was presented by La Marca et al. (2002) and Likely synapomorphies of Dendrobatidae (as Vences et al. (2003b), but the relationships optimized on our topology) from those mor- of those genera to other dendrobatids are un- phological characters reported by Haas clear. Aromobates has been hypothesized to (2003) are (1) insertion of m. rectus cervicis be the monotypic sister group of all other on proximal ceratobranchialia III and IV dendrobatids (Myers et al., 1991), but syna- (Haas 39.2); (2) adrostral cartilage present pomorphies shared with Mannophryne and but small (Haas 90.1); (3) cartilaginous roof- Nephelobates, also from the northern Andes, ing of the cavum cranii formed by taeniae cast doubt on that claim. No molecular evi- tecti medialis only (Haas 96.5); (4) larvae dence has been presented for this taxon. picked up at oviposition site and transported Among the ``aposematic'' taxa, only Phyl- to body of water adhering to dorsum of adult lobates is strongly corroborated as monophy- (Haas 137.1); (5) amplectic position cephalic letic (Myers et al., 1978; Myers, 1987; (Haas 139.2); (6) guiding behavior (Haas 142.1); (7) ®rmisterny (Haas 144.1); and (8) 30 As noted earlier, our recognition of Ameerega terminal phalanges T-shaped (Haas 156.2). Bauer, 1986, as a senior synonym of Epipedobates My- Some of these characters may ultimately ers, 1987, follows the recommendation of Walls (1994). be found to be synapomorphies of Dendro- Ranitomeya Bauer, 1988, and Oophaga Bauer, 1994, are nomenclaturally valid names, but insofar as they have batoidea, because Thoropa has not been eval- not achieved common usage, and our sampling did not uated for these characters. address their monophyly or placement, we exclude them The systematics of dendrobatids is cur- from this discussion. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 213

Clough and Summers, 2000; Vences et al., Atelopoda Fitzinger, 1843: 32. Type genus: Ate- 2000b; Widmer et al., 2000). No synapo- lopus DumeÂril and Bibron, 1841. morphy is known for Ameerega, and it is Phryniscidae GuÈnther, 1858b: 346. Type genus: likely paraphyletic or polyphyletic with re- Phryniscus Wiegmann, 1834. spect to Allobates, Colostethus, Cryptophyl- Adenomidae Cope, 1861 ``1860'': 371. Type ge- nus: Adenomus Cope, 1861. lobates, and Phobobates. Schulte (1989) and Dendrophryniscina JimeÂnez de la Espada, 1871 Myers et al. (1991) rejected Allobates and ``1870'': 65. Type genus: Dendrophryniscus Phobobates on the basis of errors in the anal- JimeÂnez de la Espada, 1871 ``1870''. ysis of behavior, lack of evidence, unac- Platosphinae FejeÂrvaÂry, 1917: 147. Type genus: counted character con¯ict, incorrect charac- Platosphus d'Isle, 1877 (fossil taxon considered ter coding, and creation of paraphyly in to be in this synonymy because Platosophus ϭ Ameerega (as also found by Clough and Bufo [sensu lato]). Summers, 2000; Vences et al., 2000b; Santos Bufavidae FejeÂrvaÂry, 1920: 30. Type genus: Bu- favus Portis, 1885 (fossil taxon considered to et al., 2003; Vences et al., 2003b), but many ϭ authors continue to recognize them. In ad- be in this synonymy because Bufavus Bufo [sensu lato]). dition, Phobobates was found to be mono- Tornierobatidae Miranda-Ribeiro, 1926: 19. Type phyletic by Vences et al. (2000b) but para- genus: Tornierobates Miranda-Ribeiro, 1926. phyletic by Clough and Summers (2000). Nectophrynidae Laurent, 1942: 6. Type genus: Similarly, Minyobates may or may not be Nectophryne Buchholz and Peters, 1875. nested within Dendrobates (Silverstone, Stephopaedini Dubois, 1987 ``1985'': 27. Type 1975; Myers, 1982, 1987; Myers and Bur- genus: Stephopaedes Channing, 1978. rowes, 1987; Jungfer et al., 1996; Clough and Summers, 2000; Jungfer et al., 2000). IMMEDIATELY MORE INCLUSIVE TAXON: Likewise, although neither study recognized [460] Agastorophrynia new taxon. Minyobates, it was found to be monophyletic SISTER TAXON: [461] Dendrobatoidea by Santos et al. (2003) but polyphyletic by Cope, 1865. Vences et al. (2003b). Cryptophyllobates is RANGE: Cosmopolitan in temperate and the most recently named genus, but it is tropical areas except for the Australo-Papuan monotypic, and its relationship to other den- region, Madagascan, Seychelles, and New drobatids is unclear. Zealand. Dif®culties in understanding the phyloge- CONTENT: Adenomus Cope, 1861 ``1860''; ny of dendrobatid frogs are compounded by Altiphrynoides Dubois, 1987 ``1986'' (in- the taxonomic problems that surround many cluding Spinophrynoides Dubois, 1987 nominal species and under appreciation of ``1986''; see Systematic Comments); Amie- species diversity (Grant and Rodriguez, tophrynus new genus (see Systematic Com- 2001). Sixty-nine valid species were named ments and appendix 7); Anaxyrus Tschudi, over the past decade (more species than were 1845 (see Systematic Comments and appen- known in 1960), 55 of which were referred dix 7); Andinophryne Hoogmoed, 1985; An- to Colostethus. Many nominal species sonia Stoliczka, 1870; Atelophryniscus throughout Dendrobatidae are likely com- McCranie, Wilson, and Williams, 1989; Ate- posed of multiple cryptic species awaiting lopus DumeÂril and Bibron, 1841; Bufo Lau- diagnosis (e.g., Caldwell and Myers, 1990; renti, 1768 (see Systematic Comments and Grant and Rodriguez, 2001; Grant, 2002), appendix 7); Bufoides Pillai and Yazdani, but the rapid increase in recognized diversity 1973; Capensibufo Grandison, 1980; is not unaccompanied by error, and critical Chaunus Wagler, 1828 (see Systematic evaluation of the limits of nominal taxa will Comments and appendix 7); Churamiti undoubtedly result in some number of these Channing and Stanley, 2002; Cranopsis being placed in synonymy (e.g., Coloma, Cope, 1875 ``1876'' (see Systematic Com- 1995; Grant, 2004). ments and appendix 7); Crepidophryne Cope, 1889; Dendrophryniscus JimeÂnez de [469] FAMILY: BUFONIDAE GRAY, 1825 la Espada, 1871 ``1870''; Didynamipus An- Bufonina Gray, 1825: 214. Type genus: Bufo Lau- dersson, 1903; Duttaphrynus new genus (see renti, 1768. appendix 7); Epidalea Cope, 1865 (see Sys- 214 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 tematic Comments and appendix 7); Frostius unexamined by us; Graybeal and Cannatella, Cannatella, 1986; Ingerophrynus new ge- 1995) lacks a Bidder's organ and because our nus; Laurentophryne Tihen, 1960; Lepto- molecular data place Melanophryniscus ®rm- phryne Fitzinger, 1843; Melanophryniscus ly as the sister taxon of remaining bufonids, Gallardo, 1961; Mertensophryne Tihen, the presence of a Bidder's organ is a syna- 1960 (including Stephopaedes Channing, pomorphy not of Bufonidae, but of branch 1979 ``1978''; see Systematic Comments and 470, Bufonidae excluding Melanophryniscus appendix 7); Metaphryniscus SenÄaris, Ayar- (and presumably Truebella). Larval charac- zaguÈena, and Gorzula, 1994; Nannophryne ters (from Haas, 2003) that are synapomor- GuÈnther, 1870 (see Systematic Comments phies of bufonids excluding Melanophrynis- and appendix 7); Nectophryne Buchholz and cus (and possibly Truebella) are (1) ramus

Peters, 1875; ``Nectophrynoides'' Noble, mandibularis (cranial nerve V3) runs through 1926 (see Systematic Comment); Nimba- the m. levator mandibulae externus group phrynoides Dubois, 1987 ``1986''; Oreo- (Haas 65.1); (2) dorsal connection from pro- phrynella Boulenger, 1895; Osornophryne cessus muscularis to commissura quadrato- Ruiz-Carranza and HernaÂndez-Camacho, orbitalis (Haas 78.2); (3) eggs deposited in 1976; Parapelophryne Fei, Ye, and Jiang, strings (Haas 141.1, diversely modi®ed high- 2003; Pedostibes GuÈnther, 1876 ``1875''; Pe- er up in the bufonid tree). Atlantal cotyles lophryne Barbour, 1938; Peltophryne Fitzin- juxtaposed (J.D. Lynch, 1971, 1973) is also ger, 1843; Phrynoidis Fitzinger, 1843 (see a likely synapomorphy of this taxon. Systematic Comments and appendix 7); SYSTEMATIC COMMENTS: As evidenced by Poyntonophrynus new genus (see Systematic our results Bufo is wildly paraphyletic with Comments and appendix 7); Pseudobufo a number of other nominal genera (as docu- Tschudi, 1838; Pseudepidalea new genus mented by Graybeal, 1997). Our sampling (see appendix 7); Rhaebo Cope, 1862 (see has likely hardly scratched the surface of this Systematic Comments and appendix 7); problem, and we hope that subsequent work Rhamphophryne Trueb, 1971; Fit- will continue to add to the evidence so far zinger, 1826 (see Systematic Comments and presented so that a more universal resolution appendix 7); Schismaderma Smith, 1849; may be reached. A complete remedy of the Truebella Graybeal and Cannatella, 1995; polyphyly/paraphyly of Bufo is beyond the Vandijkophrynus new genus (see Systematic scope of this study, although we take limited Comments and appendix 7); Werneria Po- actions to start this inevitable process. We che, 1903; ``Wolterstorf®na'' Mertens, 1939 could place all of the names that are demon- (see Systematic Comments). strably derived from ``Bufo'' into the syn- CHARACTERIZATION AND DIAGNOSIS: Several onymy of Bufo, thereby providing a mono- of the larval characters in our analysis (from phyletic taxonomy. However, because much Haas, 2003) optimize as synapomorphies of of this paraphyly was understood in 1972 Bufonidae: (1) diastema in larval lower lip (various papers in Blair, 1972a), it is clear papillation (Haas 9.1); (2) m. diaphragmato- that social inertia is standing in the way of praecordialis absent (Haas 25.0); (3) lateral progress. We judge that progress will require ®bers of m. subarcualis rectus II±IV invade the partition of ``Bufo'' into more informa- interbranchial septum IV (Haas 29.1); (4) tive natural units. processus anterolateralis of crista parotica A recent study on New World Bufo by absent (Haas 66.0); (5) larval lungs rudimen- Pauly et al. (2004) had not appeared when tary or absent (Haas 133.0, also in Ascaphus we were designing our sampling strategy. and some Litoria). Graybeal and Cannatella That work provides additional guidance in (1995) noted the fusion of the basal process our development of an improved taxonomy, of the palatoquadrate with the squamosal although the study differs from ours in ana- (Baldauf, 1959), although they noted that not lytical methods and assumptions, taxon sam- enough taxa had been evaluated to ensure pling, and amount of data involved (2.5 kb that this is the appropriate level of optimi- of mtDNA in the study by Pauly et al., 2004, zation of this character. and ca. 3.7 kb/terminal of mtDNA and Because Melanophyniscus (and Truebella, nuDNA in our study). Our results are shown 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 215 in ®gure 50 and 60; the results of Pauly et Bufo asper group on the basis of morpholo- al. (2004) are shown in ®gure 68, and a com- gy, while Liu et al. (2000) allied it with the parison of the taxa held in common by the B. melanostictus group on the basis of mo- two studies is shown in ®gure 69. Both stud- lecular evidence. For the present we accept ies found the position of Bufo margaritifer its assignment to Duttaphrynus (the Bufo me- to be remarkable. The difference is that we lanostictus group). Fei et al. (2005) regarded think further resolution should come from Torrentophryne to be part of this clade, a additional data and denser sampling rather conclusion not supported either by the study than from invoking one from among a re- of Liu et al. (2000) or by our analysis, which stricted set of published models of molecular place Torrentophryne in Bufo (sensu stricto). evolution to explain the issue away. We restrict Phrynoidis to the Bufo asper On the basis of our data analysis, as well group. We also suggest that some of the char- as other information (e.g., Pauly et al., 2004), acters that optimize to this branch in our tree we can partition the following hypothesized (appendix 5) are synapomorphies of Phry- monophyletic units out of ``Bufo'' (®g. 70): noidis. (1) [476] Rhaebo Cope, 1862 (type spe- (3) Rhinella Fitzinger, 1826: 39 (type spe- cies: Bufo haematiticus Cope, 1862). We rec- cies by monotypy: Bufo [Oxyrhynchus] pro- ognize the species of the Bufo guttatus boscideus Spix, 1824). We apply this name group, the sister group of all bufonids except to the Bufo margaritifer group (see appendix Melanophryniscus, Atelopus, Osorphophry- 6 for nomenclatural comment and appendix ne, and Dendrophryniscus (see ®gs. 50, 60), 7 for content). The most recent morphologi- as Rhaebo (see appendix 5 for molecular cal characterization of the group (as the Bufo synapomorphies, appendix 6 for nomencla- typhonius group) was by Duellman and tural comment, and appendix 7 for content) Schulte (1992), although their diagnoses ex- on the basis of their lack of cephalic crests, plicitly refer to overall similarity, not syna- their yellowish-orange skin secretions (white pomorphy. Hass et al.'s (1995) study of im- in other nominal Bufo; R.W. McDiarmid, munological distances found the group to be personal commun.), presence of an omoster- monophyletic, but their outgroup samples num (otherwise found, among bufonids, only were limited to Bufo marinus and B. spinu- in Nectophrynoides and Werneria [J.D. losus. Baldissera et al. (1999) provided evi- Lynch, 1973: 146], Capensibufo [Grandison, dence (restricted to R. margaritifer) from the 1981], and the Cranopsis valliceps group [J. nucleolar organizer region (NOR) that Rhi- R. Mendelson, III, personal commun.]), and nella may be distantly related to Chaunus. hypertrophied testes (Blair, 1972c, 1972d), Most species of Rhinella have distinctive and which in combination differentiate Rhaebo extremely expanded postorbital crests in old- from all other bufonids. (See appendix 6 for er adult females, although this does not ap- note under Bufonidae on this name.) pear to be the rule, so the diagnosis of the (2) Phrynoidis Fitzinger, 1843: 32 (type group needs re®nement. Should Rhinella Fit- species: Bufo asper Gravenhorst, 1829). Be- zinger, 1826, be found to be nested within cause it is more closely related to Pedostibes Chaunus Wagler, 1828, the name Rhinella than to other ``Bufo'', we recognize the Bufo will take precedence for the inclusive group. asper group (see appendix 7 for content) as Given our taxon sampling, we cannot rule Phrynoidis. Inger (1972) provided morpho- out the possibility that Rhinella and Rham- logical differentia that serve to distinguish phophryne are not reciprocally monophylet- Phrynoidis from other bufonid taxa. Which ic. However, although placing all the in- of the suggested characters is synapomorphic volved species in a single genus would min- is not obvious, and additional morphological imize the risk that we are wrong, we believe work is needed. Further, the monophyly of such caution to be counter-productive. Also, this taxon with respect to Pedostibes, and from a more pragmatic position, this would possibly to other unsampled genera, is an require all species currently placed in Rham- open question. The relationship of Bufo gal- phophryne to be transferred to Rhinella eatus to this taxon is arguable. Dubois and based only on the possibility of nonmono- Ohler (1999) provisionally allocated it to the phyly, not evidence. The genera may be di- 216 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 68. Maximum-likelihood tree of predominantly New World Bufonidae suggested by Pauly et al. (2004) on the basis of 2,370 bp (730 informative sites) of mitochondrial DNA (12S, tRNAVal, and 16S). Alignment was done under Clustal (Thompson et al., 1997; cost functions not disclosed) then modi®ed manually. Gaps were considered to be missing data and the substitution model assumed for the maximum-likelihood analysis was GTR ϩ⌫ϩI. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 217

Fig. 69. Comparison of our bufonid parsimony results, via terminals held in common (see ®g. 50, 60) with those of Pauly et al. (2004) (®g. 68). Taxa whose relative placement differs substantially between the two studies are in boldface. agnosed by the number and size of eggs fer from all species of Rhinella in possessing (many and small in Rhinella; few and large a conspicuously elongate snout, reduced in Rhamphophryne); by the adductor man- number of vertebrae, and vocal sacs with slit- dibulae musculature (both the m. adductor like openings. Likewise, most (but not all) mandibulae posterior subexternus and m. ad- species of Rhinella differ from all species of ductor mandibulae externus super®cialis [``S Rhamphophryne in possessing protuberant ϩ E'' of Starrett in J.D. Lynch, 1986] present vertebral spines, greatly expanded alate post- in Rhinella; only m. adductor mandibulae orbital crests, and a leaf-like dorsal pattern. subexternus [``E'' of Starrett in J.D. Lynch, Many questions remain regarding the re- 1986] present in Rhamphophryne); by the lationships between these and other New thigh musculature (m. adductor longus pre- World bufonids. For example, Crepidophry- sent in Rhinella, absent in Rhamphophryne); ne epiotica possesses the same jaw muscu- by liver morphology (trilobed with left side lature and liver morphology as Rhampho- larger than right side in Rhinella, bilobed phryne and shares large, unpigmented eggs, with right side massive, conspicuously larger similar hand and foot morphology, and ab- than left side in Rhamphophryne); and by ex- sence of the ear, suggesting it may be closely tensively webbed hands and feet in Rham- related to Rhamophophryne. However, the phophryne (T. Grant, personal obs.). Most morphological results of Graybeal (1997; (but not all) species of Rhamphophryne dif- data undisclosed) suggest that Crepidophry- 218 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Fig. 70. Generic changes suggested for bufonid taxa that we studied. This ®gure shows our terminals and the new genera, but the text should be consulted for additional generic changes that involve species not addressed in our phylogenetic analysis. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 219 ne is imbedded within Cranopsis. Similarly, of the B. melanostictus group, although this, Andinophryne is characterized as possessing too, is not consistent with our molecular ev- an omosternum, anteriorly ``®rmisternal'' idence. and posteriorly ``arciferal'' pectoral girdle (5) Epidalea Cope, 1864 (type species: (for pectoral girdle morphology see Kaplan, Bufo calamita Laurenti, 1768) is available 2004), a complete ear, partially webbed for Bufo calamita (see appendix 6 for no- hands, elongate paratoid glands, and lacking menclatural comment and appendix 7 for the m. adductor longus of the thigh (Hoog- content). We had hoped to associate the name moed, 1989b). However, none of these char- Epidalea Cope, 1864, through its type (Bufo acters is unique or clearly derived relative to calamita Laurenti, 1768), to the Bufo viridis likely relatives (e.g., Rhamophryne or Rhi- group. However, association of Bufo calam- nella), and their relationships require further ita with the Bufo viridis group is seemingly investigation. based solely on overall similarity (Inger, (4) [491] Ingerophrynus new genus (type 1972). The results based on DNA sequences species: Bufo biporcatus Gravenhorst, 1829; presented by Graybeal (1997; ®g. 25) do not etymology: Robert F. Inger ϩ Greek: phry- place B. calamita and B. viridis as closest nos [toad]). This name commemorates the relatives. Because the phylogenetic evidence extensive contributions of Robert F. Inger to so far published (Graybeal, 1997) does not the herpetology of tropical Asia and the Sun- suggest that B. calamita is a member of the das, as well as to the systematics of Asian B. viridis group (but see caveat regarding bufonids. The topology described by our ex- Graybeal's data in ``Review of Current Tax- emplars Bufo celebensis, B. galeatus, B. div- onomy''), we place them in separate genera ergens, and B. biporcatus suggests a major as an interim measure. We expect that, as bu- clade of tropical Asian bufonids. We pre- fonid phylogenetics become better under- sume that this clade contains all species of stood, the name Epidalea will be attached to the Bufo biporcatus group (see appendix 7 a larger group than just this one species. for content) in addition to B. celebensis GuÈn- (6) Pseudepidalea new genus (type spe- ther, 1859 ``1858'', and B. galeatus GuÈnther, cies: Bufo viridis Laurenti, 1768; etymology: 1864. Inger (1972) provided differentia that in reference to the overall morphological distinguish the Bufo biporcatus group from similarity of members of the ``Bufo'' viridis the remaining Bufo, although it is not obvi- group to Epidalea calamita; see appendix 7 ous which of these characters are synapo- for content). Liu et al. (2000) presented weak morphies. We also suggest that our branch evidence that Bufo raddei is not part of the 491 (see appendix 5) contains several molec- Bufo viridis complex (their exemplars being ular synapomorphies that distinguish this Bufo oblongus danatensis and B. viridis). For clade from all others. this reason we regard B. raddei as being only Association of Bufo celebensis and B. gal- provisionally assigned to this genus. A set of eatus with the Bufo biporcatus group as parts differentia provided by Martin (1972) will of Ingerophrynus rests entirely on molecular serve to distinguish this group from other bu- evidence (summarized in appendix 5), al- fonid taxa, although, as in many such diag- though we expect that some of the characters nostic summaries, we cannot identify which that differentiate the B. biporcatus group characters are apomorphies and which are from other ``Bufo'' also apply to these two plesiomorphic. We do, however, suggest that species. Bufo celebensis had not previously the molecular synapomorphies provided in been associated with any other species of appendix 5 will serve to diagnose this taxon. Bufo, so our hypothesis of relationship is (7) Duttaphrynus new genus (type spe- novel and suggests that Ingerophrynus sits cies: Bufo melanostictus Schneider, 1799; et- astride Wallace's Line. ymology: S.K. Dutta ϩ Greek: phrynos Dubois and Ohler (1999) provisionally al- [toad]) re¯ects the contributions to herpetol- located B. galeatus to the B. asper group (ϭ ogy by Sushil Kumar Dutta, noted Indian Phrynoidis), but this allocation is not consis- herpetologist). We erect this generic name for tent with our molecular evidence. Liu et al. the Bufo melanostictus group as de®ned by (2000) placed B. galeatus as the sister taxon Inger (1972) and subsequent authors. Al- 220 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 though decisive evidence for the monophyly letic Torrentophryne to be nested within the of Duttaphrynus is currently lacking, we hy- otherwise monophyletic Bufo bufo group, pothesize that the group is monophyletic and which is consistent with our results. We suggest that detailed analysis of this group therefore follow Liu et al. (2000) in placing and close relatives will document this. Mor- Torrentophryne in the synonymy of Bufo phological differentia provided by Inger (sensu stricto). Clearly, our taxon sampling (1972) serve to distinguish this group from is insuf®cient to allocate all species of re- other ``Bufo'', although which characters are maining ``Bufo'' to identi®ed clades and, as apomorphies and which are plesiomorphies we suggest later, we think that ``Bufo'' spe- remains unknown. We also suggest that at cies not allocated to this or other nominal least some of the molecular synapomorphies clades should be associated with this generic for ``Bufo'' melanostictus in our tree are syn- name in quotation marks pending resolution apomorphies for Duttaphrynus (see appendix of their phylogeny. 5, for Bufo melanostictus). (10) Vandijkophrynus new genus (type (8) Peltophryne Fitzinger, 1843 (type spe- species: Bufo angusticeps Smith, 1848; ety- cies: Bufo peltocephalus Tschudi, 1838, by mology: E. Van Dijk ϩ Greek: phrynos original designation) is a monophyletic ra- [toad], commemorating Eddie Van Dijk, not- diation within nominal ``Bufo'' and was most ed South African herpetologist and indefati- recently synonymized with Bufo by Pramuk gable tadpole specialist). (See appendix 7 for (2000). In the most recent study of the rela- content and new combinations.) We erect this tionships of this group, Pramuk (2000) ana- genus for the Bufo angusticeps group as dif- lyzed morphological characters and mtDNA ferentiated by Tandy and Keith (1972; ex- sequence data and found Peltophryne (as the cluding Bufo/Capensibufo tradouwi and C. Bufo peltocephalus group) to be most closely rosei, which do not have the distinctive re- related to the American Bufo granulosus ticulate dorsal pattern of the core group and group (see also Pregill, 1981; Pramuk, 2000; are placed phylogenetically far away in our Pramuk et al., 2001). Nevertheless, Pramuk analysis) and by Cunningham and Cherry (2000) rooted her cladogram on the Bufo re- (2004). Our discovery of the exemplar of this gularis group and otherwise had relatively group, B. angusticeps, as the sister taxon of sparse outgroup taxon sampling. Our data in- Stephopaedes is consistent with the results of dicate strongly that the Bufo peltocephalus Cunningham and Cherry (2004). Should group is not closely related to the Bufo gran- Vandijkophrynus be found to be synonymous ulosus group or any other , but with Poyntonophrynus (see below), we select is, instead, the sister taxon of the African tax- Vandijkophrynus to have priority under the on Schismaderma, which was not included in provisions of Article 24.2.1 (Rule of First previous studies of Peltophryne. The biogeo- Revisor) of the International Code of Zoo- graphic track suggested by this ®nding in- logical Nomenclature (ICZN, 1999). vites further work. We therefore resurrect (11) Mertensophryne Tihen, 1960 (type Peltophryne (see appendix 7 for content) for species: Bufo micranotis rondoensis Lover- the Bufo peltocephalus group, as distantly re- idge, 1942). We suggest that at least some of lated to other Neotropical toads. (See the no- the molecular synapomorphies (appendix 5) menclatural comment in appendix 6.) that optimize to our Stephopaedes anotis are (9) [499] Bufo Laurenti, 1768 (type spe- synapomorphies for a larger Mertensophry- cies: Rana vulgaris Laurenti, 1768, by sub- ne. Complicating discussion of phylogeny in sequent designation of Tschudi, 1838: 50). the vicinity of Stephopaedes is Mertenso- We restrict the generic name Bufo (sensu phryne and the Bufo taitanus group (see ap- stricto) to the monophyletic Bufo bufo group pendix 7 for content), which is a group of of Inger (1972) and subsequent authors (see African toads that lack tympani and colu- appendix 7 for content). Inger (1972) sug- mellae; that frequently show digit reduction gested morphological differentia for this tax- (Tandy and Keith, 1972); and that have been on that separate it from other bufonid taxa, been suggested to be related to Capensibufo although their polarity remains to be docu- (Tandy and Keith, 1972). Graybeal and Can- mented. Liu et al. (2000) found a paraphy- natella (1995) and Graybeal (1997) suggest- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 221 ed that Stephopaedes and Mertensophryne (2004). Poytonophrynus is characterized by are nested within at least some component of lacking a tarsal fold (a presumed apomor- the B. taitanus group; Cunningham and phy), having parotoid glands indistinct and Cherry (2004) arrived at similar conclusions. ¯attened (Poynton, 1964a), and the tympa- MuÈller et al. (2005) described the tadpole of num being small but distinct (Tandy and B. taitanus and reported that it has a crown Keith, 1972). We did not study any member that encircles the eyes as in Stephopaedes but of this group, but on the basis of the DNA is not so well developed. In no studies have sequence results presented by Cunningham the relationships of Mertensophryne, Stepho- and Cherry (2004), it is a monophyletic paedes, and the Bufo taitanus group been group, closely related to Mertensophryne evaluated with adequate taxon sampling, and (sensu lato) and Vandijkophrynus. See ap- the questions of relationship have remained pendix 7 for content and new combinations. recognized (e.g., Tandy and Keith, 1972) but See appendix 7 for content and new combi- unresolved for more than 30 years. What is nations. known is that the Bufo taitanus group, Mer- (13) [506] Amietophrynus new genus tensophryne, and Stephopaedes lack colu- (type species: Bufo regularis Reuss, 1833; et- mellae (convergently in the clades composed ymology: Jean-Louis Amiet, an in¯uential of [1] Wolterstorf®na, Werneria, Nectophry- herpetologist of West Africa, ϩ Greek: phry- ne, and likely Laurentophryne [Tihen, 1960; nos [toad]). We erect this taxon for all Afri- Grandison, 1981]; [2] Didynamipus; and [3] can 20-chromosome ``Bufo'' discussed by Capensibufo rosei), and likely form a mono- Cunningham and Cherry (2004; the clade phyletic group. Furthermore, Stephopaedes, subtended by our branch 506), as well as the Mertensophryne, and at least one member of 22-chromosome ``Bufo'' imbedded within the Bufo taitanus group (B. taitanus;H. this clade (the Bufo pardalis group of Cun- MuÈller et al., 2005) have accessory respira- ningham and Cherry, 2004). This includes tory structures on the head of the larva. toads previously included by various authors (Nevertheless, differences among these struc- in the Bufo blanfordi group, B. funereus tures suggest the possibility of nonhomolo- group, B. kerinyagae group, B. latifrons gy; Dubois, 1987 ``1985''; Poynton and group, B. lemairii group, B. maculatus Broadley, 1988.) Mertensophryne is current- group, B. pardalis group, B. perreti group, ly monotypic, and Stephopaedes contains B. regularis group, B. superciliaris group, three species. Our action to promote further and B. tuberosus group. Although at least research is to place the Bufo taitanus group, some of these groups are monophyletic, we Mertensophryne, and Stephopaedes into an do not recognize species groups within Amie- enlarged Mertensophryne, retaining Stepho- tophrynus at this time, because several of the paedes as a subgenus, in order not to lose existing groups are monotypic (e.g., B. le- recognition of this monophyletic group. (See mairii) or clearly nonmonophyletic (e.g., B. appendix 7 for new combinations.) Loss of regularis group). We think that recognition the middle ear is synapomorphic at this level of species groups should follow a more and, although larvae are unknown for several densely sampled study of the Amietophrynus members of the Bufo taitanus group, we sus- and near relatives. Although not previously pect that the accessory repiratory structures suggested to be a member of the 20-chro- on the head of larvae is a synapomorphy as mosome group, our phylogenetic results al- well. Ongoing work by Channing and col- low us to predict that Bufo tuberosus is a 20- laborators will address this further. chromosome frog. (12) Poyntonophrynus new genus (type Beyond the 20-chromosome condition that species: Bufo vertebralis Smith, 1848; ety- is apomorphic for group (Bogart, 1968; Cun- mology: J.C. Poyton [commemorating John ningham and Cherry, 2004), molecular trans- C. Poynton, noted South African herpetolo- formations diagnose the taxon unambiguous- gist] ϩ phrynos [Greek: toad]). We recognize ly (see appendix 5). Moreover, the monophy- Poyntonophrynus for the Bufo vertebralis ly of this taxon is a testable proposition. group of Tandy and Keith (1972; cf. Poyn- Other than the Bufo pardalis group (see ton, 1964) and Cunningham and Cherry above), we have no unambiguous evidence 222 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 tying the African 22-chromosome toad taxon was considered to comprise a number groups (B. gracilipes and B. mauritanicus of casually-de®ned species groups, most of groups) or such African species of unknown which require reevaluation. Although Tschu- karyotype such as the B. pentoni group and di (1845) provided an erroneous South B. arabicus group to any of the African (or American type locality for the type species, other) bufonid groups. Additional evidence it was recognized as early as 1882 (Boulen- and study will be needed to resolve their ger, 1882) that Anaxyrus melancholicus placement, which very clearly is not within Tschudi, 1845, is a junior synonym of the Bufo (sensu stricto). For the moment, we Mexican Bufo compactilis Wiegmann, 1834. merely place the generic name ``Bufo'' in This was most recently detailed by Pramuk quotation marks in combination with these and Mendelson (2003). (See appendix 7 for species to denote their formal exclusion from content and new and revived combinations.) Bufo (sensu stricto). A partial junior synonym of Anaxyrus is (14) Nannophryne GuÈnther, 1870 (type Cope (1863: 50). Under the provi- species: GuÈnther, sions of the ``Principle of First Revisor'' 1870, by monotypy). We resurrect the name (Art. 24; ICZN, 1999) we designate Bufo Nannophryne for Bufo variegatus (GuÈnther, cognatus Say, 1823, as the type species of 1870). Although we did not include this tax- Incilius to solidify this synonymy, which oth- on in our analysis, the molecular evidence erwise could have been assigned through one provided by Pauly et al. (2004) suggests of the originally included species to threaten strongly that this taxon, like Rhinella (the the priority of Cranopsis. Bufo margaritifer group), is only distantly re- (16) [519] Cranopsis Cope, 1875 ``1876'' lated to other New World ``Bufo''. Martin (type species: Bufo fastiodosus Cope, 1875 (1972) provided osteological differentia that ``1876''). We apply the name Cranopsis to serve to diagnose the taxon among ``Bufo'', the predominantly Middle American taxon but its exact phylogenetic position among subtended by branch 519. Although we know bufonids remains to be determined. Prior to of no morphological synapomorphy for this Pauly et al. (2004), some authors placed B. taxon, species within it generally exhibit a variegatus near the B. spinulosus group (e.g., distinctive appearance. Nevertheless, see ap- Blair, 1972c), whereas others (e.g., Cei, pendix 5 for molecular synapomorphies. This 1980) have declined to place it in any species group is composed of the former Bufo val- group. Pauly et al. (2004) placed it far from liceps group and allies. See appendix 7 for the B. spinulosus group, and attaching near content and new and revived combinations. the base of the bufonid exemplars that they (17) [522] Chaunus Wagler, 1828 (type studied. It remains possible that Nannophry- species: Chaunus marmoratus Wagler, 1828 ne will be found to be most closely related [ϭ Bufo granulosus Spix, 1824]). We rec- to Rhaebo, in which case Rhaebo will take ognize the predominantly South American nomenclatural precedence for the larger taxon subtended by branch 522 as Chaunus. group. No morphological characters are known to (15) [513] Anaxyrus Tschudi, 1845 (type diagnose this group, which is diagnosed species: Anaxyrus melancholicus Tschudi, completely on the basis of molecular data 1845 [ϭ Bufo compactilis Wiegmann, (see appendix 5, branch 522). Rhamphophry- 1833]). We recognize the North American ne and Rhinella may well be found to be clade of ``Bufo'' subtended by branch 513 nested within Chaunus (see Graybeal, 1997: (see appendix 5) as the genus Anaxyrus her ®g. 13; Pauly et al., 2004), in which case, Tschudi, 1845. We are unaware of any mor- Rhinella Fitzinger, 1826, will take prece- phological synapomorphy for this group, al- dence, but evidence has yet to be produced though, with exceptions, they do have a dif- to support this synonymy without recourse to ferent look and feel than the predominantly accepting a speci®c model of molecular evo- Middle-American (Cranopsis) and South- lution (Pauly et al., 2004). American (Chaunus) taxa. Recognition of Pauly et al. (2004) suggested on the basis this taxon is consistent with our results and of fewer data, more analytical assumptions, those of Pauly et al. (2004). Formerly, this but denser sampling that the Bufo margari- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 223 tifer group (see below) is imbedded within tus. But, because we did not study that spe- this group. This remains an open question, cies, and because its sole reason for being but we suggest that decisive resolution will placed outside of Nectophrynoides is its loss require denser taxon sampling and more data, of columella, a character strongly contingent not additional analytical assumptions. on immediate outgroups, we refrain from this There are several other groups of ``Bufo'' action until the appropriate phylogenetic and various individual species we have not comparisons can be made. addressed because we did not include any of In addition, the following monotypic gen- them in our analysis and because there is no era have been named since the publication of substantial published evidence on their phy- Graybeal and Cannatella (1995) and Gray- logenetic placement. All of these we simply beal (1997): Churamiti Channing and Stan- treat as incertae sedis within Bufonidae, ley, 2002, and Parapelophryne Fei, Ye, and tacked to the generic label ``Bufo'' (see ap- Jiang, 2003. Neither obviously renders any pendix 7 for a list). The reader will note that other taxon paraphyletic. Clearly, a detailed the bulk are Asian taxa, residing in geo- revision of Bufonidae without reference to graphic areas suggesting that they will be geographic boundaries is badly needed. found to be related to a number of non-Bufo genera. Only additional work will elucidate [108] RANOIDES NEW TAXON this. We think that our proposed breakup of ETYMOLOGY: Rana (Latin: frog) ϩ oides ``Bufo'' will promote more rapid progress in (Greek: having the form of). The taxon is the ®eld, because the sociological principle identical in content to the regulated super- that drives much of systematics is to show family name Ranoidea, but with an ending that other workers are wrong (Hull, 1988), change made to remove the implication that and many graduate students will certainly it is regulated by the International Code of take aim at our hypotheses. Most systema- Zoological Nomenclature (ICZN, 1999). tists recognize that, traditionally, the ®rst (See nomenclatural comment under Ranoides species to receive novel generic names have in appendix 6.) been those that are highly autapomorphic, IMMEDIATELY MORE INCLUSIVE TAXON: and subsequent authors are usually hesitant [107] Phthanobatrachia new taxon. to apply these names to more generalized SISTER TAXON: [314] Hyloides new taxon. forms. Having taken the controversial ®rst RANGE: Worldwide temperate and tropical step, we hope that other workers will step in regions, except New Zealand, most of Aus- and address the rather large number of prob- tralia, and southern South America. lems that we have identi®ed. There is much CONCEPT AND CONTENT: Ranoides new tax- work to be done in bufonids, and we intend on is a monophyletic group composed of our taxonomic proposal to serve as a frame- [109] Allodapanura new taxon and [180] work that will guide additional studies. Natatanura new taxon. We do not ®nd any compelling reason to CHARACTERIZATION AND DIAGNOSIS: Haas maintain the sister monotypic genera Alti- (2003) suggested the following characters phrynoides Dubois, 1987 ``1986'' and Spi- that we regard as synapomorphies of our nophrynoides Dubois, 1987 ``1986''. Gran- Ranoides: (1) insertion of m. rectus cervicis dison (1981) and Graybeal and Cannatella on proximal ceratobranchialia III and IV (1995) showed these African toads to be each (Haas 39.2); (2) ramus mandibularis (cranial other's closest relatives. Acting as First Re- nerve V3) is either posterior (ventral) to m. visor, we consider Altiphrynoides Dubois, levator mandibulae externus group or runs 1987 ``1986'', to be a senior synonym of Spi- through itÐa change from being anterior nophrynoides Dubois, 1987 ``1986''. (See (dorsal) to the externus group (Haas 65.0/1); appendix 7 for the single new combination.) and (3) ®rmisternal shoulder girdle (epicor- ``Nectophrynoides'' cryptus in their tree (®g. acoids are fully fused along their length; 26) is not part of a monophyletic group with Haas 144.2; convergent in Dendrobatidae). other Nectophrynoides. We were tempted to J.D. Lynch (1973: 146) suggested that an os- name a new genus for Nectophrynoides cryp- si®ed omosternum is a synapomorphy of 224 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

``Ranoidea'' (our Ranoides, excluding Mi- Liem, 1970). In addition, Tyler (1971a) sug- crohylidae and Brevicipitidae). This may be, gested that the presence of the m. cutaneous but there are alternative optimizations. pectoris could be a synapomorphy of Rano- Among others, the ossi®ed omosternum may ides, although with several reversals. have been gained at the level of Ranoides and lost independently in Microhylidae and [109] ALLODAPANURA NEW TAXON Brevicipitidae; gained at the level of Rano- ETYMOLOGY: Allodapos- (Greek: strange, ides, lost at Allodapanura, and regained at foreign, or belonging to another kind) ϩ an- Laurentobatrachia; or gained independently oura (Greek: without a tail, i.e., frog), refer- in Laurentobatrachia, Natatanura, and Hem- encing the exotic diversity of morphotypes isotidae. (See also appendix 5, branch 108, in this taxon. for molecular synapomorphies.) IMMEDIATELY MORE INCLUSIVE TAXON: SYSTEMATIC COMMENTS: Ranoides in our [108] Ranoides new taxon. sense is coextensive with the Recent content SISTER TAXON: [180] Natatanura new tax- of the superfamily Ranoidea Ra®nesque, on. 1814, of Dubois (2005). RANGE: North and South America; sub-Sa- A preliminary survey of literature (Liem, haran Africa; India and Korea to northern 1970; Tyler, 1972, 1982; Burton, 1986, Australia. 1998b) as well as examination of a few ex- CONCEPT AND CONTENT: Allodapanura new emplars of selected genera of several families taxon is a monophyletic group composed of suggests another likely synapomorphy of [110] Microhylidae GuÈnther, 1858 (1843), Ranoides, worthy of additional investigation. and [143] Afrobatrachia new taxon. Anteromedially differentiated elements of CHARACTERIZATION AND DIAGNOSIS: Mor- the m. intermandibularis are present in Ar- phological characters in our analysis (from throleptidae, Brevicipitidae, Cacosterninae Haas, 2003) that are synapomorphies are (1) (Pyxicephalidae), Ceratobatrachidae, Hemi- m. tympanopharyngeus present (Haas 20.1); sotidae, Hyperoliidae, Microhylidae, Pty- and (2) arcus subocularis round in cross sec- chadenidae (however, absent in Hildebrand- tion (Haas 82.2). In addition, absence of the tia), Petropedetidae, Phrynobatrachidae, and palatine bone in adults (Haas 146.0; a rever- are absent in Alytidae, Batrachophrynidae sal from the acosmanuran condition), may (although present in Batrachophrynus brach- optimize on this branch (to reappear on the ydactylus), Bombinatoridae, Heleophrynidae, branch subtending Afrobatrachia), or, alter- Limnodynastidae, Myobatrachidae, Peloba- natively, the palatine may be lost in Micro- tidae, Sooglossidae, and Hemiphractidae hylidae and independently in Xenosyneuni- (Beddard, 1908 ``1907'', 1911; Tyler, 1972; tanura. Similarly, the presence of palatal Tyler and Duellman, 1995; Burton, 1998b). folds may optimize on this branch and be This taxonomic distribution suggest that the reversed in Laurentobatrachia, or may appear presence of differentiated elements of the m. twice, once on the branch subtending Micro- intermandibularis is a synapomorphy of Ran- hylidae as well as on the branch subtending oides. Many details about the morphological Xenosyneunitanura. Regardless, the primary diversity and taxonomic distribution of this evidence for the recognition of this taxon is character remain unknown and several in- molecular (see appendix 5). stances of homoplasy are known within Hy- loides (see Tyler, 1971b, 1971c, 1972; [110] FAMILY: MICROHYLIDAE GUÈ NTHER, 1858 Burton, 1998b, and Tyler and Duellman, (1843) 1995, for examples within Noblebatrachia), and there are possibly multiple subsequent Hylaedactyli Fitzinger, 1843: 33. Type genus: Hy- transformations within Natatanura. (This laedactylus DumeÂril anbd Bibron, 1841. Gastrophrynae Fitzinger, 1843: 33. Type genus: character does not seem to occur in at least Gastrophryne Fitzinger, 1843. some Dicroglossidae [exemplars of Occidoz- Micrhylidae GuÈnther, 1858b: 346. Type genus: yga, Euphlyctis, Nannophrys] or Nyctiba- Micrhyla DumeÂril and Bibron, 1841 (an incor- trachidae [Lankanectes, Nyctibatrachus], but rect subsequent spelling of Microhyla Tschudi, is present in Mantellidae and Rhacophoridae; 1838). 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 225

Asterophrydidae GuÈnther, 1858b: 346. Type ge- 1986; Altigius Wild, 1995; Arcovomer Car- nus: Asterophrys Tschudi, 1838. valho, 1954; Chiasmocleis MeÂhely, 1904; Kalophrynina Mivart, 1869: 289. Type genus: Gastrophrynoides Noble, 1926; Glyphoglos- Kalophrynus Tschudi, 1838. sus GuÈnther, 1869 ``1868''; Hyophryne Car- Xenorhinidae Mivart, 1869: 286. Type genus: Xe- valho, 1954; Hypopachus Keferstein, 1867; norhina Peters, 1863. Dyscophidae Boulenger, 1882: 179. Type genus: Kalophrynus Tschudi, 1838; Dyscophus Grandidier, 1872. Parker, 1934; Micryletta Dubois, 1987; Cophylidae Cope, 1889: 248. Type genus: Cophy- Myersiella Carvalho, 1954; Otophryne Bou- la Boettger, 1880. lenger, 1900; Paradoxophyla Blommers- Genyophrynidae Boulenger, 1890: 326. Type ge- SchloÈsser and Blanc, 1991; Phrynella Bou- nus: Genyophryne Boulenger, 1890. lenger, 1887; Phrynomantis Peters, 186731; Rhombophryninae Noble, 1931: 529. Type genus: Ramanella Rao and Ramanna, 1925; Relic- Rhombophryne Boettger, 1880. tivomer Carvalho, 1954; Stereocyclops Cope, Sphenophryninae Noble, 1931: 531. Type genus: 1870 ``1869''; Synapturanus Carvalho, Sphenophryne Peters and Doria, 1878, by mon- 1954; Syncope Walker, 1973; Uperodon Du- otypy. meÂril and Bibron, 1841. (See Systematic Melanobatrachinae Noble, 1931: 538. Type ge- Comments.) nus: Melanobatrachus Beddome, 1878. Kaloulinae Noble, 1931: 538. Type genus: Kal- CHARACTERIZATION AND DIAGNOSIS: A large oula Gray, 1831. number of morphological characters in our Hoplophryninae Noble, 1931: 538±539. Type ge- analysis (from Haas, 2003) are synapomor- nus: Hoplophryne Barbour and Loveridge, phies of Microhylidae: (1) keratodonts absent 1928. in larvae (Haas 3.0); (2) keratinized jaw Scaphiophryninae Laurent, 1946: 337. Type ge- sheaths absent in larvae (Haas 6.0); (3) vena nus: Scaphiophryne Boulenger, 1882. caudalis dorsalis present in larvae (Haas Pseudohemisiinae Tamarunov, 1964a: 132. Type 14.1); (4) spiracle position median posterior genus: Pseudohemisus Mocquard, 1895. (Haas 18.2); (5) m. geniohyoideus origin in Otophryninae Wassersug and Pyburn, 1987: 166. larvae from connective tissue lateral to glan- Type genus: Otophryne Boulenger, 1900. dula thyroidea (Haas 19.4); (6) m. interhyoi- Phrynomantini Burton, 1986: 405±450. Type ge- nus: ``Phrynomantis Peters, 1867''. deus posterior in larvae extensive and strong- Barygenini Burton, 1986: 405±450. Type genus: ly developed (Haas 24.2); (7) m. diaphrag- Barygenys Parker, 1936. matopraecordialis absent in larvae (Haas Callulopini Dubois, 1988a: 3. Type genus: Cal- 25.0); (8) lateral ®bers of m. subarcualis rec- lulops Boulenger, 1888. tus II±IV invade interbranchial septum IV musculature in larvae (Haas 29.1); (9) m. su- IMMEDIATELY MORE INCLUSIVE TAXON: barcualis rectus II±IV split into medial and [109] Allodapanura new taxon. lateral separate muscles (Haas 30.1); (10) m. SISTER TAXON: [143] Afrobatrachia new subarcualis rectus I portion with origin from taxon. ceratobranchial III absent (Haas 35.0); (11) RANGE: North and South America; East ventral portion of the m. subarcualis rectus I and South Africa; India and Korea to north- inserts laterally on ceratohyal (Haas 36.1); ern Australia. (12) origin of m. suspensoriohyoideus from CONTENT: [135] Asterophryninae GuÈnther, posterior palatoquadrate (Haas 46.1); (13) m. 1858 (including Genyophryninae Boulenger, interhyoideus and m. intermandibularis in 1890), [118] Cophylinae Cope, 1889, Dys- close proximity (Haas 47.0); (14) m. man- cophinae Boulenger, 1882, [121] Gastro- dibulolabialis inserting exclusively on carti- phryninae Fitzinger, 1843, [130] Microhyli- lago labialis inferior (Haas 49.1); (15) m. le- nae GuÈnther, 1858 (1843), Scaphiophryninae vator mandibulae internus anterior (Haas Laurent, 1946, as well as several nominal genera unassigned to subfamily either be- 31 We realize, of course, that Phrynomantis Peters, cause we did not study them and assignment 1867, is the sole member of Phrynomerinae Noble, 1931. But, beyond the autapomorphic intercalary pha- to subfamily based on published evidence is langeal elements, we have only weak evidence for its not possible, or because they fall outside of placement. In this case, recognition of a monotypic sub- existing subfamilies: Adelastes Zweifel, family serves no purpose. 226 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

58.2); (16) m. levator mandibulae longus monophyletic taxonomy we propose the fol- originates exclusively from arcus subocularis lowing taxonomic changes: (1) place Aster- (Haas 60.2); (17) profundus and super®cialis ophryinae and Genyophryninae in one sub- portions of m. levator mandibulae longus not family, Asterophryinae (following Savage, overlapping and parallel (Haas 62.1); (18) ra- 1973); (2) restrict Dyscophinae to Dyscophus mus mandibularis (cranial nerve V3) between (also following Savage, 1973) and transfer portions of m. levator mandibulae longus Calluella from Dyscophinae to Microhyli- muscle (Haas 64.1); (19) processus suboticus nae; (3) retain Cophylinae, but note that it quadrati present (Haas 76.1); (20) partes cor- appears to be imbedded within a cluster of pores forming medial body (Haas 87.2); (21) ``microhyline'' genera that, once their phy- distal end of cartilago meckeli expanded and logeny is better resolved, may require some ¯attened with no fossa (Haas 94.2); (22) hy- reconstitution of Cophylinae; and (4) parti- pobranchial plates fused (Haas 107.1); (23) tion Microhylinae into a New World group, commissura proximalis I present (Haas Gastrophryninae, and an Old World group, 109.1); (24) processus branchialis closed Microhylinae, with several genera left incer- (Haas 114.1); (25) accessory longitudinal tae sedis until they can be adequately studied bars of cartilage dorsal to ceratobranchialia or placed in a more densely sampled frame- II and III present (Haas 120.1); (26) posterior work. Another group of genera (i.e., Kalo- margin of ventral velum discontinuous (Haas phrynus, Synapturanus, Phrynomantis, Mi- 129.1); (27) glottis position posterior (Haas cryletta) is left incertae sedis, as well, al- 130.1); (28) nostrils closed in larval stages though the phylogenetic structure we ob- (Haas 131.1); (29) branchial food traps di- tained among these taxa is instructive and vided and crescentic (Haas 135.1); and (30) points to new questions for systematists to eggs ¯oating (Haas 141.2). Although most of address. Nevertheless, our obtained structure these characters will survive denser taxon suggests that the biogeography of Microhy- sampling, the placement of some of them is lidae is complex and old. currently ambiguous inasmuch as some of Our data show that the former ``Micro- the characters listed could actually be sitting hylinae'' (sensu lato) is heterogenous mix- on branches from which Synapturanus and ture of basal taxa (e.g., Synapturanus, Mi- Kalophrynus are derived. cryletta) and two distantly related clades with Presence of palatal folds is optimization- which we have associated the names Micro- dependent. Presence of palatal folds may be hylinae (Asia) and Gastrophryninae (Ameri- convergent in Microhylidae and Xenosyneu- cas). There is no published evidence that nitanura, or a synapomorphy of Allodapan- would allow us to allocate any of the un- ura and lost in Laurentobatrachia. studied Asian taxa to Microhylinae or to any SYSTEMATIC COMMENTS: The obtained phy- other position in our cladogram beyond their logenetic structure of Microhylidae surprised being microhylids. Similarly, although we us as we expected Scaphiophryninae to form assume that such taxa as Hypopachus are in the sister taxon of the remaining microhylids, Gastrophryninae, our molecular results are so because the scaphiophrynine tadpole mor- incongruent with results from morphology phology (Blommers-SchloÈsser, 1975; Haas, (e.g., Zweifel, 1986; Donnelly et al., 1990; 2003), is annectant in many ways between Wild, 1995) that we hesitate to conjecture. the ranid and more typical microhylid con- Morphological characters that are candi- dition. As in several other parts of the tree, dates as synapomorphies of [134] Dyscophi- the density of our taxon sampling was inad- nae ϩ Asterophryninae ϩ Scaphiophryninae equate to address all problems in microhylid ϩ Microhylinae clade are (1) double-layered systematics, and we intend our results to dermis (Haas 13.1, also in Hemisus and Kas- guide more thorough studies. Rafael de Sa sina); (2) anterior insertion of m. subarcualis and collaborators have begun such a study, rectus II±IV on ceratobranchial I (Haas and we anticipate further revision of micro- 37.0); and (3) partes corpores forming medial hylid systematics as a result. For this reason body (Haas 87.2). we leave several taxa unnamed and unad- Because the nominal subfamilies of Mi- dressed. As an initial step toward an entirely crohylidae are large and morphologically dis- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 227 parate, we include separate accounts for the place completely within the egg capsule, al- nominal subfamilies. though others (e.g., Cophylinae, some Gas- trophryninae) are endotrophic and nidicolous [135] SUBFAMILY: ASTEROPHRYINAE (Blommers-SchloÈsser, 1975). (See appendix GUÈ NTHER, 1858 5 for molecular synapomorphies.) Asterophrydidae GuÈnther, 1858b: 346. Type ge- SYSTEMATIC COMMENTS: Former Geny- nus: Asterophrys Tschudi, 1838. ophryninae is paraphyletic with respect to Xenorhinidae Mivart, 1869: 286. Type genus: Xe- the old Asterophryinae, and for this reason norhina Peters, 1863. the two nominal taxa were synonymized in Genyophrynidae Boulenger, 1890: 326. Type ge- ``Results''. Parker (1934) noted Genyophry- nus: Genyophryne Boulenger, 1890. New syn- ninae (as Sphenophryninae) to be procoelous onym. and Asterophryinae as diplasiocoelous, and Sphenophryninae Noble, 1931: 531. Type genus: this clearly in¯uenced later authors (e.g., Sphenophryne Peters and Doria, 1878, by mon- Zweifel, 1972) in retaining a distinction be- otypy. New synonym. tween the nominal subfamilies. The place- Phrynomantini Burton, 1986: 405±450. Type ge- nus: ``Phrynomantis Peters, 1867''. ment in our tree of Australo-Papuan Aster- Barygenini Burton, 1986: 405±450. Type genus: ophryinae (sensu lato) as the sister taxon of Barygenys Parker, 1936. the Madagascan Dyscophinae is a remark- Callulopini Dubois, 1988a: 3. Type genus: Cal- able biogeographic signature. lulops Boulenger, 1888. Burton (1986: 443) provided evidence that Xenorhina is paraphyletic with respect to Xe- IMMEDIATELY MORE INCLUSIVE TAXON: nobatrachus, the latter differing only in lack- [134] unnamed taxon. ing large odontoids on the vomeropalatine. SISTER TAXON: Dyscophinae Boulenger, Zweifel (1972) provided no evidence for the 1882. monophyly of Xenorhina. On the basis of RANGE: Southern Philippines, Sulawesi, these works we consider them to be syno- and Lesser Sunda Islands and Moluccas east- nyms, with Xenorhina being the older name wards through New Guinea and satellite is- (see appendix 7 for new combinations). lands to Australia. Burton (1986: 443) also noted that ``Manto- CONTENT: Albericus Burton and Zweifel, phryne'' and ``Hylophorbus'' are dubiously 1995; Aphantophryne Fry, 1917 ``1916''; monophyletic, so we place these names in Asterophrys Tschudi, 1838; Austrochaperina quotation marks until their monophyly can Fry, 1912; Barygenys Parker, 1936; Callu- be substantiated. Although Burton (1986) lops Boulenger, 1888; Choerophryne Kam- provided a number of morphological char- pen, 1914; Cophixalus Boettger, 1892; Cop- acters and a character matrix, no one so far iula MeÂhely, 1901; Genyophryne Boulenger, has analyzed these data phylogenetically. 1890; Hylophorbus Macleay, 1878; Liophry- ne Boulenger, 1897; Mantophryne Boulen- [118] SUBFAMILY: COPHYLINAE COPE, 1889 ger, 1897; Oreophryne Boettger, 1895; Ox- ydactyla Kampen, 1913; Pherohapsis Zwei- Cophylidae Cope, 1889: 248. Type genus: Cophy- fel, 1972; Sphenophryne Peters and Doria, la Boettger, 1880. Rhombophryninae Noble, 1931: 529. Type genus: 1878; Xenorhina Peters, 1863 (including Xe- Rhombophryne Boettger, 1880. nobatrachus Peters and Doria, 1878; see ap- pendix 7 for new combinations). IMMEDIATELY MORE INCLUSIVE TAXON: CHARACTERIZATION AND DIAGNOSIS: None [116] unnamed taxon. of the morphological characters in our anal- SISTER TAXON: [117] An unnamed taxon in ysis apply to this taxon because as direct de- our analysis composed of our exemplars Ho- velopers they were not part of the tadpole plophryne Barbour and Loveridge, 1928 study by Haas (2003). Among microhylids, (Melanobatrachinae Noble, 1931) and Ra- only Asterophryinae and Myersiella (Micro- manella Rao and Ramanna, 1925 (formerly hylinae; Izecksohn et al., 1971; Zweifel, of ``Microhylinae''). Together these are the 1972; Thibaudeau and Altig, 1999) exhibit sister taxon of [121] Gastrophryninae Fitzin- direct development, the development taking ger, 1843. 228 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

RANGE: Madagascar. phryne (see appendix 7 for the species name CONTENT: Anodonthyla MuÈller, 1892; Co- changes that this causes). Andreone et al. phyla Boettger, 1880; Madecassophryne Gui- (2004 ``2005'') hesitated to take this step be- beÂ, 1974; Platypelis Boulenger, 1882; Pleth- cause they did not feel there was suf®cient odontohyla Boulenger, 1882 (see Systematic statistical support for their maximum-likeli- Comments); Rhombophryne Boettger, 1880 hood conclusion. They did, however, note (see Systematic Comments and appendix 7); that their parsimony tree arrived at the same Stumpf®a Boettger, 1881. conclusion. We therefore think that it is bet- CHARACTERIZATION AND DIAGNOSIS: None ter to recognize two clades that might be of the morphological characters in our anal- found to be each other's closest relatives ysis optimizes on this branch; because our when more data are added to the analysis, morphological characters were largely de- than to retain a taxon, ``Plethodontohyla'' rived from larvae, and cophylines (as tradi- (sensu lato) for which the preponderance of tionally de®ned) are endotrophic and nidic- data does not support its monophyly. There olous. Nevertheless, endotrophy is a syna- are a number of species, nominally in Pleth- pomorphy at this level. Also, cophylines odontohyla, but not treated by Andreone et have unfused sphenethmoids, which appear al. (2004 ``2005''). We retain those in Pleth- as paired elements (Parker, 1934), otherwise odontohyla, although some of may be found found convergently in Dyscophus (Dysco- to be members of Rhombophryne. phinae) and Calluella (Microhylinae). (See appendix 5 for molecular synapomorphies on SUBFAMILY: DYSCOPHINAE BOULENGER, 1882 this branch [118].) Dyscophidae Boulenger, 1882: 179. Type genus: SYSTEMATIC COMMENTS: The association by Dyscophus Grandidier, 1872. our molecular data of Cophylinae (Madagas- IMMEDIATELY MORE INCLUSIVE TAXON: car) with our exemplars Hoplophryne (East [134] unnamed taxon. Africa) and Ramanella (India) is suggestive. SISTER TAXON: [135] Asterophryinae GuÈn- Madagascar±India is a repeated pattern in ther, 1858. biogeography, as is an apparently later con- RANGE: Madagascar. nection of India±Africa (e.g., Chiromantis in CONTENT: Dyscophus Grandidier, 1872. ϩ Africa Chirixalus in Asia [Rhacophori- CHARACTERIZATION AND DIAGNOSIS: Haas ϩ dae]; Petropedetes Arthroleptides in Af- (2003) suggested the following larval char- rica and Indirana in India [Petropedetidae]). acters that are presumed synapomorphies of The association of Gastrophryninae with this the taxon: (1) ramus mandibularis (cranial overall clade also speaks to a standard bio- nerve V3) runs through the m. levator man- geographic pattern, that of South America± dibulae externus group (Haas 65.1); and (2) Madagascar. free basihyal absent (Haas 105.0). Andreone et al. (2004 ``2005'') provided SYSTEMATIC COMMENT: Our data reject the considerable DNA sequence evidence that association of Calluella with Dyscophinae Plethodontohyla is polyphyletic (not para- (Blommers-SchloÈsser, 1976), which instead phyletic as suggested in the original publi- place Calluella deeply within Microhylinae. cation; see ®g. 33). As noted by Andreone et This is not surprising, inasmuch as the only al. (2004 ``2005'') the name Plethodontohyla characteristics suggested to ally Calluella Boulenger, 1882 (type species: Callula no- with Dyscophinae are apparent plesiomor- tosticta Gunther, 1877) adheres to his Pleth- phies (e.g., presence of teeth, diplasiocoelous odontohyla Group 1. Their second group of vertebral column, large vomer). The molec- ``Plethodontohyla'' falls into a monophyletic ular synapomorphies supporting a relation- group with Rhombophryne testudo. Rhom- ship of this taxon to Asterophryinae (branch bophryne Boettger, 1880, is substantially old- 134, appendix 5) is novel. er than the next older name for this taxon, Mantiphrys Mocquard, 1901 (type species: [121] SUBFAMILY: GASTROPHRYNINAE Mantiphrys laevipes Mocquard, 1895), and FITZINGER, 1843 to provide a monophyletic taxonomy, this in- Gastrophrynae Fitzinger, 1843: 33. Type genus: clusive taxon should be known as Rhombo- Gastrophryne Fitzinger, 1843. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 229

IMMEDIATELY MORE INCLUSIVE TAXON: SUBFAMILY: MELANOBATRACHINAE NOBLE, [115] unnamed taxon. 1931 SISTER TAXON: [116] unnamed taxon. Melanobatrachinae Noble, 1931: 538. Type ge- RANGE: Southern United States south to nus: Melanobatrachus Beddome, 1878. Argentina. Hoplophryninae Noble, 1931: 538±539. Type ge- CONTENT: Ctenophryne Mocquard, 1904; nus: Hoplophryne Barbour and Loveridge, Dasypops Miranda-Ribeiro, 1924; Derma- 1928. tonotus MeÂhely, 1904; Elachistocleis Parker, IMMEDIATELY MORE INCLUSIVE TAXON: 1927; Gastrophryne Fitzinger, 1843; Hamp- [117] unnamed taxon. tophryne Carvalho, 1954; Nelsonophryne SISTER TAXON: Ramanella Rao and Ra- Frost, 1987. manna, 1925. CHARACTERIZATION AND DIAGNOSIS: Opti- RANGE: Montane and southern mization is problematic because none of the India. direct-developing microhylids were sampled CONTENT: Hoplophryne Barbour and Lov- in our morphological data set. Nevertheless eridge, 1928; Melanobatrachus Beddome, the following are candidates for being syna- 1878; Parhoplophryne Barbour and Lover- pomorphies of Gastrophryninae, although idge, 1928. they could be synapomorphies of Gastro- CHARACTERIZATION AND DIAGNOSIS: Melan- phryninae ϩ Cophylinae or some subset of obatrachinae shares two synapomorphies Gastrophryninae inasmuch as the exemplars (Parker, 1934): (1) middle and outer ear ab- on which this supposition is built are Gas- sent; (2) parasphenoid and sphenethmoid trophryne carolinensis, Hamptophryne boli- fused. viana, and ): (1) m. le- SYSTEMATIC COMMENTS: Although we pro- vator arcuum branchialium III split into two visionally retain Melanobatrachinae as an un- crossing bundles (Haas 41.1); (2) origin of tested taxon, the placement of Hoplophryne m. suspensoriohyoideus from otic capsule (our exemplar) in the general tree (see ®gs. (Haas 46.2); (3) posterolateral projections of 50 and 61) suggests that a more densely sam- the crista parotica processus otobranchialis pled analysis will provide results that render (Haas 67.2); (4) processus muscularis absent a Melanobatrachinae containing several more (Haas 79.0); (5) anterolateral base of proces- genera (such as Ramanella) than as currently sus muscularis bearing ventrolateral process composed. Hoplophryne and Parhoplophry- ne were placed in Hoplophrynine by Noble (Haas 80.1); and (6) ligamentum mandibu- (1931) on the basis of sharing the apomorphy losuprarostrale absent (Haas 127.0). of a greatly reduced ®rst ®nger. (Noble also Molecular evidence (branch 121, appendix allied these genera with Brevicipitidae on the 5) is strong that the New World microhylids basis of retaining a complete clavicle, but (with the exception of Synapturanus, and this alliance is not supported by our data.) possibly several others for which we had no Parker (1934) placed Hoplophryninae in the tissues) form a clade that is most closely re- synonymy of Melanobatrachinae (India) be- lated to the Madagascan Cophylinae. cause they share the absence of the auditory SYSTEMATIC COMMENTS: The exclusion of apparatus and fusion of the parasphenoid to Synapturanus from this taxon comes as the sphenenthmoid. We could not sample something of a surprise, inasmuch as both Melanobatrachus, but it remains possible Zweifel (1986) and Wild (1995) provided ev- that it is the sister taxon of Hoplophryninae idence for its placement within a New World and that Hoplophryne and Parhoplophryne clade. Nevertheless, neither Zweifel (1986) are African outliers of a predominantly In- nor Wild (1995) presented morphological ev- dian group. This is conjecture, however, and idence for the monophyly of the New World only more data and denser sampling will re- microhylids (of which our Gastrophryninae solve the issue. is a part). We expect that further research will show the New World microhylids to be a [130] SUBFAMILY: MICROHYLINAE GUÈ NTHER, composite of gastrophrynines, some basal 1858 (1843) taxa (e.g., Synapturanus), and possibly some Hylaedactyli Fitzinger, 1843: 33. Type genus: Hy- with relations in Asia. laedactylus DumeÂril anbd Bibron, 1841. 230 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Micrhylidae GuÈnther, 1858b: 346. Type genus: these are highly contingent on topological Micrhyla DumeÂril and Bibron, 1841 (an incor- position of Scaphiophryne: (1) keratinized rect subsequent spelling of Microhyla Tschudi, jaw sheaths present (Haas 6.1; reversal from 1838). the microhylid condition); (2) eye position Kaloulinae Noble, 1931: 538. Type genus: Kal- dorsolateral (Haas 11.0; reversal from the oula Gray, 1831. microhylid condition); (3) spiracle position IMMEDIATELY MORE INCLUSIVE TAXON: sinistral (Haas 18.1; reversal from the micro- [129] unnamed taxon. hylid condition); (4) m. interhyoideus pos- SISTER TAXON: Scaphiophryninae Laurent, terior absent (Haas 23.0; reversal from the 1946. phthanobatrachian condition); (5) m. subar- RANGE: India, China, Japan, and Korea to cualis rectus II±IV represented by a single the Philippines and Greater Sunda Islands. ¯at tract of ®bers (Haas 30.0; reversal from CONTENT: Calluella Stoliczka, 1872; the microhylid condition); (6) insertion of m. Chaperina Mocquard, 1892; Kaloula Gray, rectus cervicis on proximal ceratobranchialia 1831; Microhyla Tschudi, 1838. III and IV (Haas 39.2; reversal from micro- CHARACTERIZATION AND DIAGNOSIS: Haas hylid condition); (7) m. interhyoideus and m. (2003) examined only Kaloula within this intermandibularis well separated by a gap clade, so this is our only morphological ex- (Haas 47.1; reversal from the microhylid emplar for this subfamily, but the following condition); (8) m. mandibulolabialis inserting are candidates for being synapomorphies of in soft tissue of lip (Haas 49.0; reversal from the Microhylinae: (1) vena caudalis dorsalis microhylid condition); (9) m. levator man- absent (Haas 14.0); (2) origin of m. suspen- dibulae internus low (Haas 58.1; reversal soriohyoideus from otic capsule (Haas 46.2); from microhylid condition); (10) m. levator and (3) posterolateral projections of the crista mandibulae longus originates from posterior parotica expansive ¯at chondri®cations palatoquadrate (Haas 60.1; reversal from mi- (Haas 67.2). Nevertheless, the molecular ev- crohylid condition); (11) ramus mandibularis idence is decisive for the recognition of this (cranial nerve V3) anterior (dorsal) to the m. taxon (see appendix 5). levator mandibulae longus (Haas 64.2); (12) COMMENT: See Microhylidae account for processus suboticus quadrati absent (Haas comment on East Asian ``microhylines'' ex- 76.0; reversal from microhylid condition); cluded from this taxon because of lack of (13) arcus subocularis with irregular margin evidence to place them. (Haas 81.1; reversal of microhylid condi- tion); (14) cartilaginous roo®ng of the cavum SUBFAMILY: SCAPHIOPHRYNINAE LAURENT, cranii absent (Haas 96.0; reversal of predom- 1946 inant microhylid condition); and (15) glottis position posterior (Haas 130.0; reversal of Scaphiophryninae Laurent, 1946: 337. Type ge- microhylid condition). nus: Scaphiophryne Boulenger, 1882. Pseudohemisiinae Tamarunov, 1964a: 132. Type SYSTEMATIC COMMENTS: Ford and Canna- genus: Pseudohemisus Mocquard, 1895. tella (1993: 94±117), found no evidence for the monophyly of this taxon. Haas (2003: 50) IMMEDIATELY MORE INCLUSIVE TAXON: suggested on the basis of tadpole morphol- [129] unnamed taxon. ogy that Paradoxophyla is more closely re- SISTER TAXON: [130] Microhylinae GuÈn- lated to Phrynomantis than to the remaining ther, 1858 (1843). Scaphiophryninae, rendering the latter non- RANGE: Madagascar. monophyletic. On that basis alone, because CONTENT: Scaphiophryne Boulenger, we did not have tissues of Paradoxophyla, 1882. we transfer Paradoxophyla from Scaphio- CHARACTERIZATION AND DIAGNOSIS:Inour phryninae to incertae sedis under Microhy- topology Scaphiophryne is deeply imbedded lidae. The association (branch 129, appendix within Microhylidae, requiring a remarkable 5) of Madagascan Scaphiophryninae with number of reversals. Nevertheless, we sug- Microhylinae may suggest an Indian origin gest these reversals are likely synapomor- of Microhylinae. (See Systematic Comment phies of the taxon, while noting that most of under Cophylinae.) 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 231

[143] AFROBATRACHIA NEW TAXON which at this position in the general clado- gram is a synapomorphy. Breviceps and ETYMOLOGY: Afro- (Latin: of Africa) ϩ Hemisus also share a single median thyroid batrachos (Greek: frog), in reference to the gland (Blommers-SchloÈsser, 1993), so we predominantly African range of this taxon. presume that this, too, is a synapomorphy IMMEDIATELY MORE INCLUSIVE TAXON: joining the two taxa. Breviceps and Hemisus [109] Allodapanura new taxon. also exhibit nasal plugs (De Villiers, 1931) SISTER TAXON: [110] Microhylidae GuÈn- which may be homologous. (See also ``Char- ther, 1858 (1843). acterization and Diagnosis'' under Hemiso- RANGE: Sub-Saharan Africa, Madagascar, tidae for other characters that may optimize and the Seychelles. on this taxon.) Molecular synapomorphies CONCEPT AND CONTENT: Afrobatrachia is a are provided in appendix 5. monophyletic taxon composed of [144] Xe- nosyneunitanura new taxon and [148] Lau- [145] FAMILY: BREVICIPITIDAE BONAPARTE, rentobatrachia new taxon. 1850 CHARACTERIZATION AND DIAGNOSIS: Likely candidates for being synapomorphies are the Brevicipitina Bonaparte, 1850: 1 p. Type genus: larval characters: (1) m. transversus ventralis Breviceps Merrem, 1820. IV present (Haas 22.1); (2) posterolateral Engystomidae Bonaparte, 1850: 1 p. Type genus: projections of the crista parotica forming Engystoma Fitzinger, 1826. processus otobranchialis (Haas 67.3); (3) IMMEDIATELY MORE INCLUSIVE TAXON: processus ascendens thin (Haas 72.1); (4) [144] Xenosyneunitanura new taxon. dorsal connection from processus muscularis SISTER TAXON: Hemisotidae Cope, 1867. to ``high'' commissura quadrato-orbitalis RANGE: Sub-Saharan East Africa and (Haas 78.3); and (5) anterolateral base of southern Africa, from Ethiopia south to An- processus muscularis bearing ventrolateral gola and South Africa. process (Haas 80.1). See characterisation of CONTENT: Balebreviceps Largen and Allodapanura for additional discussion of Drewes, 1989; Breviceps Merrem, 1820; possible synapomorphies. Callulina Nieden, 1911 ``1910''; Probrevi- COMMENT: Our Afrobatrachia is identical ceps Parker, 1931; Spelaeophryne Ahl, 1924. in content to the enlarged Brevicipitidae of CHARACTERIZATION AND DIAGNOSIS: Parker Dubois (2005). (1934) noted that brevicipitids lack ossi®ed sphenethmoids, which is clearly a synapo- [144] XENOSYNEUNITANURA NEW TAXON morphy at this level. In addition, the loss of ETYMOLOGY: Xeno- (Greek: strange) ϩ the pterygoid, palatoquadrate, and m. oper- syneunitos (Greek: bed sharer) ϩ anoura cularis (De Villiers, 1931) are likely syna- (Greek: frog). In other words, the name pomorphies for this group. The extremely means ``strange bedfellows'' inasmuch as short head and direct development exhibited Hemisotidae and Brevicipitidae, although by this taxon (Parker, 1934) are also syna- cladistic nearest relatives, are dissimilar ani- pomorphies. mals. SYSTEMATIC COMMENT: Loader et al. (2004) IMMEDIATELY MORE INCLUSIVE TAXON: suggested a phylogeny of Breviceps (Spe- [143] Afrobatrachia new taxon. laeophryne ϩ (Callulina ϩ Probreviceps)); SISTER TAXON: [148] Laurentobatrachia they, like us, did not include Balebreviceps new taxon. in their analysis. On the basis of our larger RANGE: Sub-Ssaharan Africa. amount of evidence but less dense sampling, CONCEPT AND CONTENT: Xenosyneunitanu- we placed Probreviceps nearer to Breviceps ra new taxon is a monophyletic taxon con- in our tree. Nevertheless, both arrangements taining Hemisotidae Cope, 1867, and [145] con¯ict with the character of fusion of the Brevicipitidae Bonaparte, 1850. urostyle and sacrum found in Probreviceps CHARACTERIZATION AND DIAGNOSIS: Hemi- and Breviceps but not in Spelaeophryne and sotidae and Brevicipitidae share the absence Callulina (Parker, 1934), suggesting that ad- of the palatine bones (De Villiers, 1931), ditional testing is warranted. 232 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

FAMILY: HEMISOTIDAE COPE, 1867 Hyperoliidae Laurent, 1943, and [164] Ar- Hemisidae Cope, 1867: 198. Type genus: Hemisus throleptidae Mivart, 1869. GuÈnther, 1859 ``1858''. Emended to Hemisotina CHARACTERIZATION AND DIAGNOSIS: The by GuÈnther, 1870: 119. characters (from Haas, 2003) 54.1 (larval m. levator manidbulae externus in two portion), IMMEDIATELY MORE INCLUSIVE TAXON: 111.0 (commissura proximalis III absent), [144] Xenosyneunitanura new taxon. and 151.0 (intercalary elements absent) are SISTER TAXON: [145] Brevicipitidae Bona- likely synapomorphies of this group, al- parte, 1850. though because of the low density of taxon RANGE: Sub-Saharan Africa. sampling this requires additional specimen CONTENT: Hemisus GuÈnther, 1859 examination. In addition, claw-shaped ter- ``1858''. minal phalanges appear to optimize on this CHARACTERIZATION AND DIAGNOSIS: All of branch, appearing convergently in Ptychad- the characters in our analysis (from Haas, ena and several of the hyloids (Liem, 1970), 2003) that optimize on Hemisus (our only although the distribution of this character is morphological exemplar in this clade) may complicated, and further work may show that be synapomorphies of this clade, the Hemi- this optimization is mistaken. Drewes (1984) sotidae, or some subset of Hemisus: (1) dou- suggested that thyrohyals borne on cartilag- ble-layered dermis in larvae (Haas 13.1); (2) inous stalks (his character 10.1) might be a posterior dorsal process of pars alaris ex- synapomorphy, although this is optimization- panded terminally, almost rectangular in lat- dependent inasmuch as this character is not eral view (Haas 89.1); (3) larvae are guided in Leptopelis (Laurent, 1978). The external by the female from the nest to pond (Haas metatarsal tubercle is absent or poorly de- 137.1); and (4) amplexus absent (Haas veloped throughout Laurentobatrachia (Lau- 139.0). Some of these may be synapomor- rent, 1986), but the exact distribution of this phies at the level of Xenosyneunitanura in- requires veri®cation. Molecular synapomor- asmuch as Brevicipitidae was not studied by phies for this taxon are summarized in ap- Haas (2003) because they lack exotrophic pendix 5. larvae, which were the focus of Haas' study. SYSTEMATIC COMMENT: Vences and Glaw Hemisus lacks vomers, middle ear, and (2001) and Van der Meijden et al. (2005) rec- ductus lacrimosus, and exhibits fusion of ver- ognized this taxon as the epifamily Arthro- tebrae 8 and 9 (De Villiers, 1931). Further, leptoidae, and originally Laurent (1951) con- Hemisus burrows head-®rst (Channing, sidered this clade (with the possible inclusion 1995). All of these characters can safely be considered synapomorphies of Hemisotidae. of Scaphiophryninae) to be a single family, and Dubois (2005) considered our Lauren- tobatrachia to be 4 of the 6 subfamilies of [148] LAURENTOBATRACHIA NEW TAXON his Brevicipitidae. We attempted to retain fa- ETYMOLOGY: R.L. Laurent ϩ batrachia miliar usage, with the exception of moving (Greek: batrachos, frog). This name cele- Leptopelinae from Hyperoliidae to Arthro- brates the enormous contributions to amphib- leptidae. Because we think that the diversity ian systematics by the father of central Af- of this taxon has been greatly underestimat- rican herpetology and a prominent ®gure in ed, our approach leaves considerable room Argentinian herpetology, Raymond L. Lau- for more informative taxonomies as evidence rent. becomes available. IMMEDIATELY MORE INCLUSIVE TAXON: [143] Afrobatrachia new taxon. [149] FAMILY: HYPEROLIIDAE LAURENT, 1943 SISTER TAXON: [144] Xenosyneunitanura Hyperoliinae Laurent, 1943: 16. Type genus: Hy- new taxon. perolius Rapp, 1842. RANGE: Sub-Saharan Africa, Madagascar, Kassinini Laurent, 1972: 201. Type genus: Kas- and the Seychelles. sina Girard, 1853. CONTENT AND CONCEPT: Laurentobatrachia Tachycneminae Channing, 1989: 127. Type ge- is a monophyletic group composed of [149] nus: Tachycnemis Fitzinger, 1843. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 233

IMMEDIATELY MORE INCLUSIVE TAXON: taxa. We did not sample Chrysobatrachus or [143] Afrobatrachia new taxon. Callixalus and cannot guess into which SISTER TAXON: [164] Arthroleptidae Mi- group they would fall. Their association with vart, 1869. Acanthixalus in the tree of Drewes (1984) RANGE: Sub-Saharan Africa and Madagas- suggests that they might follow Acanthixalus car; Seychelles. into Kassininae, but this is merely conjecture CONTENT: Acanthixalus Laurent, 1944; and a combined study of morphology and Afrixalus Laurent, 1944; Alexteroon Perret, molecules is ongoing by Drewes and collab- 1988; Arlequinus Perret, 1988; Callixalus orators. Our results differ substantially from Laurent, 1950; Chlorolius Perret, 1988; the results of Vences et al. (2003d; ®gs. 28, Chrysobatrachus Laurent, 1951; Cryptothy- 29) with respect to the relative placement of lax Laurent and Combaz, 1950; Heterixalus several genera. This is presumably due to our Laurent, 1944; Hyperolius Rapp, 1842 (in- application of much denser sampling and cluding Nesionixalus Perret, 1976); Kassina more evidence. Girard, 1853; Kassinula Laurent, 1940; Op- The association by the molecular data of isthothylax Perret, 1966; Per- Tachycnemis (Seychelles) and Heterixalus acca, 1907; Phlyctimantis Laurent and Com- (Madagascar) is of some biogeographic in- baz, 1950; Semnodactylus Hoffman, 1939; terest. We expected Alexteroon to be imbed- Tachycnemis Fitzinger, 1843. ded within Hyperolius, but our sampling of CHARACTERIZATION AND DIAGNOSIS: One Hyperolius was insuf®cient to test this prop- larval character in our analysis that may be osition adequately. On the basis of our lim- synapomorphy of this group is (from Haas, ited exemplar selection, Alexteroon may be 2003): commissura proximalis II absent. Be- the sister taxon of Hyperolius (sensu lato). yond that, hyperoliids are unique among However, we found, as did Drewes and Wil- frogs in having a distinctive gular gland kinson (2004), that Nesionixalus is clearly (Drewes, 1984). Drewes (1984) summarized deeply imbedded in Hyperolius, but also rep- a character distribution suggesting that lack- resents a monophyletic group. We suggest ing sphincter control of the vocal slits may that Nesionixalus be treated as a subgenus of also be a synapomorphy of Hyperoliidae. Hyperolius with no coordinate taxon to im- The presence of intercalary phalangeal el- ply that the remaining species of Hyperolius ements per se is not a synapomorphy of this are a monophyletic group (see appendix 7 for group (or at least not without making as- new combinations). We expect that Chloro- sumptions of character optimization), being lius and Arlequinus will also be found to be found also in the Leptopelinae of Arthrolep- imbedded within Hyperolius, although at this tidae. Nevertheless, Drewes (1984) noted time no data can be brought to bear to test that hyperoliid and leptopeline intercalary el- this proposition. ements are histologically quite different from each other. The latter does not accept either [164] FAMILY: ARTHROLEPTIDAE MIVART, 1869 Alizarin or Alcian Blue stain, suggesting that these elements may not be homologous. Arthroleptina Mivart, 1869: 294. Type genus: Ar- throleptis Smith, 1849. SYSTEMATIC COMMENTS: The position in Astylosterninae Noble, 1927: 110. Type genus: our tree of Acanthixalus is heterodox com- Astylosternus Werner, 1898. pared with previous studies (e.g., Drewes, Leptopelini Laurent, 1972: 201. Type genus: Lep- 1984) and implies a number of reversals and topelis GuÈnther, 1859. New synonym. convergences in the morphology of hypero- liid frogs. We considered recognizing sub- IMMEDIATELY MORE INCLUSIVE TAXON: families within Hyperoliidae, corresponding [147] Laurentobatrachia new taxon. to the two major clades of exemplars, for SISTER TAXON: [149] Hyperoliidae Laurent, which the name Kassininae Laurent, 1972, is 1943. available for the Kassina±Phlyctimantis± RANGE: Sub-Saharan Africa. Acanthixalus clade and Hyperoliinae Lau- CONTENT: Arthroleptis Smith, 1849 (in- rent, 1943, for the remainder of our exemplar cluding Schoutedenella De Witte, 1921; see 234 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Systematic Comments); Astylosternus Wer- apomorphic should one be willing to make ner, 1898; Cardioglossa Boulenger, 1900; assumptions about character optimization Leptodactylodon Andersson, 1903; Lepto- and that the phalangeal elements of lepto- pelis GuÈnther, 1859; Nyctibates Boulenger, pelines and hyperoliids are homologous. 1904; Scotobleps Boulenger, 1900; Trichob- Arthroleptis renders Schoutedenella para- atrachus Boulenger, 1900. phyletic, and we therefore consider them to CHARACTERIZATION AND DIAGNOSIS: Arthro- be synonyms. Laurent and Fabrezi's (1986 leptids are small frogs exhibiting forked ``1985'') contention that Schoutedenella and omosterna that, with the exception of Arthro- Arthroleptis are not each other's closest rel- leptis, have a typically biphasic life history. atives is rejected, although the position of Like many of the taxa within Afrobatrachia, Poynton (1964a) and Poynton and Broadley many of the arthroleptids have vertical pu- (1967), that Schoutedenella are merely small pils, with the exceptions of Leptodactylodon Arthroleptis is also rejected. (Our tree sug- (quadrangular) and Arthroleptini (horizontal, gests that if size were characteristic, we except for Scotobleps). None of the morpho- would have to say that Arthroleptis are big logical characters in our analysis optimize Schoutedenella.) Our molecular data support unambiguously to this branch [164]. Regard- the notion that nominal Arthroleptis is im- less, the molecular data are decisive in sup- bedded within Schoutedenella and we place port of recognition of this group (see appen- them in synonymy. (See appendix 7 for new dix 5). and revived combinations.) Larval characters of Haas' (2003) exem- Perret (1966) suggested that Nyctibates is plar LeptopelisÐa distinct medial ossi®ca- a synonym of Astylosternus, but Amiet (1971 tion center of vertebral centra ventral to no- ``1970'', 1973 ``1972'') resurrected Nycti- tochord present (Haas 100.1)Ðmay be syn- bates on the basis of tadpole morphology be- apomorphies of Arthroleptidae, of Leptope- ing more similar to Leptodactylodon and Tri- linae, or of some subset of Leptopelis. The chobatrachus. Our molecular data support direct development of Arthroleptis is subse- recognition of Nyctibates. quently derived. SYSTEMATIC COMMENTS: We recognize two [180] NATATANURA NEW TAXON subfamilies within Arthroleptidae, [165] Leptopelinae Laurent, 1972, for Leptopelis, ETYMOLOGY: Natata- (Greek: swim) ϩ an- formerly associated with Hyperoliidae (al- oura (Greek: tailless, i.e., frog), referencing though shown to be phylogenetically distant that many of the frogs in this clade are semi- from them by Vences et al., 2003c), and aquatic. [168] Arthroleptinae Mivart, 1869, contain- IMMEDIATELY MORE INCLUSIVE TAXON: ing two tribes, [169] Astylosternini Noble, [108] Ranoides new taxon. 1931 (Leptodactylodon, Nyctibates, Trichob- SISTER TAXON: [109] Allodapanura new atrachus, and Leptodactylodon) and [172] taxon. Arthroleptini Mivart, 1869 (Arthroleptis [in- RANGE: Worldwide temperate and tropical cluding Schoutedenella], Cardioglossa, and habitats on all continents and major islands, Scotobleps). Scotobleps formerly was asso- except most of Australia and New Zealand. ciated with Astylosterninae, so its transfer to CONCEPT AND CONTENT: Natatanura is a Arthroleptini is something of a surprise (on monophyletic group composed of [181] Pty- the basis of evidence shown in appendix 5). chadenidae Dubois, 1987 ``1986'', and [183] [165] Leptopelinae Laurent, 1972, is dis- Victoranura new taxon. tinguished morphologically from its near CHARACTERIZATION AND DIAGNOSIS: Nata- neighbors by the possession of an entire, tanura is identical to the epifamily Ranoidae rather than forked, omosternum and by his- of Dubois (1992) and Ranidae (sensu lato) of tologically distinct intercalary phalangeal el- Laurent (1986). Characters in our analysis ements (Drewes, 1984). (from Haas, 2003) that are likely synapo- [168] Arthroleptinae Mivart, 1869, is not morphies of this taxon are (1) anterior inser- diagnosable via morphology, although the tion of m. subarcualis rectus II±IV on cera- absence of intercalary elements may be syn- tobranchial III (Haas 37.2); (2) commissura 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 235 proximalis II absent (Haas 110.0); and (3) Lanzarana Clarke, 1982; Ptychadena Bou- commissura proximalis III absent (Haas lenger, 1917. 111.0). CHARACTERIZATION AND DIAGNOSIS:Inour J.D. Lynch (1973) and Laurent (1986) analysis, the morphological (larval) charac- suggested that an ossi®ed metasternal style is ters that attach to the only exemplar of this a synapomorphy at this level of universality, taxon, Ptychadena, are (1) m. subarcualis but this requires corroboration inasmuch as rectus I portion with origin from ceratobran- several groups within Natatanura have carti- chial III absent (Haas 35.0); (2) partes cor- laginous metasterna (Laurent, 1986). pores medially separate (Haas 87.0); and (3) SYSTEMATIC COMMENT: Burton (1998a) eggs ¯oat as a surface ®lm (Haas 141.2). Be- noted that several genera of Natatanura share cause of our limited sampling for morphol- the presence of an extra slip of the m. ¯exor ogy, it is possible that these characters do not teres digiti IV, which is ventral to the m. apply to Hildebrandtia or Lanazarana;itis transversus metacarpus II: Altirana, Aubria, also possible that they apply only to a subset Ceratobatrachus, Conraua, Hildebrandtia, of Ptychadena. Only denser sampling will Mantella, Mantidactylus, Petropedetes, Pty- tell. chadena, Pyxicephalus, and Rana, but not in Other features that are likely synapomor- Batrachylodes, Cacosternum, Discodeles, phies, although originally suggested in a Laliostoma, Meristogenys, Micrixalus, Nan- somewhat different outgroup structure nophrys, Nanorana, Natalobatrachus, Nyc- (Clarke, 1981), are (1) otic plate absent or tibatrachus, Occidozyga, Palmatorappia, rudimentary; (2) (neo)palatines absent; (3) Platymantis,orStrongylopus (with many point overlap of the medial ramus of the pter- taxa not examined). If this character is opti- ygyoid and the anterior lateral border of the mized on our most parsimonious tree, the im- parasphenoid ala in an anterior±posterior plication is that this character arose at least plane; (4) clavicles reduced and well-sepa- six times, of which the following is one of rated at midline; (5) sternal style a short several equally parsimonious arrangements: compact bony element; (6) eight presacral (1) Ceratobatrachus; (2) in the branch sub- and sacral vertebrae fused (also in some Lith- tending Conraua ϩ Petropedetes, and there- obates); and (7) dorsal protuberance on ilium fore likely to be in Indirana and Arthrolep- not or only slightly differentiated from dorsal tides); (3) Ptychadenidae (Hildebrantia, Pty- prominence, which is smooth surfaced and chadena, and presumably in Lanzarana); (4) con¯uent with a well-developed ilial crest. Pyxicephalini (Pyxicephalus and Aubria); (5) SYSTEMATIC COMMENT: See Systematic Altirana (ϭ part of Nanorana); (6) Aglaioan- Comments under Natatanura. Our association ura (Rhacophoroidea ϩ Ranidae). Neverthe- of Hildebrandtia and Lanzarana with this less, considerably more specimens of more taxon rests on the morphological data anal- taxa need to be examined before the opti- ysis of Clarke (1981), who suggested a num- mization of this feature can con®dently be ber of synapomorphies for the group (see considered settled. above).

[181] FAMILY: PTYCHADENIDAE DUBOIS, 1987 [183] VICTORANURA NEW TAXON ``1986'' ϩ Ptychadenini Dubois, 1987 ``1985'': 55. Type ge- ETYMOLOGY: Victor (Latin: conqueror) nus: Ptychadena Boulenger, 1917. anoura (Greek: tailless; i.e., frog), alluding to the remarkable success of this taxon world- IMMEDIATELY MORE INCLUSIVE TAXON: wide. [180] Natatanura new taxon. IMMEDIATELY MORE INCLUSIVE TAXON: SISTER TAXON: [183] Victoranura new tax- [180] Natatanura new taxon. on. SISTER TAXON: [181] Ptychadenidae Du- RANGE: Sub-Saharan tropical and subtrop- bois, 1987 ``1986''. ical Africa; Seychelles and Madagascar. RANGE: Worldwide continents and major CONTENT: Hildebrandtia Nieden, 1907; islands in temperate and tropical regions, ex- 236 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 cept southern Australia, the Seychelles, and lossini Anderson, 1871) is rejected by our New Zealand. evidence. CONCEPT AND CONTENT: Victoranura is a Dubois (1992) placed Batrachylodes out- monophyletic group composed of [184] Cer- side of his Ceratobatrachini, because, unlike atobatrachidae Boulenger, 1884, and [189] the more typical members of Ceratobatrachi- Telmatobatrachia new taxon. nae, it lacks a forked omosternum. Neverthe- CHARACTERIZATION AND DIAGNOSIS: None less, Batrachylodes does have endotrophic of the morphological characters in our anal- larvae (Thibaudeau and Altig, 1999), and our ysis diagnose on this taxon, although the mo- molecular evidence places Batrachylodes lecular data are decisive (see appendix 5 for ®rmly within the ceratobatrachine clade. summary of molecular synapomorphies). Roelants et al. (2004) provided molecular evidence suggesting that Ingerana is in Oc- [184] FAMILY: CERATOBATRACHIDAE cidozyginae rather than Ceratobatrachinae, BOULENGER, 1884 but this is not corroborated by our denser Ceratobatrachidae Boulenger, 1884: 212. Type ge- taxonomic sampling and larger amount of nus: Ceratobatrachus Boulenger, 1884. data, which place Ingerana in the more con- Platymantinae Savage, 1973: 354. Type genus: ventional location in Ceratobatrachidae and Platymantis GuÈnther, 1859. as the sister taxon of the remaining genera within Ceratobatrachinae. Like Roelants et IMMEDIATELY MORE INCLUSIVE TAXON: al. (2004), we did not evaluate species of the [183] Victoranura new taxon. nominal subgenus Liurana, a taxon that Du- SISTER TAXON: [189] Telmatobatrachia new bois (1987 ``1985'') erected as a subgenus of taxon. Ingerana, but subsequently was recognized RANGE: Western China (Xizang and Yun- by some workers as a genus (Fei et al., 1997) nan); , adjacent Thailand and pen- and later (Dubois, 2005, without discussion) insular Malaysia; Philippines, Borneo; New as a synonym of Taylorana (ϭ Limnonectes). Guinea; Admiralty, Bismarck, and Solomon Liurana is reported to be differentiated from Islands; Fiji; Palau. Ingerana by condition of the ®nger disc (ab- CONTENT: Batrachylodes Boulenger, 1887; sent in Liurana, present in Ingerana) and Ceratobatrachus Boulenger, 1884; Discode- median lingual papilla (present in Liurana, les Boulenger, 1918; Ingerana Dubois, 1987 absent in Ingerana; Dubois, 1987 ``1985''), ``1986''; Palmatorappia Ahl, 1927 ``1926''; but some species of Liurana possess small Platymantis GuÈnther, 1858. ®nger discs (Zhao and Li, 1984; Fei et al., CHARACTERIZATION AND DIAGNOSIS: None 2005), and the condition of the tongue is of the morphological characters in our anal- known for only two of the ®ve species of ysis optimize as synapomorphies of this tax- Ingerana (Smith, 1930; Inger, 1954, 1966). on, although all ceratobatrachids are charac- We treat Liurana as a synonym of Ingerana, terized by large eggs and direct development pending evidence being published to sub- (Noble, 1931). Many of the species have ex- stantiate Dubois' (2005) assertion of its panded toe tips, but this is likely plesiom- placement in Limnonectini (Dicroglossidae). orphic at this level of universality. Molecular synapomorphies for the clade are summa- [189] TELMATOBATRACHIA NEW TAXON rized in appendix 5. SYSTEMATIC COMMENT: Dubois (1987 ETYMOLOGY: Telmato- (Greek: of a marsh) ``1985'', 1992) placed his Ceratobatrachiini ϩ batrachos (Greek: frog), referencing the Boulenger, 1884, within a larger Dicroglos- preference of these frogs for wet microhab- sinae Anderson, 1871. The subsequent im- itats. plication of Dubois et al. (2001) that Cera- IMMEDIATELY MORE INCLUSIVE TAXON: tobatrachidae (his Ceratobatrachinae) is of [183] Victoranura new taxon. uncertain relationship to Dicroglossinae was SISTER TAXON: [184] Ceratobatrachidae justi®ed inasmuch as an inclusive Dicroglos- Boulenger, 1884. sinae (including Ceratobatrachiini Boulenger, RANGE: Worldwide continents and major 1884, Conrauini Dubois, 1992, and Dicrog- islands in temperate and tropical environ- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 237 ments, except for southern South America, cluding Madagascar, New Zealand, Sey- Madagascar, New Zealand, and most of Aus- chelles, and Australia except for the far tralia. north. CONCEPT AND CONTENT: Telmatobatrachia CONCEPT AND CONTENT: Ametrobatrachia is is a monophyletic taxon composed of [190] a monophyletic taxon composed of [192] Af- Micrixalidae Dubois, Ohler, and Biju, 2001, ricanura new taxon and [220] Saukrobatra- and [191] Ametrobatrachia new taxon. chia new taxon. CHARACTERIZATION AND DIAGNOSIS: None CHARACTERIZATION AND DIAGNOSIS: None of the morphological characters in our anal- of the morphological characters in our anal- ysis optimize on the branch subtending this ysis optimize as synapmorphies of this taxon. taxon although our molecular data decisively Nevertheless, the molecular data are deci- support its recognition. (See appendix 5 for sive. (See appendix 5 for summary of mo- listing of molecular synapomorphies.) lecular synapomorphies for this taxon.)

[190] FAMILY: MICRIXALIDAE DUBOIS, OHLER, [192] AFRICANURA NEW TAXON AND BIJU, 2001 ϩ Micrixalinae Dubois et al., 2001: 54. Type genus: ETYMOLOGY: Afric- (Latin: of Africa) Micrixalus Boulenger, 1888. anoura (Greek: tailless, i.e., frog). IMMEDIATELY MORE INCLUSIVE TAXON: IMMEDIATELY MORE INCLUSIVE TAXON: [191] Ametrobatrachia new taxon. [189] Telmatobatrachia new taxon. SISTER TAXON: [220] Saukrobatrachia new SISTER TAXON: [191] Ametrobatrachia new taxon. taxon. RANGE: Sub-Saharan Africa. RANGE: India. CONTENT: [193] Phrynobatrachidae Lau- CONTENT: Micrixalus Boulenger, 1888. rent, 1941 ``1940'', and [200] Pyxicephalo- CHARACTERIZATION AND DIAGNOSIS: None idea Bonaparte, 1850. of the morphological characters in our anal- CHARACTERIZATION AND DIAGNOSIS: None ysis optimize on this taxon and the decisive of the morphological characters in our anal- evidence for its recognition is entirely mo- ysis optimize on this taxon. Nevertheless, lecular (see appendix 5 for summary). Unlike molecular data are decisive. (See appendix 5 Ptychadenidae, Ceratobatrachidae, and basal- for summary of molecular transformation as- ly in Ametrobatrachia, the omosternum is un- sociated with this taxon.) forked in Micrixalidae (Dubois et al., 2001), SYSTEMATIC COMMENT: The existence of which at this level of universality is a syna- this taxon had not been suspected prior to the pomorphy of the group as is the low kera- publication of Van der Meijden et al. (2005), todont formula 1/0 (Dubois et al., 2001). The although it certainly meets biogeographic ex- presence of digital discs in Micrixalinae is pectations. likely a plesiomorphy at this level of univer- sality. [193] FAMILY: PHRYNOBATRACHIDAE LAURENT, 1941 ``1940'' [191] AMETROBATRACHIA NEW TAXON Hemimantidae Hoffmann, 1878: 613. Type genus: ETYMOLOGY: Ametros (Greek: beyond ϩ Hemimantis Peters, 1863. measure) batrachos (Greek: frog), denot- Phrynobatrachinae Laurent, 1941 ``1940'': 79. ing the enormity of this taxon in terms of Type species: Phrynobatrachus GuÈnther, 1862. species and with respect to the enormous numbers of questions that remain about its IMMEDIATELY MORE INCLUSIVE TAXON: internal phylogenetic structure. [192] Africanura new taxon. IMMEDIATELY MORE INCLUSIVE TAXON: SISTER TAXON: [200] Pyxicephaloidea Bon- [189] Telmatobatrachia new taxon. aparte, 1850. SISTER TAXON: [190] Micrixalidae Dubois, RANGE: Sub-Saharan Africa. Ohler, and Biju, 2001. CONTENT: Ericabatrachus Largen, 1991 RANGE: Worldwide in temperate and trop- (see Systematic Comments); Phrynobatra- ical continental areas and major islands, ex- chus GuÈnther, 1862 (including Dimorphog- 238 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 nathus Boulenger, 1906, and Phrynodon chus as currently arrayed, but at present we Parker, 1935; see Systematic Comments). cannot reject the possibility that it is the sis- CHARACTERIZATION AND DIAGNOSIS: Phry- ter taxon of Phrynobatrachus. We presume nobatrachids are small terrestrial and semi- that Dubois' (2005) association of Ericaba- aquatic frogs with poorly understood species trachus with his Phrynobatrachinae is based boundaries and with a typically biphasic life on similar reasoning although he provided no history, with eggs laid in water. Like many justi®cation for this inclusion. members of Ranoides, phrynobatrachids fre- quently have T-shaped terminal phalanges, [200] SUPERFAMILY: PYXICEPHALOIDEA although they lack digital discs. They usually BONAPARTE, 1850 retain an outer metatarsal tubercle (Laurent, IMMEDIATELY MORE INCLUSIVE TAXON: 1986) and are characterized by a tarsal tu- [192] Africanura new taxon. bercle (Channing, 2001) that is distinctive SISTER TAXON: [193] Phrynobatrachidae and may be synapomorphic. Phrynobatra- Laurent, 1941 ``1940''. chus species exhibit a median lingual tuber- RANGE: Sub-Saharan Africa. cle (Grant et al., 1997), which may be syn- CONTENT: [201] Petropedetidae Noble, apomorphic, although this needs to be care- 1931, and [209] Pyxicephalidae Bonaparte, fully surveyed. Its presence also in Indirana, 1850. Arthroleptides, and Petropedetes suggests CHARACTERIZATION AND DIAGNOSIS: Al- that it may be synapomorphic at a more gen- though no morphological characters in our eral level. study optimize to this branch, our molecular Nevertheless, none of the morphological data are decisive. See appendix 5 for sum- characters in our analysis optimize on this mary of molecular synapomorphies. taxon, although the molecular data are deci- COMMENT: This taxon is highy heteroge- sive in recognition of this taxon. (See appen- nous morphologically, at least with respect to dix 5 for listing of molecular synapomor- overall appearance. Nevertheless, the molec- phies for this taxon.) ular evidence is strong, and the taxon should SYSTEMATIC COMMENTS: Our data show survive additional testing. that Phrynobatrachus is paraphyletic with re- spect to Phrynodon and Dimorphognathus. [201] LFAMILY: PETROPEDETIDAE NOBLE, 1931 Surprisingly, Amiet (1981) suggested a close Petropedetinae Noble, 1931: 520. Type genus: Pe- relationship of Phrynodon with Petropedetes tropedetes Reichenow, 1874. (Petropedetidae) to the exclusion of Phry- Ranixalini Dubois, 1987 ``1985'': 66. Type genus: nobatrachus. Our data do not support this re- Ranixalus Dubois, 1986. New synonym. lationship and because this nominal genus Conrauini Dubois, 1992: 314. Type genus: Con- and Dimorphognathus are both monotypic raua Nieden, 1908. New synonym. and imbedded within Phrynobatrachus,we Indiraninae Blommers-SchloÈsser, 1993: 211. Type place Phrynodon and Dimorphognathus into genus: Indirana Laurent, 1986. New synonym. the synonymy of Phrynobatrachus, which af- IMMEDIATELY MORE INCLUSIVE TAXON: ter this action is monophyletic. Nevertheless, [200] Pyxicephaloidea Bonaparte, 1850. Phrynobatrachus remains one of the larger SISTER TAXON: [209] Pyxicephalidae Bon- taxonomic problems in Africa in terms of aparte, 1850. species boundaries and infrageneric clades. It RANGE: South India; tropical West and will yield its secrets only with a considerable East Africa. amount of morphological, behavioral, and CONTENT: Arthroleptides Nieden, 1911 molecular work. (See appendix 7 for new ``1910''; Conraua Nieden, 1908; Indirana and revivied combinations caused by these Laurent, 1986; Petropedetes Reichenow, synonymies.) Our inclusion in Phrynoba- 1874. trachidae of Ericabatrachus Largen, 1991 CHARACTERIZATION AND DIAGNOSIS: Petro- (not studied by us) rests on the original pub- pedetidae is heterogeneous morphologically, lication, which suggests that Ericabatrachus with forked omosterna. No morphological is ``Phrynobatrachus-like''. Likely, it will be synapomorphies are evident to us, although found to be imbedded within Phrynobatra- the molecular data are decisive. (See appen- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 239 dix 5 for molecular synapomorphies for this Amiet and Perret, 1969; Inger et al., 1984; taxon.) Drewes et al., 1989). SYSTEMATIC COMMENTS: The association of Adults of Arthroleptides, Indirana, and Indirana (India), Conraua (tropical West Af- Petropedetes also share characters whose po- rica; Ethiopia and Eritrea), and Arthrolep- larity is less clear. Males of most Petrope- tides ϩ Petropedetes (tropical West Africa; detes and Arthroleptides, and males of Indi- Tanzania and Kenya) at ®rst surprised us, rana (where they are known) share the pres- even though we had expected the undiagnos- ence of femoral glands of variable size and able Petropedetidae (sensu lato, now distrib- the presence of spicules around the margins uted among Petropedetidae, Phrynobatrachi- of jaw and/or chin in the pectoral area dae, and Dicroglossidae) to be obliterated. (Amiet, 1973; Inger et al., 1984; Perret, The stream-dwelling larvae of Arthrolep- 1984; Dubois, 1986 ``1985''; Klemens, 1998; tides and stream-dwelling and arboreal tad- however spicules are absent in Petropedetes poles of Indirana are amazingly similar parkeri [Amiet, 1983], and femoral glands (compare Altig and Johnston, 1989, and are absent in A. yakusini [Channing et al., Channing et al., 2002b, with Annandale and 2002b]). Note that spicules around the mar- Rao, 1918) in having elongate tails with very gins of jaw and/or chin and pectoral area, low caudal ®ns, large bulging eyes, a dor- occur also in Conraua and in at least several soventrally ¯attened body, and a laterally phrynobatrachids as rede®ned here (Perret, compressed jaw sheath with prominent lat- 1966). Until this character can be widely as- eral processes (Annandale, 1918; Rao, 1920; sessed its level of generality remains un- Amiet and Perret, 1969; Inger et al., 1984; known. Dubois, 1986 ``1985''; Drewes et al., 1989; Dubois (1987 ``1985'') proposed the rec- Channing et al., 2002b). Only larvae of Pe- ognition of the tribe Ranixalini (later treated tropedetes natator and P. palmipes have as a subfamily by Dubois, 1992), for the gen- been fully described (Lamotte and Zuber-Vo- era Nannophrys, Nyctibatrachus, and Indi- geli, 1954; Lamotte et al., 1959; Lamotte and rana on the basis of the presence of femoral Lescure, 1989), but some super®cial refer- glands in males of Nyctibatrachus and Indi- ences to morphology or behavior are avail- rana (unknown in Nannophrys), and the able for the larvae of P. cameronensis (Bou- morphological proximity of Nannophrys and lenger, 1906 ``1905''; Lawson, 1993), P. Nyctibatrachus was noted by Clarke (1981). newtoni (Perret, 1966; Amiet and Perret, Nannophrys and Indirana further share the 1969; Lawson, 1993), and P. parkeri and P. modi®cations of larval morphology associ- johnstoni (Amiet and Perret, 1969; Amiet, ated with semiterrestrial life that were men- 1983; Lawson, 1993). Drewes et al. (1989) tioned earlier (Kirtisinghe, 1958). From a noted inconsistencies in the description of morphological perspective, the evidence sup- the larva of P. palmipes. Regardless, from porting the monophyly of Nannophrys ϩ In- the comments or illustrations presented by dirana is the same as that favoring a rela- the authors mentioned above, larvae of Pe- tionship among Indirana, Arthroleptides, and tropedetes seem to have the same morpho- Petropedetes. As discussed earlier, other logical peculiarities as do those of Arthrolep- characters of still unclear polarity that could tides and Indirana. The only exception is the further support this hypothesis are the pres- larva of P. natator, which has an abdominal ence of femoral glands and spicules around disc and an oral disc that is proportionally the margins of jaw and/or chin and pectoral larger, with conspicuous lateral folds, and area. jaw sheaths that are not compressed laterally Petropedetes and Arthroleptides have (Lamotte and Zuber-Vogeli, 1954; Lamotte large digital discs, a long metasternal style, and Lescure, 1989). and T-shaped terminal phalanges. Indirana In transforming larvae of Arthroleptides, has Y-shaped terminal phalanges (Laurent, Indirana, and Petropedetes, the hind legs are 1986), which may be synapomorphic with large and seem to develop precociously, on the T-shaped terminal phalanges of Petro- a different growth trajectory from the front pedetes ϩ Arthroleptides although in our to- legs (Annandale, 1918; Lamotte et al., 1959; pology the simple terminal phalanges of 240 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Conraua presumably represent the apomor- SYSTEMATIC COMMENTS: This morphologi- phy. Roelants et al. (2004) suggested that In- cally heterogeneous taxon is coherent geo- dirana would ®nd its closest relatives in In- graphically. Although the association of dia. However, inasmuch as these authors did these genera was only noted recently (Van not include any African taxa in their analysis, der Meijden et al., 2005), much of the earlier it was impossible for them to detect a rela- taxonomy was based on very general notions tionship with African taxa. Van der Meijden of overall similarity, which are signi®cantly et al. (2005) placed Indirana as the sister tax- in¯uenced by perceptions of body size. The on of our Dicroglossinae. They also placed association of Afrana and Strongylopus (for- Conraua outside of a clade composed of Pe- merly in Ranini of Dubois, 1992) with An- tropedetes ϩ Pyxicephalinae, in both cases hydrophryne, Arthroleptella, Cacosternum, on the basis of fewer data and more analyt- and Natalobatrachus (formerly of Phryno- ical assumptions. Additional data or denser batrachidae [Petropedetidae] of Dubois, taxon sampling may rearrange these taxa, but 1992), and with Pyxicephalus and Aubria (in at present our molecular data are decisive Pyxicephalinae of Dubois, 1992), was some- and, as discussed earlier, they are consistent thing of a surprise (at least for us, as this was with the distribution of various larval and before Van der Meijden et al., 2005, ap- adult characteristics. peared), although no evidence beyond over- all similarity ever supported the older tax- [209] FAMILY: PYXICEPHALIDAE BONAPARTE, onomy. We still have three ``¯avors'' of frogs 1850 in this group: those that look like Rana (Af- Pyxicephalina Bonaparte, 1850: 1. Type genus: rana and Strongylopus); those that are stocky Pyxicephalus Tschudi, 1838. and big (Pyxicephalus and Aubria); and Phrynopsinae Noble, 1931: 518. Type genus: those that are generally small and have not Phrynopsis Pfeffer, 1893. attracted from systematists the attention they Cacosterninae Noble, 1931: 540. Type genus: Ca- deserve (the remainder). The absence of a costernum Boulenger, 1887. median lingual process may be synapo- Tomopternini Dubois, 1987 ``1985'': 56. Type ge- morphic, as this feature is present in Petro- nus: Tomopterna DumeÂril and Bibron, 1841. pedetidae and Phrynobatrachidae (Grant et New synonym. al., 1997). Dubois (2005), anticipating the IMMEDIATELY MORE INCLUSIVE TAXON: publication of Van der Meijden et al. (2005), [200] Pyxicephaloidea Bonaparte, 1850. recognized this taxon as a subfamily of Ran- SISTER TAXON: [201] Petropedetidae Noble, idae, Pyxicephalinae, which we recognize as 1931. a family. RANGE: Sub-Saharan Africa. Within Pyxicephalidae, we recognize two CONTENT: Amietia Dubois, 1987 ``1986'' subfamilies: [210] Pyxicephalinae Bonapar- (including Afrana Dubois, 1992, see Sys- te, 1850 (Pyxicephalus and Aubria) and tematic Comments); Anhydrophryne Hewitt, [212] Cacosterninae Noble, 1931 (for the re- 1919; Arthroleptella Hewitt, 1926; Aubria maining genera). Pyxicephalinae is united by Boulenger, 1917; Cacosternum Boulenger, the following synapomorphies: (1) skull ex- 1887; Microbatrachella Hewitt, 1926; Na- ostosis; (2) occipital canal present; (3) zy- talobatrachus Hewitt and Methuen, 1912; gomatic ramus much longer than otic ramus, Nothophryne Poynton, 1963; Poyntonia articulating with the postorbital process of Channing and Boycott, 1989; Pyxicephalus the pars facialis of the maxilla; and (4) strong Tschudi, 1838; Strongylopus Tschudi, 1838; overlap of the medial ramus of the pterygoid Tomopterna DumeÂril and Bibron, 1841. and the parasphenoid ala (Clarke, 1981). Ca- CHARACTERIZATION AND DIAGNOSIS: Al- costerninae in our sense is not united by any though we know of no morphological syna- morphological feature that we can identify pomorphies for this group, the molecular ev- with any certainty, although the molecular idence is decisive in support of this branch. data are decisive (see appendix 5). (See appendix 5 for molecular synapomor- We place Afrana Dubois, 1992, into the phies of this taxon; also see Systematic Com- synonymy of [218] Amietia Dubois, 1987 ments.) ``1986'', to resolve the paraphyly of Afrana. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 241

No characteristics of ``Afrana'' or Amietia of the morphological characters in our anal- reject this placement. ysis optimize on this taxon although the mo- Clearly, our data do not support the notion lecular data decisively support recognition of (Poynton, 1964a) that Cacosternum is close- this taxon. (See appendix 5 for molecular ly related to Phrynobatrachus. Our associa- transformations.) We recognized two sub- tion of Microbatrachella, Nothophryne, and families within Dicroglossidae, which are Poyntonia with this clade is provisional, discussed in separate accounts because of the based on the assertion by Blommers-SchloÈs- size and complexity of discussion. ser (1993) that these genera are allied by re- duced ossi®cation of the omosternal style [225] SUBFAMILY: DICROGLOSSINAE and procoracoid clavicular bar. ANDERSON, 1871 Dicroglossidae J. Anderson, 1871: 38. Type ge- [220] SAUKROBATRACHIA NEW TAXON nus: Dicroglossus GuÈnther, 1860. Limnonectini Dubois, 1992: 315. Type genus: ETYMOLOGY: Saukro- (Latin: graceful, Limnonectes Fitzinger, 1843. pretty) ϩ batrachos (Greek: frog), referenc- Paini Dubois, 1992: 317. Type genus: Paa Du- ing the beauty of many of the species in- bois, 1975. cluded in this clade. IMMEDIATELY MORE INCLUSIVE TAXON: IMMEDIATELY MORE INCLUSIVE TAXON: [221] Dicroglossidae Anderson, 1871. [191] Ametrobatrachia new taxon. SISTER TAXON: [222] Occidozyginae Fei, SISTER TAXON: [192] Africanura new tax- Ye, and Huang, 1991 ``1990''. on. RANGE: Northwestern and sub-Saharan Af- RANGE: Eurasia, Africa, and Madagascar, rica; southern Arabian Peninsula; Pakistan, to northern Australia; North and Central- Afghanistan, India, Sri Lanka, and , America to central South America. through southern China (including part of CONCEPT AND CONTENT: Saukrobatrachia Xizang) and Indochina to the islands of the new taxon is a monophyletic taxon com- Sunda Shelf; Japan. posed of [221] Dicroglossidae Anderson, CONTENT: Annandia Dubois, 1992 (see 1871, and [244] Aglaioanura new taxon. Systematic Comments); Eripaa Dubois, CHARACTERIZATION AND DIAGNOSIS: Al- 1992 (see Systematic Comments); Euphlyc- though no morphological characters that we tis Fitzinger, 1843; ``Fejervarya'' Bolkay, are aware of optimize on this branch, the mo- 1915 (see Systematic Comments); Hoploba- lecular data are decisive in support of this trachus Peters, 1863; Limnonectes Fitzinger, taxon. (See appendix 5 for listing of molec- 1843 (including Taylorana Dubois, 1987 ular synapomorphies.) ``1986''); Minervarya Dubois, Ohler, and Biju, 2001; Nannophrys GuÈnther, 1869 [221] FAMILY: DICROGLOSSIDAE ANDERSON, 1871 ``1868''; Nanorana GuÈnther, 1896 (includ- ing Altirana Stejneger, 1927; Chaparana IMMEDIATELY MORE INCLUSIVE TAXON: Bourret, 1939; and Paa Dubois, 1975; see [220] Saukrobatrachia new taxon. Systematic Comments); Ombrana Dubois, SISTER TAXON: [244] Aglaioanura new tax- 1992 (see Systematic Comments); Quasipaa on. Dubois, 1992; Sphaerotheca GuÈnther, 1859 RANGE: Northwestern and sub-Saharan Af- ``1858''. rica; southern Arabian Peninsula; Pakistan, CHARACTERIZATION AND DIAGNOSIS: Al- Afghanistan, India, Sri Lanka, and Nepal, though the molecular evidence is decisive for through southern China (including part of the existence of Dicroglossinae, we are Xizang) and Indochina to Japan and the Phil- aware of no morphological synapomorphies ippines; islands of the Sunda Shelf as far as that optimize to this branch. (See Systematic Flores. Comments.) Appendix 5 shows the molecu- CONTENT: [225] Dicroglossinae Anderson, lar transformations associated with this tax- 1871, and [222] Occidozyginae Fei, Ye, and on. Huang, 1991 ``1990''. SYSTEMATIC COMMENTS: Within Dicroglos- CHARACTERIZATION AND DIAGNOSIS: None sinae Anderson, 1871, we recognize two 242 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 monophyletic tribes, [226] Limnonectini Du- pending the publication of evidence, we re- bois, 1992, for Limnonectes (including as gard these as monotypic genera of uncertain synonyms Elachyglossa Anderson, 1916; placement within Dicroglossidae (see appen- Taylorana Dubois, 1987), and [232] Dicrog- dix 7 for combinations). lossini Anderson, 1871, for the remaining Previous authors (Dubois and Ohler, 2000; genera, Annandia,``Fejervarya'' (see be- Dubois et al., 2001; Grosjean et al., 2004) low), Nanorana (including Chaparana and demonstrated that Sphaerotheca and Fejer- Paa), Quasipaa, Sphaerotheca, Nannophrys, varya are closely related. Our data permit us Euphlyctis, and Hoplobatrachus. (Evidence to go further and suggest strongly that rec- for both is listed in appendix 5.) This agrees ognition of Sphaerotheca (as well as Eu- with several other phylogenetic analyses that phlyctis, Hoplobatrachus, and Nannophrys) used DNA evidence (e.g., Bossuyt and Mil- renders Fejervarya sensu Dubois and Ohler inkovitch, 2000; Emerson et al., 2000b; Mar- (2000) paraphyletic, as does a group com- mayou et al., 2000; Vences et al., 2000c; Ko- posed of Nannophrys, Euphlyctis, and Ho- such et al., 2001; Grosjean et al., 2004; Roe- plobatrachus. J. M. Hoyos (in Dubois and lants et al., 2004; Jiang et al., 2005; Jiang Ohler, 2000) suggested that Fejervarya does and Zhou, 2005), although our expanded tax- have a morphological synapomorphy: ven- on sampling and data altered some relation- trolateral edge of the m. pectoralis pars ab- ships within Dicroglossini. dominalis slightly attached to muscles that As noted in ``Results'', our results are are dorsal relative to it, which results in a strongly congruent with those of Jiang et al. dark ventrolateral line from axilla to groin, (2005), especially when the rooting point is especially visible in live specimens. This corrected by our larger outgroup sampling needs to be veri®ed with reference to the (see ®g. 64). Because their analysis provided condition in Sphaerotheca and the other sat- DNA sequence evidence unrejected by mor- ellite genera as well as to assure that this is phological synapomorphies, we take their re- universal in Fejervarya and not just in some sults at face value: Nanorana as they viewed subset of the nominal genus. Serious system- it is imbedded within a paraphyletic ``Paa'', atic and nomenclatural issues impede reso- and ``Chaparana'' is polyphyletic with the lution of this paraphyly. The most important two components both imbedded within is that there are many species of nominal Fe- ``Paa''. Nevertheless, they provided evi- jervarya that we did not study, and there may dence that their Group 1 (composed of nom- be several species of frogs masquerading un- inal Paa, Nanorana, and Chaparana, and ex- der the name (Du- cluding Quasipaa), is monophyletic. Group bois and Ohler, 2000). Because our exemplar 1 is characterized by paired patches of spines of Fejervarya limnocharis is from Vietnam on the chest (Jiang et al., 2005), which may and the type locality of this same nominal not be synapomorphic but distinguishes this taxon is Java, we are reluctant to assume too taxon morphologically from Quasipaa. The much about the phylogenetic placement of F. oldest name for Group 1 is Nanorana GuÈn- limnocharis sensu stricto. Ongoing research ther, 1896. (See appendix 7 for the name by Dubois and Ohler (cited in Dubois and changes that extend from the synonymy of Ohler, 2000) should provide some resolution Chaparana Bourret, 1939, and Paa Dubois, in the near future to this problem. In the in- 1975, with Nanorana GuÈnther, 1896.) An- terim we recommend using quotation marks nandia Dubois, 1992, and Ombrana Dubois, around the name ``Fejervarya'' to denote the 1992, were originally named as subgenera of paraphyly of this taxon. Chaparana, and Eripaa Dubois, 1992, was We reaf®rm that placement of Limnonec- originally named as a subgenus of Paa. None tes limborgi in the monotypic genus Taylor- of these three taxa were included, discussed, ana renders Limnonectes paraphyletic and or even mentioned in the study of Jiang et therefore continue the synonymy of Taylor- al. (2005). Without discussion, Dubois ana with Limnonectes, following Inger (2005) transferred Annandia into Limnonec- (1996) and Emerson et al. (2000a). Emerson tini. The placement of these taxa in Dicrog- et al. (2000a) and Evans et al. (2004) pro- lossinae is presumably not controversial, so vided considerable evidence that Elachyglos- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 243 sa (formerly Bourretia) renders Limnonectes [244] AGLAIOANURA NEW TAXON paraphyletic as well. We therefore reject the ETYMOLOGY: Aglaio- [Greek: splendid or use of subgeneraÐat least as currently for- noble] ϩ anoura [Greek: tailless, i.e., frog]. mulatedÐwithin Limnonectes, even though IMMEDIATELY MORE INCLUSIVE TAXON: some authors (e.g., Delorme et al., 2004) [220] Saukrobatrachia new taxon. have retained their use even though they mis- SISTER TAXON: [221] Dicroglossidae An- lead about evolutionary relationship. derson, 1871. Although Minervarya exhibits the ``Fejer- RANGE: Eurasia, Africa, and Madagascar, varyan line'' (of Dubois and Ohler, 2000; see to northern Australia; the Americas exclud- Dubois et al., 2001), it was not included in ing southern South America. our study, so we are unable to make any CONCEPT AND CONTENT: Aglaioanura is a comments about its position in the tree. Our monophyletic group composed of [245] Rha- inclusion of Minervarya in Dicroglossinae is cophoroidea Hoffman, 1932 (1858), and obviously provisional; additional study is [269] Ranoidea Ra®nesque, 1814. needed. CHARACTERIZATION AND DIAGNOSIS:Onthe basis of our few exemplars for morphology [222] SUBFAMILY: OCCIDOZYGINAE FEI, YE, (Chiromantis xerampelina, Rhacophorus AND HUANG, 1991 ``1990'' pardalis, Rana nigrovittata, and Rana tem- poraria) the following characters are sug- Occydozyginae Fei et al., 1991 ``1990'': 123. gested as possibly synapomorphies of this Type genus: Occidozyga Kuhl and Van Hasselt, group: (1) functional larval m. levator man- 1822. dibulae lateralis absent (Haas 56.0); and (2) terminal phalanges bifurcated T-shape or Y- IMMEDIATELY MORE INCLUSIVE TAXON: shaped (Haas 156.2; reversed in several lin- [221] Dicroglossidae Anderson, 1871. eages of Ranidae). (Molecular synapomor- SISTER TAXON: [225] Dicroglossinae An- phies are provided in appendix 5.) derson, 1871. RANGE: Southern China (Guangxi, Yun- [245] SUPERFAMILY: RHACOPHOROIDEA nan, and ), Thailand, Indochina, Ma- HOFFMAN, 1932 (1858) laya, Greater and Lesser Sunda Islands as far as Flores, and Philippines. IMMEDIATELY MORE INCLUSIVE TAXON: CONTENT: Occidozyga Kuhl and Hasselt, [244] Aglaioanura new taxon. 1822 (including Phrynoglossus Peters, 1867; SISTER TAXON: [269] Ranoidea Ra®nesque, see Systematic Comments). 1814. RANGE: Tropical sub-Saharan Africa; Mad- CHARACTERIZATION AND DIAGNOSIS: Al- though the molecular data are decisive (see agascar; South India and Sri Lanka; Japan; northeastern India to eastern China south appendix 5), Occidozyginae has other syna- through the Philippines and Greater Sundas; pomorphies: (1) aquatic larvae with a kera- Sulawesi. todont formula of 0/0; and (2) a lateral line CONTENT: [246] Mantellidae Laurent, system that persists into adulthood (absent in 1946, and [253] Rhacophoridae Hoffman, Occidozyga lima; Dubois et al., 2001; con- 1932 (1858). vergent in Euphlyctis: Dicroglossinae). CHARACTERIZATION AND DIAGNOSIS: See SYSTEMATIC COMMENTS: Our data demon- Rhacophoridae. One character in our analysis strate that Phrynoglossus (which retains the de®nitely optimizes on this taxon: intercalary lateral line system into adulthood) is para- element present (Haas 151.1). Channing phyletic with respect to Occidozyga (which (1989) also suggested the following as syn- does not). We therefore agree with Inger apomorphies: (1) only one slip of the m. ex- (1996) that Phrynoglossus is a synonym of tensor digitorum communis longus, inserting Occidozyga (the senior name), providing a on distal portion of fourth metatarsal; and (2) monophyletic Occidozyga. (See appendix 7 outermost slip of the m. palmaris longus in- for new and revived combinations resulting serting on the proximolateral rim of the apo- from this synonymy.) neurosis palmaris. Ford and Cannatella 244 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(1993) also suggested that bifurcate terminal and Vences, 1994). They share with their sis- phalanges are a synapomorphy of this taxon, ter taxon, Rhacophoridae, intercalary phalan- although this character may optimize at a geal elements. more general level inasmuch as expanded toe Laurent (1986: 764) distinguished mantel- tips seem to optimize on or near Aglaioan- lids from rhacophorids solely on basis of the ura. third carpal being fused with the fourth and SYSTEMATIC COMMENTS: Our study puts to ®fth in rhacophorids, but being free in man- rest whether mantellids and rhacophorids are tellids (this feature is likely synapomorphic sister taxa (e.g., Emerson et al., 2000b) or at this level of universality). Nevertheless, mantellids are imbedded in some way within this feature has not been adequately assayed, the rhacophorids (Liem, 1970). Whether they so at present the molecular evidence is par- should be considered mutual subfamilies of ticularly decisive in distinguishing this as a a larger Rhacophoridae (ϭ Rhacophoroidea monophyletic group that forms the sister tax- in our use) is not a scienti®c proposition. We on of Rhacophoridae. None of the morpho- follow the usage of Glaw and Vences (e.g., logical characters in our analysis optimize on Vences et al., 2002; Vallan et al., 2003; this taxon. (Molecular transformations are Vences et al., 2003a; Vences and Glaw, listed in appendix 5.) 2004). SYSTEMATIC COMMENTS: Vences and Glaw (2001) recognized three subfamilies on the [246] FAMILY: MANTELLIDAE LAURENT, 1946 basis of molecular data arranged phylogenet- ically: Laliostominae (Boophinae ϩ Mantel- Mantellinae Laurent, 1946: 336. Type genus: linae). We consider Mantellinae and Lalios- Mantella Boulenger, 1882. Boophinae Vences and Glaw, 2001: 88. Type ge- tominae of Vences and Glaw (2001) to be nus: Boophis Tschudi, 1838. tribes within a larger subfamily [248] Man- Laliostominae Vences and Glaw, 2001: 88. Type tellinae, this subfamily forming the sister tax- genus: Laliostoma Glaw, Vences, and BoÈhme, on of [247] Boophinae. Aglyptodactylus and 1998. Laliostoma are in [249] Laliostomini, and within Boophini, only Boophis, and [252] IMMEDIATELY MORE INCLUSIVE TAXON: Mantella and [251] ``Mantidactylus'' are in [245] Rhacophoroidea Hoffman, 1932 [250] Mantellini. Although ``Mantidactylus'' (1858). is clearly paraphyletic with respect to Man- SISTER TAXON: [253] Rhacophoridae Hoff- tella (e.g., Vences and Glaw, 2001), our lim- man, 1932 (1858). ited taxon sampling did not reveal this. It RANGE: Madagascar. should be noted that there are many nominal CONTENT: Aglyptodactylus Boulenger, subgenera that require reformulation as well 1919 ``1918''; Boophis Tschudi, 1838; Lal- (Raxworthy, Grant, and Faivovich, in prep- iostoma Glaw, Vences, and BoÈhme, 1998; aration). For instance, Vences et al. (2002) Mantella Boulenger, 1882; ``Mantidactylus'' revised the species of the ``Mantidactylus'' Boulenger, 1895. subgenus Laurentomantis and presented ev- CHARACTERIZATION AND DIAGNOSIS: Man- idence in their resulting tree of the paraphyly tellids are small to medium-size terrestrial or of ``Mantidactylus'' with respect to Mantella, arboreal frogs, predominantly found in semi- the paraphyly of the subgenus Brygoomantis, arid to wet forested habitats. Although most and the polyphyly of and Ge- are drab or cryptically colored, species of phyromantis, as well as a lack of evidence Mantellini in particular are brightly colored. for either paraphyly or monophyly of Pan- Life history is varied, from the usual biphasic danusicola. Much remains to be done, and life history with aquatic eggs and feeding we cannot recommend the use of subgenera tadpoles (Boophis) to nidicolous larvae (e.g., within ``Mantidactylus'' until the inconsis- many Mantidactylus). At least some (e.g., tency of taxonomy with phylogeny is ad- Mantidactylus eiselti) have direct develop- dressed within that group. ment. Most species lay eggs away from wa- Laurent, 1943, was ter, in some cases in a suspended nest from placed in the synonymy of Philautus by R.F. which the tadpoles drop into water (Glaw Inger (In Frost, 1985). This was accepted by 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 245

Dubois (1999b: 5) although the assignment dialis only (Haas 96.5); (5) free basihyal ab- to Mantellidae by Laurent (1986) has not sent (Haas 105.0); (6) commissura proximal- been directly challenged through discussion is II present (Haas 110.1); and (7) commis- of evidence. A second look is warranted. sura proximalis III present (Haas 111.1). SYSTEMATIC COMMENTS: Taxonomic deci- [253] FAMILY: RHACOPHORIDAE HOFFMAN, sions taken here are guided by our results 1932 (1858) (®gs. 50, 65), the DNA sequence study of Polypedatidae GuÈnther, 1858b: 346. Type genus: J.A. Wilkinson et al. (2002; ®g. 48) and the Polypedates Tschudi, 1838. essentially data-free tree of Delorme et al. Rhacophoridae Hoffman, 1932: 581. Type genus: (2005; ®g. 49), which was presented along Rhacophorus Kuhl and Van Hasselt, 1822. with a system of morphological differentia Philautinae Dubois, 1981: 258. Type genus: Phi- that delimited a number of monophyletic and lautus Gistel, 1848. paraphyletic groups, seemingly without ref- Buergeriinae Channing, 1989. Type genus: Buer- erence to the tree itself. Results of the three geria Tschudi, 1838. have basic agreements. IMMEDIATELY MORE INCLUSIVE TAXON: Buergeriinae Channing, 1989, may be rec- [244] Rhacophoroidea. ognized for Buergeria and Rhacophorinae SISTER TAXON: [246] Mantellidae. Hoffman, 1932 (1858), for the remaining RANGE: Tropical sub-Saharan Africa; rhacophorines, as was suggested by Chan- South India and Sri Lanka; Japan; northeast- ning (1989) and as diagnosed by J.A. Wil- ern India to eastern China south through the kinson et al. (2002). We cannot subscribe to Philippines and Greater Sundas; Sulawesi. the tribal taxonomy of Delorme et al. (2005) CONTENT: Aquixalus Delorme, Dubois, because their Philautini is not monophyletic Grosjean, and Ohler, 2005 (see Systematic on their own ®gure (®g. 49), and because the Comments); Buergeria Tschudi, 1838; Chi- evidence in support of their tree was largely romantis Peters, 1854 (including Chirixalus undisclosed. Boulenger, 1893; see Systematic Comments); On the basis of our results, and the studies Feihyla new genus (see Systematic Com- of J.A. Wilkinson et al. (2002) and Delorme ments); Kurixalus Ye, Fei, and Dubois, 1999 et al. (2005), two problems of generic delim- (see Systematic Comments); Nyctixalus itation appear to persist in the taxonomy. The Boulenger, 1882; Philautus Gistel, 1848; Po- ®rst of these, the paraphyly/polyphyly of lypedates Tschudi, 1838; Rhacophorus Kuhl ``Rhacophorus'' is beyond the scope of this and Hasselt, 1822; Theloderma Tschudi, paper; more taxa need to be analyzed before 1838. this problem can be addressed. The second CHARACTERIZATION AND DIAGNOSIS: Al- problem is that nominal ``Chirixalus'' seem- though a few groups are primarily terrestrial, ingly falls into four generic units. We can rhacophorids are predominantly treefrogs, help correct the problems surrounding the sharing with basal ranids expanded digital polyphyly/paraphyly ``Chirixalus'', although pads and with mantellids the characteristic of the phylogenetic position of many species of intercalary phalangeal elements. Most spe- both ``Chirixalus'' and nominal Philautus cies have T-shaped terminal phalanges. Sev- needs to be evaluated. eral larval characters that optimized on this (1) Kurixalus Fei, Ye, and Dubois (in Fei, branch may actually be synapomorphies of 1999). As noted in ``Results'', we apply this Rhacophoroidea, or some part of Rhaco- name to a taxon that includes K. eif®ngeri phoridae: (1) anterior insertion of m. subar- and K. idiootocus, which is diagnosed by our cualis rectus II±IV on ceratobranchial II molecular evidence (see appendix 5, branch (Haas 37.1); (2) larval m. levator mandibulae 256). We provisionally include K. verruco- externus present as two portions (profundus sus, which Delorme et al. (2005), without ev- and super®cialis; Haas 54.1); (3) posterior idence or discussion, ®gured as the sister tax- dorsal process of pars alaris expanded ter- on of Kurixalus eif®ngeri ϩ K. idiootocus. minally, almost rectangular in lateral view (These authors included idiootocus and ver- (Haas 89.1); (4) cartilaginous roo®ng of the rucosus without discussion in their new poly- cavum cranii composed of taeniae tecti me- phyletic/paraphyletic ``Aquixalus'', even as 246 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 they illustrated these species as being in an other species with this taxon will require exclusive monophyletic group with Kurixal- considerable additional work. us eif®ngeri). Under this concept, there are Although ``Chirixalus'' palpebralis has currently no identi®ed morphological syna- been demonstrated to be phylogenetically pomorphies of Kurixalus, because the pur- distinct (J.A. Wilkinson et al., 2002; Delorme ported synapomorphies associated with Ku- et al., 2005) and deserving a new generic rixalus eif®ngeri (well-developed prepollex name, the status of presumably closely relat- and oophagus tadpoles) are not exhibited in ed species ``Chirixalus'' romeri and ``C.'' K. idiootocus or K. verrucosus (Kuramoto ocellatus of the ``Philautus'' palpebralis and Wang, 1987; Ziegler and Vences, 2002; group of Fei, 1999) remains an open ques- Matsui and Orlov, 2004). Excluding ``Aquix- tion, although no evidence so far has sug- alus'' idiootocus and ``A.'' verrucosus from gested that these species form a monophy- ``Aquixalus'', we suggest, renders Aquixalus letic group. Morphological evidence provid- (sensu stricto) monophyletic (see below), if ed by Delorme et al. (2005) differentiating we assume that the tree of Delorme et al. their Rhacophorini (including ``Chirixalus'' (2005) survives testing by evidence. palpebralis on their tree) and Philautini (a (2) Feihyla new genus (type species: Phi- paraphyletic group that on their tree includes ϭ lautus palpebralis Smith, 1924. Etymology: ``Philautus'' gracilipes [ Aquixalus graci- Fei Liang ϩ hyla [Greek: vocative form of lipes]), suggests that Aquixalus (including Hylas, a traditional generic root for treefrogs] ``Chirixalus'' gracilipes) is not close to Feih- to commemorate the extensive contributions yla (see discussion below under Aquixalus). to Chinese herpetology by Fei Liang). J.A. (3) Chiromantis Peters, 1854, and Chirix- Wilkinson et al. (2002) found his exemplar alus Boulenger, 1893. A third unit is the of the ``Philautus'' palpebralis group of Fei cluster of species paraphyletic with respect (1999), ``Chirixalus'' palpebralis,tobethe to Chiromantis. The paraphyly of Chirixalus sister taxon of a group composed of all rha- (sensu stricto) with respect to Chiromantis was not a surprise to us. J.A. Wilkinson et cophorids except Buergeria. Delorme et al. al. (2002) had suggested that Chirixalus do- (2005) placed ``Chirixalus'' palpebralis in riae is the sister taxon of Chiromantis, and their Rhacophorini, which otherwise corre- that Chirixalus vittatus is close to Polype- sponds to a monophyletic group recovered dates (compare their results with ours, which by us and by J.A. Wilkinson et al. (2002). In are based on substantially more data). We fact, this is the major point of disagreement place Chirixalus Boulenger, 1893, into the between J.A. Wilkinson et al. (2002) and De- synonymy of Chiromantis Peters, 1854, to lorme et al. (2005). What is clear is that correct this paraphyly. (See appendix 7 for ``Chirixalus'' palpebralis is not in a mono- new combinations that extend from this phyletic group with Chirixalus (sensu stric- change and appendix 5 for molecular syna- to), nor obviously associated closely with pomorphies.) any other generic grouping. For this reason (4) Aquixalus Delorme, Dubois, Grosjean, we have named Feihyla to recognize its dis- and Ohler, 2005. We recognize a monophy- tinctiveness. We cannot construe Feihyla to letic Aquixalus (i.e., Aquixalus sensu Delor- the ``Philautus'' palpebralis group of Fei me et al., 2005, but excluding ``Aquixalus'' (1999) because the diagnosis of this group is idiootocus and ``Aquixalus'' verrucosus; that insuf®cient to distinguish it from many other is, without the molecular synapomorphies of species outside of China (i.e., Fei, 1999, branch 256Ðsee above). Delorme et al diagnosed his ``Philautus'' palpebralis group (2005) diagnosed this taxon (although we do as ``Philautus'' from China, with an X not know which of the listed species they or )( shape on the dorsum and lacking vo- actually evaluated for these characters), but merine teeth), such as Aquixalus gracilipes our exclusion of Kurixalus idiootocus (and and A. supercornutus; see discussion below). provisionally K. verrucosus) from Aquixalus We therefore diagnose Feihyla by the char- on the basis of the molecular synapomor- acters for the species ``Philautus'' palpe- phies that place Kurixalus distant from bralis provided by Fei (1999). Association of Aquixalus should render Aquixalus mono- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 247 phyletic if the tree provided by Delorme et rixalus verrucosus, so this diagnosis must be al. (2005) is correct. We suggest, on the basis largely or entirely based on plesiomorphies, of the tree provided by Delorme et al (2005), with the nominal subgenus Aquixalus being that the morphological similarities shared by those members of Aquixalus that do not share Kurixalus and Aquixalus are plesiomorphic. the apomorphies of Gracixalus. Detailed We follow the recognition by Delorme et analysis of disclosed evidence is necessary. al. (2005) of a putatively monophyletic sub- genus Gracixalus for ``Philautus'' gracilipes [269] SUPERFAMILY: RANOIDEA RAFINESQUE, Bourret, 1937, and ``Philautus'' supercor- 1814 nutus Orlov, Ho, and Nguyen, 2004 (not IMMEDIATELY MORE INCLUSIVE TAXON: studied by us). The morphological diagnosis [244] Aglaioanura new taxon. of Gracixalus (spines on the upper , SISTER TAXON: [245] Rhacophoroidea rictal gland connected to the mouth, foot Hoffman, 1932 (1858). very thin, two outer palmar tubercles, white RANGE: Worldwide temperate and tropical spot on snout tip of tadpole, ®ve pairs of pre- environments, except for southern Australia, lingual papillae on the tadpole, crescent- New Zealand, Seychelles, and southern shaped crest on the tadpole) purportedly sep- South America. arates it from the nominate subgenus Aquix- CONTENT: [270] Nyctibatrachidae Blom- alus, but the absence of adequate published mers-SchloÈsser, 1993, and [272] Ranidae Raf- tadpole descriptions suggest that this diag- inesque, 1814. nosis should be treated as provisional (Bain CHARACTERIZATION AND DIAGNOSIS: Mor- and Nguyen, 2004; Matsui and Orlov, 2004; phological synapomorphies for Ranidae (see Delorme et al., 2005). Although Gracixalus below) may actually optimize at this level. can be separated from Feihyla palpebralis Regardless, the molecular data are decisive (the latter in parentheses): snout triangularly in support of this taxon (appendix 5). pointed (obtusely pointed); skin translucent (not translucent); small white tubercles along [270] FAMILY: NYCTIBATRACHIDAE the head, anal region, and large conical tu- BLOMMERS-SCHLOÈ SSER, 1993 bercles on upper eyelid (all absent), these characters do not unambiguously separate Nyctibatrachinae Blommers-SchloÈsser, 1993: 211. Gracixalus from ``P.'' romeri,``P.'' ocella- Type genus: Nyctibatrachus Boulenger, 1882. tus, the other members of the ``P.'' palpe- Lankanectinae Dubois and Ohler, 2001: 82. Type genus: Lankanectes Dubois and Ohler, 2001. bralis group of Fei (1999). The placement of New synonym. these two species, as well as higher level re- lationships will be dependent upon a rigorous IMMEDIATELY MORE INCLUSIVE TAXON: phylogenetic analysis. [269] Ranoidea Ra®nesque, 1814. Although we cannot reject the putative SISTER TAXON: [272] Ranidae Ra®nesque, monophyly of the subgenus Aquixalus (in- 1814. cluding the type species A. odontotarsus,as RANGE: Sri Lanka and India. well as A. ananjevae, A. baliogaster, A. bis- CONTENT: Nyctibatrachus Boulenger, acculus, A. carinensis, and A. naso; modi®ed 1882; Lankanectes Dubois and Ohler, 2001. from Delorme et al., 2005), we do not see CHARACTERIZATION AND DIAGNOSIS: None any reason to recognize it, either, until the of our analyzed morphology optimizes on relevant phylogenetic data are published by this branch, although the molecular data are the original authors. According to Delorme decisive. See appendix 5 for list of unambig- et al. (2005), the morphological diagnosis of uous molecular synapomorphies. Aquixalus (webbing on feet not extending to SYSTEMATIC COMMENTS: Nyctibatrachidae toes, rictal gland not connected to mouth, in our sense brings two genera together, Nyc- foot very thick, one outer palmar tubercle, tibatrachus, with a median lingual process concavity on tadpole snout in lateral view, (unknown polarity), digital discs present four pairs of prelingual papillae in tadpole, (plesiomorphic), femoral glands present (un- median crest in tadpole triangular shaped, known polarity), and lateral line system not 180±240 eggs per clutch) also applies to Ku- persisting into adulthood (plesiomorphic), 248 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 and Lankanectes, with no median lingual Amerana Dubois, 1992; Aurorana Dubois, process, digital discs absent, femoral glands 1992; Pseudoamolops Jiang, Fei, Ye, Zeng, absent, and lateral line system persisting into Zhen, Xie, and Chen, 1997; and Pseudorana adulthood (Dubois et al., 2001). They are ar- Dubois, 1992); Sanguirana Dubois, 1992; ranged in a single family to avoid the taxo- Staurois Cope, 1865; Sylvirana Dubois, nomic redundancy of having monotypic (and 1992 (including Papurana Dubois, 1992, therefore uninformative) family-group and Tylerana Dubois, 199232). (See System- names. atic Comments.) CHARACTERIZATION AND DIAGNOSIS: Al- [272] FAMILY: RANIDAE RAFINESQUE, 1814 though Haas (2003) included only two ranids in his study, Sylvirana nigrovittata and Rana Ranaridia Ra®nesque, 1814: 102. Type genus: Ranaridia Ra®nesque, 1814 temporaria, characters that optimize on their Limnodytae Fitzinger, 1843: 31. Type genus: Lim- subtending branch are candidates as syna- nodytes DumeÂril and Bibron, 1841. pomorphies for Ranidae: (1) posterolateral Amolopsinae Yang, 1991a: 172. Type genus: projections of the crista parotica absent (Haas Amolops Cope, 1865. 67.0); and (2) processus branchialis closed (Haas 114.1). Denser sampling should test IMMEDIATELY MORE INCLUSIVE TAXON: this proposition. These characters may actu- [269] Ranoidea Ra®nesque, 1814. ally optimize on Ranoides. Regardless, the SISTER TAXON: [270] Nyctibatrachidae molecular data are decisive (see appendix 5). Blommers-SchloÈsser, 1993. SYSTEMATIC COMMENTS: As noted in ``Re- RANGE: Temperate and tropical Africa and sults'', Batrachylodes is transferred de®ni- Eurasia through Indonesia to northern Aus- tively to Ceratobatrachidae and Amietia (in- tralia, North America, Central America, and cluding Afrana) and Strongylopus are trans- northern South America. ferred to Pyxicephalidae. For discussion of CONTENT: Amolops Cope, 1865 (including these taxa see those familial accounts. Amo Dubois, 1992); Babina Thompson, As noted in the ``Review of Current Tax- 1912 (including Nidirana Dubois, 1992); onomy'', the sections and subsections of Clinotarsus Mivart, 1869; Glandirana Fei, ``Rana'' (sensu lato) provided by Dubois 32 Ye, and Huang, 1991 ``1990'' (including (1992) do not inform about evolutionary re- Rugosa Fei, Ye, and Huang, 1991 ``1990''); lationships, so for this discussion and the tax- Hydrophylax Fitzinger, 1843 (including Am- onomic remedies we suggest, we will focus nirana Dubois, 1992, and Chalcorana Du- on genera and subgenera. The discussion that bois, 1992); Hylarana Tschudi, 1838; Huia follows addresses the generic taxonomy that Yang, 1991 (including Eburana Dubois, we recommend (moving from top to bottom 1992; Bamburana Fei, Ye, Jiang, Xie, and of Ranidae [new taxonomy] in ®gure 71, al- Huang, 2005; Odorrana Fei, Ye, and Huang, though addressing other genera and problems 1991 ``1990''); Humerana Dubois, 1992; in passing). Lithobates Fitzinger, 1843 (including Aquar- Staurois Cope, 1865: We accept Staurois ana Dubois, 1992; Pantherana Dubois, as a genus, although we note that evidence 1992; Sierrana Dubois, 1992; Trypheropsis for this taxon's monophyly is equivocal and Cope, 1868; Zweifelia Dubois, 1992); Mer- requires testing. The traditional diagnosis of istogenys Yang, 1991; Nasirana Dubois, StauroisÐdigital discs broader than long; T- 1992; Pelophylax Fitzinger, 1843; Pterorana shaped terminal phalanges with horizontal Kiyasetuo and Khare, 1986; Pulchrana Du- arm longer than longitudinal arm; outer bois, 1992; Rana Linnaeus, 1758 (including metatarsals separated to base but webbed; nasals small separated from each other and 32 Dubois (1999a: 91) considered Glandirana Fei, Ye, frontoparietal; omosternal style not forked and Huang, 1991, to have priority over Rugosa Fei, Ye, (Boulenger, 1918); and lacking a raised ab- and Huang, 1991, and Sylvirana Dubois, 1992, to have dominal sucker disc on larva (Inger, 1966)Ð priority over Papurana Dubois, 1992, and Tylerana Du- bois, 1992, under the provisions of Article 24.2 (``Prin- are plesiomorphic for Ranidae. Although ciple of First Revisor'') of the International Code of some larval characters are thought to be Zoological Nomenclature (ICZN, 1999). common among species of Staurois (tadpole 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 249

Fig. 71. Generic changes suggested for ranid taxa that we studied. This is not exhaustive and the Systematic Comments under Ranidae in ``A Taxonomy of Living Amphibians'' should be consulted for additional taxonomic changes. with deep, cup-like labial parts; upper lip of the large number of ranid species whose oral disc with two continuous rows of papil- adults are morphologically similar to those of lae; lower lip with one broad continuous Staurois, but whose larvae remain unde- band of papillae; Inger, 1966), the diagnostic scribed. value of these characters is unknown due to [274] Hylarana Tschudi, 1838: We asso- 250 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 ciate our exemplars of Hylarana Tschudi, which are reported to not bear humeral 1838 (H. erythraea, the type species, and H. glands33) is done on the assumption that taipehensis), as well as of ``Sylvirana'' some of the molecular apomorphies of this guentheri, with the generic name Hylarana. taxon are synapomorphies of Hydrophylax in Although these two units were assigned, re- the sense of including the species that we did spectively, to the no-humeral-gland (Hylar- not study. On the basis of evidence presented ana) and humeral-gland subsections (Hydro- by Matsui et al. (2005), we place Chalcorana phylax) of Dubois (1992), our data suggest hosii in our Huia. Chalcorana is likely strongly that the humeral gland is convergent broadly polyphyletic, but without evidence in ``S.'' guentheri and Sylvirana (sensu stri- of the remainder's placement we provision- co) or that the presence of the structure has ally regard them as close to Chalcorana been missed in a widespread way because of chalconota, the type-species of Chalcorana. the lack of detailed morphological study (in- We could have retained Chalcorana as a ge- cluding dissections). Hylarana (including nus, but it is clear that, as data emerge, the ``Sylvirana'' guentheri and H. macrodactyla, species in this nominal taxon will be as- the third species of Hylarana sensu Dubois, signed to Hydrophylax, Sylvirana, and likely 1992) lacks dermal glands in the larvae, a others as well. This is not a satisfactory so- character that appears to optimize on the sis- lution to the problem of trying to sort ter branch of Hylarana. The vocal sac con- through this morass, but it is the only prac- dition is variable among species of Hylarana, tical solution available to us at present. with ``S.'' guentheri possessing gular pouch- We retain Humerana Dubois, 1992, and es and H. taipehensis and H. erythraea lack- Pulchrana Dubois, 1992, as nominal genera ing gular pouches. This character is highly only because we did not study these humeral- homoplastic throughout the ranid portion of gland-bearing genera. Future work should our tree. We take the molecular apomorphies test the hypothesis that the remaining species for branch 274 (appendix 5) to be synapo- of the ``humeral-gland group'' constitute a morphies of Hylarana. monophyletic unit. The results of Matsui et We are unable to diagnose Hylarana on al. (2005; ®g. 46) suggest that Humerana ul- the basis of morphology. We did not study, timately will be assigned to Hylarana. and so cannot document, the phylogenetic [280] Sylvirana Dubois, 1992: Our results position of H. macrodactyla. Thus, our as- demonstrate the polyphyly of nominal Syl- sociation of this species with Hylarana re- virana (see discussion of ``S.'' guentheri un- quires testing. Similarly, we do not know der discussion of Hylarana) and the para- which other species may be included in this phyly of the major group of nominal Sylvir- historically ambiguously diagnosed genus. ana (including its type species, S. nigrovit- [278] Hydrophylax Fitzinger, 1843 (in- tata). To remedy the demonstrated polyphyly cluding Amnirana Dubois, 1992, and Chal- of Sylvirana, we transfer ``S.'' guentheri into corana Dubois, 1992): We associate our ex- Hylarana Tschudi, 1838 (see above). To re- emplars of humeral-gland-bearing genera lieve the paraphyly of remaining Sylvirana, (Hydrophylax and Amnirana), as well as the we place Papurana Dubois, 1992, and Ty- imbedded Chalcorana, with the generic lerana Dubois, 1992, into the synonymy of name Hydrophylax Fitzinger, 1843. Chan- Sylvirana Dubois, 1992. Although it is clear ning (2001) had already considered the Af- on the basis of molecular data that ``S.'' rican member of Hydrophylax (H. galamen- guentheri is not in the clade containing S. sis)tobeinAmnirana, along with other Af- nigrovittata (the type species of Sylvirana), rican Hylarana-like frogs. Our association of it is also not clear how many species of nom- the type species of Hydrophylax, H. mala- barica (unstudied by us), with the clade of 33 Possession of humeral glands can be a dif®cult studied terminals requires testing, of course, characteristic to assess due to level of development, and as does the association of the unstudied their presence may be apparent only on dissection. Therefore, any statement that humeral glands are absent members of these nominal taxa. The associ- really requires that a dissection has been made. Dubois ation of unstudied members of Amnirana, (1992) did not mention whether he had made such dis- Hydrophylax, and Chalcorana (some of sections. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 251 inal Sylvirana are associated with ``S.'' Kiyasetuo and Khare, 1986: We provision- guentheri. We take the most falsi®able po- ally retain Sanguirana Dubois, 1992, and sitionÐthat only ``S.'' guentheri is far from Pterorana Kiyasetuo and Khare, 1986 (both Sylvirana nigrovittataÐand suggest that unstudied by us) as genera, owing to the am- careful study is needed. biguous nature of their putative synapomor- Meristogenys Yang, 1991, Clinotarsus Mi- phies (both genera are Hylarana-like forms). vart, 1869, and Nasirana Dubois, 1992: Our Sanguirana sanguinea (type species of San- results place Meristogenys as the sister taxon guirana) has a tadpole with characters shared of Clinotarsus (as found by Roelants et al., with Meristogenys, Clinotarsus, and Altir- 2004; ®g. 35), and far from both Amolops ana: a moderate to high number of labial ker- and Huia, to which it was considered to be atodont rows (4±6/4±5); upper lip with di- closely related by Yang (1991b) and Dubois vided keratodont rows; and dermal glands on (1992). Besides the molecular evidence, Cli- the head and body; and ventral portions of notarsus shares several larval characters with the body and tail ®ns (Alcala and Brown, Meristogenys: (1) dermal glands on the ¯ank; 1982). Pterorana khare (tadpole unknown) is (2) increased numbers of rows of labial ker- distinguished from other ranid frogs by the atodonts (5±9/5±10 in Meristogenys and 6± large, ¯eshy folds on the ¯anks and thighs 8/6±8 in Clinotarsus; over 1±5/2±8 in Amo- and over the vent that extend away from the lops and Huia; Boulenger, 1920: 132±133; body when the frog is under water (Kiyase- Chari, 1962; Yang, 1991b; Hiragond et al., tuo and Khare, 1986). 2001); and (3) upper labial keratodont rows Amolops Cope, 1865, and Amo Dubois, with a medial gap. Unlike Clinotarsus, but 1992: The phylogenetic association of Amo- like Amolops, Huia, and (super®cially) Pseu- lops, Meristogenys, and Huia (Yang, 1991b; doamolops, Meristogenys have a raised ab- Dubois, 1992), as noted in ``Results'' and in dominal sucker in the larvae (Kuramoto et the discussion above of Meristogenys, was al., 1984; Yang, 1991b; Jiang et al., 1997). rejected. Further, the association of Pseu- Clinotarsus lacks the obvious synapomor- doamolops Jiang et al., 1997, suggested by phies associated with Meristogenys (a raised, Kuramoto et al. (1984) and Fei et al. (2000) sharply de®ned abdominal sucker in the lar- is also rejected, suggesting that in each case vae, ribbed jaw sheaths, and upper or both the ventral sucker on the larvae is nonho- jaw sheaths divided (Yang, 1991b). Because mologous and should be considered indepen- most of the species of Meristogenys, like dently apomorphic in each lineage. Kura- most Hylarana-like species (sensu lato), moto et al. (1984) provided morphological have not been sampled and may be involved evidence that the ventral sucker disc on the with this group, we retain both Clinotarsus larvae of Amolops is not homologous with and Meristogenys as genera. that of ``Pseudorana'' sauteri: the edge of Nasirana alticola (not studied by us) may the disc is sharply de®ned in Amolops (not be allied with Clinotarsus, as their larvae so in sauteri); the m. diaphragmatobranchial- share two possible synapomorphies: (1) large is medialis engages the ¯oor of the sucker to size; and (2) supracaudal glands (Grosjean et generate negative pressure in Amolops (mus- al., 2003). Furthermore, Nasirana shares cle does not communicate with sucker in with Meristogenys and Clinotarsus other lar- sauteri); and inframarginal U-shaped band of val characters of uncertain polarity: multiple keratinized material on the sucker in Amo- (3±7) medially divided upper labial kerato- lops (absent in sauteri). Regardless, Kura- dont rows; high numbers of labial keratodont moto et al. (1984) suggested a close relation- rows (7±8: 7±8); and presence of dermal ship of sauteri to Amolops. glands on the ¯anks of the body (Yang, The status of Amo Dubois, 1992 (not stud- 1991b; Hiragond et al., 2001; Grosjean et al., ied by us), is arguable. Dubois (1992) sug- 2003). Nasirana can be distinguished from gested that Amo is unique among Amolops in all other frogs by a ¯eshy prominence on the having axillary glands in both sexes and an snout of the male. As with Clinotarsus,we outer metatarsal tubercle (a character ple- provisionally retain Nasirana as a genus. siomorphic at the base of the ranids), but the Sanguirana Dubois, 1992, and Pterorana outer metatarsal tubercle is nevertheless pre- 252 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 sent in Amolops nepalicus34 and A. torrentis low glands; axillary glands and distal femo- (after Yang, 1991b). Amo lacks the charac- ral glands densely packed, forming a roll; teristics of both Huia and Meristogenys (tibia and intermittent longitudinal ridges, densely elongate; having lateral dermal glands on the covered with small tubercles on the dorsum larvae; high number of larval keratodont (Fei et al., 1991 ``1990''). It shares with Pe- rows on the lower lip) but otherwise shares lophylax a very low number of labial kera- one apomorphy with Amolops (sensu stricto) todont rows in larvae (likely plesiomorphic in our topology: ®rst metacarpal greater than on our topology). Jiang and Zhou (2005; half the length of the second. So, rather than their ®g. 1), with different taxon sampling, suggest that a sucker developed on the venter found Glandirana to be the sister taxon of of the larvae ®ve times in ranids (rather than Rugosa (not studied by us, but placed by Du- the four events currently required by our to- bois, 1992, in his section Pelophylax), and pology) we regard Amo as a synonym of phylogenetically distant from their samples Amolops. of Pelophylax (P. hubeiensis and P. nigro- We found nominal Amolops to be poly- maculata). phyletic (®gs. 50, 65). In this case, the larva Glandirana and Rugosa share the follow- of Amolops chapaensis is unknown (Yang, ing characteristics that appear to be synapo- 1991b), and that species had been assigned morphic (on our tree and on that of Jiang and to Amolops on the basis of having an adult Zhou, 2005): entire body of tadpole covered morphology more similar to Amolops than to in glands; digital discs absent in adults; and Hylarana (i.e., no humeral glands and pres- dorsum densely covered with longitudinal, ence of gular pouches in males; after Inger, tubercular skin ridges in adults (Stejneger, 1966: 257), rather than its having the larval 1907: 123±126; Okada, 1966; Ting and T'sai, synapomorphies of Amolops. We transfer this 1979; Fei et al., 1991 ``1990''; Fei et al., species out of Amolops and into another ge- 2005: 132±138). There are morphological nus below. (See discussion of Huia, Odor- differences between the two genera (Okada, rana, and Eburana). Although we obtain 1966; Fei et al., 1991 ``1990''; Fei et al., Amolops as the sister taxon of Pelophylax, 2005: 132±138; Stejneger, 1907: 123±126; we are unaware of any morphological syna- Ting and T'sai, 1979): sternal cartilage pomorphy uniting these groups (see appen- forked posteriorly in Glandirana [deeply dix 5, branch 287). notched in Rugosa]; toes half-webbed, reach- [288] Pelophylax Fitzinger, 1843: We re- ing the second subarticular tubercle on toe strict the generic name Pelophylax to the IV in Glandirana [fully webbed to beyond subgenus Pelophylax of Dubois (1992). We second subarticular tubercle on toe IV in Ru- are unaware of any morphological synapo- gosa]; skin densely covered in granular yel- morphy for this group, although the molec- low glands, as well as axillary and distal ular data are seemingly decisive (see appen- femoral glands densely packed, forming a dix 5, branch 288). roll in Glandirana [prominent glands only Glandirana Fei, Ye, and Huang, 1991 behind tympanum in Rugosa]). However, ``1990'', and Rugosa Fei, Ye, and Huang, none of these characters is obviously in con- 1991 ``1990'': Glandirana minima is the sole ¯ict with Glandirana ϩ Rugosa forming a species in its nominal genus (formerly a sub- monophyletic group. In light of this evi- genus of the section Hylarana, subsection dence, we recognize this clade as one genus, Hylarana: Dubois, 1992). It is diagnosed by Glandirana, placing Rugosa into synonomy. having skin densely covered in granular yel- Rugosa rugosa, the type species of Rugosa, should be included in subsequent phyloge- 34 Dubois (2000: 331; 2004a: 176) suggested, on the netic analysis to test this hypothesis. basis of examination of the holotype, this taxon is syn- [291] Babina Thompson, 1912, and Nidi- onymous with but did not provide rana Dubois, 1992: Nidirana Dubois, 1992, any discussion regarding the differences itemized in the has been associated with Babina Thompson, original description or the diagnostic differences noted by Yang (1991b). Dubois (2004a: 176) subsequently 1912 (unstudied by us) on the basis of two criticized Anders (2002) for retaining Amolops nepalicus characters: presence of a large suprabrachial without providing a detailed discussion of the issue. gland in breeding-condition males, and egg 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 253 deposition in water-®lled nests of terrestrial raised, sharply de®ned abdominal sucker in burrows or open puddles (Pope, 1931: 536± the tadpole (Yang, 1991b; see discussion of 538; C.-C. Liu, 1950: 258±260; Kuramoto, Meristogenys and Amolops above). Beyond 1985; Dubois, 1992: 154±156; Chou, 1999: this structure, the only characters uniting 398±399). Babina is further diagnosable Huia with Amolops and Meristogenys are from Nidirana on the basis of the male hav- ventral and postorbital glands of the larvae. ing a spine on the prepollex (absent in Ni- None of these characters is present in Odor- dirana; Okada, 1966; Kuramoto, 1985; rana grahami, the only other member of this Chou, 1999). Nidirana, however, has no clade whose tadpole is known. characters that suggest that it is monophylet- We know of no morphological synapo- ic with respect to Babina (Dubois, 1992; morphy that unites this clade (branch 292), Chou, 1999). For this reason, although a sub- but our molecular data are decisive for its genus Babina (the group with the large pre- being a monophyletic group (see appendix pollical spine) could be employed, the name 5). We therefore apply a single generic name. Nidirana applies to no monophyletic group The oldest available name from this group of that can be identi®ed at this time. We there- species is Huia Yang, 1991b (published 18 fore transfer all members of Dubois' subge- February, 1991; the publication containing nus Nidirana to the genus Babina. Odorrana did not appear until at least March [292] Huia Yang, 1992, Odorrana Fei, Ye, of 1991; Fei et al., 1991 ``1990''). We there- and Huang, 1991 ``1990'', Bamburana Fei et fore place ``Amolops'' chapaensis; Eburana al., 2005, ``Amolops'' chapaensis, and Ebur- Dubois, 1992; and Odorrana Fei, Ye, and ana Dubois, 1992: Although our molecular Huang, 1991 ``1990'', into the synonymy of evidence capturing this clade of Himalayan Huia Yang, 1991. and Southeast Asian cascade-dwelling spe- We recognize that this taxonomy is prob- cies is unambiguous (see appendix 5, branch lematic for two reasons. First, we did not in- 292), insuf®cient sampling, the lack of mor- clude any of the types of the nominal genera phological data, and the concomitant taxo- in this study. Thus, the assigned name may nomic confusion surrounding these taxa pre- be inappropriate. Indeed, Huia nasica may sented us with a signi®cant taxonomic chal- not be closely related to Huia cavitympanum lenge. ``Amolops'' chapaensis is embedded Boulenger, 1893 (the type species of Huia in our Huia±Eburana±Odorrana clade, but and not studied by us). The association with its assignment to Amolops was done on the Huia nasica of a tadpole with a raised, sharp- basis of overall similarity (see discussion in ly de®ned abdominal sucker and ventral and Amolops section), and it is clearly not part of postorbital glands of the larvae was based on that genus. There is no known morphological one specimen (C.-C. Liu and Hu, 1961). synapomorphy linking species of Odorrana, Yang (1991b) cast doubt on this assignment as its purported synapomorphy, colorless when he reported that a ``tadpole from Men- spines on chest of the male, is also known in yang assigned to H. nasica by Liu and Hu Huia nasica (B.L. Stuart and Chan-ard, (1961), is certainly Huia even if not larval 2005) and species of at least two other gen- H. nasica''. Our grouping of H. nasica with- era (i.e., some Chalcorana and at least Ba- in a clade of Odorrana and Eburana might bina caldwelli [R. Bain, personal obs.]), and be evidence that nasica is not a member of is absent in many species of Odorrana sensu Huia. And second, our small sample size (4 Fei et al. (1991 ``1990''; see discussion in species, only 2 of which have known tad- ``Review of Current Taxonomy''). Similarly, poles) from this large, undiagnosed group of there is no evidence suggesting that Eburana species (minimum 36 species; Frost, 2004) is monophyletic, because its putative syna- may speak to an oversimpli®cation of the re- pomorphy, unpigmented eggs, is shared by lationships among these taxa. at least some species of three other genera Whereas both of these problems are real (e.g., Chalcorana, Odorrana, Amolops; see concerns, this decision, as with all of our tax- discussion in ``Review of Current Taxono- onomic decisions, is a hypothesis based on my''). Huia (sensu stricto) represents a third the preponderance of the available evidence, example in our tree of convergence of a which we prefer to taxonomic decisions 254 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 based on similarity groupings. As this entire the abdominal suction cup on the larvae. section of former Rana seems to have avoid- This structure is found in sauteri alone, al- ed detailed study, we suggest that a concerted though in a less-developed form than in effort to amass the necessary comparative Amolops, Meristogenys, and Huia (sensu morphological and molecular data is needed, stricto; Jiang et al., 1997). Tanaka-Ueno et and we interpret our results as identifying al. (1998a) suggested on the basis of 587 bp key areas for further study and not as a de- of mtDNA that sauteri is imbedded within cisive resolution of these problems. the brown frog clade (Dubois' subgenus [296] Rana Linnaeus, 1758 (including Au- Rana). Our results corroborate this. Unlike rorana Dubois, 1992, Amerana Dubois, Amolops, Meristogenys, and Huia, both 1992, Pseudoamolops Jiang, Fei, Ye, Zeng, Pseudorana and Pseudoamolops lack dermal Zhen, Xie, and Chen, 1997, and Pseudorana glands on the larvae, which might be a syn- Fei, Ye, and Huang, 1991 ``1990''): To ren- apomorphy, although we do not know the der a monophyletic grouping, we place Pseu- condition of this feature in the Rana tempor- dorana and Pseudoamolops as junior syno- aria group. For our taxonomy, we relegate nyms of Rana, because they are both embed- Pseudoamolops and Pseudorana to the syn- ded within the same clade as Rana tempor- onymy of Rana, which is decisively diag- aria (the type species of Rana). The nosable on the basis of molecular data (ap- abdominal sucker disc of the tadpole of pendix 5, branch 296). Pseudoamolops is not homologous with [301] Lithobates Fitzinger, 1843 (includ- those of Amolops, Huia, and Meristogenys, ing Aquarana Dubois, 1992, Pantherana Du- all of which are distant from each other in bois, 1992, Sierrana Dubois, 1992, Trypher- our tree. opsis Cope, 1868, and ``Rana'' sylvatica): Because Amerana ϩ Aurorana form the Because of the phylogenetic propinquity of sister taxon of our exemplars of a clade with Aquarana Dubois, 1992, Lithobates Fitzin- Rana temporaria, we also place both of these ger, 1843, Pantherana Dubois, 1992, Sier- genera as junior synonyms of Rana (sensu rana Dubois, 1992, Trypheropsis Cope, stricto) to render a monophyletic group. 1868, ``Rana'' sylvatica, and Zweifelia Du- These frogs are unusual among American bois, 1992 (the latter not studied by us, but ``Rana'', but otherwise similar to members placed phylogenetically in this group by Hil- of Rana (sensu stricto) in retaining an outer lis and Wilcox, 2005; ®g. 44), we place these metatarsal tubercle. taxa into their own genus, for which the old- Dubois (1992) recognized Pseudorana as est available name is Lithobates Fitzinger, including Rana sangzhiensis, Rana sauteri, 1843. Therefore, we consider Lithobates to and R. weiningensis, characterized as lacking be a genus, within which we place Aquar- dermal glands in the larvae (likely a syna- ana, Trypheropsis, Sierrana, Zweifelia, and pomorphy at this level of universality) and Pantherana as junior synonyms. Absence of having a labial keratodont row formula of 4± an outer metatarsal tubercle is a morpholog- 7/5±8, an abdominal sucker in the larvae (al- ical synapomorphy. (For species affected by though not as well-developed as in Amo- this nomenclatural change see Frost, 2004, lops), digit I longer than digit II (likely ple- and appendix 7). siomorphy), toe pads present on digit I and We considered recognizing Aquarana for toe IV; metatarsal tubercle present (plesiom- the former R. clamitans/R. catesbeiana orphy), dorsolateral folds present; no gular group; Lithobates for the former R. palmipes pouches in males; and a chevron-shaped group; Pantherana for the R. pipiens group; mark on the anterior dorsum. Subsequently, and Zweifelia for the former R. pustulosa/R. Jiang et al. (1997) partitioned Pseudorana, tarahumarae group. However, this would with P. weiningensis staying in Pseudorana have necessitated naming a new monotypic along with johnsi and sangzhiensis, but sau- genus for Rana sylvatica. Hillis and Wilcox teri being transferred to Pseudoamolops on (2005) also suggested, on the basis of a gen- the basis of several features. The most dis- erally more limited study, but much more tinctive feature is that Pseudorana (contra densely sampled within ``Rana'' than ours, the diagnosis of Dubois, 1992) actually lacks that ``Rana'' sylvatica is the sister taxon of 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 255

Aquarana. We found it to be the sister taxon thanks: Lisa Gugenheim and Merrily Sterns of the (old) Pantherana±Sierrana±Lithoba- (Of®ce of Government Relations) and Diane tes±Typheropsis clade. However, this result Bynum and Barbara Green (Of®ce of Grants is weakly corroborated (due to the variable and Fellowships) were unstinting in their placement of ``R.'' sylvatica; this branch has support. Eleanor Sterling (Center for Biodi- a Bremer value of 1 and jackknife frequency versity and Conservation), initiated and pro- of 52%), and the results of Hillis and Wilcox moted ®eldwork in Vietnam and Bolivia that (2005) therefore deserve further careful con- resulted in the acquisition of many of the sideration. What does seem to be highly cor- valuable tissue samples used in this study. roborated by both our data and those of Hillis Angelique Corthals and Julie Feinstein and Wilcox (2005) is that, excluding the spe- (AMNH Monell Cryo-Collection) cooperat- cies formerly assigned to Amerana and Au- ed in last-minute tissue requests and acces- rorana, all North American species currently sions. Leo Smith provided advice and sup- assigned to Rana form a clade. To recognize port regarding the vagaries of POY. Ho Ling this and to underscore the fact that the spe- Poon (Center for Biodiversity and Conser- cies on the West Coast are more closely re- vation) provided timely assistance with trans- lated to Eurasian species than to other North lations of Chinese literature. Mary DeJong American species, we recognize the com- was invaluable in providing library assis- pletely American group as Lithobates. (See tance. In the AMNH Herpetology Depart- appendix 7 for new combinations and con- ment, Iris Calderon and Dawn Skala dealt tent.) Hillis and Wilcox (2005) provided sev- skillfully with the large demands placed on eral new names for various clades within them by this and related projects. Denny Di- Lithobates, but inasmuch as these were not veley provided extensive editorial and library associated with organismal characteristics support. that purport to delimit them, they are nomina Enormous assistance and encouragement nuda. was also provided from formal and informal reviewers. Maureen Donnelly, David Gower, ACKNOWLEDGMENTS Robert F. Inger, Roy W. McDiarmid, Joseph Mendelson III, Jay M. Savage, and Tom A. We thank the National Aeronautic and Titus read the entire manuscript, caught Space Administration (NASA) for signi®cant many errors, and provided invaluable insight support of computational and molecular bi- and suggestions; their efforts are deeply ap- ology at the American Museum of Natural preciated. Paul Chippindale read the sala- History. This support (NASA grants NAG5- mander sections, caught errors, and provided 12333 and NAG5-8443 to Frost and NAG5- timely advice. Jeffery A. Wilkinson, provid- 13028 to Wheeler) allowed the continued de- ed welcome advice and comments on the velopment of necessary algorithms, the soft- various sections relevant to rhacophorid sys- and hardware for massively parallel compu- tematics. Richard Mayden was a great source tation of large phylogenetic trees, the of counsel and encouragement during the de- large-scale acquisition of molecular data that velopment of this study. elucidate our understanding of the evolution Grant and Frost acknowledge NSF grant and distribution of life on planet Earth, as DEB-0309226, which allowed development well as the student involvement so necessary of many of the primers used in this study and to the success of this venture. This support many of the sequences used both in the sup- went far to assuring the success of this in- ported dendrobatid research as well as in this ternational community project and we are study. During the course of this study Grant deeply grateful. Regardless, any opinions, was supported by an AMNH Graduate Stu- ®ndings, and conclusions or recommenda- dent Fellowship, a Columbia University Cen- tions expressed in this material are those of ter for Environmental Research and Conser- the authors and do not necessarily re¯ect the vation Faculty Fellowship, and NASA grant views of the National Aeronautics and Space NAG5-13028. Administration. Faivovich and Frost acknowledge NSF Many people in the AMNH deserve grant DEB-0407632 which supported devel- 256 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 opment of primers and sequences used in this Haas acknowledges support by the Deut- study as well as the supported hylid research. sche Forschungsgemeinschaft Grant Ha Faivovich acknowledges the timely support 2323/2-1. of a Theodore Roosevelt Memorial grant. De Sa acknowledges NSF grants DEB- During the course of this study Faivovich 0342918 and 9815787 which provided sup- was supported by an AMNH Graduate Stu- port for ®eld work and Leptodactylus re- dent Fellowship and NASA grant NAG5- search that concomitantly furthered the de- 13028. Faivovich and Blotto thank Santiago velopment of this study. Nenda, Guido Corallo, Andres Sehinkman, Campbell gratefully acknowledges the and Diego Baldo for ®eld assistance. support of NSF grants DEB-0102383 and Bain acknowledges NSF grant DEB- 9705277, which allowed the acquisition of 9870232 to the Center for Biodiversity and many of the Middle American and tissue Conservation (CBC/AMNH) for ®nancial samples used in this paper, as well as ®eld support as well as the generous support of and collection assistance by Dwight Law- The John D. and Catherine T. MacArthur son, Brice Noonan, Eric Smith, and Paul Us- Foundation. Collecting and export permits tach. for Vietnam amphibians were granted by the Channing acknowledges the Tanzania Forestry Protection Department, Ministry of Commission for Science and Technology Re- Agriculture and Rural Development, Viet- search Permit 2002-319-ER-99-40, which nam (export permit numbers 31±98, 102±98, provided ®eld support for the collection of 340±99, 341±99, and 174±00). Thanks are genetic samples in Tanzania. also due to Le Xuan Canh, Nguyen Quang Donnellan thanks Michael Mahony (Uni- Truong, Ho Thu Cuc, and Khuat Dang Long versity of Newcastle), Steve Richards (South of the Institute of Ecology and Biological Australian Museum), Allen Allison (Bernice Resources, Hanoi, and Melina Laverty, CBC/ P. Bishop Museum), Dale Roberts (Univer- AMNH, for cooperation and assistance in all sity of Western Australia), Michael Tyler aspects of Vietnam ®eldwork. Tissues of Bo- (University of Adelaide), and Ken Aplin livian amphibians were collected on expedi- (Western Australian Museum) for access to tions supported by the CBC/AMNH and the critical tissues, ®eld support, and courtesies Center for Environmental Research and Con- extended to him that furthered this study. servation at Columbia University, New York, Raxworthy acknowledges NSF grant in collaboration with the Museo de Historia Natural Noel-Kempff Mercado, Santa Cruz, DEB-9984496, National Geographic Society Bolivia, and ColeccioÂn Boliviana de la Fau- grant 5396-94, and grants from Earthwatch na, La Paz. Collection permits for Bolivian which provided ®eld support for acquisition material were granted by el Ministerio de of important genetic samples from Madagas- Desarrollo Sostenible y Plani®cacion de Bo- car. livia. Nussbaum and Raxworthy gratefully ac- Haddad gratefully acknowledges Biota- knowledge support from NSF grants DEB- FAPESP (01/13341-3) and CNPq for ®nan- 9024505, 9322600, and 9625873, which pro- cial support. Exportation permits of Brazilian vided funds for ®eld research and acquisition samples were issued by CITES Lic. 081968 of tissues. BR; AutorizacËoÄes de Acesso e de Remessa Nussbaum acknowledges NSF grants de Amostras de Componentes do PatrimoÃnio DEB-0070485, 9625873, and 9917453, GeneÂtico numbers 02001002851/2004; which provided funds for the acquisition of 02001.002669/2004; permits for collection genetic samples from Madagascar and the were issued by IBAMA/RAN, licencËas 057/ Seychelles. Nussbaum also thanks Greg 03 and 054/05. Haddad thanks the following Schneider for efforts in developing and for ®eld support in the acquisition of relevant maintaining the tissue collections at UMMZ; tissues: Antonio P. Almeida, JoaÄo L. Gapar- and Ronn Altig, Michael J. Pfrender, and Ed- ini, Jose P. Pombal, Jr., Luis O.M. Giasson, mund D. Brodie II, Jr. for help with ®eld MarõÂlia T.A. Hartmann, and Paulo C.A. work in China, Madagascar, SaÄo TomeÂ, and Garcia. Seychelles. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 257

Moler thanks Barry Mansell for ®eld as- SaÄo Paulo, Brazil), JoseÂNuÂnÄez N. (Instituto sistance. de ZoologõÂa, Universidad Austral de Chile, Drewes thanks the NSF for grant DBI- Valdivia, Chile), Wade Ryberg (Washington 9876766, that helped support the CAS frozen University, St. Louis), Elizabeth Scott tissue collection, which proved to be an in- (Transvaal Museum, Pretoria, South Africa), valuable resource for this study. Tom A. Titus (University of Oregon, Eu- Green thanks for funding the National Sci- gene), Jens V. Vindum (California Academy ences and Engineering Research Council of Sciences, San Francisco), David B. Wake (NSERC) of Canada. (Museum of Vertebrate Zoology, University For access to critical tissues and other of California, Berkeley), and Jorge Williams courtesies with respect to this study and (Museo de la Plata, Buenos Aires, Argenti- closely related ones we thank Stevan J. Ar- na). nold (Oregon State University, Corvallis), J.W. Arntzen (National Museum of Natural REFERENCES History, Leiden), Christopher Austin, Robb Abel, O. 1919. Die StaÈmme der Wirbeltiere. Klas- T. Brum®eld, Donna Dittman, and Frederick se Amphibia. Berlin and Leipzig: Walter de Sheldon (Louisiana State University Muse- Gruyter. um of Zoology, Baton Rouge), Boris Blotto Abourachid, A., and D.M. Green. 1999. Origins (Museo Argentino de Ciencias Naturales of the frog kick? Alternate-leg swimming in ``Bernardino Rivadavia'', Buenos Aires, Ar- primitive frogs, families Leiopelmatidae and gentina), Rafe Brown, William E. Duellman, Ascaphidae. Journal of Herpetology 33: 657± and John Simmons (University of Kansas 663. Museum of Natural History and Biodiversity Adler, K.A. 1989. Herpetologists of the past. In K.A. Adler (editor), Contributions to the his- Center, Lawrence), Marius Burger (Univer- tory of herpetology. Contributions to Herpetol- sity of the Western Cape, Bellville, South Af- ogy, no. 5. Ithaca, NY: Society for the Study of rica), Janalee Caldwell (Sam Noble Amphibians and Reptiles: 5±143. Oklahoma Museum of Natural History, Nor- Ahl, E. 1930. Neuere Erkenntnisse uÈber die sys- man), Paul Chippindale, Paul Franklin, Eric tematische Einteilung der Amphibian. Sitzungs- Smith, and Paul Ustach (University of Texas berichte der Gesellschaft Naturforschender at Arlington), Bruce L. Chrisman (Albuquer- Freunde zu Berlin 1930: 78±85. que), Maureen Donnelly (Florida Interna- Alcala, A.C., and W.C. Brown. 1982. Reproduc- tional University, Miami), Robert N. Fisher tive biology of some species of Philautus (Rha- cophoridae) and other Philippine anurans. Phil- and Brian Yang (U.S. Fish & Wildlife Ser- ippine Journal of Biology 11: 203±226. vice, San Diego), Ron Gagliardo (Atlanta Alford, R.A., and S.J. Richards. 1999. Global am- Botanical Garden), Frank Glaw (Zoologische phibian declines: a problem in applied ecology. StaÈatssammlung MuÈnchen), Martin Henzl Annual Review of Ecology and Systematics 30: (Wien, Austria), Caren Goldberg (University 133±165. of Idaho, Moscow), Andrew Holycross (Ar- Allard, M.W., M.M. Miyamoto, L. Jarecki, F. izona State University, Tempe), Robert Inger, Kraus, and M.R. Tennant. 1992. DNA system- Alan Resetar, and Harold Voris (Field Mu- atics and evolution of the artiodactyl family Bovidae. Proceedings of the National Academy seum, Chicago), Cornelya KluÈtsch (Zoolo- of Sciences of the United States of America 89: gisches Forschungsinstitut und Museum Al- 3972±3976. exander Koenig, Bonn, Germany), Dwight Altig, R.I., and G.F. Johnston. 1989. Guilds of an- Lawson (Zoo Atlanta), Karen Lips (Southern uran larvae: relationships among developmen- Illinois University, Carbondale), Steve Gotte, tal modes, morphologies, and habitats. Herpe- W. Ronald Heyer, Roy W. McDiarmid, Rob- tological Monographs 3: 81±109. ert Reynolds, and Addison Wynn (National Altig, R.I., and R.W. McDiarmid. 1999. Diversity: Museum of Natural History, Washington, familial and generic characterizations. In R.W. McDiarmid and R. Altig (editors), Tadpoles: D.C.), Robert W. Murphy (Royal Ontario the biology of anuran larvae: 295±337. Chica- Museum, Toronto, Canada), Brice Noonan go: University of Chicago Press. (Brigham Young University, Provo), Paulo Amiet, J.-L. 1970. Morphologie et deÂveloppement NuõÂn (Museu de Zoologia, Universidade de de la larve de Leptodactylodon ventrimarmor- 258 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

atus (Boulenger) (amphibien, anoure). Annales found in the plains of India. Records of the In- de la Faculte des Sciences du Cameroun. Ya- dian Museum 15: 24±40. ounde 4: 53±71. Anonymous. 1956. Opinion 417. Rejection for Amiet, J.-L. 1971 ``1970''. Les batraciens oro- nomenclatorial purposes of volume 3 (Zoolo- philes du Cameroun. Annales de la Faculte des gie) of the work by Lorenz Oken entitled Sciences du Cameroun. Yaounde 5: 83±102. ``Okens Lehrbuch der Naturgeschichte'' pub- Amiet, J.-L. 1973 ``1972''. Compte rendu d'une lished in 1815±1816. Opinions and Declara- mission batachologique dans le Nord-Came- tions Rendered by the International Commis- roun. Annales de la Faculte des Sciences du sion on Zoological Nomenclature 14: 1±42. Cameroun. Yaounde 12: 63±78. Anonymous. 1977. Opinion 1071. Emendation Amiet, J.-L. 1973. Caracteres diagnostiques de under the plenary powers of Liopelmatina to Petropedetes perreti, nov. sp. et notes sur les Leiopelmatidae (Amphibia, Salientia). Bulletin autres espeÁces camerounaises du genre (amphi- of Zoological Nomenclature 33: 167±169. biens anoures). Bulletin de l'Institut Fondamen- Anonymous. 1990. Opinion 1604. Ichthyophiidae tal d'Afrique Noire, SeÂrie A, Sciences Naturel- Taylor, 1968 (Amphibia, Gymnophiona): con- les 35: 462±474. served. Bulletin of Zoological Nomenclature Amiet, J.-L. 1981. Ecologie, ethologie et devel- 47: 166±167. oppement de Phrynodon sandersoni Parker, Anonymous. 1996. Opinion 1830. Caeciliidae 1939 (Amphibia, Anura, Ranidae). Amphibia- Kolbe, 1880 (Insecta, Psocoptera): spelling Reptilia 2: 1±13. emended to Caeciliusidae, so removing the Amiet, J.-L. 1983. Une espeÁce meconnue de Pe- homonymy with Caeciliidae Ra®nesque, 1814 tropedetes du Cameroun: Petropedetes parkeri (Amphibia, Gymnophiona). Bulletin of Zoolog- n. sp. (Amphibia, Anura, Ranidae, Phrynoba- ical Nomenclature 53: 68±69. trachinae). Revue Suisse de Zoologie 90: 457± Aplin, K.P., and M. Archer. 1987. Recent advanc- 468. es in systematics with a new syn- Amiet, J.-L. 1989. Quelques aspects de la biologie cretic classi®cation. In M. Archer (editor), vol. des amphibiens anoures du Cameroun. AnneÂe 1. Possums and opossums: studies in evolution: Biologique. Paris 28: 73±116. xv±lxxii. Chipping Norton, Australia: Surrey Amiet, J.-L., and J.-L. Perret. 1969. Contributions Beatty. aÁ la faune de la reÂgion de Yaounde (Cameroun Archey, G. 1922. The habitat and life history of II. Amphibiens anoures. Annales de la Faculte Liopelma hochstetteri. Records of the Canter- des Sciences du Cameroun. Yaounde 1969: bury Museum 2: 59±71. 117±137. Archibald, J.D. 1994. Metataxon concepts and as- Anders, C.C. 2002. Class Amphibia (amphibians). sessing possible ancestry using phylogenetic In H.H. Schleich and W. KaÈstle (editors), Am- systematics. Systematic Biology 43: 27±40. phibians and reptiles of Nepal: biology, system- Ardila-Robayo, M.C. 1979. Status sistematico del atics, ®eld guide: 133±340. Ruggell: A.R.G. genero Geobatrachus Ruthven, 1915 (Amphib- Gantner. ia: Anura). Caldasia 12: 383±495. Anderson, J. 1871. A list of the reptilian accession Austin, J.D., S.C. Lougheed, K. Tanner, A.A. to the Indian Museum, Calcutta from 1865 to Chek, J.P. Bogart, and P.T. Boag. 2002. A mo- 1870, with a description of some new species. lecular perspective on the evolutionary af®ni- Journal of the Asiatic Society of Bengal 40: ties of an enigmatic Neotropical frog, Allophry- 12±39. ne ruthveni. Zoological Journal of the Linnean Anderson, J.S. 2001. The phylogenetic trunk: Society. London 134: 335±346. maximal inclusion of taxa with missing data in BaÂez, A.M., and N.G. Basso. 1996. The earliest an analysis of the Lepospondyli (Vertebrata, known frogs of the of South America: Tetrapoda). Systematic Biology 50: 170±193. review and cladistic appraisal of their relation- Andreone, F., M. Vences, D.R. Vieites, F. Glaw, ships. MuÈnchner Geowissenschaftliche Abhan- and A. Meyer. 2004 ``2005''. Recurrent ecolog- dlungen. Reihe A. Geologie und PalaÈontologie ical adaptations revealed through a molecular 30: 131±158. analysis of the secretive cophyline frogs of BaÂez, A.M., and L.A. Pugener. 2003. Ontogeny Madagascar. and Evo- of a new Paleogene pipid frog from southern lution 34: 315±322. South America and xenopodinomorph evolu- Annandale, N. 1918. Some undescribed tadpoles tion. Zoological Journal of the Linnean Society. from the hills of southern India. Records of the London 139: 439±476. Indian Museum 15: 17±23. BaÂez, A.M., and L. Trueb. 1997. Redescription of Annandale, N., and C.R.N. Rao. 1918. The tad- the Paleogene Shelania pascuali from Patagon- poles of the families Ranidae and Bufonidae ia and its bearing on the relationships of fossil 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 259

and Recent pipoid frogs. Scienti®c Papers. Nat- family Pelobatidae. Proceedings of the Zoolog- ural History Museum, University of Kansas 4: ical Society of London 1907: 871±911. 1±41. Beddard, F.E. 1911. Contributions to the anatomy Bain, R.H., A. Lathrop, R.W. Murphy, N.L. Orlov, of the Anura. Proceedings of the Zoological So- and T.C. Ho. 2003. Cryptic species of a cascade ciety of London 1911: 393±412. frog from Southeast Asia: taxonomic revisions Bell, B.D., and R.J. Wassersug. 2003. Anatomical and descriptions of six new species. American features of Leiopelma embryos and larvae: im- Museum Novitates 3417: 1±60. plication for anuran evolution. Journal of Mor- Bain, R.H., and Q.T. Nguyen. 2004. Herpetofauna phology 256: 160±170. diversity of Ha Giang Province in northeastern Benton, M.J. 2000. Stems, nodes, crown clades, Vietnam, with descriptions of two new species. and rank-free lists: is Linnaeus dead? Biologi- American Museum Novitates 3453: 1±42. cal Reviews of the Cambridge Philosophical Baldauf, R.J. 1959. Morphological criteria and Society 75: 633±648. their use in showing bufonid phylogeny. Jour- Berthold, A.A. 1827. Latreille's natuÈrliche Fami- nal of Morphology 104: 527±560. lien der Thierreichs aus dem FranzoÈsischen mit Baldissera, F.A., Jr., R.F. Batistic, and C.F.B. Had- Anmerkungen und ZusaÈtzen von Dr. Arnold dad. 1999. Cytotaxonomic considerations with Adoph Berthold. Weimar: Landes-Industrie the description of two new NOR locations for Comptoir. South American toads, genus Bufo (Anura: Bu- Biju, S.D., and F. Bossuyt. 2003. New frog family fonidae). Amphibia-Reptilia 20: 413±420. from India reveals an ancient biogeographical Barbour, T., and A. Loveridge. 1928. A compar- link with the Seychelles. Nature. London 425: ative study of the herpetological faunae of the 711±714. Uluguru and Usambara Mountains, Tanganyika Bishop, S.C. 1943. Handbook of salamanders. Ith- aca: Comstock Publishing Company. Territory, with descriptions of new species. Me- Blair, W.F. (editor), 1972a. Evolution in the genus moires of the Museum of Comparative Zoology Bufo. Austin: University of Texas Press. 50: 85±265. Blair, W.F. 1972b. Bufo of North and Central Barrio, A. 1954. SistemaÂtica, morfologõÂa y re- America. In W.F. Blair (editor), Evolution in the produccion de (Peters) y genus Bufo: 93±101. Austin: University of Tex- Pseudopaludicola falcipes (Hensel). Physis. as Press. Buenos Aires 20: 379±389. Blair, W.F. 1972c. Evidence from hybridization. In Barrio, A. 1963. Consideraciones sobre compor- W.F. Blair (editor), Evolution in the genus Bufo: tamiento y ``grito agresivo'' propio de algunas 196±243. Austin: University of Texas Press. especies de Ceratophrynidae (Anura). Physis. Blair, W.F. 1972d. Characteristics of the testes. In Buenos Aires 24: 143±148. W.F. Blair (editor), Evolution in the genus Bufo: Barrio, A. 1968. Revision del genero Lepidoba- 324±328. Austin: University of Texas Press. trachus Budgett (Anura, Ceratophrynidae). Blaustein, A.R., and J.M. Kiesecker. 2002. Com- Physis. Buenos Aires 27: 445±454. plexity in conservation: lessons from the global Barrio, A. 1977. Aportes para la elucidacion del decline of amphibian populatons. Ecology Let- ``status'' taxonomico de ters 5: 597±608. Tschudi y Pleurodema kriegi (Muller) (Am- Blommers-SchloÈsser, R.M.A. 1975. Observations phibia, Anura, Leptodactylidae). Physis. Buen- on the larval development of some Malagasy os Aires 37: 311±331. frogs, with notes on their ecology and biology Barrio, A., and P. Rinaldi de Chieri. 1971. Con- (Anura: Dyscophinae, Scaphiophryninae and tribucioÂn al esclarecimiento de la posicioÂn tax- Cophylinae). Beaufortia 24: 7±26. o®leÂtica de algunos batracios argentinos me- Blommers-SchloÈsser, R.M.A. 1976. Chromosomal diante el anaÂlisis cariotõÂpico. Physis. Buenos analysis of twelve species of Microhylidae (An- Aires 30: 673±685. ura) from Madagascar. Genetica 46: 199±210. Barrio-AmoroÂs, C.L. 2004. Amphibians of Vene- Blommers-SchloÈsser, R.M.A. 1993. Systematic zuela. Systematic list, distribution and referenc- relationships of the Mantellinae Laurent 1946 es. Revista de EcologõÂa Latino-Americana 9: 1± (Anura Ranoidea). Ethology Ecology & Evo- 48. lution 5: 199±218. Bauer, L. 1986. A new genus and a new speci®c Blommers-SchloÈsser, R.M.A., and C.P. Blanc. name in the dart frog family (Dendro- 1991. Amphibiens (premiere partie). Faune de batidae, Anura, Amphibia). Ripa. Netherlands Madagascar 75: 1±379. 1986(November): 1±12. Blommers-SchloÈsser, R.M.A., and C.P. Blanc. Beddard, F.E. 1908 ``1907''. Contributions to the 1993. Amphibiens (deuxieme partie). Faune de knowledge of the anatomy of the batrachian Madagascar 75: 385±530. 260 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Bogart, J.P. 1968. Chromosome number differ- phibians as indicators of early Tertiary ``out-of- ence in the amphibian genus Bufo: the Bufo re- India'' dispersal of vertebrates. Science 292: gularis group. Evolution 22: 42±45. 93±95. Bogart, J.P. 2003. Genetics and systematics of hy- Boulenger, G.A. 1882. Catalogue of the Batrachia brid species. In D.M. Sever (editor), Reproduc- Salientia s. Ecaudata in the collection of the tive biology and phylogeny of Urodela (Am- British Museum, 2nd ed. London: Taylor and phibia): 109±134. En®eld, New Hampshire: Francis. Science Publishers, Inc. Boulenger, G.A. 1884. Diagnoses of new reptiles Bogart, J.P., and M. Tandy. 1981. Chromosome and batrachians from the Solomon Islands, col- lineages in African ranoid frogs. Monitore lected and presented to the British Museum by Zoologico Italiano. Nuova Serie, Supplemento H.B. Guppy, Esq., M.B., H.M.S. ``Lark''. Pro- 15: 55±91. ceedings of the Zoological Society of London Boie, H. 1828. Bemerfungen uÈber die Ubtheilun- 1884: 210±213. gen im natuÈrlichen Systeme und deren Char- Boulenger, G.A. 1890. Second report on additions acteristit. Isis von Oken 21: 351±363. to the batrachian collection in the Natural-His- Bokermann, W.C.A. 1965. Notas soÃbre as espeÂ- tory Museum. Proceedings of the Zoological cies de Thoropa Fitzinger (Amphibia, Lepto- Society of London 1890: 323±328. dactylidae). Anais da Academia Brasileira de Boulenger, G.A. 1892 ``1891''. A synopsis of the CieÃncias. Rio de Janeiro 37: 525±537. tadpoles of the European batrachians. Proceed- BolõÂvar-G., W., T. Grant, and L.A. Osorio. 1999. ings of the Zoological Society of London 1891: Combat behavior in and 593±627. other centrolenid frogs. Alytes 16: 77±83. Boulenger, G.A. 1906 ``1905''. Report on the ba- Bolkay, S.J. 1919. Osnove uporedne osteologije trachians collected by the late L. Fea in West anurskih batrahija [Elements of the comparative Africa. Annali del Museo Civico di Storia Na- osteology of the tailless batrachians]. Glasnika turale di Genova. Serie 3, 2: 157±172. Zemaljskog Muzeja u Bosni i Hercegovini. Sa- Boulenger, G.A. 1918. Remarks on the batrachian rejevo 31: 275±357. genera Cornufer, Tschudi, Platymantis, Gthr., Bonacum, J., J. Stark, and E. Bonwich. 2001. Simomantis, g.n., and Staurois, Cope. Annals PCR methods and approaches. In R. DeSalle, and Magazine of Natural History, Series 9, 1: G. Giribet, and W.C. Wheeler (editors), Tech- 372±375. niques in molecular systematics and evolution: Boulenger, G.A. 1920. A monograph of the South 302±328. Boston: BirkhaÈuser. Asian, Papuan, Melanesian and Australian Bonaparte, C.L.J.L. 1839. Iconographia della fau- frogs of the genus Rana. Records of the Indian na italica per le quattro classi degli animali ver- Museum 20: 1±226. tebrati. Tomo II. Amphibi. Fascicolo 26. Roma: Boycott, R.C. 1982. On the taxonomic status of Salviucci. Heleophryne regis Hewitt, 1909 (Anura: Lep- Bonaparte, C.L.J.L. 1840. Prodromus systematis todactylidae). Annals of the Cape Provincial herpetologiae. Nuovi Annali delle Scienze Na- Museums. Natural History 14: 89±108. turali. Bologna 4: 90±101. Bremer, K. 1994. Branch support and tree stabil- Bonaparte, C.L.J.L. 1845. Specchio generale dei ity. 10: 295±304. sistemi erpetologico, an®biologico ed ittiologi- Brongniart, A.T. 1800a. Essai d'une classi®cation co. Milano: Coi Tipi di Luigi di Giacomo Pi- naturelle des reptiles. Iere Partie. Bulletin des rola. Sciences, par La SocieÂte Philomathique 2: 81± Bonaparte, C.L.J.L. 1850. Conspectus systema- 82. tum. Herpetologiae et amphibiologiae. Editio Brongniart, A.T. 1800b. Essai d'une classi®cation altera reformata. Leiden: Brill. naturelle des reptiles. II Partie. Bulletin des Sci- Bonett, R.M., and P.T. Chippindale. 2004. Speci- ences, par La SocieÂte Philomathique 2: 89±91. ation, phylogeography and evolution of life his- Brown, W.C. 1952. The amphibians of the Solo- tory and morphology in plethodontid salaman- mon Islands. Bulletin of the Museum of Com- ders of the Eurycea multiplicata complex. Mo- parative Zoology 107: 3±64. lecular Ecology 13: 1189±1203. Brummitt, R.K. 2002. How to chop up a tree. Tax- Bossuyt, F., and M.C. Milinkovitch. 2000. Con- on 51: 31±41. vergent adaptive radiations in Madagascan and Burton, T.C. 1986. A reassessment of the Papuan Asian ranid frogs reveal covariation between subfamily Asterophryninae (Anura: Microhyli- larval and adult traits. Proceedings of the Na- dae). Records of the South Australian Museum tional Academy of Sciences of the United 19: 405±450. States of America 97: 6585±6590. Burton, T.C. 1998a. Pointing the way: the distri- Bossuyt, F., and M.C. Milinkovitch. 2001. Am- bution and evolution of some characters of the 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 261

®nger muscles of frogs. American Museum Carpenter, J.M. 2003. Critique of pure folly. Bo- Novitates 3229: 1±13. tanical Review 69: 79±92. Burton, T.C. 1998b. Variation in the hand and su- Carroll, R.L. 2000a. The lissamphibian enigma. In per®cial throat musculature of Neotropical lep- H. Heatwole and R.L. Carroll (editors), Am- todactylid frogs. Herpetologica 54: 53±72. phibian biology, vol. 4. , the evo- Burton, T.C. 2004. Muscles and pes of hylid frogs. lutionary history of amphibians: 1270±1273. Journal of Morphology 260: 209±233. Chipping Norton, Australia: Surrey Beatty. Caccone, A., M.C. Milinkovitch, V. Sbordoni, and Carroll, R.L. 2000b. and the origin of J.R. Powell. 1994. Molecular biogeography: us- caecilians. In H. Heatwole and R.L. Carroll (ed- ing the Corsica-Sardinia microplate disjunction itors), Amphibian biology, vol. 4. Paleontology, to calibrate mitochondrial rDNA evolutionary the evolutionary history of amphibians: 1402± rates in mountain newts (Euproctus). Journal of 1411. Chipping Norton, Australia: Surrey Beat- Evolutionary Biology 7: 227±245. ty & Sons. Caldwell, J.P., and C.W. Myers. 1990. A new poi- Carroll, R.L., and P.J. Currie. 1975. Microsaurs as son frog from Amazonian Brazil, with further possible apodan ancestors. Zoological Journal revision of the quinquevittatus group of Den- of the Linnean Society. London 57: 229±247. drobates. American Museum Novitates 2988: Carroll, R.L., A. Kunst, and K. Albright. 1999. 1±21. Vertebral development and amphibian evolu- Cannatella, D.C. 1985. A phylogeny of primitive tion. Evolution and Development 1: 36±48. frogs (Archeobatrachia). Ph.D. dissertation. De- Case, S.M. 1978. Biochemical systematics of partment of Systematics and Ecology, Univer- members of the genus Rana native to western sity of Kansas, Lawrence. North America. Systematic Zoology 27: 299± Cannatella, D.C. 1989. On the monophyly of dis- 311. Cei, J.M. 1970. La posicioÂn ®letica de Telmato- coglossoid frogs. Fortschritte der Zoologie/Pro- biinae, su discusioÂn reciente y signi®cado cri- gress in Zoology 35: 230±231. tico de algunos imunotests. Acta Zoologica Lil- Cannatella, D.C., and D.M. Hillis. 1993. Amphib- loana 27: 181±192. ian relationships: phylogenetic analysis of mor- Cei, J.M. 1972. Bufo of South America. In W.F. phology and molecules. Herpetological Mono- Blair (editor), Evolution in the genus Bufo: 82± graphs 7: 1±7. 92. Austin: University of Texas Press. Cannatella, D.C., and D.M. Hillis. 2004. Amphib- Cei, J.M. 1980. Amphibians of Argentina. Moni- ians: leading a life of slime. In J. Cracraft and tore Zoologico Italiano. Nuova Serie, Monogra- M.J. Donoghue (editors), Assembling the tree phia 2: 1±609. of life: 430±450. Oxford, U.K.: Oxford Uni- Channing, A. 1989. A re-evaluation of the phy- versity Press. logeny of Old World treefrogs. South African Cannatella, D.C., D.M. Hillis, P.T. Chippindale, Journal of Zoology 24: 116±131. L.A. Weigt, A.S. Rand, and M.J. Ryan. 1998. Channing, A. 1995. The relationship between Phylogeny of frogs of the Physalaemus pustu- Breviceps (Anura: Microhylidae) and Hemisus losus species group, with an examination of (Hemisotidae) remains equivocal. Journal of data incongruence. Systematic Biology 47: the Herpetological Association of Africa 44: 311±335. 55±57. Cannatella, D.C., and L. Trueb. 1988. Evolution Channing, A. 2001. Amphibians of central and of pipoid frogs: intergeneric relationships of the southern Africa. Ithaca, NY: Cornell University aquatic frog family Pipidae (Anura). Zoological Press. Journal of the Linnean Society. London 94: 1± Channing, A. 2003. Ghost frogs (Heleophryni- 38. dae). In W.E. Duellman (editor), Grzimek's an- Cantino, P.D., H.N. Bryant, K. de Queiroz, M.J. imal life encyclopedia, 2nd ed. Vol. 6. Am- Donoghue, T. Eriksson, D.M. Hillis, and M.S.Y. phibians: 131±134. Detroit: Gale Group. Lee. 1999. Species names in phylogenetic no- Channing, A., D. Moyer, and M. Burger. 2002. menclature. Systematic Biology 48: 790±807. Cryptic species of sharp-nosed reed frogs in the Cantino, P.D., R.G. Olmstead, and S.J. Wagstaff. complex: advertisement 1997. A comparison of phylogenetic nomencla- call differences. African Zoology 36: 91±99. ture with the current system: a botanical case Channing, A., D.C. Moyer, and K.M. Howell. study. Systematic Biology 46: 313±331. 2002. Description of a new torrent frog in the Caramaschi, U., and J.P. Pombal, Jr. 2001. Bary- genus Arthroleptides from Tanzania (Amphibia, cholos savagei: a junior synonym of Paludicola Anura, Ranidae). Alytes 20: 13±27. ternetzi, with notes on development. Journal of Chari, V.K. 1962. A description of the hitherto Herpetology 35: 357±360. undescribed tadpole of, and some ®eld notes on 262 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

the Rana malabarica (Bibr.). Jour- Paraguay, Vermejo and Uraguay [sic] Rivers, nal of the Bombay Natural History Society 59: by Capt. Thos. J. Page, U.S.N.; and of those 71±76. procured by Lieut. N. Michler, U.S. Top. Eng., Chen, L.Q., R.W. Murphy, A. Lathrop, A. Ngo, Commander of the expedition conducting the N.L. Orlov, T.C. Ho, and I.L.M. Somorjai. survey of the Atrato River. Proceedings of the 2005. Taxonomic chaos in Asian ranid frogs: Academy of Natural Sciences of Philadelphia an initial phylogenetic resolution. Herpetologi- 14: 346±359. cal Journal 15: 231±243. [Not seen prior to De- Cope, E.D. 1863. On Trachycephalus, Scaphiopus cember, 2005.] and other Batrachia. Proceedings of the Acad- Chen, N., J.X. Ma, D.W. Corson, E.S. Hazard, and emy of Natural Sciences of Philadelphia 15: R.K. Crouch. 1996. Molecular of a rho- 43±54. dopsin gene from salamander rods. Investiga- Cope, E.D. 1865. Sketch of primary groups of tions in Ophthalmology and Vision Science 37: Batrachia s. Salientia. Natural History Review. 1907±1913. New Series 5: 97±120. Chippindale, P.T., R.M. Bonett, A.S. Baldwin, and Cope, E.D. 1866. On the structure and distribution J.J. Wiens. 2004. Phylogenetic evidence for a of the genera of the arciferous Anura. Journal major reversal of life-history evolution in pleth- of the Academy of Natural Sciences of Phila- odontid salamanders. Evolution 58: 2809± delphia. Series 2, 6: 67±112. 2822. Cope, E.D. 1867. On the families of the raniform Chou, W.-H. 1999. A new frog of the genus Rana Anura. Journal of the Academy of Natural Sci- (Anura: Ranidae) from China. Herpetologica ences of Philadelphia. Series 3, 6: 189±206. 55: 389±400. Cope, E.D. 1869. A review of the species of Clarke, B.T. 1981. Comparative osteology and Plethodontidae and Desmognathidae. Proceed- evolutionary relationships in the African Rani- ings of the Academy of Natural Sciences of nae (Anura: Ranidae). Monitore Zoologico It- Philadelphia 21: 93±118. aliano. Nuova Serie, Supplemento 15: 285±331. Cope, E.D. 1875. Check-list of North American Clough, M.E., and K. Summers. 2000. Phyloge- Batrachia and Reptilia; with a systematic list of netic systematics and biogeography of the poi- the higher groups, and an essay on geographical son frogs: evidence from mitochondrial DNA distribution based on specimens contained in sequences. Biological Journal of the Linnean the U.S. National Museum. Bulletin of the Society 70: 515±540. United States National Museum 1: 1±104. Cogger, H.G., E.E. Cameron, and H.M. Cogger. Cope, E.D. 1880. Geology and paleontology. 1983. Zoological catalogue of Australia, vol. 1. American Naturalist 14: 609±610. Amphibia and Reptilia. Canberra: Australian Cope, E.D. 1887. The hyoid structure in the am- Government Publishing Service. blystomid salamanders. American Naturalist Colgan, D.J., A. McLauchlan, G.D.F. Wilson, S.P. 21: 88. Livingston, G.D. Edgecombe, J. Macaranas, Cope, E.D. 1889. Batrachia of North America. and G. Cassis. 1999. Histone H3 and U2 sn- Bulletin of the United States National Museum RNA DNA sequences and molecular 34: 5±525. evolution. Australian Journal of Zoology 46: Couper, P. 1992. Hope for our missing frogs. 419±437. Wildlife Australia. Brisbane 1992: 10±11. Collins, J.P., and A. Storfer. 2003. Global am- Crawford, A.J. 2003. Huge populations and old phibian declines: sorting the hypotheses. Di- species of Costa Rican and Panamanian dirt versity and Distributions 9: 89±98. frogs inferred from mitochondrial and nuclear Coloma, L.A. 1995. Ecuadorian frogs of the ge- gene sequences. Molecular Ecology 12: 2525± nus Colostethus (Anura: Dendrobatidae). Uni- 2540. versity of Kansas Museum of Natural History, Crawford, A.J., and E.N. Smith. 2005. Cenozoic Miscellaneous Publications 87: 1±72. biogeography and evolution in direct-develop- Cope, E.D. 1859. On the primary divisions of the ing frogs of Central America (Leptodactylidae: Salamandridae, with descriptions of two new Eleutherodactylus) as inferred from a phylo- species. Proceedings of the Academy of Natu- genetic analysis of nuclear and mitochondrial ral Sciences of Philadelphia 11: 122±128. genes. Molecular Phylogenetics and Evolution Cope, E.D. 1861 ``1860''. Descriptions of reptiles 35: 536±555. from tropical America and Asia. Proceedings of Cunningham, M., and M.I. Cherry. 2000. Mito- the Academy of Natural Sciences of Philadel- chondrial DNA divergence in southern African phia 12: 368±374. bufonids: are species equivalent entities? Afri- Cope, E.D. 1862. Catalogues of the reptiles ob- can Journal of Herpetology 49: 9±22. tained during the explorations of the Parana, Cunningham, M., and M.I. Cherry. 2004. Molec- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 263

ular systematics of African 20-chromosome Annales du MuseÂum d'Histoire Naturelle. Paris toads (Anura: Bufonidae). Molecular Phyloge- 4: 233±296, 4 pl. netics and Evolution 32: 671±685. de Carvalho, M.R., F.A. Bockman, D.S. Amorim, Cuvier, G.L.C.F.D. 1829. Le reÁgne animal distri- M. de Vivo, M. de Toledo-Piza, N.A. Menezes, bue d'apreÁs son organisation, pour servir de J.L. De Figueiredo, R.M.C. Castro, A.C. Gill, base aÁ l'histoire naturelle des animaux et J.D. McEachran, L.J.V. Compagno, R.C. Schel- d'introduction aÁ l'anatomie compareÂe. Nou- ly, R. Britz, J.C. Lundberg, R.P. Vari, and G. velle ed., revue et augmenteÂe par P.A. Latreille, Nelson. 2005. Revisiting the taxonomic imped- vol. 2. Paris: Deterville. iment. Science 307: 353. Daltry, J.C., and G.N. Martin. 1997. Rediscovery De la Riva, I., and J.D. Lynch. 1997. New species of the black narrow mouthed frog, Melanoba- of Eleutherodactylus from Bolivia (Amphibia: trachus indicus Beddome, 1878. Hamadryad Leptodactylidae). Copeia 1997: 151±157. 22: 57±58. Delorme, M., and A. Dubois. 2001. Une nouvelle Darst, C.R., and D.C. Cannatella. 2004. Novel re- espeÁce de Scutiger du Bhutan, et quelques re- lationships among hyloid frogs inferred from marques sur la classi®cation subgeÂneÂrique du 12S and 16S mitochondrial DNA sequences. genre Scutiger (Megophryidae, Leptobrachi- Molecular Phylogenetics and Evolution 31: inae). Alytes 19: 141±153. 462±475. Delorme, M., A. Dubois, S. Grosjean, and A. Ohl- da Silva, H.R. 1998. Phylogenetic relationships of er. 2005. Une nouvelle classi®cation generique the family Hylidae with emphasis on the rela- et subgenerique de la tribu des Philautini (Am- tionships within the subfamily Hylinae (Am- phibia, Anura, Ranidae, Rhacophorinae). Bul- phibia: Anura). Ph.D. dissertation. Department letin Mensuel de la SocieÂte LinneÂenne de Lyon of Sytematics and Ecology, University of Kan- 74: 165±171. Delorme, M., A. Dubois, J. Kosuch, and M. sas, Lawrence. Vences. 2004. Molecular phylogenetic relation- da Silva, H.R., and J.R. Mendelson, III. 1999. A ships of Lankanectes corrugatus from Sri Lan- new organ and sternal morphology in toads ka: endemism of South Asian frogs and the (Anura: Bufonidae): descriptions, taxonomic concept of monophyly in phylogenetic studies. distribution, and evolution. Herpetologica 55: Alytes 22: 53±64. 114±126. del Pino, E.M., and B. Escobar. 1981. Embryonic Davies, M.M. 2003a. Australian ground frogs stages of Gastrotheca riobambae (Fowler) dur- (Limnodynastidae). In W.E. Duellman (editor), ing maternal incubation and comparison with Grzimek's animal life encyclopedia, 2nd ed. development with other marsupial frogs. Jour- Vol. 6. Amphibians: 139±146. Detroit: Gale nal of Morphology 167: 277±295. Group. de Queiroz, K. 1988. Systematics and the Dar- Davies, M.M. 2003b. Australian toadlets and wa- winian Revolution. Philosophy of Science 55: ter frogs (Myobatrachidae). In W.E. Duellman 238±259. (editor), Grzimek's animal life encyclopedia, de Queiroz, K., and J.A. Gauthier. 1992. Phylo- 2nd ed. Vol. 6. Amphibians: 147±154. Detroit: genetic taxonomy. Annual Review of Ecology Gale Group. and Systematics 23: 449±480. Dawood, A., A. Channing, and J.P. Bogart. 2002. de Queiroz, K., and J.A. Gauthier. 1994. Toward A molecular phylogeny of the frog genus To- a phylogenetic system of biological nomencla- mopterna in southern Africa: examining spe- ture. Trends in Ecology and Evolution 9: 27± cies boundaries with mitochondrial 12S rRNA 31. sequence data. Molecular Phylogenetics and de SaÂ, R.O., W.R. Heyer, and A. Camargo. 2005. Evolution 22: 407±413. A phylogenetic analysis of Vanzolinius (Am- De Bavay, J.M. 1993. The developmental stages phibia, Anura, Leptodactylidae): taxonomic and of the , Kyarranus sphagnicol- life history implications. Arquivos do Museu ous Moore (Anura, Myobatrachidae). Austra- Nacional do Rio de Janeiro, In press. lian Journal of Zoology 41: 151±201. de SaÂ, R.O., and D.M. Hillis. 1990. Phylogenetic de Blainville, H.M.D. 1816. Prodrome d'une nou- relationships of the pipid frogs Xenopus and velle distribution systematique du regne animal. Silurana: an integration of ribosomal DNA and Bulletin de la SocieÂte Philomathique de Paris. morphology. Molecular Biology and Evolution Series 3, 3: 113±124. 7: 365±376. de Blainville, H.M.D. 1835. Description de quel- de SaÂ, R.O., and C.C. Swart. 1999. Development ques espeÁces de reptiles de la Californie, preÂ- of the suprarostral plate of pipoid frogs. Journal ceÂdeÂe de l'analyse d'un systeÁme geÂneÂral of Morphology 240: 143±153. d'erpeÂtologie et d'amphibiologie. Nouvelles De Villiers, C.G.S. 1929. The development of a 264 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

species of Arthroleptella from Jonkershoek, Dubois, A. 1981. Liste des genres et sous-genres Stellenbosch. South African Journal of Science nominaux de Ranoidea (amphibiens anoures) 26: 481±510. du monde, avec identi®cation de leurs espeÁces De Villiers, C.G.S. 1931. UÈ ber den SchaÈdelbau types; consequences nomenclaturales. Monitore des . Anatomischer Anzeiger Zoologico Italiano. Nuova Serie, Supplemento 72: 164±178. 15: 225±284. D'Haese, C.A. 2003. Morphological appraisal of Dubois, A. 1982. Les notions de genre, sous- Collembola phylogeny with special emphasis genre et groupe d'espeÁces en zoologie aÁ la lu- on Poduromorpha and a test of the aquatic or- mieÁre de la systeÂmatique eÂvolutive. Monitore igin hypothesis. Zoologica Scripta 32: 563± Zoologico Italiano. Nuova Serie, Supplemento 586. 16: 9±65. Diaz, N.F., J. Valencia, and M. Sallaberry. 1983. Dubois, A. 1983. Classi®cation et nomenclature Life history and phylogenetic relationships of supragenerique des amphibiens anoures. Bul- Insuetophrynus acarpicus (Anura: Leptodactyl- letin Mensuel de la SocieÂte LinneÂenne de Lyon idae). Copeia 1983: 30±37. 52: 270±276. Donnelly, M.A., and M.L. Crump. 1998. Potential Dubois, A. 1984a. Miscellanea nomenclatorica effects of on two Neotropical batrachologica (V). Alytes 3: 111±116. amphibian assemblages. Climatic Change 39: Dubois, A. 1984b. La nomenclature suprageÂneÂ- 541±561. rique des amphibiens anoures. MeÂmoires du Donnelly, M.A., R.O. de SaÂ, and C. Guyer. 1990. MuseÂum National d'Histoire Naturelle. Paris. Description of the tadpoles of Gastrophryne SeÂrie A, Zoologie 131: 1±64. pictiventris and Nelsonophryne aterrima (An- Dubois, A. 1985. Miscellanea nomenclatorica ba- ura: Microhylidae), with a review of morpho- trachologica (VII). Alytes 4: 61±78. logical variation in free-swimming microhylid Dubois, A. 1986 ``1985''. Diagnose preÂliminaire larvae. American Museum Novitates 2796: 1± d'un nouveau genre de Ranoidea (amphibiens, 19. anoures) du sud de l'Inde. Alytes 4: 113±118. Donoghue, M.J. 1985. A critique of the biological Dubois, A. 1987 ``1985''. Miscellanea taxinomica species concept and recommendations for a batrachologica (I). Alytes 5: 7±95. phylogenetic alternative. Bryologist 88: 172± Dubois, A. 1987 ``1986''. Living amphibians of 181. the world: a ®rst step towards a comprehensive Drewery, G.E., and K.L. Jones. 1976. A new ovo- checklist. Alytes 5: 99±149. viviparous frog, Eleutherodactylus jasperi Dubois, A. 1987. Discoglossidae GuÈnther, 1858 (Amphibia, Anura, Leptodactylidae) from (Amphibia, Anura): proposed conservation. Al- . Journal of Herpetology 10: 161± ytes 6: 56±68. 165. Dubois, A. 1988a. Miscellanea nomenclatorica Drewes, R.C. 1984. A phylogenetic analysis of batrachologica (XVII). Alytes 7: 1±5. the Hyperoliidae (Anura): treefrogs of Africa, Dubois, A. 1988b. The genus in zoology: a con- Madagascar, and the Seychelles Islands. Occa- tribution to the theory of evolutionary system- sional Papers of the California Academy of Sci- atics. MeÂmoires du MuseÂum National ences 139: 1±70. d'Histoire Naturelle. Paris. SeÂrie A, Zoologie Drewes, R.C., R.I. Altig, and K.M. Howell. 1989. 140: 1±122. Tadpoles of three frog species endemic to the Dubois, A. 1992. Notes sur la classi®cation des forest of the Eastern Arc Mountains, Tanzania. Ranidae (amphibiens anoures). Bulletin Men- Amphibia-Reptilia 10: 435±443. suel de la SocieÂte LinneÂenne de Lyon 61: 305± Drewes, R.C., and J.A. Wilkinson. 2004. The Cal- 352. ifornia Academy of Sciences Gulf of Guinea Dubois, A. 1995. Keratodont formulae in anuran Expedition (2001) I. The taxonomic status of tadpoles: proposals for a standardization. Jour- the genus Nesionixalus Perret, 1976 (Anura: nal of Zoological Systematics and Evolutionary Hyperoliidae), treefrogs of SaÄo Tome and PrõÂn- Research 33: 1±15. cipe, with comments on the genus Hyperolius. Dubois, A. 1999a. Miscellanea nomenclatorica Proceedings of the California Academy of Sci- batrachologica. 19. Notes on the nomenclature ences 55: 395±407. of Ranidae and related groups. Alytes 17: 81± Dubois, A. 1980. Notes sur la systematique et la 100. repartition des amphibiens anoures de Chine et Dubois, A. 1999b. South Asian Amphibia: a new des regions avoisinantes IV. Classi®cation ge- frontier for taxonomists. Journal of South Asian nerique et subgenerique des Pelobatidae Me- Natural History 4: 1±11. gophryinae. Bulletin Mensuel de la SocieÂte Dubois, A. 2000. The in¯uence of man on the LinneÂenne de Lyon 49: 469±482. distribution of amphibians in the Himalayas of 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 265

Nepal: an example of critical evaluation of bio- todactylidae). In W.E. Duellman (editor), Grzi- geographical data. In G. Miehe and Y. Zhang mek's animal life encyclopedia, 2nd ed. Vol. 6. (editors), Environmental changes in High Asia. Amphibians: 155±171. Detroit: Gale Group. Proceedings of an international symposium at Duellman, W.E., and P. Gray. 1983. Developmen- the University of Marburg, Faculty of Geogra- tal biology and systematics of the egg-brooding phy, 29 May to 1 June 1997, under the auspices hylid frogs, genera Flectonotus and Fritziana. of the Unesco. Marburger Geographische Herpetologica 39: 333±359. Schriften 135: 326±345. Duellman, W.E., and M.S. Hoogmoed. 1984. The Dubois, A. 2003. True frogs (Ranidae). In W.E. taxonomy and phylogenetic relationships of the Duellman (editor), Grzimek's animal life en- hylid frog genus Stefania. University of Kansas cyclopedia, 2nd ed. Vol. 6. Amphibians: 245± Museum of Natural History, Miscellaneous 264. Detroit: Gale Group. Publications 75: 1±39. Dubois, A. 2004a. Amphibians of Nepal: a few Duellman, W.E., and R. Schulte. 1992. Descrip- words of caution. Alytes 21: 174±180. tion of a new species of Bufo from northern Dubois, A. 2004b. The higher nomenclature of Peru with comments on phenetic groups of recent amphibians. Alytes 22: 1±14. South American toads (Anura: Bufonidae). Dubois, A. 2005. Amphibia Mundi. 1.1. An er- Copeia 1992: 162±172. gotaxonomy of Recent amphibians. Alytes 23: Duellman, W.E., and L. Trueb. 1986. Biology of 1±24. amphibians. New York: McGraw-Hill. Dubois, A., and R. GuÈnther. 1982. Klepton and Duellman, W.E., and A. Veloso M. 1977. Phylog- synklepton: two new evolutionary systematics eny of Pleurodema (Anura: Leptodactylidae): a categories in zoology. Zoologische JahrbuÈcher. biogeographic model. Occasional Papers of the Abteilung fuÈr Systematik, OÈ kologie und Geo- Museum of Natural History, University of Kan- graphie. Jena 109: 290±305. sas 64: 1±46. Dubois, A., and A. Ohler. 1998. A new species of DumeÂril, A.H.A. 1863. Catalogue meÂthodique de Leptobrachium (Vibrissaphora) from northern la collection des batraciens du MuseÂum Vietnam, with a review of the taxonomy of the d'Histoire Naturelle de Paris. MeÂmoires de la genus Leptobrachium (Pelobatidae, Megophyi- SocieÂte ImpeÂriale des Sciences Naturelles de nae). Dumerilia 4: 1±32. Cherbourg 9: 295±321. Dubois, A., and A. Ohler. 1999. Asian and Ori- DumeÂril, A.M.C. 1806. Zoologie analytique, ou ental toads of the Bufo melanostictus, Bufo sca- meÂthode naturelle de classi®cation des ani- ber and Bufo stejnegeri groups (Amphibia, An- maux, rendue plus facile aÁ l'dide de tableaux ura): a list of available and valid names and synoptiques. Paris: Allais. redescription of some name-bearing types. Dunn, E.R. 1920. Notes on two Paci®c coast Am- Journal of South Asian Natural History 4: 133± bystomidae. Proceedings of the New England 180. Zoological Club 7: 55±59. Dubois, A., and A. Ohler. 2000. Systematics of Dunn, E.R. 1922. The sound-transmitting appa- Fejervarya limnocharis (Gravenhorst, 1829) ratus of salamanders and the phylogeny of the (Amphibia, Anura, Ranidae) and related spe- Caudata. American Naturalist 56: 418±487. cies. 1. Nomenclatural status and type-speci- Dunn, E.R. 1939. Bathysiredon, a new genus of mens of the nominal species Rana limnocharis salamanders, from Mexico. Notulae Naturae of Gravenhorst, 1829. Alytes 18: 15±50. the Academy of Natural Sciences of Philadel- Dubois, A., and A. Ohler. 2001. A new genus for phia 36: 1. an aquatic ranid (Amphibia, Anura) from Sri Dutta, S.K., K. Vasudevan, M.S. Chaitra, K. Lanka. Alytes 19: 81±106. Shanker, and R.-K. Aggarwal. 2004. Jurassic Dubois, A., A. Ohler, and S.D. Biju. 2001. A new frogs and the evolution of amphibian endemism genus and species of Ranidae (Amphibia, An- in the western Ghats. Current Science. Banga- ura) from south-western India. Alytes 19: 53± lore 86: 211±216. 79. Echeverria, D.D. 1998. Aspectos de la reproduc- Duellman, W.E. 1975. On the classi®cation of cion in-vitro del desarrollo larval de Melano- frogs. Occasional Papers of the Museum of phryniscus stelzneri (Weyenbergh, 1875) (An- Natural History, University of Kansas 42: 1± ura, Bufonidae), con comentarios acerca del or- 14. gano de Bidder. Alytes 15: 158±170. Duellman, W.E. 2001. Hylid frogs of Middle Edwards, J.L. 1976. Spinal nerves and their bear- America, 2nd ed. 2 vol. Contributions to Her- ing on salamander phylogeny. Journal of Mor- petology, no. 18. Ithaca, NY: Society for the phology 148: 305±328. Study of Amphibians and Reptiles. Elias, P., and D.B. Wake. 1983. Nyctanolis pernix, Duellman, W.E. 2003. Leptodactylid frogs (Lep- a new genus and species of plethodontid sala- 266 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

mander from northwestern Guatemala and chia arborõÂcola Allophryne ruthveni Gaige, , Mexico. In A. Rhodin and K. Miyata 1926. Cuadernos de HerpetologõÂa 14: 47±59. (editors), Advances in herpetology and evolu- Faivovich, J. 2002. A cladistic analysis of Scinax tionary biology. Essays in honor of Ernest E. (Anura: Hylidae). Cladistics 18: 367±393. Williams: 1±12. Cambridge, MA: Museum of Faivovich, J., C.F.B. Haddad, P.C.A. Garcia, D.R. Comparative Zoology, Harvard University. Frost, J.A. Campbell, and W.C. Wheeler. 2005. Emerson, S.B., and D. Berrigan. 1993. System- Systematic review of the frog family Hylidae, atics of Southeast Asian ranids: multiple origins with special reference to Hylinae: a phyloge- of voicelessness in the subgenus Limnonectes netic analysis and taxonomic revision. Bulletin (Fitzinger). Herpetologica 49: 22±31. of the American Museum of Natural History Emerson, S.B., R.F. Inger, and D.T. Iskandar. 294: 1±240. 2000a. Molecular systematics and biogeogra- Farris, J.S. 1973. On comparing the shapes of tax- phy of the fanged frogs of Southeast Asia. Mo- onomic trees. Systematic Zoology 22: 50±54. lecular Phylogenetics and Evolution 16: 131± Farris, J.S., A.G. Kluge, and M.F. Mickevich. 142. 1979. Paraphyly of the Rana boylii species Emerson, S.B., C.M. Richards, R.C. Drewes, and group. Systematic Zoology 28: 627±634. K.M. Kjer. 2000b. On the relationships among Farris, J.S., A.G. Kluge, and M.F. Mickevich. ranoid frogs: a review of the evidence. Herpe- 1982a. Phylogenetic analysis, the monothetic tologica 56: 209±230. group method, and myobatrachid frogs. Sys- Estes, R. 1965. Fossil salamanders and salaman- tematic Zoology 31: 317±327. der origins. American Zoologist 5: 319±334. Farris, J.S., A.G. Kluge, and M.F. Mickevich. Estes, R. 1970. New fossil pelobatid frogs and a 1982b. Immunological distance and the phylo- review of the genus . Bulletin of genetic relationships of the Rana boylii species the Museum of Comparative Zoology 139: group. Systematic Zoology 31: 479±491. 239±340. Fatio, V. 1872. Faune des verteÂbreÂs de la Suisse, Estes, R. 1981. Handbuch der PalaÈoherpetologie/ vol. III. Histoire naturelle des reptiles et des Encyclopedia of paleoherpetology. Part 2. batraciens. Paris: J.-P. Bailliere. Gymnophiona, Caudata. Stuttgart: Gustav Fi- Fei, L. 1999. [Atlas of amphibians of China.] scher. Zhengzhou: Henan Press of Science and Tech- Estes, R., K. de Queiroz, and J.A. Gauthier. 1988. nology. [In Chinese.] Phylogenetic relationships within . In R. Estes and G.K. Pregill (editors), Phyloge- Fei, L., and C. Ye. 2000. A new hynobiid subfam- netic relationships of the families, essays ily with a new genus and new species of Hy- commemorating Charles L. Camp: 119±281. nobiidae from West China. Cultum Herpetolo- Stanford, CA: Stanford University Press. gica Sinica 8: 64±70. [In Chinese with English Estes, R., and O.A. Reig. 1973. The early fossil abstract.] record of frogs: a review of the evidence. In J. Fei, L., and C. Ye. 2001. [The color handbook of Vial (editor), Evolutionary biology of the an- the amphibians of Sichuan.] Chengdu, China: urans: contemporary research on major prob- Sichuan Forestry Department, Sichuan Associ- lems: 11±63. Columbia: University of Missouri ation of Wildlife Conservation, and Chengdu Press. Institute of Biology. [In Chinese.] Evans, B.J., R.M. Brown, J.A. McGuire, J. Su- Fei, L., C. Ye, and Y. Huang. 1991 ``1990''. [Key priatna, N. Andayani, A.C. Diesmos, D.T. Is- to Chinese amphibians.] Chongqing, China: kandar, D.J. Melnick, and D.C. Cannatella. Publishing House for Scienti®c and Technolog- 2004. Phylogenetics of fanged frogs: testing ical Literature. [In Chinese.] biogeographical hypotheses at the interface of Fei, L., C. Ye, and Y. Huang. 1997. Taxonomic the Asian and Australian fauna zones. System- studies of the genus Liurana of China including atic Biology 52: 794±819. descriptions of a new species (Amphibia: Ran- Evans, B.J., D.J. Melnick, D.C. Cannatella, J. Su- idae). Cultum Herpetologica Sinica 6±7: 75±80. priatna, N. Andayani, and M.I. Setiadi. 2003. Fei, L., C. Ye, and J. Jiang. 2000. A new genus Monkeys and toads de®ne areas of endemism of the subfamily AmolopinaeÐPseudoamo- on Sulawesi. Evolution 57: 1436±1443. lops, and its relationship to related genera. Acta Evans, S.E., and D. Sigogneau-Russell. 2001. A Zoologica Sinica 46: 19±26. stem-group caecilian (Lissamphibia: Gymno- Fei, L., C. Ye, J. Jiang, X. Feng, and Y. Huang. phiona) from the Lower of North 2005. [An illustrated key to Chinese amphibi- Africa. Palaeontology 44: 259±274 ans.] Chongqing: Sichuan Publishing House of Fabrezi, M., and J.A. Langone. 2000. Los carac- Science and Technology. [In Chinese.] teres morfoloÂgicos del controvertido Neobatra- Fei, L., C. Ye, and C. Li. 2001. Descriptions of 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 267

two new species of the genus Odorrana in Chi- clades of frogs. Herpetological Monographs 7: na. Acta Zootaxonomica Sinica 26: 108±114. 94±117. FejeÂrvaÂry, G.J. 1917. Anoures fossiles des couch- Formas, J.R., and N.D. Espinoza. 1975. Karyo- es preÂglaciares de PuÈspoÈkfuÈrdoÈ en Hongrie. logical relatonships of frogs of the genus Tel- FoÈldtani KoÈzloÈny 47: 141±172. matobufo (Anura: Leptodactylidae). Herpeto- FejeÂrvaÂry, G.J. 1920. Remarques sur la position logica 31: 429±432. systematique des genres Bufavus et Ranavus. Formas, J.R., E. Pugin, and B. Jorquera. 1975. La Annales Historico-Naturales Musei Nationalis identidad del batracio chileno Heminectes rufus Hungarici 18: 28±30. Philippi, 1902. Physis. Buenos Aires 34: 147± FejeÂrvaÂry, G.J. 1921. Kritische Bemerkungen zur 157. Osteologie, Phylogenie und Systematik der An- Frost, D.R. (editor), 1985. Amphibian species of uren. Archiv fuÈr Naturgeschichte. Abteilung A the world. A taxonomic and geographical ref- 87: 1±30. erence. Lawrence, KS: Association of System- FejeÂrvaÂry, G.J. 1923. Ascaphidae, a new family atics Collections and Allen Press. of the tailless batrachians. Annales Historico- Frost, D.R. 2004. Amphibian species of the world: Naturales Musei Nationalis Hungarici 20: 178± an online reference. Version 3.0. New York: 181. American Museum of Natural History. [Elec- Feller, A.E., and S.B. Hedges. 1998. Molecular tronic database available at http://re- evidence for the early history of living amphib- search.amnh.org/herpetology/amphibia/in- ians. Molecular Phylogenetics and Evolution 9: dex.html.] 509±516. Frost, D.R., and A.G. Kluge. 1994. A consider- Fink, W.L. 1990. Review of: The genus in zool- ation of epistemology in systematic biology, ogy: a contribution to the theory of evolution- with special reference to species. Cladistics 10: 259±294. ary systematics, by Alain Dubois. Mem. du Frost, D.R., M.T. Rodrigues, T. Grant, and T.A. Museum national d'Histoire Naturelle, Serie A, Titus. 2001. Phylogenetics of the lizard genus Zoology, Tome 140. 1988. Quarterly Review of Tropidurus (Squamata: Tropiduridae: Tropidur- Biology 65: 79±80. inae): direct optimization, descriptive ef®cien- Fischer von Waldheim, G. 1813. Zoognosia tabulis cy, and sensitivity analysis of congruence be- synopticis illustrata, in usum prñlectionum Aca- tween molecular data and morphology. Molec- demiñ Imperialis Medico-Chirurgicñ Mosquen- ular Phylogenetics and Evolution 21: 352±371. sis edita. 3rd ed. Vol. 1. Moscow: Nicolai Ser- Gadow, H. 1901. The Cambridge natural history, geidis Vsevolozsky. vol. 8. Amphibia and reptiles. London: Mac- Fitzinger, L.J.F.J. 1826. Neue Classi®cation der millan and Co. Reptilien nach ihren natuÈrlichen Verwandt- Gallardo, J.M. 1965. A proposito de los Lepto- schaften nebst einer Verwandtschafts-Tafel und dactylidae (Amphibia, Anura). PapeÂis Avulsos einem Verzeichnisse der Reptilien-Sammlung de Zoologia. SaÄo Paulo 17: 77±87. des k.k. zoologisch Museum's zu Wien. Wien: Gao, K., and N.H. Shubin. 2001. J.G. Heubner. salamanders from northern China. Nature. Lon- Fitzinger, L.J.F.J. 1835. Entwurf einer systema- don 410: 574±577. tischen Anordnung der SchildkroÈten nach den Gao, K., and N.H. Shubin. 2003. Earliest known GrandsaÈtzen der natuÈrlichen Methode. Annalen crown group salamanders. Nature. London 422: des Naturhistorischen Museums in Wien 1: 424±429. 105±128. Gao, K., and Y. Wang. 2001. Mesozoic anurans Fitzinger, L.J.F.J. 1843. Systema reptilium. Fas- from Liaoning Province, China, and phyloge- ciculus primus. Wien: BraumuÈller et Seidel. netic relationships of archaeobatrachian anuran Ford, L.S. 1990. The phylogenetic position of poi- clades. Journal of 21: son-dart frogs (Dendrobatidae): reassessment of 460±476. the neobatrachian phylogeny with commentary GarcõÂa-ParõÂs, M., D.R. Buchholz, and G. Parra- on complex character systems. Ph.D. disserta- Olea. 2003. Phylogenetic relationships of Pe- tion. Department of Systematics and Ecology, lobatoidea re-examined using mtDNA. Molec- University of Kansas, Lawrence. ular Phylogenetics and Evolution 28: 12±23. Ford, L.S. 1993. The phylogenetic position of the GarcõÂa-ParõÂs, M., A. Monton, and M.A. Alonso- dart-poison frogs (Dendrobatidae) among an- Zarazaga. 2004a. ApeÂndice 1. Nomenclatura: urans: an examination of the competing hy- lista de sinoÂnimos y combinaciones. In M. potheses and their characters. Ethology Ecolo- GarcõÂa-ParõÂs, A. Montori, and P. Herrero (edi- gy & Evolution 5: 219±231. tors), Fauna Iberica, vol. 24. Amphibia. Lis- Ford, L.S., and D.C. Cannatella. 1993. The major samphibia: 589±602. Madrid: Museo Nacional 268 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

de Ciencias Naturales and Consejo Superior de primers which have ampli®ed DNA in amphib- Investigaciones Cientõ®ca. ians successfully. Molecular Phylogenetics and GarcõÂa-ParõÂs, M., A. Montori, and P. Herrero. Evolution 11: 163±199. 2004b. Fauna Iberica, vol. 24. Amphibia. Lis- Goin, C.J., O.B. Goin, and G.R. Zug. 1978. Intro- samphibia. Madrid: Museo Nacional de Cien- duction to herpetology, 3rd ed. San Francisco: cias Naturales and Consejo Superior de Inves- W.H. Freeman and Co. tigaciones Cientõ®ca. Goldfuss, G.A. 1820. Handbuch der Zoologie. GarcõÂa-ParõÂs, M., and D.B. Wake. 2000. Molecu- Zweite Abtheilung. NuÈrnburg: Johann Leon- lar phylogenetic analysis of relationships of the hard Schrag. tropical salamander genera Oedipina and No- Goloboff, P.A. 1993±1999. NONA. Version 2.9. totriton, with descriptions of a new genus and TucumaÂn, Argentina: Computer software dis- three new species. Copeia 2000: 42±70. tributed by the author. Gardiner, B.G. 1982. Tetrapod classi®cation. Zoo- Goloboff, P.A. 1999. Analyzing large data sets in logical Journal of the Linnean Society. London reasonable times: solutions for composite opti- 74: 207±232. ma. Cladistics 15: 415±428. Gardner, J.D. 2001. Monophyly and af®nities of Goloboff, P.A., and J.S. Farris. 2001. Methods of albanerpetontid amphibians (Temnospondyli; quick consensus estimation. Cladistics 17: S26± Lissamphibia). Zoological Journal of the Lin- S34. nean Society. London 131: 309±352. Goloboff, P.A., J.S. Farris, and K.C. Nixon. 2003. Gardner, J.D. 2002. Monophyly and intra-generic T.N.T.: Tree analysis using new technology, relationships of (Lissamphibia; version 1.0. Program and documentation avail- Albanerpetontidae). Journal of Vertebrate Pa- able from the authors and at www.zmuc.dk/ leontology 22: 12±22. public/phylogeny. Gauthier, J.A., D.C. Cannatella, K. de Queiroz, Good, D.A., and D.B. Wake. 1992. Geographic A.G. Kluge, and T. Rowe. 1989. Tetrapod phy- variation and speciation in the torrent salaman- logeny. In B. Fernholm, K. Bremer, and H. ders of the genus Rhyacotriton (Caudata: Rhy- Jornvall (editors), The hierarchy of life: 337± acotritonidae). University of California Publi- 353. New York: Elsevier. cations in Zoology 126: 1±91. Gauthier, J.A., A.G. Kluge, and T. Rowe. 1988a. Gower, D.J., A. Kupfer, O.V. Oommen, W. Him- Amniote phylogeny and the importance of fos- stedt, R.A. Nussbaum, S.P. Loader, B. Pres- sils. Cladistics 4: 105±209. Gauthier, J.A., A.G. Kluge, and T. Rowe. 1988b. swell, H. MuÈller, S.B. Krishna, R. Boistel, and The early evolution of the Amniota. In M.J. M. Wilkinson. 2002. A molecular phylogeny of Benton (editor), The phylogeny and classi®ca- ichthyophiid caecilians (Amphibia: Gymno- tion of the tetrapods, vol. 1. Amphibians, rep- phiona: Ichthyophiidae): out of India or out of tiles, birds: 103±155. Systematics Association South East Asia? Proceedings of the Royal So- Special vol. 23. New York: Academic Press. ciety of London. Series B, Biological Sciences Ghiselin, M.T. 1966. On psychologism in the log- 269: 1563±1569. ic of taxonomic controversies. Systematic Zo- Gower, D.J., and M. Wilkinson. 2002. Phallus ology 15: 207±215. morphology in caecilians (Amphibia, Gymno- Giaretta, A.A., and R.J. Sawaya. 1998. Second phiona) and its systematic utility. Bulletin of species of Psyllophryne (Anura: Brachycephal- the Natural History Museum. London. Zoology idae). Copeia 1998: 985±987. Series 68: 143±154. Glaw, F., and M. Vences. 1994. A ®eldguide to Grandison, A.G.C. 1981. Morphology and phy- the amphibians and reptiles of Madagascar, 2nd logenetic position of the West African Didyn- ed., including mammals and freshwater ®sh. amipus sjoestedti Andersson, 1903 (Anura, Bu- KoÈln: Moos Druck. fonidae). Monitore Zoologico Italiano. Nuova Glaw, F., and M. Vences. 2002. A new sibling Serie, Supplemento 15: 187±215. species of the anuran subgenus Grant, T. 1998. Una nueva especie de Colostethus from Madagascar (Amphibia: Mantellidae: del grupo edwardsi de Colombia. Revista de la Mantidactylus) and its molecular phylogenetic Academia Colombiana de Ciencias Exactas, relationships. Herpetological Journal. London Fisicas y Naturales 22: 423±428. 12: 11±20. Grant, T. 2002. Testing methods: the evaluation of Goebel, A.M., J.M. Donnelly, and M.E. Atz. discovery operations in evolutionary biology. 1999. PCR primers and ampli®cation methods Cladistics 18: 94±111. for 12S ribosomal DNA, the control region, cy- Grant, T. 2004. On the identities of Colostethus tochrome oxidase I, and cytochrome b in bu- inguinalis (Cope, 1868) and C. panamensis fonids and other frogs, and an overview of PCR (Dunn, 1933), with comments on C. latinasus 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 269

(Cope, 1863) (Anura: Dendrobatidae). Ameri- frogs of the family Sooglossidae. Herpetologica can Museum Novitates 3444: 1±24. 44: 113±119. Grant, T., and F. Castro-Herrera. 1998. The cloud Green, D.M., T.F. Sharbel, R.A. Hitchmough, and forest Colostethus (Anura, Dendrobatidae) of a C.H. Daugherty. 1989. Genetic variation in the region of the Cordillera Occidental of Colom- genus Leiopelma and relationships to other bia. Journal of Herpetology 32: 378±392. primitive frogs. Zeitschrift fuÈr Zoologische Grant, T., E.C. Humphrey, and C.W. Myers. 1997. Systematik und Evolutionsforschung 27: 65± The median lingual process of frogs: a bizarre 79. character of Old World ranoids discovered in Grif®ths, I. 1954. On the ``otic element'' in Am- South American dendrobatids. American Mu- phibia Salientia. Proceedings of the Zoological seum Novitates 3212: 1±40. Society of London 124: 3±50. Grant, T., and A.G. Kluge. 2004. Transformation Grif®ths, I. 1959a. The phylogenetic status of the series as an ideographic character concept. Cla- Sooglossinae. Annals and Magazine of Natural distics 20: 23±31. History, Series 13, 2: 626±640. Grant, T., and L.O. Rodriguez. 2001. Two new Grif®ths, I. 1959b. The phylogeny of Sminthillus species of frogs of the genus Colostethus (Den- limbatus and the status of the Brachycephalidae drobatidae) from Peru and a redescription of C. (Amphibia Salientia). Proceedings of the Zoo- trilineatus (Boulenger, 1883). American Mu- logical Society of London 132: 457±487. seum Novitates 3355: 1±24. Grif®ths, I. 1963. The phylogeny of the Salientia. Gray, J.E. 1825. A synopsis of the genera of rep- Biological Reviews of the Cambridge Philo- tiles and Amphibia, with a description of some sophical Society 38: 241±292. new species. Annals of Philosophy. Series 2, 10 Grosjean, S., M. Perez, and A. Ohler. 2003. Mor- 193±217. phology and buccopharyngeal anatomy of the tadpole of Rana (Nasirana) alticola (Anura: Gray, J.E. 1850a. Catalogue of the specimens of Ranidae). Raf¯es Bulletin of Zoology 51: 101± Amphibia in the collection of the British Mu- 107. seum. Part. II. Batrachia Gradientia, etc. Lon- Grosjean, S., M. Vences, and A. Dubois. 2004. don: Spottiswoodes and Shaw. Evolutionary signi®cance of oral morphology Gray, J.E. 1850b. Description of a new genus of in the carnivorous tadpoles of tiger frogs, genus batrachians from Swan River. By Dr. H. Schle- Hoplobatrachus (Ranidae). Biological Journal gel, Curator of the Royal Zoological Museum, of the Linnean Society 81: 171±181. Leyden. (Extracted from a letter to J.E. Gray, GuÈnther, A. 1858a. Neue Batrachier in der Samm- Esq.). Proceedings of the Zoological Society of lung des britischen Museums. Archiv fuÈr Na- London 1850: 9±10. turgeschichte 24: 319±328. Graybeal, A. 1995. Phylogenetic relationships and GuÈnther, A. 1858b. On the systematic arrange- evolution of bufonid frogs based on molecular ment of the tailless batrachians and the struc- and morphological characters. Ph.D. disserta- ture of Rhinophrynus dorsalis. Proceedings of tion. Department of Integrative Biology, Uni- the Zoological Society of London 1858: 339± versity of California, Berkeley. 352. Graybeal, A. 1997. Phylogenetic relationships of GuÈnther, A. 1859 ``1858''. Catalogue of the Ba- bufonid frogs and tests of alternate macroevo- trachia Salientia in the collection of the British lutionary hypotheses characterizing their radi- Museum. London: Taylor and Francis. ation. Zoological Journal of the Linnean Soci- GuÈnther, A. 1870. Reptilia (1869). Zoological Re- ety. London 119: 297±338. cord 6: 105±122. Graybeal, A., and D.C. Cannatella. 1995. A new Haas, A. 1995. Cranial features of dendrobatid taxon of Bufonidae from Peru, with descrip- larvae (Amphibia: Anura: Dendrobatidae). tions of two new species and a review of the Journal of Morphology 224: 241±264. phylogenetic status of supraspeci®c bufonid Haas, A. 2003. Phylogeny of frogs as inferred taxa. Herpetologica 51: 105±131. from primarily larval characters (Amphibia: Green, D.M. 2005. The ecology of : Anura). Cladistics 19: 23±90. population ¯uctuation and decline in amphibi- Haas, A., and S.J. Richards. 1998. Correlations of ans. Biological Conservation 111: 331±343. cranial morphology, ecology, and evolution in Green, D.M., and D.C. Cannatella. 1993. Phylo- Australian suctorial tadpoles of the genera Li- genetic signi®cance of the amphicoelous frogs, toria and Nyctimystes (Amphibia: Anura: Hy- Ascaphidae and Leiopelmatidae. Ethology lidae: Pelodryadinae. Journal of Morphology Ecology & Evolution 5: 233±245. 238: 109±141. Green, D.M., R.A. Nussbaum, and D. Yang. 1988. Haddad, C.F.B., and C.P.A. Prado. 2005. Repro- Genetic divergence and heterozygosity among ductive modes in frogs and their unexpected di- 270 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

versity in the of Brazil. Bio- Hedges, S.B., R.A. Nussbaum, and L.R. Maxson. Science 55: 207±217. 1993. Caecilian phylogeny and biogeography Haeckel, E.H.P.A. 1866. Generelle Morphologie inferred from mitochondrial DNA sequences of der Organismen, vol. 2. Berlin: Georg Reimer. the 12S rRNA and 16S rRNA genes (Amphib- Hall, J.A., and J.H. Larsen, Jr. 1998. Postembry- ia: Gymnophiona). Herpetological Monographs onic ontogeny of the spadefoot toad, Scaphio- 7: 64±76. pus intermontanus (Anura: Pelobatidae): skel- Hedges, S.B., and L.L. Poling. 1999. A molecular etal morphology. Journal of Morphology 238: phylogeny of reptiles. Science 283: 998±1001. 179±244. Hennig, W. 1966. Phylogenetic systematics. Chi- Halliday, T. 2005. Diverse factors in¯uencing am- cago: University of Illinois Press. phibian population declines. In M.J. Lannoo Henrici, A.C. 1991. Chelomophrynus bayi (Am- (editor), Amphibian declines: the conservation phibia, Anura, Rhinophrynidae), a new genus status of United States species: 3±9. Berkeley: and species from the Middle Eocene of Wyo- University of California Press. ming: ontogeny and relationships. Annals of Hallowell, E. 1856. Description of several species the Carnegie Museum 60: 97±144. of Urodela, with remarks on the geographical Hertwig, S., R.O. de SaÂ, and A. Haas. 2004. Phy- distribution of the Caducibranchiata division of logenetic signal and the utility of 12S and 16S these and their classi®cation. Proceed- mtDNA in frog phylogeny. Journal of Zoolog- ings of the Academy of Natural Sciences of ical Systematics and Evolutionary Research 42: Philadelphia 8: 6±11. 2±18. Hanken, J., and D.B. Wake. 1982. Genetic differ- Hewitt, J. 1919. Anhydrophryne rattrayi, a re- entiation among plethodontid salamanders (ge- markable new frog from Cape Colony. Record nus Bolitoglossa) in Central and South Ameri- of the Albany Museum. Grahamstown 3: 182± ca: implications for the South American inva- 189. sion. Herpetologica 38: 272±287. Heyer, W.R. 1970. Studies on the frogs of the ge- Harris, D.J. 2001. Reevaluation of 16S ribosomal nus Leptodactylus (Amphibia: Leptodactyli- RNA variation in Bufo (Anura: Amphibia). Mo- dae). VI. Biosystematics of the melanonotus lecular Phylogenetics and Evolution 19: 326± group. Natural History Museum of Los Angeles 329. County Contributions in Science 191: 1±48. Hass, C.A., J.F. Dunski, L.R. Maxson, and M.S. Heyer, W.R. 1975. A preliminary analysis of the Hoogmoed. 1995. Divergent lineages within intergeneric relationships of the frog family the Bufo margaritifera complex (Amphibia: Leptodactylidae. Smithsonian Contributions in Anura; Bufonidae) revealed by albumin im- Zoology 199: 1±55. munology. Biotropica 27: 238±249. Heyer, W.R. 1998. The relationships of Leptodac- Hay, J.M., I. Ruvinsky, S.B. Hedges, and L.R. tylus diedrus (Anura, Leptodactylidae). Alytes Maxson. 1995. Phylogenetic relationships of 16: 1±24. amphibian families inferred from DNA se- Heyer, W.R. 1999. A new genus and species of quences of mitochondrial 12S and 16S ribo- frog from Bahia, Brazil (Amphibia: Anura: somal RNA genes. Molecular Biology and Leptodactylidae) with comments on the zoo- Evolution 12: 928±937. geography of the Brazilian campos rupestres. Hayes, M.P., and P.H. Starrett. 1981 ``1980''. Proceedings of the Biological Society of Wash- Notes on a collection of centrolenid frogs from ington 112: 19±30. the Colombia Choco. Bulletin of the Southern Heyer, W.R. 2003. Viewpoint: -B and California Academy of Science 79: 89±96. Amphibia. BioScience 53: 540±541. Hedges, S.B. 1989. Evolution and biogeography Heyer, W.R., and R.I. Crombie. 1979. Natural his- of West Indian frogs of the genus Eleuthero- tory notes on Craspedoglossa stejnegeri and dactylus: slow-evolving loci and the major Thoropa petropolitana (Amphibia: Salientia, groups. In C.A. Woods (editor), Biogeography Leptdactylidae). Journal of the Washington of the West Indies: past, present, and future: Academy of Sciences 69: 17±20. 305±370. Gainesville, FL: Sandhill Heyer, W.R., and D.S.S. Liem. 1976. Analysis of Press. the intergeneric relationships of the Australian Hedges, S.B. 1994. Molecular evidence for the frog family Myobatrachidae. Smithsonian Con- origin of birds. Proceedings of the National tributions in Zoology 233: 1±29. Academy of Science of the United States of Highton, R. 1997. Geographic protein variation America 91: 2621±2624. and speciation in the Plethodon dorsalis com- Hedges, S.B., and L.R. Maxson. 1993. A molec- plex. Herpetologica 53: 345±356. ular perspective on lissamphibian phylogeny. Highton, R. 1998. Is Ensatina eschscholtzii a ring Herpetological Monographs 7: 27±42. species? Herpetologica 54: 254±278. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 271

Highton, R. 1999. Geographic protein variation dae). Zoologische Verhandelingen. Leiden 250: and speciation in the salamanders of the Pleth- 1±32. odon cinereus group with the description of Hoogmoed, M.S., and J. Lescure. 1984. A new two new species. Herpetologica 55: 43±90. genus and two new species of minute leptodac- Highton, R., and R.B. Peabody. 2000. Geographic tylid frogs from northern South America, with protein variation and speciation in salamanders comments upon Phyzelaphryne (Amphibia: An- of the Plethodon jordani and Plethodon gluti- ura: Leptodactylidae). Zoologische Mededelin- nosus complexes in the southern Appalachian gen. Leiden 58: 85±115. Mountains with the description of four new Houlahan, J.E., C.S. Findlay, B.R. Schmidt, A.H. species. In R.C. Bruce, R.G. Jaeger, and L.D. Meyer, and S.L. Kuzmin. 2000. Quantitative Houck (editors), The biology of plethodontid evidence for global amphibian population de- salamanders: 31±93. New York: Kluwer Aca- clines. Nature. London 404: 752±755. demic/Plenum Publishers. Hu, S., E. Zhao, and C.-C. Liu. 1966. A herpe- Hillis, D.M., and S.K. Davis. 1986. Evolution of tological survey of the Tsinling and Ta-Pa Shan ribosomal DNA: ®fty million years of recorded region. Acta Zoologica Sinica 18: 57±89. [In history in the frog genus Rana. Evolution 40: Chinese with English abstract.] 1275±1288. Hu, S., E. Zhao, and C.-C. Liu. 1973. A survey Hillis, D.M., and M.T. Dixon. 1991. Ribosomal of amphibians and reptiles in Kweichow prov- DNA: molecular evolution and phylogenetic in- ince, including a herpetofaunal analysis. Acta ference. Quarterly Review of Biology 66: 411± Zoologica Sinica 19: 149±181. 453. Hull, D.L. 1988. Science as a process: an evolu- Hillis, D.M., and T.P. Wilcox. 2005. Phylogeny of tionary account of the social and conceptual de- the New World true frogs (Rana). Molecular velopment of science. Chicago: University of Phylogenetics and Evolution 34: 299±314. Chicago Press. Hiragond, N.C., B.A. Shanbhag, and S.K. Saida- Hutchinson, M.N., and L.R. Maxson. 1987. Phy- pur. 2001. Description of the tadpole of a logenetic relationships among Australian tree stream breeding frog, Rana curtipes. Journal of frogs (Anura: Hylidae: Pelodryadinae): an im- Herpetology 35 166±168. munological approach. Australian Journal of Hoffman, A.C. 1932. Researches relating to the Zoology 35: 61±74. validity of the South African Polypedatidae Huxley, T.H. 1863. Description of Anthracosau- (Rhacophoridae) as an autonomous family of rus russelli, a new labyrinthodont from the the Anura. South African Journal of Science Lanarkshire coal ®eld. Quarterly Journal of the 29: 562±583. Geological Society of London 19: 56±68. Hoffman, A.C. 1935. Die sistematiese posiesie Inger, R.F. 1954. Systematics and zoogeography van Heleophryne. SooÈlogiese Navorsing van of Philippine Amphibia. Fieldiana. Zoology 33: die Nasionale Museum. Bloemfontein 1: 1±2. 183±531. Hoffmann, C.K. 1878. Die Klassen und Ordnun- Inger, R.F. 1966. The systematics and zoogeog- gen des Thier-Reichs wissenschaftlich darges- raphy of the Amphibia of Borneo. Fieldiana. telldt in Wort und Bild, vol. 6 (part 2). Klassen Zoology 52: 1±402. und Ordnungen der Amphibien wissenschaf- Inger, R.F. 1967. The development of a phylogeny tlich dargestelldt in Wort un Bild. Leipzig and of frogs. Evolution 21: 369±384. Heidelberg: C.F. Winter. Inger, R.F. 1972. Bufo of Eurasia. In W.F. Blair Hogg, J. 1838. On the classi®cation of the Am- (editor), Evolution in the genus Bufo: 102±118. phibia. Annals of Natural History 1: 152. Austin: University of Texas Press. Hogg, J. 1839a. On the classi®cation of the Am- Inger, R.F. 1996. Commentary on a proposed clas- phibia. Magazine of Natural History, New Se- si®cation of the family Ranidae. Herpetologica ries 3: 265±274. 52: 241±246. Hogg, J. 1839b. On the classi®cation of the Am- Inger, R.F., H.B. Shaffer, M. Koshy, and R. Bakde. phibia (continued from page 274). Magazine of 1984. A report on a collection of amphibians Natural History, New Series 3: 367±378. and reptiles from the Ponmudi, , South Hoogmoed, M.S. 1989a. South American bufon- India. Journal of the Bombay Natural History ids (Amphibia: Anura: Bufonidae), an enigma Society 81: 406±427. for taxonomists. Treballs d'Ictiologia i Herpe- International Commission of Zoological Nomen- tologia 2: 167±180. clature. 1999. International code of zoological Hoogmoed, M.S. 1989b. On the identity of some nomenclature, 4th ed. London: International toads of the genus Bufo from Ecuador, with ad- Trust for Zoological Nomenclature. ditional remarks on Andinophryne colomai Iordansky, N.N. 1996. Evolution of the muscula- Hoogmoed, 1985 (Amphibia: Anura: Bufoni- ture of the jaw apparatus in the Amphibia. Ad- 272 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

vances in Amphibian Research in the Former tionships among Chinese ranids inferred from Soviet Union 1: 3±26. sequence data set of 12S and 16S rDNA. Her- Iwabe, N., Y. Hara, Y. Kumazawa, K. Shibamoto, petological Journal. London 15: 1±8. Y. Saito, T. Miyata, and K. Kazutaka. 2005. Sis- JimeÂnez de la Espada, M. 1871 ``1870''. Fauna ter-group relationship of turtles to the bird-croc- neotropicalis species quaedam nondum cogni- odilian clade revealed by nuclear DNA-coded tae. Jornal de ScieÃncias, MathemaÂticas, Physi- . Molecular Biology and Evolution 22: cas e Naturaes. LisboÄa 3: 57±65. 810±813. Jockusch, E.L., and D.B. Wake. 2002. Falling Izecksohn, E. 1971. NoÃvo geÃnero e nova espeÂcie apart and merging: diversi®cation of slender de Brachycephalidae do Estado do Rio de Ja- salamanders (Plethodontidae: Batrachoseps)in neiro, Brasil (Amphibia, Anura). Boletim do the American West. Biological Journal of the Museu Nacional, Rio de Janeiro. Nova SeÂrie, Linnean Society 76: 361±391. Zoologia 280: 1±12. Jockusch, E.L., K.P. Yanev, and D.B. Wake. 2001. Izecksohn, E. 1988. Algumas consideracËoÄes soÃbre Molecular phylogenetic analysis of slender sal- ogeÃnero Euparkerella, coma descricËaÄo de treÃs amanders, genus Batrachoseps (Amphibia: novas espeÂcies (Amphibia, Anura, Leptodactyl- Plethodontidae), from central coastal California idae). Revista Brasileira de Biologia 48: 59±74. with descriptions of four new species. Herpe- Izecksohn, E., J.J. Jim, S.T. de Albuquerque, and tological Monographs 15: 54±99. W.F. de MendoncËa. 1971. ObservacËoÄes soÃbre o Jungfer, K.-H., H. Birkhahn, V. KuÈlpmann, and K. desenvolvimento e os haÂbitos de Myersiella Wassmann. 1996. Haltung und Fortp¯anzung subnigra (Miranda-Ribeiro). Arquivos do Mu- von Dendrobates fulguritus Silverstone, 1975, seu Nacional. Rio de Janeiro 43: 69±73. mit Anmerkungen zur Gattung Minyobates My- Jackman, T.R., G. Applebaum, and D.B. Wake. ers, 1987. Herpetofauna. Weinstadt 15: 19±27. 1997. Phylogenetic relationships of bolitoglos- Jungfer, K.-H., S. LoÈtters, and D. JoÈrgens. 2000. sine salamanders: a demonstration of the effects Der kleinste PfeilgiftfroscheÐeine neue Den- of combining morphological and molecular drobates-Art aus West-Panama. Herpetofauna. data sets. Molecular Biology and Evolution 14: Weinstadt 22: 11±18. 883±891. Jurgens, J.D. 1971. The morphology of the nasal Janies, D., and W.C. Wheeler. 2001. Ef®ciency of region of the Amphibia and its bearing on the parallel direct optimization. Cladistics 17: S71± phylogeny of the group. Annals of the Univer- S82. sity of Stellenbosch 46: 1±146. Janke, A., D. Erpenbeck, M. Nilsson, and U. Ar- Kaiser, H., L.A. Coloma, and H.M. Gray. 1994. nason. 2001. The mitochondrial genomes of the A new species of Colostethus (Anura: Dendro- iguana (Iguana iguana) and the caiman (Cai- batidae) from Martinique, French Antilles. Her- man crocodylus): implications for amniote phy- petologica 50: 23±32. logeny. Proceedings of the Royal Society of Kaplan, M. 1994. Analysis of some long-standing London. Series B, Biological Sciences 268: controversies concerning the pectoral girdle of 623±631. Atelopus (Bufonidae) using ontogenetic studies. Jenkins, F.A., and D.M. Walsh. 1993. An early Journal of Herpetology 28: 128±131. Jurassic caecilian with limbs. Nature. London Kaplan, M. 1995. On the presence of overlap dur- 365: 246±250. ing the development of the pectoral girdle of Jiang, J., A. Dubois, A. Ohler, A. Tillier, X. Chen, Colostethus subpunctatus (Amphibia: Anura) F. Xie, and M. StoÈck. 2005. Phylogenetic rela- and its relevance in the classi®cation of Den- tionships of the tribe Paini (Amphibia, Anura, drobatidae. Journal of Herpetology 29: 300± Ranidae) based on partial sequences of mito- 304. chondrial 12S and 16S rRNA genes. Zoological Kaplan, M. 1997. A new species of Colostethus Science. Tokyo 22: 353±362. from the Sierra Nevada de Santa Marta (Co- Jiang, J., L. Fei, C. Ye, X. Zeng, M. Zhen, F. Xie, lombia) with comments on intergeneric rela- and Y. Chen. 1997. Studies on the taxonomics tionships within the Dendrobatidae. Journal of of species of Pseudorana and discussions of the Herpetology 31: 369±375. phylogenetical relationships with its relative Kaplan, M. 2000. The pectoral girdles of Rana genera. Cultum Herpetologica Sinica 6±7: 67± rugulosa (Ranidae) and Nesomantis thomasseti 74. (Sooglossidae). Herpetologica 56: 188±195. Jiang, J., and K. Zhou. 2001. Evolutionary rela- Kaplan, M. 2001. On the relevance of the char- tionships among Chinese ranid frogs inferred acter ``absence of epicoracoid horns'' in the from mitochondrial DNA sequences of 12S systematics of anurans. Alytes 19: 196±204. rRNA gene. Acta Zoologica Sinica 47: 38±44. Kaplan, M. 2002. Histology of the anteroventral Jiang, J., and K. Zhou. 2005. Phylogenetic rela- part of the breast-shoulder apparatus of Bra- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 273

chycephalus ephippium (Brachycephalidae) Knauer, F.K. 1883. Naturgeschichte der Lurche. with comments on the validity of the genus (Amphibiologie.) Eine umfassendere Darle- Psyllophryne (Brachycephalidae). Amphibia- gung unserer Kenntnisse von dem anatomisch- Reptilia 23: 225±227. en Bau, der Entwicklung und systematischen Kaplan, M. 2004. Evaluation and rede®nition of Eintheilung der Amphibien, & c. Zweite Aus- the states of anuran pectoral girdle architecture. gabe. Wien and Leipzig: A. Pilchler's Witwe & Herpetologica 60: 84±97. Sohn. Keller, R.A., R.N. Boyd, and Q.D. Wheeler. 2003. KoÈhler, J. 2000. New species of Eleutherodactylus The illogical basis of Phylogenetic Nomencla- (Anura: Leptodactylidae) from cloud forest of ture. Botanical Review 69: 93±110. Bolivia. Copeia 2000: 516±520. King, M., M.J. Tyler, M.M. Davies, and D. King. Kojima, J.-I. 2003. Apomorphy-based de®nition 1979. Karyotypic studies on Cyclorana and as- also pinpoints a node, and PhyloCode names sociated genera of Australian frogs. Australian prevent effective communication. Botanical Journal of Zoology 27: 699±708. Review 69: 44±58. Kirsch, J.A., A.W. Dickerman, O.A. Reig, and Kokubum, M.N., de Carvalho, and A.A. Giaretta. M.S. Springer. 1991. DNA hybridization evi- 2005. Reproductive ecology and behaviour of dence for the Australasian af®nity of the Amer- a species of Adenomera (Anura, Leptodactyli- ican marsupial Dromiciops australis. Proceed- nae) with endotrophic tadpoles: systematic im- ings of the National Academy of Sciences of plications. Journal of Natural History. London the United States of America 88: 10465±10469. 39: 1745±1758. Kirtisinghe, P. 1958. Some hitherto undescribed Kosuch, J., M. Vences, A. Dubois, A. Ohler, and anuran tadpoles. Ceylon Journal of Science 1: W. BoÈhme. 2001. Out of Asia: mitochondrial 171±176. DNA evidence for an oriental origin of tiger Kiyasetuo and M.K. Khare. 1986. A new genus frogs, genus Hoplobatrachus. Molecular Phy- of frog (Anura: Ranidae) from Nagaland at the logenetics and Evolution 21: 398±407. northeastern hills of India. Asian Journal of Ex- Kress, W.J., and P. DePriest. 2001. What's in a ploration and Science 1: 12. Phylocode name? Science 292: 6. Kjer, K.M. 1995. Use of rRNA secondary struc- Kuramoto, M. 1985. A new frog (genus Rana) ture in phylogenetic studies to identify homol- from the Yaeyama Group of the Ryukyu Is- ogous positions: an example of alignment and lands. Herpetologica 41: 150±158. data presentation from frogs. Molecular Phy- Kuramoto, M. 1990. A list of chromosome num- logenetics and Evolution 4: 314±330. bers of anuran amphibians. Bulletin of Fukuoka Klein, S.L., R.L. Strausberg, L. Wagner, J. Pon- University of Education 39: 83±127. tius, S.W. Clifton, and P. Richardson. 2002. Ge- Kuramoto, M., and C.-S. Wang. 1987. A new rha- netic and genomic tools for Xenopus research: cophorid treefrog from Taiwan, with compari- the NIH Xenopus initiative. Developmental Dy- sons to Chirixalus eif®ngeri (Anura, Rhaco- namics 225: 384±391. phoridae). Copeia 1987: 931±942. Klemens, M.W. 1998. The male nuptial charac- Kuramoto, M., C.-S. Wang, and H.-T. Yu. 1984. teristics of Arthroleptides martiensseni Nieden, Breeding, larval morphology and experimental an endemic torrent frog from Tanzania's East- hybridization of Taiwanese brown frogs, Rana ern Arc Mountains. Herpetological Journal. longicrus and R. sauteri. Journal of Herpetol- London 8: 35±40. ogy 18: 387±395. Kluge, A.G. 1966. A new pelobatine frog from Kwon, A.S., and Y.H. Lee. 1995. Comparative the lower of South Dakota with a dis- spermatology of anurans with special referenc- cussion of the evolution of the Scaphiopus- es to phylogeny. In B.G.M. Jamieson, J. Ausio, Spea complex. Natural History Museum of Los and J.-L. Justine (editors), Advances in sper- Angeles County Contributions in Science 113: matozoal phylogeny and taxonomy. MeÂmoires 1±26. du MuseÂum National d'Histoire Naturelle. Par- Kluge, A.G. 2005. Taxonomy in theory and prac- is. SeÂrie A, Zoologie 166: 321±332. tice, with arguments for a new phylogenetic La Marca, E., M. Vences, and S. LoÈtters. 2002. system of taxonomy. In M.A. Donnelly, B.I. Rediscovery and mitochondrial relationships of Crother, C. Guyer, M.H. Wake, and M.E. White the dendrobatid frog Colostethus humilis sug- (editors), Ecology and evolution in the tropics: gest parallel colonization of the Venezuelan a herpetological perspective: 7±47. Chicago: Andes by poison frogs. Studies on Neotropical University of Chicago Press. Fauna and Environment 37: 233±240. Kluge, A.G., and J.S. Farris. 1969. Quantitative Lamotte, M. 1961. Contribution aÁ l'eÂtude des ba- phyletics and the evolution of anurans. System- traciens de l'ouest africain XII. Les formes lar- atic Zoology 18: 1±32. vaires de Cardioglossa leucomystax Blgr. Bul- 274 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

letin de l'Institut Fondamental d'Afrique Noire of Urodela (Amphibia): 31±108. En®eld, NH: 23: 211±216. Science Publishers. Lamotte, M., and J. Lescure. 1989. Les teÃtards Lataste, F. 1879. E tude sur le Discoglossus pictus rheÂophiles et hygropeÂtriques de l'Ancien et du Otth. Actes de la SocieÂte LinneÂenne de Bor- Noveau Monde. Annales des Sciences Naturel- deaux, SeÂrie 4, 3: 275±342. les. Zoologie. Paris. Serie 13, 10: 111±122. Lathrop, A. 1997. Taxonomic review of the me- Lamotte, M., J.-L. Perret, and S. Dzieduszycka. gophryid frogs (Anura: Pelobatoidea). Asiatic 1959. Contribution aÁ l'eÂtude des batraciens de Herpetological Research 7: 68±79. l'ouest africain. IX. Les formes larvaires de Pe- Lathrop, A. 2003. Asian toadfrogs (Megophryi- tropedetes palmipes, Conraua goliath et Acan- dae). In W.E. Duellman (editor), Grzimek's an- thixalus spinosus. Bulletin de l'Institut Fran- imal life encyclopedia, 2nd ed. Vol. 6. Am- cËaise d'Afrique Noire. SeÂrie A, Sciences Na- phibians: 109±117. Detroit: Gale Group. turelles 21: 762±776. Latreille, P.A. 1800. Histoire naturelle des sala- Lamotte, M., and F. Xavier. 1972. Les amphibiens mandres de France, preÂceÂdeÂe d'un tableau anoures a developpement direct d'Afrique. Ob- meÂthodique des autres reptiles indigeÁnes. Paris: servations sur la biologie de Nectophrynoides Imprimerie de Crapelet. tornieri (Roux). Bulletin de la SocieÂte Zoolo- Latreille, P.A. 1825. Familles naturelles du reÁgne gique de France 97: 413±428. animal, exposeÂes succinctement et dans un or- Lamotte, M., and M. Zuber-Vogeli. 1954. Contri- dre analytique, avec l'indication de leurs bution aÁ l'eÂtude des batraciens de l'ouest afri- genres. Paris: J.B. BaillieÁre. cain. III. Les formes larvaires de Astylosternus Laurent, R.F. 1941 ``1940''. Contribution aÁ diadematus et Petropedetes natator. Bulletin de l'osteÂologie et aÁ la systeÂmatique des ranides af- l'Institut FrancËaise d'Afrique Noire. SeÂrie A, ricains. PremieÁre note. Revue de Zoologie et de Sciences Naturelles 16: 1222±1233. Botanique Africaines 34: 74±96. Langone, J.A. 1994. Ranas y sapos del Uruguay Laurent, R.F. 1941. Contribution aÁ l'osteÂologie et (reconocimientos y aspectos bioloÂgicos). Mu- aÁ la systematique des ranides africains. Deux- seo Damaso Antonio LarranÄaga, Serie de Di- ieÁme note. Revue de Zoologie et de Botanique vulgacioÂn 5: 1±123. Africaines 34: 192±234. Largen, M.J. 1991. A new genus and species of Laurent, R.F. 1942. Note sur les procoeliens ®r- petropedetine frog (Amphibia, Anura, Ranidae) misternes (Batrachia Anura). Bulletin du MuseÂe from high altitude in the mountains of Ethiopia. Royal d'Histoire Naturelle de Belgique 18: 1± Tropical Zoology. Firenze 4: 139±152. 20. Largen, M.J., and R.C. Drewes. 1989. A new ge- Laurent, R.F. 1943. Contribution a l'osteologie et nus and species of brevicipitine frog (Amphib- a la systematique des rhacophorides non afri- ia, Anura, Microhylidae) from high altitude in caines. Bulletin du MuseÂe Royal d'Histoire Na- the mountains of Ethiopia. Tropical Zoology. turelle de Belgique 19: 1±16. Firenze 2: 13±30. Laurent, R.F. 1946. Mises au point dans la tax- Larson, A. 1991. A molecular perspective on the onomie des ranides. Revue de Zoologie et de evolutionary relationship of the salamander Botanique Africaines 39: 336±338. families. In M.K. Hecht, B. Wallace, and R.J. Laurent, R.F. 1951. Sur la necessite de supprimer MacIntyre (editors), Evolutionary biology, vol. la famille des Rhacophoridae mais de creÂer cel- 25: 211±277. New York: Plenum Publishing le des Hyperoliidae. Revue de Zoologie et de Corporation. Botanique Africaines 45: 116±122. Larson, A., and W. Dimmick. 1993. Phylogenetic Laurent, R.F. 1954. Remarques sur le genre relationships of the salamander families: an Schoutedenella. Annales du MuseÂe Royal du analysis of congruence among morphological Congo Belge. Nouvelle SeÂrie in Quarto. Sci- and molecular characters. Herpetological ences Zoologiques. Tervuren 1: 34±40. Monographs 7: 77±93. Laurent, R.F. 1961. Notes on some South African Larson, A., D.B. Wake, L.R. Maxson, and R. amphibians. Publications de l'UniversiteÂde Highton. 1981. A molecular phylogenetic per- l'E tat aÁ Elisabethville. Lubumbashi 1: 197±209. spective on the origins of morphological nov- Laurent, R.F. 1972. The morphology, systematics, elties in the salamanders of the tribe Pletho- and evolution of the Old World treefrogs (Rha- dontini (Amphibia, Plethodontidae). Evolution cophoridae and Hyperoliidae) [Review]. Cop- 35: 405±422. eia 1972: 198±201. Larson, A., D.W. Weisrock, and K.H. Kozak. Laurent, R.F. 1973. The natural classi®cation of 2003. Phylogenetic systematics of salamanders the Arthroleptinae (Amphibia, Hyperoliidae). (Amphibia: Urodela), a review. In D.M. Sever Revue de Zoologie et de Botanique Africaines (editor), Reproductive biology and phylogeny 87: 666±678. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 275

Laurent, R.F. 1975. BiogeÂographie et liaisons in- Lavilla, E.O., and J.M. Cei. 2001. Amphibians of tercontinentales au course du MeÂsozoõÈque. MeÂ- Argentina. A second update, 1987±2000. Mu- moires du MuseÂum National d'Histoire Natu- seo Regionale di Scienze Naturali Monogra®e. relle. Paris. SeÂrie A, Zoologie 88: 176±191. Torino 28: 1±177. Laurent, R.F. 1978. L'appareil hyoidien des As- Lawson, D.P. 1993. The reptiles and amphibians tylosterninae et des Arthroleptinae (Amphibia). of Korup National Park Project, . Revue Zoologique Africaine 92: 233±240. Herpetological Natural History 1: 27±90. Laurent, R.F. 1980 ``1979''. Esquisse d'une phy- Lebedkina, N.S. 2004. Evolution of the amphibian logeneÁse des anoures. Bulletin de la SocieÂte skull. Advances in Amphibian Research in the Zoologique de France 104: 397±422. Former Soviet Union 9: 1±265. Laurent, R.F. 1983. Heterogeneidad del genero Lescure, J., V. Marty, C. Marty, F. Starace, M.A. Batrachophrynus Peters (Leptodactylidae). Thomay, and F. Letellier. 1995. Contribution aÁ Acta Zoologica Lilloana 37: 107±113. l'eÂtude des amphibiens de Guyane francËaise. X. Laurent, R.F. 1984a. Heterogeneidad de la familia Les Phyllomedusa (Anura, Hylidae). Revue Caeciliidae (Amphibia-Apoda). Acta Zoologica FrancËaise d'Aquariologie, HerpeÂtologie 22 35± Lilloana 37: 199±200. 50. Laurent, R.F. 1984b. La phylogenese des Rano- Lescure, J., S. Renous, and J.-P. Gasc. 1986. Prop- idea et le cladisme. Alytes 3: 97±111. osition d'une nouvelle classi®cation des am- Laurent, R.F. 1986. Sous classe des lissamphi- phibiens gymnophiones. MeÂmoires de la SocieÂ- biens (Lissamphibia). In P. Grasse and M. Del- te Zoologique de France 43: 145±177. sol (editors), Traite de zoologie. Anatomie, sys- Leuckart, F.S. 1821. Einiges uÈber die ®schartigen tematique, biologie, vol. 14. Batraciens, fasc. 1- Amphibien. Isis von Oken 9: 257±265. B: 594±797. Paris: Masson. Li, W.-H., and Z.-Y. Wang. 1985. Karyotype of Laurent, R.F. 1986 ``1985''. Sur la classi®cation Rana livida. Acta Herpetologica Sinica 4: 56. et la nomenclature des amphibiens. Alytes 4: [In Chinese with English abstract.] 119±120. Licht, L.E. 2003. Shedding light on ultraviolet ra- Laurent, R.F., and M. Fabrezi. 1986 ``1985''. Le diation and amphibian embryos. BioScience carpe des Arthroleptinae. Alytes 4: 85±93. 53: 551±561. Laurenti, J.N. 1768. Specimen medicum, exhibens Liem, D.S.S. 1970. The morphology, systematics, synopsin reptilium emendatum cum experimen- and evolution of the Old World treefrogs (Rha- tis circa venena et antidota Reptilium Austria- corum. Wien. cophoridae and Hyperoliidae). Fieldiana. Zo- Laurin, M. 1998a. The importance of global par- ology 57: i±vii, 1±145. simony and historical bias in understanding tet- Linnaeus, C. 1758. per regna tria rapod evolution. Part I. Systematics, middle ear naturae, secundum classes, ordines, genera, evolution and jaw suspension. Annales des Sci- species, cum characteribus, differentiis, synon- ences Naturelles. Zoologie et Biologie Anima- ymis, locis, 10th ed. Stockholm. le. Paris. Serie 13, 1998: 1±42. Lips, K.R., P.A. Burrowes, J.R. Mendelson III, Laurin, M. 1998b. The importance of global par- and G. Parra-Olea. 2005. Amphibian declines simony and historical bias in understanding tet- in Latin America: a synthesis. Biotropica 37: rapod evolution. Part II. Vertebral centrum, cos- 222±228. tal ventilation, and paedomorphosis. Annales Littlejohn, M.J. 1963. The breeding biology of the des Sciences Naturelles. Zoologie et Biologie Philoria frosti Spencer. Pro- Animale. Paris. Serie 13, 1998: 99±114. ceedings of the Linnaean Society of New South Laurin, M. 1998c. A reevaluation of the origin of Wales 88: 273±276. pentadactyly. Evolution 52: 1476±1482. Liu, C.-C. 1950. Amphibians of western China. Laurin, M. 2002. Tetrapod phylogeny, amphibian Fieldiana. Zoology Memoires 2: 1±397 ϩ 10 origins, and the de®nition of the name Tetra- pl. poda. Systematic Biology 51: 364±369. Liu, C.-C., and S. Hu. 1961. [Tailless amphibians Laurin, M., and R.R. Reisz. 1997. A new per- of China.] Shanghai: Science Press. [In Chi- spective on tetrapod phylogeny. In S.S. Sumida nese.] and K.L.M. Martin (editors), Amniote origins: Liu, C.-C., and S. Hu. 1962. A herpetological re- completing the transition to land: 9±59. San Di- port of Kwangsi. Acta Zoologica Sinica 14: 73± ego: Academic Press. 104. [In Chinese with English abstract and Laurin, M., R.R. Reisz, and M. Girondot. 2000. translations of descriptions.] Caecilian viviparity and amniote origins: a re- Liu, W., A. Lathrop, J. Fu, D. Yang, and R.W. ply to Wilkinson and Nussbaum. Journal of Murphy. 2000. Phylogeny of East Asian bufon- Natural History. London 34: 311±315. ids inferred from mitochondrial DNA sequenc- 276 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

es (Anura: Amphibia). Molecular Phylogenetics search on major problems: 133±182. Columbia: and Evolution 14: 423±435. University of Missouri Press. Loader, S.P., D.J. Gower, K.M. Howell, N. Dog- Lynch, J.D. 1975. A review of the Andean lep- gart, M.-O. RoÈdel, R.O. de SaÂ, B.L. Cohen, and todactylid frog genus Phrynopus. Occasional M. Wilkinson. 2004. Phylogenetic relationships Papers of the Museum of Natural History, Uni- of African microhylid frogs inferred from DNA versity of Kansas 35: 1±51. sequences of mitochondrial 12S and 16S rRNA Lynch, J.D. 1976. The species groups of the South genes. Organisms, Diversity & Evolution 4: American frogs of the genus Eleutherodactylus 227±235. (Leptodactylidae). Occasional Papers of the Lombard, R.E., and S.S. Sumida. 1992. Recent Museum of Natural History, University of Kan- progress in understanding early tetrapods. sas 61: 1±24. American Zoologist 32: 609±622. Lynch, J.D. 1978a. A new eleutherodactyline frog Lombard, R.E., and D.B. Wake. 1986. Tongue from the Andes of northern Colombia. Copeia evolution in the lungless salamanders, family 1978: 17±21. Plethodontidae IV. Phylogeny of plethodontid Lynch, J.D. 1978b. A re-assessment of the tel- salamanders and the evolution of feeding dy- matobiine leptodactylid frogs of PatagoÂnia. Oc- namics. Systematic Biology 35: 532±551. casional Papers of the Museum of Natural His- Loveridge, A. 1957. Check list of the reptiles and tory, University of Kansas 72: 1±57. amphibians of East Africa (Uganda; Kenya; Lynch, J.D. 1980. A new species of Barycholos Tanganyika; Zanzibar). Bulletin of the Museum from Estado GoiaÂs, Brasil (Amphibia, Anura, of Comparative Zoology 117: 153±362. Leptodactylidae) with remarks on related gen- LoÈytynoja, A., and M.C. Milinkovitch. 2000. era. Bulletin du Museum National d'Histoire SOAP, cleaning multiple alignments from un- Naturelle. Paris. Section A, Zoologie, Biologie stable blocks, version 1.0. Bioinformatics 17: et Ecologie Animales 2: 289±302. 573±574. Lynch, J.D. 1982a. Two new species of poison- LoÈytynoja, A., and M.C. Milinkovitch. 2003. A dart frogs (Colostethus) from Colombia. Her- hidden Markov model for progressive multiple petologica 38: 366±374. alignments. Bioinformatics 19: 1505±1513. Lynch, J.D. 1982b. Relationships of the frogs of Lutz, B. 1954. AnfõÂbios anuros do Distrito Fed- the genus Ceratophrys (Leptodactylidae) and eral/The frogs of the Federal District of Brazil. their bearing on hypotheses of Pleistocene for- MemoÂrias do Instituto Oswaldo Cruz. Rio de est refugia in South America and punctuated Janeiro 52: 155±197 (Portuguese), 219±238 equilibrium. Systematic Zoology 31: 166±179. (English). Lynch, J.D. 1986. The de®nition of the Middle Lutz, B. 1968. Taxonomy of the Neotropical Hy- American clade of Eleutherodactylus based on lidae. Pearce-Sellards Series. Texas Memorial jaw musculature (Amphibia: Leptodactylidae). Museum 11: 3±26. Herpetologica 42: 248±258. Lutz, B. 1969. AdaptacËoÄes, especializacËoÄes e lin- Lynch, J.D. 1989. Intrageneric relationships of hagens nos anuros neotropicais. Acta Zoologica mainland Eleutherodactylus (Leptodactylidae). Lilloana 24: 267±292. I. A review of the frogs assigned to the Eleuth- Lynch, J.D. 1969. Program. Final Ph.D. exami- erodactylus discoidalis species group. Milwau- nation. Department of Zoology, University of kee Public Museum Contributions in Biology Kansas, Lawrence. and Geology 79: 1±25. Lynch, J.D. 1971. Evolutionary relationships, os- Lynch, J.D. 1994. A new species of high-altitude teology, and zoogeography of leptodactyloid frog (Eleutherodactylus: Leptodactylidae) from frogs. University of Kansas Museum of Natural the Cordillera Oriental of Colombia. Revista de History, Miscellaneous Publications 53: 1±238. la Academia Colombiana de Ciencias Exactas, Lynch, J.D. 1972a. Generic partitioning of the FõÂsicas y Naturales 19: 195±203. South American leptodactyloid frog genus Eup- Lynch, J.D. 1996. The relationships of the His- sophus Fitzinger, 1843 (sensu lato). Bulletin of paniolan frogs of the subgenus Pelorius the Southern California Academy of Science (Eleutherodactylus: Leptodactylidae). In R. 71: 2±11. Powell and R.W. Henderson (editors), Contri- Lynch, J.D. 1972b. Two new species of frogs butions to West Indian herpetology: a tribute to (Eleutherodactylus: Leptodactylidae) from the Albert Schwartz: 141±155. Contribution to paÂramos of northern Ecuador. Herpetologica Herpetology, no. 12. Ithaca, NY: Society for the 28: 141±147. Study of Amphibians and Reptiles. Lynch, J.D. 1973. The transition from archaic to Lynch, J.D. 2000. The relationships of an ensem- advanced frogs. In J.L. Vial (editor), Evolution- ble of Guatemalan and Mexican frogs (Eleuth- ary biology of the anurans: contemporary re- erodactylus: Leptodactylidae: Amphibia). Re- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 277

vista de la Academia Colombiana de Ciencias dae: Plethodontini): phylogenetic analysis of an Exactas, FõÂsicas y Naturales 24: 129±156. old and rapid radiation. Molecular Phylogenet- Lynch, J.D., and W.E. Duellman. 1997. Frogs of ics and Evolution 18: 174±188. the genus Eleutherodactylus in western Ecua- Maienschein, J. 1994. Cutting edges cut both dor. Systematics, ecology, and biogeography. ways. Biology and Philosophy 9: 1±24. University of Kansas Natural History Museum, Mannen, H., and S.S.-L. Li. 1999. Molecular ev- Special Publication 23: 1±236. idence for a clade of turtles. Molecular Phylo- Lynch, J.D., and H.L. Freeman. 1966. Systematic genetics and Evolution 13: 144±148. status of a South American frog, Allophryne Manzano, A.S., and E.O. Lavilla. 1995. Myolog- ruthveni Gaige. University of Kansas Publica- ical peculiarities in Rhinoderma darwini (An- tions, Museum of Natural History 17: 493±502. ura: Rhinodermatidae). Journal of Morphology Lynch, J.D., and R.W. McDiarmid. 1987. Two 224: 125±129. new species of Eleutherodactylus (Amphibia: Manzano, A.S., S.A. Moro, and V. Abdala. 2003. Anura: Leptodactylidae) from Bolivia. Proceed- The depressor mandibulae muscle in Anura. ings of the Biological Society of Washington Alytes 20: 93±131. 100: 337±346. Marmayou, J., A. Dubois, A. Ohler, E. Pasquet, Lynch, J.D., and P.M. Ruiz-Carranza. 1982. A and A. Tillier. 2000. Phylogenetic relationships new genus and species of poison-dart frog in the Ranidae (Amphibia, Anura). Independent (Amphibia: Dendrobatidae) from the Andes of origin of direct development in the genera Phi- northern Colombia. Proceedings of the Biolog- lautus and Taylorana. Comptes Rendus de ical Society of Washington 95: 557±562. l'Academie des Sciences. Series 3, Life Scienc- Lynch, J.D., and P.M. Ruiz-Carranza. 1997 es Paris 323: 287±297. ``1996''. A remarkable new centrolenid frog Martin, A.A. 1967. Australian anuran life histo- from Colombia with a review of nuptial ex- ries: some evolutionary and ecological aspects. crescences in the family. Herpetologica 52: In A.H. Weatherly (editor), Australian inland 525±535. and their fauna: eleven studies: 175± Lynch, J.F., and D.B. Wake. 1978. A new species 191. Canberra: Australian National University of Chiropterotriton (Amphibia: Caudata) from Press. Baja Verapaz, Guatemala, with comments on Martin, R.F. 1972. Evidence from osteology. In relationships among Central American mem- W.F. Blair (editor), Evolution in the genus Bufo: bers of the genus. Natural History Museum of 37±70. Austin: University of Texas Press. Los Angeles County Contributions in Science Martin, W.F. 1972. Evolution of vocalization in 294: 1±22. the genus Bufo. In W.F. Blair (editor), Evolution Lynch, J.F., D.B. Wake, and S.-Y. Yang. 1983. in the genus Bufo: 279±309. Austin: University Genic and morphological differentiation in of Texas Press. Mexican Pseudoeurycea (Caudata: Plethodon- Matsui, M. 1994. A taxonomic study of the Rana tidae). Copeia 1983: 884±894. narina complex, with description of three new Macey, J.R., J.L. Strasburg, J.A. Brisson, V.T. species (Amphibia: Ranidae). Zoological Jour- Vredenburg, M. Jennings, and A. Larson. 2001. nal of the Linnean Society. London 111: 385± Molecular phylogenetics of western North 415. American frogs of the Rana boylii species Matsui, M., and N.L. Orlov. 2004. A new species group. Molecular Phylogenetics and Evolution of Chirixalus from Vietnam (Anura: Rhaco- 19: 131±143. phoridae). Zoological Science. Tokyo 21: 671± Macey, J.R. 2005. Plethodontid salamander mi- 676. tochondrial genomics: a parsimony evaluation Matsui, M., H. Ota, M.W.-N. Lau, and A. Boga- of character con¯ict and implications for his- dek. 1995. Cytotaxonomic studies of three ra- torical biogeography. Cladistics 21: 194±202. nid species (Amphibia: Anura) from Hong Maglia, A.M. 1998. Phylogenetic relationships of Kong. Japanese Journal of Herpetology/ Ha- extant pelobatoid frogs (Anura: Pelobatoidea): chu-Ryoseiruigaku Zasshi 16: 12±17. evidence from adult morphology. Scienti®c Pa- Matsui, M., T. Shimada, H. Ota, and T. Tanaka- pers. Natural History Museum, University of Ueno. 2005. Multiple invasions of the Ryukyu Kansas 10: 1±19. Archipelago by Oriental frogs of the subgenus Maglia, A.M., L.A. Pugener, and L. Trueb. 2001. Odorrana with phylogenetic reassessment of Comparative development of anurans: using the related subgenera of the genus Rana. Mo- phylogeny to understand ontogeny. American lecular Phylogenetics and Evolution 37: 733± Zoologist 41: 538±551. 742. [Seen as an electronic/pdf preprint avail- Mahoney, M.J. 2001. Molecular systematics of able from the publisher.] Plethodon and Aneides (Caudata: Plethodonti- Maxson, L.R., R. Highton, and D.B. Wake. 1979. 278 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Albumin evolution and its phylogenetic impli- Fraser and H.-D. Sues (editors), In the shadow cations in the plethodontid salamander genera of the dinosaurs: 5±23. New York: Cambridge Plethodon and Ensatina. Copeia 1979: 502± University Press. 508. Milner, A.R. 2000. Mesozoic and Tertiary Cau- Maxson, L.R., D.P. Ondrula, and M.J. Tyler. 1985. data and Albanerpetontidae. In H. Heatwole An immunological perspective on evolutionary and R.L. Carroll (editors), Amphibian biology, relationships in Australian frogs of the hylid vol. 4. Paleontology, the evolutionary history genus Cyclorana. Australian Journal of Zoolo- of amphibians: 1412±1444. Chipping Norton, gy 33: 17±22. Australia: Surrey Beatty. Maxson, L.R., and D.B. Wake. 1981. Albumin Min, M.-S., S.-Y. Yang, R.M. Bonett, D.R. Viei- evolution and its phylogenetic implications in tes, R.A. Brandon, and D.B. Wake. 2005. Dis- the plethodontid salamander genera Pseudoeu- covery of the ®rst Asian plethodontid salaman- rycea and Chiropterotriton. Herpetologica 37: der. Nature. London 435: 87±90. 109±117. Miranda-Ribeiro, A., de. 1923. Os hylodideos do Mayer, A.F.J.K. 1849. System des Thier-Reiches Museu Paulista. Revista do Museu Paulista. oder Eintheilung der Thiere nach einem Prin- SaÄo Paulo 13: 825±846 (reprint pages 3±24). cip, entworfen. Verhandlungen des Naturhisto- Miranda-Ribeiro, A., de. 1926. Notas para servi- rischen Vereines der Preussischen Rheinlande rem ao estudo dos Gymnobatrachios (Anura) 6: 177±210. brasileiros. Arquivos do Museu Nacional do McDiarmid, R.W., and R.I. Altig. 1999. Research Rio de Janeiro 27: 1±227. materials and techniques. In R.W. McDiarmid Mivart, S.G. 1869. On the classi®cation of the and R. Altig (editors), Tadpoles: the biology of anurous batrachians. Proceedings of the Zoo- anuran larvae: 7±23. Chicago: University of logical Society of London 1869: 280±295. Chicago Press. Mookerjee, H.K. 1931. On the development of the McDiarmid, R.W., and S.J. Gorzula. 1989. As- vertebral column of Anura. Philosophical pects of the reproductive ecology and behavior Transactions of the Royal Society of London of the tepui toads, genus Oreophrynella (Anura, 219: 165±196. Bufonidae). Copeia 1989: 445±451. Morrison, D.A., and J.T. Ellis. 1997. Effects of McGowan, G.J., and S.E. Evans. 1995. Albaner- nucleotide sequence alignment on phylogeny petontid amphibians from the Cretaceous of estimation: a case study of 18S rDNAs of Ap- Spain. Nature. London 373: 143±145. icomplexa. Molecular Biology and Evolution Meegaskumbura, M., F. Bossuyt, R. Pethiyagoda, 14: 428±441. K. Manamendra-Arachchi, M. Bahir, M.C. Mil- Mueller, R.L., J.R. Macey, M. Jaekel, D.B. Wake, inkovitch, and C.J. Schneider. 2002. Sri Lanka: and J.L. Boore. 2004. Morphological homopla- an amphibian hot spot. Science 298: 379. sy, life history evolution, and historical bioge- Meinhardt, D.J., and J.R. Parmalee. 1996. A new ography of plethodontid salamanders inferred species of Colostethus (Anura: Dendrobatidae) from complete mitochondrial genomes. Pro- from Venezuela. Herpetologica 52: 70±77. ceedings of the National Academy of Sciences Mendelson, J.R., III, H.R. da Silva, and A.M. of the United States of America 101: 13820± Maglia. 2000. Phylogenetic relationships 13825. among marsupial frog genera (Anura: Hylidae: MuÈller, H., G.J. Measey, and P.K. Malonza. 2005. Hemiphractinae) based on evidence from mor- Tadpole of Bufo taitanus (Anura: Bufonidae) phology and natural history. Zoological Journal with notes on its systematic signi®cance and of the Linnean Society. London 128: 125±148. life history. Journal of Herpetology 39: 138± Merrem, B. 1820. Versuch eines Systems der Am- 141. phibien/ Tentamen systematis amphibiorum. MuÈller, J. 1831. An einer jungen Coecilia hypo- Marburg: Johann Christian Krieger. cyanea im Museum der Naturgeschichte. Isis Milner, A.R. 1988. The relationships and origin von Oken 1831: 710±711. of the living amphibians. In M.J. Benton (edi- MuÈller, J. 1832. BeitraÈge zur Anatomie und Na- tor), The phylogeny and classi®cation of the turgeschichte der Amphibien. I. Ueber die na- tetrapods, vol. 1. Amphibians, reptiles, birds: tuÈrliche Eintheilung der Amphibien. Zeitschrift 59±102. Systematics Association Special Vol- fuÈr Physiologie 4: 190±275, pl. 18±22. ume 23. New York: Academic Press. MuÈller, K. 2004. PRAPÐcomputation of Bremer Milner, A.R. 1993. The relatives of lis- support for large data sets. Molecular Phylo- samphibians. Herpetological Monographs 7: 8± genetics and Evolution 31: 780±782. 27. Myers, C.W. 1982. Spotted poison frogs: descrip- Milner, A.R. 1994. Late and Jurassic am- tions of three new Dendrobates from western phibians: fossil record and phylogeny. In N.C. Amazonia, and resurrection of a lost species 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 279

from ``Chiriqui''. American Museum Novitates Noble, G.K. 1926. The pectoral girdle of the bra- 2721: 1±23. chycephalid frogs. American Museum Novita- Myers, C.W. 1987. New generic names for some tes 230: 1±14. Neotropical poison frogs (Dendrobatidae). Pa- Noble, G.K. 1927. The value of life history data peÂis Avulsos de Zoologia. SaÄo Paulo 36: 301± in the study of the evolution of the Amphibia. 306. Annals of the New York Academy of Sciences Myers, C.W., and P.A. Burrowes. 1987. A new 30: 31±128. poison frog (Dendrobates) from Andean Co- Noble, G.K. 1929. The adaptive modi®cations of lombia, with notes on a lowland relative. Amer- the arboreal tadpoles of Hoplophryne and the ican Museum Novitates 2899: 1±17. torrent tadpoles of Staurois. Bulletin of the Myers, C.W., J.W. Daly, and B. Malkin. 1978. A American Museum of Natural History 58: 291± dangerously toxic new frog (Phyllobates) used 334. by Embera indians of western Colombia, with Noble, G.K. 1931. The biology of the Amphibia. discussion of blowgun fabrication and dart poi- New York: McGraw-Hill. soning. Bulletin of the American Museum of Noble, G.K., and P.G. Putnam. 1931. Observa- Natural History 161: 307±366. tions on the life history of Ascaphus truei Ste- Myers, C.W., and L.S. Ford. 1986. On Atopo- jneger. Copeia 1931: 97±101. phrynus, a recently described frog wrongly as- Nussbaum, R.A. 1976. Geographic variation and signed to the Dendrobatidae. American Muse- systematics of salamanders of the genus Di- um Novitates 2843: 1±15. camptodon Strauch (Ambystomatidae). Miscel- Myers, C.W., A. Paolillo O., and J.W. Daly. 1991. laneous Publications. Museum of Zoology, Discovery of a defensively malodorous and University of Michigan 149: 1±94. nocturnal frog in the family Dendrobatidae: Nussbaum, R.A. 1977. Rhinatrematidae: a new phylogenetic signi®cance of a new genus and family of caecilians (Amphibia: Gymno- species from the Venezuelan Andes. American phiona). Occasional Papers of the Museum of Museum Novitates 3002: 1±33. Zoology, University of Michigan 682: 1±30. Nascimento, L.B., U. Caramaschi, and C.A.G. Nussbaum, R.A. 1979. The taxonomic status of Cruz. 2005. Taxonomic review of the species the caecilian genus Uraeotyphlus Peters. Oc- group of the genus Physalameus Fitzinger, casional Papers of the Museum of Zoology, 1826 with revalidation of the genera Engysto- University of Michigan 687: 1±20. mops Jimenez de la Espada, 1872 and Eupem- Nussbaum, R.A. 1980. Phylogenetic implications phix Steindachner, 1836 (Amphibia, Anura, of amplectic behavior in sooglossid frogs. Her- Leptodactylidae). Arquivos do Museu Nacional petologica 36: 1±5. do Rio de Janeiro 63: 297±320. Nussbaum, R.A. 1982. Heterotopic bones in the Nicholls, G.E. 1916. The structure of the vertebral hindlimbs of frogs of the families Pipidae, Ran- column in the Anura Phaneroglossa and its im- idae and Sooglossidae. Herpetologica 38: 312± portance as a basis of classi®cation. Proceed- 320. ings of the Linnaean Society of London 128: Nussbaum, R.A., A.P. Jaslow, and J. Watson. 80±92. 1982. Vocalization in frogs of the family Soog- Niklas, K.J. 2001. Taxing debate for taxonomists. lossidae. Journal of Herpetology 16: 198±203. Science 292: 2249±2250. Nussbaum, R.A., and M. Wilkinson. 1989. On the Nixon, K.C. 1999. The parsimony ratchet, a new classi®cation and phylogeny of caecilians (Am- method for rapid parsimony analysis. Cladistics phibia: Gymnophiona), a critical review. Her- 15: 407±414. petological Monographs 3: 1±42. Nixon, K.C. 1999±2002. WinClada. Version 1.0. Nussbaum, R.A., and M. Wilkinson. 1995. A new Ithaca, New York: Computer software distrib- genus of lungless tetrapod: a radically divergent uted by the author. caecilian (Amphibia: Gymnophiona). Proceed- Nixon, K.C., and J.M. Carpenter. 2000. On the ings of the Royal Society of London. Series B, other ``phylogenetic systematics''. Cladistics Biological Sciences 261: 331±335. 16: 298±318. Ohler, A., S.R. Swan, and J.C. Daltry. 2002. A Nixon, K.C., J.M. Carpenter, and D.W. Stevenson. recent survey of the amphibian fauna of the 2003. The PhyloCode is fatally ¯awed and the Cardamom Mountains, Southwest Cambodia ``Linnaean'' system can be easily ®xed. Botan- with descriptions of three new species. Raf¯es ical Review 69: 111±120. Bulletin of Zoology. Singapore 50: 465±481. Noble, G.K. 1924. A new spadefoot toad from the Okada, Y. 1966. Fauna Japonica. Anura (Amphib- Oligocene of Mongolia with a summary of the ia). Tokyo: Biogeographical Society of Japan. evolution of the Pelobatidae. American Muse- Olsson, L., and J. Hanken. 1996. Cranial neural- um Novitates 132: 1±15. crest migration and chondrogenic fate in the 280 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

oriental ®re-bellied toad Bombina orientalis: Parra-Olea, G., M. GarcõÂa-ParõÂs, and D.B. Wake. de®ning the ancestral pattern of head develop- 2004. Molecular diversi®cation of salamanders ment in anuran amphibians. Journal of Mor- of the tropical American genus Bolitoglossa phology 299: 105±120. (Caudata: Plethodontidae) and its evolutionary Oppel, M. 1810. Second memoire sur las classi- and biogeographical implications. Biological ®cation des reptiles. Annales du MuseÂum Journal of the Linnean Society 81: 325±346. d'Histoire Naturelle 16: 394±418. Parra-Olea, G., and D.B. Wake. 2001. Extreme Orlov, N.L., L.N. Ngat, and T.C. Ho. 2003. A new morphological and ecological homoplasy in species of cascade frog from North Vietnam tropical salamanders. Proceedings of the Na- (Ranidae, Anura). Russian Journal of Herpetol- tional Academy of Sciences of the United ogy 10: 123±134. States of America 98: 7888±7891. Orton, G.L. 1949. Larval development of Necto- Parsons, T.S., and E.E. Williams. 1963. The rela- phrynoides tornieri (Roux), with comments on tionships of the modern Amphibia: a re±ex- direct development in frogs. Annals of the Car- amination. Quarterly Review of Biology 38: negie Museum 31: 257±274. 26±53. Orton, G.L. 1953. The systematics of vertebrate Passmore, N.I., and V.C. Carruthers. 1979. South larvae. Systematic Zoology 2: 63±75. African frogs. Johannesburg: Witwatersrand Orton, G.L. 1957. The bearing of larval evolution University Press. on some problems in frog classi®cation. Sys- Patterson, C., and D.E. Rosen. 1977. Review of tematic Zoology 6: 79±86. the ichthyodectiform and other Mesozoic tele- Page, L.M, H.L. Bart, Jr., R. Beaman, L. Bohs, ost ®shes and the theory and practice of clas- L.T. Deck, V.A. Funk, D. Lipscomb, M.A. Ma- sifying fossils. Bulletin of the American Mu- res, L.A. Prather, J. Stevenson, Q.D. Wheeler, seum of Natural History 158: 85±172. J.B. Wooley, and D.W. Stevenson. 2005. LIN- Pauly, G.B., D.M. Hillis, and D.C. Cannatella. NE: legacy infrastructure network for natural 2004. The history of Nearctic colonization: mo- environments. Champaign, IL: Illinois Natural lecular phylogenetics and biogeography of the History Survey. Nearctic toads (Bufo). Evolution 58: 2517± Palma, R.E., and A.E. Spotorno. 1999. Molecular 2535. systematics of marsupials based on the rRNA Peixoto, O.L. 1995. AssociacËaÄo de anuros a bro- 12S mitochondrial gene: the phylogeny of di- meliaÂceas na mata AtlaÃntica. Revista de Univ- delphimorphia and the living fossil microbioth- ersidade Federal Rural do Rio de Janeiro. SeÂrie eriid Dromiciops gliroides Thomas. Molecular CieÃncias da Vida 17: 75±83. Phylogenetics and Evolution 13: 525±535. Pennisi, E. 2001. Linnaeus's last stand? Science Palumbi, S.R., A. Martin, S. Romano, W.O. Mc- 291: 2304±2307. Millan, L. Stice, and G. Grabawski. 1991. The Perret, J.-L. 1966. Les amphibiens du Cameroun. simple fool's guide to PCR, version 2.0. Uni- Zoologische JahrbuÈcher. Abteilung fuÈr Syste- versity of Hawaii, Honolulu: Privately pub- matik, OÈ kologie und Geographie. Jena 93: lished, compiled by S. Palumbi. 289±464. Papavero, N., J. Llorente-Bousquets, and J.M. Perret, J.-L. 1977. Les Hylarana (amphibiens, Abe. 2001. Proposal of a new system of no- ranideÂs) du Cameroun. Revue Suisse de Zool- menclature for phylogenetic systematics. Ar- ogie 84: 841±868. quivos de Zoologia. SaÄo Paulo 36: 1±145. Perret, J.-L. 1984. Identi®cation des syntypes de Parker, H.W. 1934. A monograph of the frogs of Petropedetes obscurus Ahl, 1924 (Amphibia, the family Microhylidae. London: Trustees of Phrynobatrachinae), conserveÂs au museÂum de the British Museum. Berlin. Bulletin de la SocieÂte NeuchaÃteloise des Parker, H.W. 1940. The Australasian frogs of the Sciences Naturelles 107: 167±170. family Leptodactylidae. Novitates Zoologicae. Peters, W.C.H. 1862. UÈ ber die Batrachier-Gattung Tring 42: 1±106. Hemiphractus. Monatsberichte der KoÈniglichen Parra-Olea, G. 2002. Molecular phylogenetic re- Preussische Akademie des Wissenschaften zu lationships of Neotropical salamanders of the Berlin 1862: 144±152. genus Pseudoeurycea. Molecular Phylogenetics Philippe, H. 1993. MUST: management utilities and Evolution 22: 234±246. for sequences and trees. Nucleic Acids Re- Parra-Olea, G., M. GarcõÂa-ParõÂs, and D.B. Wake. search 21: 5264±5272. 2002. Phylogenetic relationships among the sal- Phillips, A., D. Janies, and W.C. Wheeler. 2000. amanders of the Bolitoglossa macrinii species Multiple sequence alignment in phylogenetic group (Amphibia: Plethodontidae), with de- analysis. Molecular Phylogenetics and Evolu- scriptions of two new species from tion 16: 317±330. (Mexico). Journal of Herpetology 36: 356±366. Pickett, K.M. 2005. The new and improved 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 281

PhyloCode, now with types, ranks, and even West Indian toads (Anura: Bufonidae). Molec- polyphyly: a conference report from the First ular Phylogenetics and Evolution 20: 294±301. International Phylogenetic Nomenclature Meet- Pramuk, J.B., and J.R. Mendelson, III. 2003. An- ing. Cladistics 21: 79±82. axyrus melanocholicus Tschudi: synonym of Platnick, N.I. 1977. Cladograms, phylogenetic the Mexican taxon Bufo compactilis Wiegmann trees, and hypothesis testing. Systematic Zool- (Anura: Bufonidae). Southwestern Naturalist ogy 26: 438±443. 48: 676±680. Pombal, J.P., Jr. 1999. OviposicËaÄo e desenvolvi- Pregill, G.K. 1981. Cranial morphology and the mento de Brachycephalus ephippium (Spix) evolution of West Indian toads: resurrection of (Anura, Brachycephalidae). Revista Brasileira the genus Peltophryne (Fitzinger). Copeia de Zoologia 16: 967±976. 1981: 273±285. Pombal, J.P., Jr., and C.F.B. Haddad. 1999. Frogs Procter, J.B. 1925. Notes on the nests of some of the genus Paratelmatobius (Anura: Lepto- African frogs. Proceedings of the Zoological dactylidae) with descriptions of two new spe- Society of London 1925: 909±910. cies. Copeia 1999: 1014±1026. Pugener, L.A., A.M. Maglia, and L. Trueb. 2003. Pope, C.H. 1931. Notes on amphibians from Fu- Revisiting the contribution of larval characters kien, Hainan and other parts of China. Bulletin to an analysis of phylogenetic relationships of of the American Museum of Natural History basal anurans. Zoological Journal of the Lin- 61: 397±611. nean Society. London 139: 129±155. Posada, D., and K.A. Crandall. 1998. Modeltest: Pusey, H.K. 1943. On the head of the liopelmid testing the model of DNA substitution. Bioin- frog, Ascaphus truei I. The chondrocranium, formatics 14: 817±818. , arches, and muscles of a partly grown lar- Post, T.J., and T. Uzzell. 1981. The relationships va. Quarterly Journal of Microscopical Science of Rana sylvatica and the monophyly of the 84: 106±185. Rana boylii group. Systematic Zoology 30: Pyburn, W.F. 1970. Breeding behavior of the leaf- 170±180. frogs Phyllomedusa callidryas and Phyllome- Poynton, J.C. 1964a. Amphibia of southern Afri- dusa dacnicolor in Mexico. Copeia 1970: 209± ca: a faunal study. Annals of the Natal Museum 218. 17: 1±334. Pytel, B.A. 1986. Biochemical systematics of the Poynton, J.C. 1964b. Amphibia of the Nyasa-Lu- eastern North American frogs of the genus angwa region of Africa. Senckenbergiana Biol- ogica 45: 193±225. Rana. Herpetologica 42: 273±282. Poynton, J.C. 1976. Classi®cation and the Arthro- Rabb, G.B. 1960. On the unique sound production leptinae. Revue Zoologique Africaine 90: 215± of the Surinam toad, Pipa pipa. Copeia 1960: 220. 368±369. Poynton, J.C. 2003. Arthroleptis troglodytes and Ra®nesque, C.S. 1814. Fine del prodromo the content of Schoutedenella (Amphibia: An- d'erpetologia siciliana. Specchio delle Scienze, ura: Arthroleptidae). African Journal of Her- o, Giornale Enciclopedico di Sicilia 2: 102± petology 52: 49±51. 104. Poynton, J.C., and D.G. Broadley. 1967. A new Ra®nesque, C.S. 1815. Analyse de nature, ou tab- species of Probreviceps (Amphibia) from Rho- leau de l'universe et des corps organiseÂs. Pa- desia. Arnoldia. Zimbabwe 3: 1±3. lermo: Jean Barravecchia. Poynton, J.C., and D.G. Broadley. 1985. Amphib- Rambaut, A. 1995. Se-Al. Sequence alignment ia Zambesiaca 2. Ranidae. Annals of the Natal editor, version 1.d1. University of Oxford, Museum 27: 115±181. U.K.: Computer software distributed by the au- Poynton, J.C., and D.G. Broadley. 1988. Amphib- thor. ia Zambesiaca, 4. Bufonidae. Annals of the Na- Rao, C.R.N. 1920. Some South Indian batrachi- tal Museum 29: 447±490. ans. Journal of the Bombay Natural History So- Pramuk, J.B. 2000. Prenasal bones and snout mor- ciety 27: 119±127. phology in West Indian bufonids and the Bufo Read, K., J.S. Keogh, I.A. Scott, J.D. Roberts, and granulosus species group. Journal of Herpetol- P. Doughty. 2001. Molecular phylogeny of the ogy 34: 334±340. Australian frog genera Crinia, Geocrinia, and Pramuk, J.B. 2002. Combined evidence and cla- allied taxa (Anura: Myobatrachidae). Molecular distic relationships of West Indian toads (Anu- Phylogenetics and Evolution 21: 294±308. ra: Bufonidae). Herpetological Monographs 16: Regal, P.J. 1966. Feeding specializations and the 121±151. classi®cation of terrestrial salamanders. Evolu- Pramuk, J.B., C.A. Hass, and S.B. Hedges. 2001. tion 20: 392±407. Molecular phylogeny and biogeography of Reig, O.A. 1958. Proposiciones para una nueva 282 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

macrosistematica de los anuros (nota prelimi- the anuran Pipa pipa. Journal of Morphology nar). Physis. Buenos Aires 21: 109±118. 200: 300±319. Reig, O.A. 1960. Las relaciones geneÂricas del an- RoÈdel, M.-O., J. Kosuch, M. Veith, and R. Ernst. uro chileno Calyptocephalella gayi (Dum. & 2003. First record of the genus Acanthixalus Bibr.). In Actas y trabajos del Primer Congreso Laurent, 1944 from the upper Guinean rain for- Sudamericano de ZoologõÂa (La Plata, 12±24 est, West Africa, with the description of a new Octubre 1959), vol. 4: 113±147. La Plata: Com- species. Journal of Herpetology 37: 43±52. isioÂn de InvestigacioÂn Cientõ®ca de la Provincia Rodman, J.E., and J.H. Cody. 2003. The taxonom- de Buenos Aires y Consejo Nacional de Inves- ic impediment overcome: NSF's Partnerships tigaciones Cientõ®cas y TeÂcnicas. for Enhancing Expertise in Taxonomy (PEET) Reig, O.A. 1972. Macrogenioglottus and the as a model. Systematic Biology 52: 428±435. South American bufonid toads. In W.F. Blair Roelants, K., and F. Bossuyt. 2005. Archaeobatra- (editor), Evolution in the genus Bufo: 14±36. chian paraphyly and Pangaean diversi®cation Austin: University of Texas Press. of crown-group frogs. Systematic Biology 54: Richards, C.M., and W.S. Moore. 1996. A phy- 111±126. logeny for the African treefrog family Hyper- Roelants, K., J. Jiang, and F. Bossuyt. 2004. En- oliidae based on mitochondrial rDNA. Molec- demic ranid (Amphibia: Anura) genera in ular Phylogenetics and Evolution 5: 522±532. southern mountain ranges of the Indian subcon- Richards, C.M., and W.S. Moore. 1998. A molec- tinent represent ancient frog lineages: evidence ular phylogenetic study of the Old World tree- from the molecular data. Molecular Phyloge- frog family Rhacophoridae. Herpetological netics and Evolution 31: 730±740. Journal. London 8: 41±46. Romer, A.S. 1933. Vertebrate paleontology. Chi- Richards, C.M., R.A. Nussbaum, and C.J. Rax- cago: University of Chicago Press. Romer, A.S. 1945. Vertebrate paleontology, 2nd worthy. 2000. Phylogenetic relationships within ed. Chicago: University of Chicago Press. the Madagascan boophids and mantellids as Ruiz-Carranza, P.M., and J.D. Lynch. 1991a. Ran- elucidated by mitochondrial ribosomal genes. as Centrolenidae de Colombia I. Propuesta de African Journal of Herpetology 49: 23±32. una nueva clasi®cacion generica. Lozania 57: Rieppel, O., and M. de Braga. 1996. Turtles as 1±30. diapsid reptiles. Nature. London 384: 453±455. Ruiz-Carranza, P.M., and J.D. Lynch. 1991b. Ran- Risch, J.-P. 1985. The Himalayan salamander and as Centrolenidae de Colombia II. Nuevas es- its relatives: a short review of the Pleurodelinae pecies de Centrolene de la Cordillera Oriental (Amphibia, Caudata, Salamandridae). Journal y Sierra Nevada de Santa Marta. Lozania 58: of the Bengal Natural History Society. New Se- 1±26. ries 4: 139±143. Ruiz-Carranza, P.M., and J.D. Lynch. 1998. Ranas Ritgen, F.A. 1828. Versuch einer naturlichen Centrolenidae de Colombia XI. Nuevas espe- Eintheilung der Amphibien. Nova Acta Physi- cies de ranas cristal del genero Hyalinobatrach- co-medica Academiae Caesareae Leopoldino- ium. Revista de la Academia Colombiana de Carolinae Naturae Curiosorum. Halle 14: 277, Ciencias Exactas, FõÂsicas y Naturales 22: 571± 278. 586. Ritland, R.M. 1955. Studies on the post-cranial Ruta, M., M.I. Coates, and D.L.J. Quicke. 2003. morphology of Ascaphus truei. II. Journal of Early tetrapod relationships revisited. Biologi- Morphology 97: 215±282. cal Reviews of the Cambridge Philosophical RocÏek, Z. 1981 ``1980''. Cranial anatomy of frogs Society 78: 251±345. of the family Pelobatidae Stannius, 1856, with Ruvinsky, I., and L.R. Maxson. 1996. Phyloge- outlines of their phylogeny and systematics. netic relationships among bufonoid frogs (An- Acta Universitatis Carolinae. Biologica. Prague ura: Neobatrachia) inferred from mitochondrial 1980: 1±160. DNA sequences. Molecular Phylogenetics and RocÏek, Z. 1989. Developmental pattern of the eth- Evolution 5: 533±547. moidal region of the anuran skull. In H. Saint-Aubain, M.L., de. 1981. Amphibian limb Splechtna (editor), Trends in vertebrate mor- ontogeny and its bearing on the phylogeny of phology: 412±515. Stuttgart: G. Fischer. the group. Zeitschrift fuÈr Zoologische Syste- RocÏek, Z. 1990. Ethmoidal endocranial structures matik und Evolutionsforschung 19: 175±194. in primitive tetrapods: their bearing on the Salthe, S.N. 1967. The courtship patterns and the search for anuran ancestry. Zoological Journal phylogeny of the urodeles. Copeia 1967: 100± of the Linnean Society. London 99: 389±407. 117. RocÏek, Z., and M. Vesely. 1989. Development of SanchõÂz, F.B. 1984. AnaÂlisis ®logeneÂtico de la tri- the ethmoidal structures of the endocranum in bu Alytini (Anura, Discoglossidae) mediante el 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 283

estudio de su morfoestructura oÂsea. In H. Hem- and new taxa. American Museum Novitates mer and J.A. Alcover (editors), HistoÁria biooÁg- 3357: 1±48. ica del ferreret (Life history of the Mallorcan SchaÈuble, C.S., C. Moritz, and R.W. Slade. 2000. ): 61±108. Mallorca, Spain: Edi- A molecular phylogeny for the frog genus Lim- torial Moll. nodynastes (Anura: Myobatrachidae). Molecu- SanchõÂz, F.B. 1998. Encyclopedia of paleoherpe- lar Phylogenetics and Evolution 16: 379±391. tology, vol. 4. Salientia. MuÈnchen: Dr. Fried- Scheel, J.J. 1970. Notes on the biology of the Af- rich Pfeil. rican tree-toad, Nectophryne afra Buchholz & SanchõÂz, F.B., and I. de la Riva. 1993. Remarks Peters, 1875 (Bufonidae, Anura) from Fernan- on the tarsus of centrolenid frogs (Amphibia, do Poo. Revue de Zoologie et de Botanique Af- Anura). Graellsia 49: 115±117. ricaines 81: 225±236. Sankoff, D. 1975. Minimal mutation trees of se- Scheltinga, D.M., B.G.M. Jamieson, D.P. Bick- quences. SIAM Journal on Applied Mathemat- ford, A.A. Garda, S.N. Bao, and K.R. Mc- ics 28: 35±42. Donald. 2002. Morphology of the spermatozoa Sankoff, D., R.J. Cedergren, and G. Lapalme. of the Microhylidae (Anura, Amphibia). Acta 1976. Frequency of insertion-deletion, transver- Zoologica. Stockholm 83: 263±275. sion, and transition in evolution of 5S ribosom- Schmidt, K.P., and R.F. Inger. 1959. Amphibians al RNA. Journal of Molecular Evolution 7: exclusive of the genera Afrixalus and Hypero- 133±149. lius. Exploration du Parc National de San Mauro, D., D.J. Gower, O.V. Oommen, M. l'Upemba. Mission G.F. de Witte, en Collabo- Wilkinson, and R. Zardoya. 2004. Phylogeny of ration avec W. Adam, A. Janssens, L. van Meel caecilian amphibians (Gymnophiona) based on et R. Verheyen (1946±1949) 56: 1±264. complete mitochondrial genomes and nuclear Schoch, R.R., and A.R. Milner. 2004. Structure and implications of theories on the origins of RAG1. Molecular Phylogenetics and Evolution lissamphibians. In G. Arratia, M.V. Wilson, and 33: 413±427. R. Cloutier (editors), Recent advances in the San Mauro, D., M. Vences, M. Alcobendas, R. origin and early radiation of vertebrates: 345± Zardoya, and A. Meyer. 2005. Initial diversi®- 377. MuÈnchen: Pfeil. cation of living amphibians predated the break- Scholz, K.P. 1995. Zur Stammesgeschichte der up of . American Naturalist 165: 590± Salamandridae Gray, 1825. eine kladistische 599. Analyse anhand von Merkmalen aus Morphol- Santos, J.C., L.A. Coloma, and D.C. Cannatella. ogie und Balzverhalten. Acta Biologica Ben- 2003. Multiple, recurring origins of aposema- rodis 7: 25±75. tism and diet specialization in poison frogs. Schuh, R.T. 2003. The Linnaean system and its Proceedings of the National Academy of Sci- 250- persistence. Botanical Review 69: 59± ences of the United States of America 100: 79. 12792±12797. Schulte, R. 1989. Nueva especie de rana venenosa Sarasin, P., and F. Sarasin. 1890. Zur Entiwick- del genero Epipedobates registrada en la Cor- lungsgeschichte und Anatomie der ceylonisch- dillera Oriental, Departamento de San Martin. en Blindwuehle, (Epi- BoletõÂn de Lima 11: 41±46. crium glutinosum Aut.). Ergebnisse naturwis- Schwenk, K., and D.B. Wake. 1993. Prey pro- senschaftlicher Forschungen auf Ceylon in den cessing in Leurognathus marmoratus and the Jahren 1884±86. Vol. II. Heft 4. Wiesbaden: evolution of form and function in desmogna- C.W. Kreidel. thine salamanders (Plethodontidae). Biological Savage, J.M. 1973. The geographic distribution of Journal of the Linnean Society 49: 141±162. frogs: patterns and predictions. In J.L. Vial (ed- Scopoli, G.A. 1777. Introductio ad historiam na- itor), Evolutionary biology of the anurans: con- turalem, sistens genera lapidium, planatarum, et temporary research on major problems: 351± animalium hactenus detecta, caracteribus essen- 445. Columbia: University of Missouri Press. tialibus donata, in tribus divisa, subinde ad le- Savage, J.M. 1987. Systematics and distribution ges naturae. Prague: Gerle. of the Mexican and Central American rainfrogs Scott, E. 2002. Phylogenetic relationships of the of the Eleutherodactylus gollmeri group (Am- subfamily Petropedetinae Noble, 1931 (Anura: phibia: Leptodactylidae). Fieldiana. Zoology. Ranidae): a simultaneous analysis of morpho- New Series 33: 1±57. logical and molecular data. Ph.D. dissertation. Savage, J.M., and C.W. Myers. 2002. Frogs of the Department of Zoology, University of the Eleutherodactylus biporcatus group (Leptodac- Western Cape, Bellville, South Africa. tylidae) of Central America and northern South Sever, D.M. 1990. Cloacal anatomy of female sal- America, including rediscovered, resurrected, amanders of the plethodontid subfamily Des- 284 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

mognathinae (Amphibia: Urodela). Transac- Simmons, M.P., and H. Ochoterena. 2000. Gaps tions of the American Microscopical Society as characters in sequence-based phylogenetic 109: 193±204. analyses. Systematic Biology 49: 369±381. Sever, D.M. 1991a. Comparative anatomy and Sinsch, U., and N. Juraske. 1995. Reassessment phylogeny of the cloacae of salamanders (Am- of central Peruvian Telmatobiinae (genera Ba- phibia: Caudata). 1. Evolution at the family lev- trachophrynus and Telmatobius) II. Allozymes el. Herpetologica 47: 165±193. and phylogenetic relationships. Alytes 13: 52± Sever, D.M. 1991b. Comparative anatomy and 66. phylogeny of the cloacae of salamanders (Am- Sinsch, U., A.W. Salas, and V. Canales. 1995. Re- phibia: Caudata). 2. Cryptobranchidae, Hyno- assessment of central Peruvian Telmatobiinae biidae, and Sirenidae. Journal of Morphology (genera Batrachophrynus and Telmatobius)I. 207: 283±301. Morphometry and classi®cation. Alytes 13: 14± Sever, D.M. 1992. Comparative anatomy and phy- 44. logeny of the cloacae of salamanders (Amphib- Sites, J.W., Jr., M. Morando, R. Highton, F. Huber, ia: Caudata). VI. Ambystomatidae and Dicamp- and R.E. Jung. 2004. Phylogenetic relationships todontidae. Journal of Morphology 212: 305± of the endangered 322. (Plethodon shenandoah) and other salamanders Sever, D.M. 1994. Comparative anatomy and phy- of the Plethodon cinereus group (Caudata: logeny of the cloacae of salamanders (Amphib- Plethodontidae). Journal of Herpetology 38: ia: Caudata). VII. Plethodontidae. Herpetologi- 96±105. cal Monographs 7: 276±337. Slabbert, G.K., and W.A. Maree. 1945. The cra- Sever, D.M., E.A. Heinz, P.A. Lempart, and M.S. nial morphology of the Discoglossidae and its Taghon. 1990. Phylogenetic signi®cance of the bearing upon the phylogeny of the primitive cloacal anatomy of female bolitoglossine sala- Anura. Annals of the University of Stellen- manders (Plethodontidae: tribe Bolitoglossini). bosch 23A: 91±97. Herpetologica 46: 431±446. Slowinski, J.B. 1998. The number of multiple Shaffer, H.B., J.M. Clark, and F. Kraus. 1991. alignments. Molecular Phylogenetics and Evo- When molecules and morphology clash: a phy- lution 10: 264±266. logenetic analysis of the North American am- Smith, M.A. 1930. The Reptilia and Amphibia of bystomatid salamanders (Caudata: Ambysto- matidae). Systematic Zoology 40: 284±303. the Malay Peninsula. A supplement to G.A. Sharrock, G., and J. Felsenstein. 1975. Finding all Boulenger's Reptilia and Batrachia, 1912. Bul- monothetic subsets of a taxonomic group. Sys- letin of the Raf¯es Museum 3: i±xviii, 1±149. tematic Zoology 24: 373±377. [Cited as ``1969. Sokol, O. 1975. The phylogeny of anuran larvae: manuscript in preparation'' by Liem, 1970.] a new look. Copeia 1975: 1±23. Sheil, C.A., J.R. Mendelson, III, and H.R. da Sil- Sokol, O. 1977. A subordinal classi®cation of va. 2001. Phylogenetic relationships of the spe- frogs (Amphibia: Anura). Journal of Zoology. cies of Neotropical horned frogs, genus Hemi- London 182: 505±508. phractus (Anura: Hylidae: Hemiphractinae), Sokol, O. 1981. The larval chondrocranium of Pe- based on evidence from morphology. Herpeto- lodytes punctatus, with a review of tadpole logica 57: 203±214. chondrocrania. Journal of Morphology 169: Shubin, N.H., and F.A. Jenkins. 1995. An early 161±183. Jurassic jumping frog. Nature. London 377: Starrett, P.H. 1973. Evolutionary patterns in larval 49±52. morphology. In J.L. Vial (editor), Evolutionary Silva, A.P.Z., C.F.B. Haddad, and S. Kasahara. biology of the anurans: contemporary research 2003. Chromosome banding in Macrogenioglot- on major problems: 251±271. Columbia: Uni- tus alipioi Carvalho, 1946 (Amphibia, Anura, versity of Missouri Press. Leptodactylidae), with comments on its taxo- Steindachner, F. 1867. Amphibien. In Reise der nomic position. Boletim do Museu Nacional, oÈsterreichischen Fregatte Novara um die Erde Rio de Janeiro. Nova SeÂrie, Zoologia. 499: 1±9. in den Jahren 1857, 1858, 1859 unter den Ba- Silverstone, P.A. 1975. A revision of the poison- fehlen des Commodore B. von WuÈllerstorf-Ur- arrow frogs of the genus Dendrobates Wagler. bair, Zoologischer Theil. 1: 1±98. Wien: Kais- Natural History Museum of Los Angeles Coun- erlich-KoÈnigliche Hof- und Staatsdruckerei; in ty Science Bulletin 21: 1±55. Commission bei K. Gerold's Sohn. Simmons, M.P. 2004. Independence of alignment Steinfartz, S., U.W. Hwang, D. Tautz, M. OÈ z, and and tree search. Molecular Phylogenetics and M. Veith. 2002. Molecular phylogeny of the Evolution 31: 874±879. salamandrid genus Neurergus: evidence for an 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 285

intrageneric switch of reproductive biology. D.M. Hillis, C. Moritz, and B.K. Mable (edi- Amphibia-Reptilia 23: 419±431. tors), Molecular systematics: 407±514. Sunder- Stejneger, L. 1907. Herpetology of Japan and ad- land, MA: Sinauer Associates. jacent territory. Bulletin of the United States Tamarunov, L.P. 1964a. Salientia. In Y.A. Orlov National Museum 58: i±xx, 1±577. (editor), Osnovy paleontologii: spravochnik Stejneger, L., and T. Barbour. 1917. A check list dlia paleontologov i geologov SSSR, vol. 12. of North American amphibians and reptiles. Amphibia±Aves: 125±133. Moscow: Nauka. Cambridge, MA: Harvard University Press. Tamarunov, L.P. 1964b. Lepospondyli. In Y.A. Stephenson, E.M., E.S. Robinson, and N.G. Ste- Orlov (editor), Osnovy paleontologii: spravo- phenson. 1974. Inter-speci®c relationships of chnik dlia paleontologov i geologov SSSR, vol. Leiopelma (Amphibia: Anura): further karyo- 12. Amphibia±Aves: 144±164. Moscow: Nau- logical evidence. Experientia 30: 1248±1250. ka. Stephenson, N.G. 1951. Observations on the de- Tanaka-Ueno, T., M. Matsui, S.-L. Chen, O. Tak- velopment of the amphicoelous frogs, Leiopel- enaka, and H. Ota. 1998a. Phylogenetic rela- ma and Ascaphus. Journal of the Linnean So- tionships of brown frogs from Taiwan and Ja- ciety of London. Zoology 42: 18±28. pan assessed by mitochondrial cytochrome b Stoll, N. 1961. Introduction. In International gene sequences (Rana: Ranidae). Zoological Commission of Zoological Nomenclature (edi- Science. Tokyo 15: 283±288. tors), International code of zoological nomen- Tanaka-Ueno, T., M. Matsui, T. Sato, S. Takenaka, clature adopted by the XV International Con- and O. Takenaka. 1998b. Phylogenetic relation- gress of Zoology: vii±xvii. London: Interna- ships of brown frogs with 24 chromosomes tional Trust for Zoological Nomenclature. from Far East Russia and Hokkaido assessed Stuart, B.L., and T. Chan-ard. 2005. Two new by mitochondrial cytochrome b gene sequences Huia (Amphibia: Ranidae) from Laos and Thai- (Rana: Ranidae). Zoological Science. Tokyo land. Copeia 2005: 279±289. 15: 289±294. Stuart, S.N., J.S. Chanson, N.A. Cox, B.E. Young, Tandy, M., and R. Keith. 1972. Bufo of Africa. In A.S.L. Rodrigues, D.L. Fischman, and R.W. W.F. Blair (editor), Evolution in the genus Bufo: Waller. 2004. Status and trends of amphibian 119±170. Austin: University of Texas Press. declines and worldwide. Science Taylor, E.H. 1951. Two new genera and a new 306: 1783±1786. Sumida, M., A. Allison, and M. Nishioka. 2000a. family of tropical American frogs. Proceedings Evolutionary relationships among 12 species of the Biological Society of Washington 64: belonging to three genera of the family Micro- 33±40. hylidae in Papua New Guinea revealed by al- Taylor, E.H. 1968. The caecilians of the world: a lozyme analysis. Biochemical Systematics and taxonomic review. Lawrence: University of Ecology 28: 721±736. Kansas Press. Sumida, M., Y. Kondo, Y. Kanamori, and M. Ni- Taylor, E.H. 1969a. A new family of African shioka. 2002. Inter- and intraspeci®c evolution- Gymnophiona. University of Kansas Science ary relationships of the rice frog Rana limno- Bulletin 48: 297±305. charis and the allied species R. cancrivora in- Taylor, E.H. 1969b. of Gymnophiona and ferred from crossing experiements and mito- their signi®cance in the taxonomy of the group. chondrial DNA sequences of the 12S and 16S University of Kansas Science Bulletin 48: 585± rRNA genes. Molecular Phylogenetics and 687. Evolution 25: 293±305. Thibaudeau, G., and R.I. Altig. 1999. Endotrophic Sumida, M., M. Ogata, and M. Nishioka. 2000b. anurans. Development and evolution. In R.W. Molecular phylogenetic relationships of pond McDiarmid and R. Altig (editors), Tadpoles: frogs distributed in the Palearctic Region in- the biology of anuran larvae: 170±188. Chica- ferred from DNA sequences of mitochondrial go: University of Chicago Press. 12S ribosomal RNA and cytochrome b genes. Thompson, J.D., T.J. Gibson, F.J. Plewniak, F., and Molecular Phylogenetics and Evolution 16: D.G. Higgins. 1997. The ClustalX windows in- 278±285. terface: ¯exible strategies for multiple sequence Swofford, D.L. 2002. PAUP*. Phylogenetic anal- alignment aided by quality analysis tools. Nu- ysis using parsimony (*and other methods), cleic Acids Research 24 4876±4882. version 4.0 beta. Sunderland, MA: Sinauer As- Thompson, J.D., D.G. Higgins, and T.J. Gibson. sociates. 1994. CLUSTAL W: improving the sensitivity Swofford, D.L., G.J. Olsen, P.J. Waddell, and of progressive multiple sequence alignments D.M. Hillis. 1996. Phylogenetic inference. In through sequence weighting, position speci®c 286 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

gap penalties and weight matrix choice. Nu- Tschudi, J.J., von. 1838. Classi®cation der Batra- cleic Acids Research 22: 4673±4680. chier mit BeruÈcksichtigung der fossilen Thiere Tian, W., and Q. Hu. 1985. Taxonomical studies dieser Abtheilung der Reptilien. NeuchaÃtel: Pe- on the primitive anurans of the Hengduan titpierre. Mountains, with descriptions of a new subfam- Tschudi, J.J., von. 1845. Reptilium conspectus ily and subdivision of Bombina. Acta Herpe- quae in Republica Peruana reperiuntur et ple- tologica Sinica, New Series 4: 219±224. raquae observata vel collecta sunt in itinere a Tihen, J.A. 1958. Comments on the osteology and Dr. J.J. de Tschudi. Archiv fuÈr Naturgeschichte phylogeny of ambystomatid salamanders. Bul- 11: 150±170. letin of the Florida Museum of Natural History. Tyler, M.J. 1971a. The occurrence of the muscu- Biological Sciences 3: 1±50. lus cutaneous pectoris in the Anura. Herpeto- Tihen, J.A. 1960. Two new genera of African bu- logica 27: 150±152. fonids, with remarks on the phylogeny of re- Tyler, M.J. 1971b. Observations on anuran myo- lated genera. Copeia 1960: 225±233. integumental attachments associated with the Tihen, J.A. 1965. Evolutionary trends in frogs. vocal sac apparatus. Journal of Natural History American Zoologist 5: 309±318. 5: 225±231. Tilley, S.G., and J. Bernardo. 1993. Life history Tyler, M.J. 1971c. The phylogenetic signi®cance evolution in plethodontid salamanders. Herpe- of vocal sac structure in hylid frogs. University tologica 49: 154±163. of Kansas Publications. Museum of Natural Ting, H.-P., and M.-C. T'sai. 1979. A new species History 19: 319±360. of frog (Rana minimus) from Fujian Province. Tyler, M.J. 1972. Super®cial mandibular muscu- Acta Zootaxonomica Sinica 4: 297±300. [In lature, vocal sacs and the phylogeny of Austra- Chinese with English abstract.] lo-Papuan leptodactylid frogs. Records of the Titus, T.A., and A. Larson. 1995. A molecular South Australian Museum 16: 1±20. phylogenetic perspective on the evolutionary Tyler, M.J. 1979. Herpetofauna relationships of radiation of the salamander family Salamandri- South America with Australia. In W.E. Duell- dae. Systematic Biology 44: 125±151. man (editor), The South American herpetofau- Titus, T.A., and A. Larson. 1996. Molecular phy- na: its origin, evolution, and dispersal. Univer- logenetics of desmognathine salamanders (Cau- sity of Kansas Museum of Natural History, data: Plethodontidae): a reevaluation of evolu- Monograph 7: 73±106. tion in ecology, life history, and morphology. Tyler, M.J. 1982. Frogs, 2nd ed. Sydney: Collins. Systematic Biology 45: 451±472. Tyler, M.J. 1985. Reproductive modes in Austra- Trewevas, E. 1933. The hyoid and larynx of the lian Amphibia. In G. Grigg, R. Shine, and H. Anura. Philosophical Transactions of the Royal Ehman (editors), Biology of Australasian frogs Society of London, B, Biological Sciences 222: and reptiles: 265±267. Sydney: Royal Zoolog- 401±527. ical Society of New South Wales. Trontelj, P., and S. Goricki. 2003. Monophyly of Tyler, M.J. 1989. Australian frogs. Ringwood, the family Proteidae (Amphibia: Caudata) test- Victoria, Australia: Viking O'Neil. ed by phylogenetic analysis of mitochondrial Tyler, M.J., and M.M. Davies. 1978. Species- 12S rDNA sequences. Natura Croatica. Zagreb groups within the Australopapuan hylid frog 12: 113±120. genus Litoria Tschudi. Australian Journal of Trueb, L. 1993. Patterns of cranial diversity Zoology, Supplementary Series 63: 1±47. among the Lissamphibia (Amphibia, Temnos- Tyler, M.J., and W.E. Duellman. 1995. Super®cial pondyli). In J. Hanken and B.K. Hall (editors), mandibular musculature and vocal sac structure The skull: 255±343. Chicago: University of in hemiphractine hylid frogs. Journal of Mor- Chicago Press. phology 224: 65±71. Trueb, L., and D.C. Cannatella. 1986. Systemat- Uzzell, T., and T.J. Post. 1986. Rana temporaria ics, morphology, and phylogeny of genus Pipa is not a member of the Rana boylii group. Sys- (Anura: Pipidae). Herpetologica 42: 412±449. tematic Zoology 35: 414±421. Trueb, L., and R. Cloutier. 1991. A phylogenetic Vallan, D., M. Vences, and F. Glaw. 2003. Two investigation of the inter- and intrarelationships new species of the Boophis mandraka complex of the Lissamphibia (Amphibia: Temnospon- (Anura, Mantellidae) from the Andasibe region dyli). In H.P. Schultze and L. Trueb (editors), in eastern Madagascar. Amphibia-Reptilia 24: Origins of the higher groups of tetrapods: con- 305±319. troversy and consensus: 223±313. Ithaca, NY: Van der Hoeven, J. 1833. Handboek der dierkun- Cornell University Press. de, grondbeginsels der natuurlijke geschiedenis 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 287

van het dierenrijk, tweeden deels, tweede stuk. Madagascar/Diversity and endemism in Mada- Amsterdam: C.G. Sulpke. gascar: 229±242. MeÂmoires de la SocieÂteÂde Van der Meijden, A., M. Vences, S. Hoegg, and BiogeÂographie. Paris. A. Meyer. 2005. A previously unrecognized ra- Vences, M., J. Kosuch, R. Boistel, C.F.B. Haddad, diation of ranid frogs in southern Africa re- E. La Marca, and S. LoÈtters. 2003b. Convergent vealed by nuclear and mitochondrial DNA se- evolution of aposematic coloration in Neotrop- quences. Molecular Phylogenetics and Evolu- ical poison frogs: a molecular phylogenetic per- tion 37: 674±685. [Seen as an electronic/pdf spective. Organisms, Diversity & Evolution 3: preprint available from the publisher.] 215±226. Van der Meijden, A., M. Vences, and A. Meyer. Vences, M., J. Kosuch, F. Glaw, W. BoÈhme, and 2004. Novel phylogenetic relationships of the M. Veith. 2003c. Molecular phylogeny of hy- enigmatic brevicipitine and scaphiophrynine peroliid treefrogs: biogeographic origin of Mal- toads as revealed by sequences from the nucle- agasy and Seychellean taxa and re-analysis of ar Rag-1 gene. Proceedings of the Royal Soci- familial paraphyly. Journal of Zoological Sys- ety of London. B (SupplementÐBiology Let- tematics and Evolutionary Research 41: 205± ters) 271: S378±S381. 213. Van Dijk, D.E. 2001. Osteology of the ranoid bur- Vences, M., J. Kosuch, S. LoÈtters, A. Widmer, rowing African anurans Breviceps and Hemi- K.H. Jungfer, J. KoÈhler, and M. Veith. 2000b. sus. African Zoology. Pretoria 36: 137±141. Phylogeny and classi®cation of poison frogs Van Gelder, R.G. 1977. Mammalian hybrids and (Amphibia: Dendrobatidae), based on mito- generic limits. American Museum Novitates chondrial 16S and 12S ribosomal RNA gene 2635: 1±25. sequences. Molecular Phylogenetics and Evo- Veith, M., and S. Steinfartz. 2004. When non- lution 15: 34±40. monophyly results in taxonomic consequenc- Vences, M., D.R. Vieites, F. Glaw, H. Brinkmann, esÐthe case of Mertensiella within the Sala- J. Kosuch, M. Veith, and A. Meyer. 2003d. mandridae (Amphibia: Urodela). Salamandra Multiple overseas dispersal in amphibians. Pro- 40: 67±80. ceedings of the Royal Society of London. Se- Vences, M., F. Andreone, F. Glaw, and J.E. Ran- ries B, Biological Sciences 270: 2435±2442. drianirina. 2003a. Molecular and bioacoustic Vences, M., S. Wanke, G. Odierna, J. Kosuch, and divergence in Mantidactylus granulatus and M. zavona n. sp. (Anura: Mantellidae): bearings M. Veith. 2000c. Molecular and karyological for the biogeography of northern Madagascar. data on the south Asian ranid genera Indirana, African Zoology. Pretoria 38: 67±78. Nyctibatrachus and Nannophrys (Anura: Rani- Vences, M., and F. Glaw. 2001. When molecules dae). Hamadryad 25: 75±82. claim for taxonomic changes: new proposals on Wagler, J.G. 1828. VorlaÈu®ge Uebersicht des Ger- the classi®cation of Old World treefrogs (Am- uftes, sowie Untungigung feines Systema am- phibia, Anura, Ranoidea). Spixiana. MuÈnchen phibiorum. Isis von Oken 21: 859±861. 24: 85±92. Wagler, J.G. 1830. NatuÈrliches System der Am- Vences, M., and F. Glaw. 2004. Revision of the phibien, mit vorangehender Classi®cation der subgenus Chonomantis (Anura: Mantellidae: SaÈugthiere und Vogel. Ein Beitrag zur verglei- Mantidactylus) from Madagascar, with descrip- chenden Zoologie. MuÈnchen, Stuttgart and TuÈ- tion of two new species. Journal of Natural His- bingen: J.G. Cotta. tory. London 38: 77±118. Wake, D.B. 1966. Comparative osteology and Vences, M., F. Glaw, F. Andreone, R. Jesu, and G. evolution of the lungless salamanders, family Schimmenti. 2002. Systematic revision of the Plethodontidae. Memoires of the Southern Cal- enigmatic Malagasy broad-headed frogs (Lau- ifornia Academy of Sciences 4: 1±111. rentomantis Dubois, 1980) and their phyloge- Wake, D.B. 1993. Phylogenetic and taxonomic is- netic position within the endemic mantellid ra- sues relating to salamanders of the family diation of Madagascar. Bijdragen tot de Dier- Plethodontidae. Herpetologica 49: 229±237. kunde 70: 191±212. Wake, D.B., and P. Elias. 1983. New genera and Vences, M., F. Glaw, J. Kosuch, I. Das, and M. new species of Central American salamanders, Veith. 2000a. Polyphyly of Tomopterna (Am- with a review of the tropical genera (Amphibia, phibia: Ranidae) based on sequences of the mi- Caudata, Plethodontidae). Natural History Mu- tochondrial 16S and 12S rRNA genes, and eco- seum of Los Angeles County Contributions in logical biogeography of Malagasy relict am- Science. 345: 1±19. phibian groups. In W.R. LourencËo and S.M. Wake, D.B., and J.F. Lynch. 1976. The distribu- Goodman (editors), Diversite et endeÂeÂmisme aÁ tion, ecology and evolutionary history of pleth- 288 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

odontid salamanders in tropical America. Nat- Wheeler, Q.D., P.H. Raven, and E.O. Wilson. ural History Museum of Los Angeles County 2004. Taxonomy: impediment or expedient? Science Bulletin 175: 1±65. Science 303: 285. Wake, D.B., L.R. Maxson, and G.Z. Wurst. 1978. Wheeler, W.C. 1992. Extinction, sampling, and Genetic differentiation, albumin evolution, and molecular phylogenetics. In M.J. Novacek and their biogeographic implications in plethodon- Q.D. Wheeler (editors), Extinction and phylog- tid salamanders of California and southern Eu- eny: 205±215. New York: Columbia University rope. Evolution 32: 529±539. Press. Wake, M.H. 1977. Fetal maintenance and its evo- Wheeler, W.C. 1994. Sources of ambiguity in nu- lutionary signi®cance in the Amphibia: Gym- cleic acid sequence alignment. In B. Schier- nophiona. Journal of Herpetology 11: 379±386. water, B. Streit, G.P. Wagner, and R. DeSalle Wake, M.H. 1980. The reproductive biology of (editors), Molecular ecology and evolution: ap- Nectophrynoides malcolmi (Amphibia: Bufon- proaches and applications: 323±352. Basel, idae), with comments on the evolution of re- Switzerland: BirkhaÈuser. productive modes in the genus Nectophryno- Wheeler, W.C. 1995. Sequence alignment, param- ides. Copeia 1980: 193±209. eter sensitivity, and the phylogenetic analysis of Wake, M.H. 1993. Non-traditional characters in molecular data. Systematic Biology 44: 321± the assessment of caecilian phylogenetic rela- 331. tionships. Herpetological Monographs 7: 42± Wheeler, W.C. 1996. Optimization alignment: the 55. end of multiple sequence alignment in phylo- Wake, M.H., G. Parra-Olea, and J.P.-Y. Shee. genetics? Cladistics 12: 1±9. 2005. Biogeography and molecular phylogeny Wheeler, W.C. 1998. Alignment characters, dy- of certain New World caecilians. In M.A. Don- namic programing and heuristic solutions. In R. DeSalle and B. Schierwater (editors), Molecu- nelly, B.I. Crother, C. Guyer, M.H. Wake, and lar approaches to ecology and evolution, 2nd M.E. White (editors), Ecology and evolution in ed.: 243±251. Basel, Switzerland: BirkhaÈuser. the tropics: a herpetological perspective: 48± Wheeler, W.C. 1999. Fixed character states and 64. Chicago: University of Chicago Press. the optimization of molecular sequence data. Walls, J.G. 1994. Jewels of the rainforestÐpoison Cladistics 15: 379±385. frogs of the family Dendrobatidae. Neptune Wheeler, W.C. 2000. Heuristic reconstruction of City, New Jersey: T.F.H. Publications. hypothetical-ancestral DNA sequences: se- Wang, L., and T. Jiang. 1994. On the complexity quence alignment vs optimization. In R.W. of multiple sequence alignment. Journal of Scotland and R.T. Pennington (editors), Ho- Computational Biology 1: 337±348. mology and systematics: 106±113. New York: Wassersug, R.J. 1984. The Pseudohemisus tad- Taylor and Francis. pole: a morphological link between microhylid Wheeler, W.C. 2001. Homology and the optimi- (Orton Type 2) and ranoid (Orton Type 4) lar- zation of DNA sequence data. Cladistics 17: vae. Herpetologica 40: 138±148. S3±S11. Wassersug, R.J., and W.R. Heyer. 1983. Morpho- Wheeler, W.C. 2002. Optimization alignment: logical correlates of subaerial existence in lep- down, up, error, and improvements. In R. todactylid tadpoles associated with ¯owing wa- DeSalle, G. Giribet, and W.C. Wheeler (edi- ter. Canadian Journal of Zoology 61: 761±769. tors), Techniques in molecular systematics and Wassersug, R.J., and W.F. Pyburn. 1987. The bi- evolution: 55±69. Basel, Switzerland: BirkhaÈu- ology of the Pe-ret toad, ser. (Microhylidae), with special consideration of Wheeler, W.C. 2003a. Iterative pass optimization its larva and systematic relationships. of sequence data. Cladistics 19: 254±260. Zoological Journal of the Linnean Society. Wheeler, W.C. 2003b. Implied alignment: a syn- London 91: 137±169. apomorphy-based multiple sequence alignment Wei, G., N. Xu, D. Li, G. Wu, and X. Song. 1993. method. Cladistics 19: 261±268. Karyotype C-band and Ag-NORs study of three Wheeler, W.C. 2003c. Search-based optimization. stink frogs. Asiatic Herpetological Research 5: Cladistics 19: 348±355. 45±50. Wheeler, W.C., and D.S. Gladstein. 1992. Malign, Werner, F. 1896. BeitraÈge zur Kenntniss der Rep- version 1.2. New York: Computer software dis- tilien und Batrachier von Centralamerika und tributed by the authors. Chile, sowie einiger seltenerer Schlangenarten. Wheeler, W.C., D.S. Gladstein, and J. De Laet. Verhandlungen des Zoologisch-Botanischen 1996±2003. POY: Phylogeny reconstruction via Vereins in Wien 46: 344±365. optimization of DNA data. New York: Com- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 289

puter software distributed by the authors and porary Herpetology 2000: 1±14. [Electronic from the American Museum of Natural Histo- journal available at http://www.nhm.ac.uk/ ry: ftp://ftp.amnh.org/pub/molecular/poy. hosted࿞sites/ch/ch/index.htm.] Widmer, A., S. LoÈtters, and K.-H. Jungfer. 2000. Wilkinson, J.A., R.C. Drewes, and O.L. Tatum. A molecular phylogenetic analysis of the Neo- 2002. A molecular phylogenetic analysis of the tropical dart-poison frog genus Phyllobates family Rhacophoridae with an emphasis on the (Amphibia: Dendrobatidae). Naturwissenschaf- Asian and African genera. Molecular Phyloge- ten. Berlin 87: 559±562. netics and Evolution 24: 265±273. Wieczorek, A.M., A. Channing, and R.C. Drewes. Wilkinson, J.A., M. Matsui, and T. Terachi. 1996. 1998. A review of the taxonomy of the Hyper- Geographic variation in a olius viridi¯avus complex. Herpetological Jour- (Rhacophorus arboreus) revealed by PCR-aid- nal. London 8: 29±34. ed restriction site analysis of mtDNA. Journal Wieczorek, A.M., R.C. Drewes, and A. Channing. of Herpetology 30: 418±423. 2000. Biogeography and evolutionary history Wilkinson, M. 1991. Adult crown morphol- of Hyperolius species: application of molecular ogy in the Typhlonectidae (Amphibia: Gym- phylogeny. Journal of Biogeography. Oxford nophiona): a reinterpretation of variation and its 27: 1231±1243. signi®cance. Zeitschrift fuÈr Zoologische Syste- Wieczorek, A.M., R.C. Drewes, and A. Channing. matik und Evolutionsforschung 29: 304±311. 2001. Phylogenetic relationships within the Hy- Wilkinson, M. 1997. Characters, congruence and perolius viridi¯avus complex (Anura: Hypero- quality: a study of neuroanatomical and tradi- liidae), and comments on taxonomic status. tional data in caecilian phylogeny. Biological Amphibia-Reptilia 22: 155±166. Reviews of the Cambridge Philosophical Soci- Wied-Neuwied, M.A.P., Prinz zu. 1825. BeitraÈge ety 72: 423±470. zur Naturgeschichte von Brasilien, vol. 1 (Ver- Wilkinson, M., S.P. Loader, D.J. Gower, J.A. zeichniss der Amphibien). Weimar, Germany: Sheps, and B.L. Cohen. 2003. Phylogenetic re- Gr. H.S. priv. Landes-Industrie-Comptoir. lationships of African caecilians (Amphibia: Wiedersheim, R. 1877. Das Kopfskelet der Uro- Gymnophiona): insights from mitochondrial delen (Fortsetzung). Morphologisches Jahr- rRNA gene sequences. African Journal of Her- buch. Leipzig 3: 459±548. petology 52: 83±92. Wiens, J.J. 1989. Ontogeny of the skeleton of Spea bombifrons (Anura: Pelobatidae). Journal Wilkinson, M., and R.A. Nussbaum. 1996. On the of Morphology 202: 29±51. phylogenetic position of the Uraeotyphlidae Wiens, J.J., R.M. Bonett, and P.T. Chippindale. (Amphibia: Gymnophiona). Copeia 1996: 550± 2005. Ontogeny discombobulates phylogeny: 562. paedomorphosis and higher-level salamander Wilkinson, M., and R.A. Nussbaum. 1999. Evo- relationships. Systematic Biology 54: 91±110. lutionary relationships of the lungless caecilian Wild, E.R. 1995. New genus and species of Am- Atretochoana eiselti (Amphibia: Gymnophiona: azonian microhylid frog with a phylogenetic Typhlonectidae). Zoological Journal of the Lin- analysis of New World genera. Copeia 1995: nean Society. London 126: 191±223. 837±849. Wilkinson, M., J.A. Sheps, O.V. Oommen, and Wild, E.R. 1997. Description of the adult skeleton B.L. Cohen. 2002. Phylogenetic relationships and developmental osteology of the hyperossi- of Indian caecilians (Amphibia: Gymnophiona) ®ed horned frog, Ceratophrys cornuta (Anura: inferred from mitochondrial rRNA gene se- Leptodactylidae). Journal of Morphology 232: quences. Molecular Phylogenetics and Evolu- 169±206. tion 23: 401±407. Wild, E.R. 1999. Description of the chondrocra- Wilkinson, M., J. Thorley, and M.J. Benton. 1997. nium and osteogenesis of the chacoan burrow- Uncertain turtle relationships. Nature. London ing frog, Chacophrys pierotti (Anura: Lepto- 387: 466. dactylidae). Journal of Morphology 242: 229± Withgott, J. 2000. Is it ``So long, Linnaeus''? 249. BioScience 50: 646±651. Wiley, E.O. 1981. Phylogenetics. The theory and Wogel, H., P.A. Abrunhosa, and J.P. Pombal, Jr. practice of phylogenetic systematics. New 2004. Vocalizations and aggressive behavior of York: Wiley Interscience. Phyllomedusa rohdei (Anura: Hylidae). Her- Wilkinson, J.A., and R.C. Drewes. 2000. Char- petological Review 35: 239±243. acter assessment, genus level boundaries, and Wu, L., R. Xu, Q. Dong, D. Li, and J. Liu. 1983. phylogenetic analysis of the family Rhacophor- A new species of Rana and records of amphib- idae: a review and present day status. Contem- ians from Guizhou Province. Acta Zoologica 290 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Sinica 29: 66±70. [In Chinese, with English caecilians (Amphibia: Gymnophiona). Genetics summary.] 155: 765±775. Wu, S.-H. 1994. Phylogenetic relationships, high- Zardoya, R., and A. Meyer. 2001. On the origin er classi®cation and historical biogeography of of and phylogenetic relationships among living the microhyloid frogs (Lissampibia: Anura: amphibians. Proceedings of the National Acad- Brevicipitidae and Microhylidae). Ph.D. disser- emy of Sciences of the United States of Amer- tation. Department of Zoology, University of ica 98: 7380±7383. Michigan, Ann Arbor. Zhang, P., Y.Q. Chen, Y.F. Liu, H.-Y. Zhou, and Xie, F., and Z. Wang. 2000. [Review of the sys- L.H. Qu. 2003a. The complete mitochondrial tematics of pelobatids.] Cultum Herpetologica genome of the Chinese , An- Sinica 8: 356±370. [In Chinese.] drias davidianus (Amphibia: Caudata). Gene Yang, D. (editor), 1991a. [The Amphibia-fauna of 311: 93±98. Yunnan.] Beijing, China: China Forestry Pub- Zhang, P., Y.Q. Chen, H.-Y. Zhou, X.L. Wang, and lishing House. [In Chinese.] L.H. Qu. 2003b. The complete mitochondrial Yang, D. 1991b. Phylogenetic systematics of the genome of a relic salamander, Ranodon sibiri- Amolops group of ranid frogs of southeastern cus (Amphibia: Caudata) and implications for Asia and the Greater Sunda Islands. Fieldiana. amphibian phylogeny. Molecular Phylogenetics Zoology. New Series 63: 1±42. and Evolution 28: 620±626. Yang, D., and S. Li. 1980. A new species of the Zhao, E. 1994. A study on vomerine teeth pattern genus Rana from Yunnan. Zoological Research. of the genus , with revised diagnoses of Kunming 1: 261±264. Liua and Ranodon (Caudata: Hynobiidae). Yokoyama, S., H. Zhang, F.B. Radlwimmer, and Sichuan Journal of Zoology 13: 162±166. N.S. Blow. 1999. Adaptive evolution of color Zhao, E., and S.-Q. Li. 1984. A new species of vision of the Comoran coelacanth (Latimeria the genus Platymantis (Amphibia: Ranidae) chalumnae). Proceedings of the National Acad- from Xizang. Acta Herpetologica Sinica, New emy of Sciences of the United States of Amer- Series 3: 55±57. [In Chinese with English sum- ica 96: 6279±6284. mary.] Young, B.E., K.R. Lips, J.K. Reaser, R. IbaÂnÄez D., Ziegler, M., and M. Vences. 2002. The tadpole of A.W. Salas, J.R. CedenÄo, L.A. Coloma, S.R. Rhacophorus verrucosus Boulenger, 1893, Ron, E. La Marca, J.R. Meyer, A. MunÄoz, F. from Vietnam (Amphibia: Anura: Rhacophori- BolanÄos, G. Chaves, and D. Romo. 2001. Pop- ulation declines and priorities for amphibian dae). Faunistische Abhandlungen. Staatliches conservation in Latin America. Conservation Museum fuÈr Tierkunde in Dresden 22: 319± Biology 15: 1213±1223. 327. Zacj, I., and J.W. Arntzen. 1999. Phylogenetic re- Zug, G.R., L.J. Vitt, and J.P. Caldwell. 2001. Her- lationships of the European newts (genus Tri- petology: an introductory biology of amphibi- turus) tested with mitochondrial DNA sequence ans and reptiles. New York: Academic Press. data. Contributions to Zoology. Amsterdam 68: Zweifel, R.G. 1955. Ecology, distribution, and 78±81. systematics of frogs of the Rana boylei group. ZaldõÂvar-RiveroÂn, A., V. LeoÂn-Regagnon, and A. University of California Publications in Zool- Nieto-Montes de Oca. 2004. Phylogeny of the ogy 54: 207±292. Mexican coastal leopard frogs of the Rana ber- Zweifel, R.G. 1956. Two pelobatid frogs from the landieri group based on mtDNA sequences. Tertiary of North America and their relation- Molecular Phylogenetics and Evolution 30: 38± ships to fossil and recent forms. American Mu- 49. seum Novitates 1762: 1±45. Zardoya, R., and A. Meyer. 1996. Evolutionary Zweifel, R.G. 1971. Results of the Archbold Ex- relationships of the coelacanth, lung®shes, and peditions. No. 96. Relationships and distribu- tetrapods based on the 28S ribosomal RNA tion of Genyophryne thomsoni, a microhylid gene. Proceedings of the National Academy of frog of New Guinea. American Museum Nov- Sciences United States of America 93: 5449± itates 2469: 1±13. 5454. Zweifel, R.G. 1972. Results of the Archbold Ex- Zardoya, R., and A. Meyer. 1998. Complete mi- peditions. No. 97. A revision of the frogs of the tochondrial genome suggests diapsid af®nities subfamily Asterophryinae, family Microhyli- of turtles. Proceedings of the National Acade- dae. Bulletin of the American Museum of Nat- my of Sciences of the United States of America ural History 148: 411±546. 95: 14226±14231. Zweifel, R.G. 1986. A new genus and species of Zardoya, R., and A. Meyer. 2000. Mitochondrial microhylid frog from the Cerro de la Neblina evidence on the phylogenetic position of region of Venezuela and a discussion of rela- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 291

tionships among New World microhylid gen- American Museum of Natural History 253: 1± era. American Museum Novitates 2863: 1±24. 130. Zweifel, R.G. 2000. Partition of the Australopa- Zwickl, D.J., and D.M. Hillis. 2002. Increased puan microhylid frog genus Sphenophryne with taxon sampling greatly reduces phylogenetic descriptions of new species. Bulletin of the error. Systematic Biology 51: 588±598. 292 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

APPENDIX 1

VOUCHER AND DNA LOCUS INFORMATION Below we provide the specimen or tissue identi®cation (ID) numbers (or source, if ID number unavail- able), localities, GenBank numbers, and total number of base pairs (bp) analyzed for each terminal in the analysis. The locus mtDNA refers to 12S, tRNAVal, and 16S sequences, and SIA refers to the gene Seven in Absentia. Asterisks (*) mark the 85 species for which all sequence data were obtained from GenBank and not generated by us. Species for which morphological data were included from Haas (2003) are in boldface. Sequences obtained from or previously deposited in GenBank are given in bold (see appendix 2 for references); all other sequences are new. ID numbers and localities are given only for sequences generated by us. Localities for conspeci®c tissues are separated by a semicolon. Abbre- viations are: ABTC (Australian Biological Tissue Collection, South Australian Museum, Adelaide), AC (Alan Channing ®eld series), ACD (Arvin C. Deismos ®eld series), AH (Alexander Haas), AMCC (Ambrose Monell Cryo-Collection, American Museum of Natural History, New York); AMNH (Amer- ican Museum of Natural History, New York), AMS (Australian Museum, Sydney), ARBT (Adam Back- lin ®eld series, via Robert Fisher), ASU (Arizona State University, Tempe), ATH (Andrew T. Holycross ®eld series), BB (Boris Blotto ®eld series), BLC (Bruce L. Christman), BMNH (The Natural History Museum, London), BPN (Brice P. Noonan ®eld series), BY (Brian Yang), CAR (Channing collection, deposited at SAM), CAS (California Academy of Sciences, San Francisco), CFBH (CeÂlio F.B. Haddad specimen collection), CFBH-T (CeÂlio F.B. Haddad tissue collection), CG (Caren Goldberg), DMG (David M. Green ®eld series), DPL (Dwight P. Lawson ®eld series), ENS (Eric N. Smith ®eld series), FMNH (Field Museum, Chicago), IWK (Iwokrama collection ®eld series, Maureen Donnelly), IZUA (Instituto de ZoologõÂa, Universidad Austral de Chile, Valdivia), JAC (Jon- athan A. Campbell ®eld series), JF (JuliaÂn Faivovich ®eld series), JLG (JoaÄo Luiz Gasparini ®eld series), KRL (Karen R. Lips ®eld series), KU (Natural History Museum, University of Kansas, Lawrence), LSUMZ (Louisiana State University Museum of Zoology, Baton Rouge), MACN (Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires), MAD (Maureen A. Donnelly ®eld series), MB (Marius Burger ®eld series), MHNSM (Museo de Historia Natural San Marcos, Lima, Peru), MJH (Martin J. Henzl ®eld series), MLPA (Museo de la Plata, Buenos Aires, Argentina), MVZ (Museum of Vertebrate Zoology, University of California at Berkeley), MW (Mark Wilkinson ®eld series), NK (Museo de Historia Natural Noel Kempff Mercado, Santa Cruz, Bolivia), NTM (Museum and Art Galleries of the Northern Territory, Darwin, Australia), QMJ (Queensland Museum, Brisbane), RABI (Marius Burger, Rabi oil®eld, , ®eld series), RAN (Ronald A. Nussbaum ®eld series), RAX (Christopher Raxworthy ®eld series), RdS (Rafael de Sa collection), RG (Ron Gagliardo), RNF (Robert N. Fisher ®eld series), RWM (Roy W. McDiarmid ®eld series), SAM (South African Museum, Cape Town), SAMA (South Australian Museum, Adelaide), SIUC (Southern Illinois University at Carbon- dale), TAT (Tom A. Titus ®eld series), TMSA (Transvaal Museum, Pretoria, South Africa), UAZ (Herpetology Collection, University of Arizona, Tucson), USNM (National Museum of Natural History, Smithsonian Institution, Washington, D.C.), UMFS (University of Michigan Museum of Zoology, Ann Arbor, ®eld series), UMMZ (University of Michigan Museum of Zoology, Ann Arbor), UTA (University of Texas at Arlington), WAM (Western Australia Museum, Perth), WCS (Wildlife Conservation Society, New York), WR (Wade Ryberg), ZFMK (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany), and ZSM (Zoologisches Museum, MuÈnchen, Germany).

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Acanthixalus AJ437002 1283 spinosus* AF215214 AF465438 Acris crepitans LSUMZ USA, , De AY843559 DQ284107 AY844533 AY844762 AY844019 3952 H-2164 Kalb Co, powerline access, 0.1 mi W Lookout Mt Boys Camp Rd Adelotus brevis SAMA Australia, Queensland, DQ283298 DQ284307 DQ283948 DQ282800 DQ283638 3731 R39251 Nambour Adenomera MJH 3669 Peru, HuaÂnuco, RõÂo DQ283063 DQ284093 DQ283790 3063 hylaedactyla Llullapichis, Panguana 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 293

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Afrana angolensis CAS 202040 Uganda, Rukungiri DQ284258 DQ284258 DQ283597 1600 Dist, Bwindi Impenetrable National Park, Munyaga Falls trail Afrana fuscigula AMNH South Africa, Western DQ283069 DQ284105 DQ283794 DQ282909 DQ283476 4162 A144976 Cape Prov, Bainskloof, in stream at settlement at crest of pass Afrixalus fornasinii AMNH Tanzania, Morogoro, U22071 DQ284382 DQ284013 DQ282859 DQ283003 DQ283713 4289 A153277 Udzungwa Mts DQ283401 National Park, Man'gula camp site 3 on Mwaya River, 350 m, 7Њ50Ј51ЉS, 36Њ53Ј0ЉE Afrixalus pygmaeus CAS 214836 Kenya, Kili® Dist, DQ283234 DQ284263 DQ283908 DQ282765 DQ282955 DQ283602 4265 Kararacha Pond I, 03Њ24Ј54ЉS, 39Њ52Ј19.8ЉE Agalychnis RdS 537 Belize, Stann Creek AY843563 DQ284401 AY844537 AY844765 DQ283018 4017 callidryas Dist, Cockscomb Basin Wildlife Sanctuary Aglyptodactylus UMMZ Madagascar, DQ283056 DQ283785 DQ282906 DQ283469 3927 madagascariensis 198472 Toamasina, Moramanga, Mantady Park, 48.458333ЊS, 18.85ЊE Alexteroon MB 5515 Gabon, Rabi (Shell DQ283171 DQ284209 DQ283864 DQ282723 DQ282969 DQ283561 2311 obstetricans (SAM) Gabon), at Rabi 059, at trap lines 1±3, 01Њ56Ј33ЉS, 09Њ51Ј09ЉE Alexteroon UTA Cameroon, DQ283344 DQ282820 DQ283666 3135 obstetricans A44465 Southwestern Prov, vicinity Ediensoa Alligator sinensis WCS 850352 No data (WCS) NC004448 DQ283961 DQ282809 DQ283650 3848 LSUMZ Brazil, RondoÃnia, Rio DQ283045 DQ284074 DQ283774 DQ282657 DQ283465 4226 17552 Formoso, Parque Estadual Guajira- Mirim, ca. 90 km N Nova Mamore, 10Њ19ЈS, 64Њ33ЈW Allophryne MAD 1512 Guyana, Kabocali AF364511 AY844538 AY844766 3134 ruthveni camp, 101 m, AF364512 4Њ17.10ЈN, AY843564 58Њ30.56ЈW MACN Argentina, NeuqueÂn, AY843565 DQ284118 AY844539 AY844767 AY844197 4211 37942 AlumineÂ, stream 10 km W Primeros Pinos Alytes obstetricans AH Germany, ThuÈringen, DQ283112 DQ284158 AY364385 DQ282683 DQ283510 4155 Schnellbach, 725 m Ambystoma AMCC USA, Florida, DQ283184 DQ284218 2697 cingulatum 125631 Wakulla Co, 30Њ08.96ЈN, 84Њ09.23ЈW Ambystoma AMCC No data DQ283213 DQ284244 DQ283893 3025 mexicanum 105479 Ambystoma AMNH USA, Arizona, DQ283407 DQ284388 U36574 DQ282864 3422 tigrinum A164658 Cochise Co, Hwy 80, 0.5 mi N Price Canyon Rd, ca. 50 m W Hwy 80, ca. 1401 m, 31Њ38Ј15ЉN, 109Њ11Ј27ЉW 294 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Amerana muscosa BY USA, California, Los DQ283190 DQ284224 DQ283877 DQ282735 DQ282945 DQ283575 4686 Angeles Co, tributary of South Fork Big Rock Creek, 34.3775ЊN, 117.82824ЊW Amietia vertebralis AMNH Lesotho, Katse DQ283402 DQ284383 DQ282860 DQ283004 DQ283714 4382 A144977 Amnirana UTA Cameroon, DQ283368 DQ284354 DQ283989 DQ283687 3273 albilabris A44423 Southwestern Prov, Kumba±Mamfe rd, 6.4 km S Nguti Amnirana KU 290412 Ghana, Muni Lagoon, DQ283058 AY341808 AY341749 3250 galamensis Winneba, 5Њ21Ј14ЉN, 0Њ42Ј12ЉW Amolops AMNH Vietnam, Lai Chau DQ283372 DQ284358 DQ283992 DQ282837 DQ282984 DQ283690 4695 chapaensis A161439 Prov, Mt Fansipan, 1600 m Amolops AF206072 1939 hongkongensis* AF206453 AF206117 Amphiuma UMFS No data DQ283372 DQ284358 DQ283690 3410 tridactylum 10349 Andrias AJ492192 2390 davidianus* Andrias japonicus UMFS No data (Detroit Zoo; DQ283274 DQ284358 2714 11734 living animal) Aneides hardii* AY728226 2342 Anhydrophryne AF215504 500 rattrayi* Anodonthyla AJ314812 493 montana* Anotheca spinosa ENS 10039 Mexico, Oaxaca, AY843566 DQ284101 AY844540 AY844768 AY844022 AY844198 4730 IxtlaÂn de JuaÂrez, Santiago Comaltepec, Vista Hermosa Ansonia FMNH Malaysia, Sabah, DQ283341 DQ283968 DQ282817 3136 longidigitata 242550 Sipitang Dist, 3.3 km W Mendolong camp Ansonia muelleri* U52740 1200 U52784 Aphantophryne ABTC 49605 Papua New Guinea, DQ283195 DQ284228 DQ283879 DQ282739 DQ283578 4171 pansa Bolulo Aplastodiscus MACN Argentina, Misiones, AY843569 DQ284044 AY844543 AY844771 AY844025 AY844201 4746 perviridis 37791 GuaranõÂ, San Vicente, Campo Anexo INTA ``Cuartel RõÂo Victoria'' Aquarana BLC USA, New Mexico, DQ283257 DQ283926 DQ282778 DQ282959 DQ283618 4363 catesbeiana Sierra Co, Las Animas Creek, Ladder Ranch Aquarana AMCC USA, Florida, Walton DQ283185 DQ284219 DQ283872 DQ282730 DQ283570 4160 clamitans 125633 Co, Eglin Air Force Base, Range Rd 211, ca. 1.2 mi E Indigo Pond, 30Њ53.28ЈN, 86Њ53.26ЈW Aquarana grylio AMCC USA, Florida, Walton DQ283186 DQ284220 DQ283873 DQ282731 DQ283571 4160 125634 Co, Eglin Air Force Base, Range Rd 211, ca. 1.2 mi E Indigo Pond, 30Њ53.28ЈN, 86Њ53.26ЈW Aquarana AMCC USA, Florida, DQ283191 DQ284225 DQ283878 DQ282736 DQ282946 DQ283576 4693 heckscheri 125635 Wakulla Co, Smith Creek at County Rd 375 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 295

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Aquixalus AMNH Vietnam, Ha Giang DQ283051 DQ283780 2712 gracilipes A163897 Prov, Vi Xuyen, Cao Bo Commune, Mt Tay Conn Linh II, above Tham Ve Village, small forest pools ca. 1 km SW high camp, 1700 m, 22Њ45Ј27ЉN, 104Њ49Ј35ЉE Arenophryne WAM Australia, Western DQ283326 DQ284322 DQ283965 DQ283656 2268 rotunda R146480 Australia, False Entrance Well Argenteohyla MACN Argentina, Corrientes, AY843570 DQ284064 AY844544 AY844772 AY844026 AY844202 4734 siemersi pederseni 38644 Bella Vista, junction R.N. 12 at RõÂo Santa LucõÂa Arthroleptella AMNH South Africa, Western DQ283070 DQ284106 DQ283795 DQ282662 DQ282910 DQ283477 4701 bicolor A144967 Cape Prov, Landdroskop (nr peak), ca. 15 km SW Stellenbosch (by air), ca. 1450 m Arthroleptides RdS 862 Tanzania, Uluguru DQ283415 DQ284396 DQ284020 DQ282872 DQ283010 DQ283725 4729 yakusini Mts, Tegetero Village, 6Њ56Ј30ЉS, 37Њ43Ј10ЉE Arthroleptis RdS 929 Tanzania, West DQ283427 DQ284405 DQ284028 DQ283020 DQ283736 3844 tanneri Usambara Mts, Mazumbai, 04Њ48Ј46.5ЉS, 38Њ30Ј12.0ЉE Arthroleptis UTA Cameroon, DQ283081 DQ284133 DQ283803 DQ282914 DQ283483 4263 variabilis A44448 Southwestern Prov, vicinity Babong Ascaphus truei UMFS USA, Washington, AJ440760 DQ284162 DQ283514 2905 10198 Skamania Co, McCloskey Creek at Maybee Mines Rd Assa darlingtoni SAMA Australia, New South DQ283284 DQ284300 DQ283943 2128 R39233 Wales, Wiangaree Astylosternus UTA 52398 Cameroon, South DQ283349 DQ284340 DQ283976 DQ282826 DQ282974 DQ283674 4699 schioetzi Prov Atelognathus MACN Argentina, NeuqueÂn, AY843571 AY844545 AY844773 AY844027 AY844203 4408 patagonicus 37905 Catan Lil, Laguna del Burro Atelopus ¯avescens BPN 726 , N side DQ283259 DQ284282 DQ283928 DQ282780 3462 (UTA) of mt just S Kaw BPN 754 French Guiana, Grand DQ283260 DQ284283 DQ283929 DQ282781 DQ283619 4229 (UTA) Boe of Mort (circuit trail) just SSW Saul Atelopus zeteki UMFS Captive raised, Detroit DQ283252 1518 11492 Zoo (parental stock from Panama, Las Filipinas near Sora, 8Њ39.99ЈN, 80Њ0.249ЈW) Aubria subsigillata MB 5855 Gabon, at forest DQ283172 DQ284210 DQ283865 DQ282724 DQ283562 3689 (SAM) stream in Gamba DQ283173 region, 02Њ46Ј30ЉS, 10Њ00Ј59ЉE Aubria subsigillata DPL 4936 Cameroon, DQ283350 DQ284341 DQ283977 DQ282827 DQ282975 DQ283675 3880 (UTA) Southwestern Prov, DQ283351 Nguti±Bayenti rd DQ283352 Aurorana aurora ARBT 018 USA, California, Los DQ283189 DQ284223 DQ283876 DQ282734 DQ282944 DQ283574 4683 Angeles, San Francisquito Canyon, plunge pool, 34.545767ЊN, 118.51653ЊW CFBH-T 306 Brazil, Minas Gerais, DQ283094 DQ284144 DQ283810 DQ284144 DQ283496 2938 GurinhataÄ 296 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Batrachoseps AY728228 2327 attenuatus* Batrachoseps AY728221 2335 wrightorum* Batrachuperus FMNH China, Sichuan, DQ283339 DQ284330 DQ283664 3017 pinchoni 232841 Hongya Xian, 15 km SW Bin Ling, top of Wa Shan Batrachyla MACN Argentina, Chubut, AY843572 DQ284119 AY844546 AY844774 AY844028 AY844204 4726 leptopus 38008 Cushamen, Lago Puelo Batrachylodes AMS Solomon Islands, New DQ283210 DQ284242 DQ283891 DQ282753 DQ283586 4124 vertebralis R134887 Georgia, Patutiva Bolitoglossa JAC 21178 Mexico, Oaxaca, DQ283210 DQ284242 DQ282753 DQ283586 2890 rufescens Coconales±Zacatepec Hwy, 1625 m Bombina bombina AH No data DQ283250 DQ284275 DQ283920 3067 Bombina AMNH Vietnam, Ha Giang DQ283408 DQ284389 DQ284017 DQ282865 DQ283718 4171 microdeladigitora A163789 Prov, Vi Xuyen, Cao Bo Commune, Mt Tay Conn Linh II, above Tham Ve Village, 1900 m Bombina orientalis RdS No data DQ283432 DQ284032 DQ283741 3474 Bombina variegata ZSM 724/ No data DQ283249 DQ284274 DQ283919 DQ283612 3779 2000 Boophis albilabris RAX 2714 Madagascar, DQ283033 DQ284054 DQ283762 3050 Antsiranana, Ambanja, Antsahatelo Camp, Tsaratanana Reserve, 13Њ51Ј35ЉS, 48Њ51Ј59ЉE Boophis AMNH Madagascar, DQ283032 DQ284053 DQ283761 AF249168 3590 tephraeomystax A168144 Antsiranana, Ambanja, Mandrizavona Village, Ramena Valley, 13Њ48Ј3ЉS, 48Њ44Ј47ЉE Boulengerula MW 3268 Tanzania DQ283087 DQ284138 DQ282670 DQ283488 3810 uluguruensis (BMNH) Brachycephalus CFBH 2466, Brazil, SaÄo Paulo, DQ283091 DQ283808 DQ282672 DQ282919 DQ283494 4263 ephippium CFBH 2468 Campinas, Joaquim DQ283806 DQ282673 DQ282917 DQ283492 Egydio Brachytarsophrys AY236799 555 feae* Breviceps RdS 903 Tanzania, Morogoro DQ283155 DQ284397 DQ284023 DQ283013 DQ283546 4292 mossambicus Buergeria japonica UMMZ China, Taiwan, I-Lan, DQ283055 DQ283784 2710 190051 near Ran-Jeh spring Bufo alvarius ATH 499 USA, Arizona Santa DQ283269 DQ284289 DQ283933 DQ282785 DQ283625 4221 Cruz Co, 0.5 mi (by air) SW junction of I-19 and Ruby Rd Bufo amboroensis NK A5302 Bolivia, Dept Santa DQ283386 DQ284003 DQ282848 DQ283701 3894 Cruz, Prov Caballero, San JuaÂn CantoÂn, AmboroÂ, National Park, near San JuaÂn del Portrero, on RõÂo Cerro Bravo, near 17Њ50Ј08ЉS, 64Њ23Ј23ЉW, 1800± 2100 m 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Bufo andrewsi CAS 214911 China, Yunnan, Nu DQ283230 DQ284260 DQ283905 DQ282763 DQ283599 4216 Jiang Pref, small village S Gongshan, 27Њ42Ј13.7ЉN, 98Њ42Ј10.2ЉE, ca. 4760 ft Bufo angusticeps* AF220852 850 AF220899 Bufo arenarum MACN Argentina, San LuõÂs, AY843573 DQ284103 AY844547 AY844775 AY844205 4217 38639 Rte 20 between Bardas Blancas and km 330 Bufo asper FMNH Brunei, Dutong Dist, DQ283148 DQ284188 DQ283848 DQ282704 DQ282939 DQ283539 4748 248147 Tasek Merimbun, Sg Merimbun Bufo biporcatus* AY325987 2350 Bufo boreas RNF 2416 USA, California, San DQ283180 DQ284215 DQ283871 DQ283567 3821 Diego Co, Marron Valley Rd, 0.25 mi E Mine Canyon Bufo brauni RdS 952 Tanzania, East DQ283416 DQ284021 DQ282873 DQ283011 DQ283726 4420 Usambara Mts, adjacent to Amanai Nature Reserve, 05Њ07Ј38.0ЉS, 38Њ37Ј22.6ЉE Bufo bufo* AY325988 U59921 2672 Bufo camerunensis UTA Cameroon, East Prov, DQ283358 DQ284345 DQ283979 DQ282830 DQ283678 4211 A44478 ca. 35 km E Lipondji Village Bufo celebensis* AF375513 1209 AY180245 Bufo cf. arunco AMNH No data ( trade) DQ283162 DQ284200 DQ283857 DQ282715 DQ283553 4219 A168401 Bufo cognatus AMNH No data (pet trade) DQ283159 DQ284197 DQ282713 DQ283550 3902 A168396 Bufo coniferus SIUC 6913 Panama, Cocle Prov, DQ283166 DQ284204 DQ283860 DQ282719 DQ283556 4216 Parque Nacional El Cope Bufo divergens FMNH Malaysia, Sabah, DQ283149 DQ284189 DQ283849 DQ282705 DQ283540 4210 242591 Sipitang Dist, Mendelong camp, watershed Bufo galeatus AMNH Vietnam, Quang Nam DQ283376 DQ284362 DQ283995 DQ282839 DQ282987 3994 A163648 Prov, TraÄMy,TraÄ TaÄp commune, stream near Thon 2 village, 920±1060 m, 15Њ09.622ЈN, 108Њ02.427ЈE Bufo granulosus AMNH Guyana, southern DQ283332 DQ284323 DQ283966 DQ283657 3808 A139020 Rupununi Savanna, Aishalton (on Kubabawau Creek), 150 m, 2Њ28Ј31ЉN, 59Њ19Ј16ЉW Bufo guttatus AMNH Guyana, Dubulay DQ283375 DQ284361 DQ283994 DQ283693 3823 A141058 Ranch on Berbice River, 200 ft, 5Њ40Ј55ЉN, 57Њ51Ј32ЉW Bufo gutturalis RdS 873 Tanzania, Mumba DQ283436 DQ284035 DQ282890 DQ283745 3863 Village, 08Њ10Ј44.9ЉS, 31Њ51Ј47.8ЉE Bufo haematiticus SIUC 7059 Panama, Cocle Prov, DQ283167 DQ284205 DQ283861 DQ282720 DQ283557 4217 Parque Nacional El Cope 298 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Bufo latifrons UTA Cameroon, Southwest DQ283343 DQ284332 DQ283970 DQ282819 DQ283665 3310 A44695 Prov, Manyu Division, ca. 9.0 km W Bakumba Village, between Mpi River and primary forest Bufo lemur UMFS No data (Detroit Zoo) DQ283273 2424 11733 Bufo maculatus AMNH Mali, 12Њ36Ј45ЉN, DQ283388 DQ284374 DQ284005 DQ282850 DQ283703 4213 A163573 7Њ59Ј21ЉW Bufo margaritifer* AF375514 1101 AF375489 Bufo marinus MJH 3678 Peru, HuaÂnuco, RõÂo DQ283062 DQ284092 DQ283789 DQ283472 3820 Pachitea, Puerto Inca Bufo U52755 1343 mazatlanensis* U52723 Bufo AMNH Vietnam, Ha Tinh DQ283333 DQ284324 DQ283967 DQ282815 DQ283658 4237 melanostictus A161135 Prov, Huang Son Reserve, Rao An region, 200 m, 18Њ22Ј0ЉN, 105Њ13Ј13ЉE Bufo nebulifer* AY325985 2426 Bufo punctatus AMNH No data (pet trade) DQ283160 DQ284198 DQ283855 DQ282714 DQ283551 4167 A168398 Bufo quercicus AMNH USA, Florida, Walton DQ283153 DQ284192 DQ282708 DQ283544 3900 A168432 Co, Eglin Air Force Base, Range Rd 211, ca. 1.2 mi E Indigo Pond, 30Њ42Ј0ЉN, 86Њ19Ј0ЉW Bufo regularis FMNH Tanzania, Kilimanjaro DQ283163 DQ284201 DQ283858 DQ282716 DQ283554 4191 251386 Region, South Pare Mts, Chome Forest Reserve, 7 km S Bombo (by air), 4Њ20ЈS, 38Њ00ЈE, 1100 m Bufo schneideri BB 1224 Argentina, Santiago DQ283065 DQ284102 DQ283791 3068 del Estero, GuasayaÂn, DonÄa Luisa Bufo spinulosus BB 1032 Argentina, RõÂo Negro, DQ283046 DQ284077 DQ283775 DQ282658 3469 Bariloche, Pampa Linda Bufo terrestris AMNH USA, Florida, Marion DQ283158 DQ284196 DQ283854 DQ282712 DQ283549 4214 A168433 Co, 4 mi WSW Micanopy, 29Њ38.56ЈN, 82Њ20.30ЈW Bufo tuberosus UTA Cameroon, Center DQ283362 DQ283984 DQ282832 DQ283683 3856 A52375 Prov, east bank of Nyong River, vicinity 0742952, 0382754 (UTM 32N) Bufo viridis AMNH No data (pet trade) DQ283279 DQ284297 DQ283940 DQ282791 DQ283630 4220 A168402 Bufo woodhousii RNF 2417 USA, California, DQ283188 DQ284222 DQ283875 DQ282733 DQ283573 4217 Imperial Co, Winterhaven, 1.5 mi (by air) N Hwy 8 on Picacho Rd, 32.75100ЊN, 114.61596ЊW Cacosternum AC South Africa, Western DQ283258 DQ284281 DQ283927 DQ282779 DQ282960 3968 platys Cape Prov, Cape Town 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 299

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Caecilia AMNH Guyana, Dubulay DQ283406 DQ284387 DQ282863 DQ283717 2936 tentaculata A145174 Ranch on Berbice River, 200 ft., 5Њ40Ј55ЉN, 57Њ51Ј32ЉW Calluella guttulata FMNH Vietnam, Gia-Lai DQ283144 DQ284184 DQ283845 DQ282700 DQ282937 DQ283536 4714 252955 Prov, Ankhe Dist, Kannack town, Buon Luoi village, 20 km NW Kannack, Annam mts, 700±750 m, 14Њ20ЈN, 106Њ36ЈE Callulina RdS 936 Tanzania, West DQ283429 DQ284406 DQ282884 DQ283021 DQ283737 4333 kisiwamsitu Usambara Mts, Mazumbai, 04Њ48Ј46.5ЉS, 38Њ30Ј12.0ЉE Callulina kreffti* AY326068 2363 Callulops slateri* AF095339 541 Capensibufo rosei* AF220864 841 AF220911 Capensibufo AF220865 842 tradouwi* AF220912 Cardioglossa RABI 141 Gabon, Rabi (Shell DQ283176 DQ284213 DQ283868 DQ282726 DQ283565 3699 gratiosa (SAM) Gabon), Toucan Well Head, funnel trap line 1.8, 01.4750ЊS, 09.5335ЊE Cardioglossa UTA Cameroon, Southwest DQ283080 DQ284348 DQ283802 DQ282978 DQ283681 4266 leucomystax A44591; Prov, vicinity DQ283079 DQ284132 DQ283982 DQ282913 UTA Ediensoa A44585 Caudiverbera AMNH No data (pet trade) DQ283439 DQ284415 DQ284036 DQ282893 DQ283748 4222 caudiverbera A168414 Centrolene X86230 1146 geckoideum* X86264 X86298 Centrolene SIUC 7053 Panama, Cocle Prov, AY364358 AY364404 AY844776 AY844206 3871 prosoblepon El CopeÂ, Parque AY364379 Nacional ``Omar AY843574 Torrijos'' Ceratobatrachus AMS Solomon Islands, DQ283197 DQ284230 DQ283881 DQ282741 DQ283024 DQ283579 4675 guentheri R137134; Malaita, Su'u Bay; DQ283198 DQ284409 DQ284031 DQ282886 DQ283740 AMNH Solomon Islands A161634 Ceratophrys JF 929 Argentina, Santa Fe, AY843575 AY843797 AY844207 3469 cranwelli Vera, ``Las Gamas'' Chacophrys AMNH No data (pet trade) DQ283328 2421 pierottii A168435 Chalcorana FMNH Brunei, Belait Dist, DQ283139 DQ284179 DQ283840 DQ282695 DQ282933 DQ283531 4687 chalconota 248327 Labi, Sg Mendaram Chaperina fusca FMNH Malaysia, Sabah, DQ283145 DQ284185 DQ282701 DQ282938 2307 231111 Lahad Datu Dist, Danum Valley Research Center Charadrahyla UTA Mexico, Oaxaca, AY843649 DQ284100 AY844635 AY844853 AY844094 AY844272 4730 nephila A54772 Colonia Rodulfo Figueroa, El Carrizal, 1475 m Chelydra AMCC USA, New York, DQ283320 DQ282810 DQ283651 3108 serpentina 101071 Cornwall Co, , Arthurs Pond 300 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Chirixalus doriae FMNH Laos, Huaphahn Prov, DQ283135 DQ284176 DQ283836 DQ282691 DQ283528 4139 255213 Vieng Tong Dist, Phou Louey National Biodiversity Conservation Area, near Nam Puong River, 985 m, 20Њ14ЈN, 103Њ16ЈE Chirixalus vittatus FMNH Vietnam, Gia-Lai DQ283134 DQ284175 DQ283835 DQ282690 DQ283527 4140 254444 Prov, Ankhe Dist, Kannack town, Buon Luoi village, 20 km NW Kannack, Annam mts, 700±750 m, 14Њ20ЈN, 106Њ36ЈE Chiromantis AMNH Tanzania, Morogoro, AF215348 DQ284380 DQ284012 2656 xerampelina A153250 Udzungwa Mts AF458132 National Park, Njokamoni River drainage, 1100± 1200 m Choerophryne sp. ABTC 47720 Papua New Guinea, DQ283207 DQ284239 DQ283889 DQ282750 DQ283583 4156 Mt Menawa Clinotarsus AF249058 AF249117 AF249180 2110 curtipes* AF249021 Cochranella NK A 5292 Bolivia, Dept Santa AY843576 DQ284066 AY844372 AY844777 AY844029 AY844208 4730 bejaranoi Cruz, Prov Caballero, San JuaÂn CantoÂn, AmboroÂ, National Park, near San JuaÂn del Portrero, on the RõÂo Cerro Bravo, near 17Њ50Ј08ЉS, 64Њ23Ј23ЉW, 1800± 2100 m Colostethus AMNH Venezuela, Amazonas, DQ283044 DQ284073 DQ283773 DQ282656 DQ283464 4223 undulatus A159139 Cerro YutajeÂ, 1700 m, 5Њ46ЈN, 66Њ8ЈW Conraua goliath FMNH Cameroon, Ebowala DQ283132 DQ284173 DQ283833 DQ282688 DQ283525 4158 262216 area Conraua robusta UTA Cameroon, Southwest DQ283347 DQ284337 DQ283973 DQ282823 DQ282972 DQ283671 4688 A44401 Prov, plateau NW of Ntale village, ca. 700 m Cophixalus ABTC 47881 Papua New Guinea, DQ283206 DQ284238 DQ282749 DQ283582 4032 sphagnicola Wau Copiula sp. AMS Papua New Guinea, DQ283208 DQ284240 DQ282751 DQ283584 3415 R124417 Sinyarge Crinia nimbus ABTC 25300 Australia, Tasmania, DQ283299 DQ283949 DQ282801 DQ283639 3351 Haast Mts Crinia signifera SAMA Australia, New South DQ283192 DQ284226 DQ282737 2695 R40274 Wales, Watagan S.F. DQ283193 Crossodactylus MLPA 1414 Argentina, Misiones, AY843579 DQ284050 AY844552 AY844780 AY844031 3989 schmidti Aristobulo del Valle, Balneario CunÄapiru Crotaphatrema UTA Cameroon, Adamoua, DQ283353 DQ284342 DQ283676 2430 tchabalmbaboensis A51667 N face of Mt Tchabal DQ283354 Mbabo, 1950 m Cruziohyla KRL 800 Panama, Cocle Prov, AY843562 AY844536 DQ282950 AY844196 4061 calcarifer El CopeÂ, Parque Nacional ``Omar Torrijos'' Cryptobatrachus AY326050 2329 sp.* Cryptobranchus TAT USA, Arkansas, North DQ283263 DQ284286 DQ283621 2998 alleganiensis Fork White River 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 301

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Cryptothylax CAR 381 Central African DQ283170 DQ284208 DQ283863 DQ282722 DQ283560 4184 gresshof® (SAM) Republic, Pref Sangha-MbaeÂreÂ, Park National de Dzanga- Ndoki, 38.6 km 173ЊS Lidjombo, Camp 3, 02.2136ЊS, 16.0312ЊE Cryptotriton AF199196 507 alvarezdeltoroi* Ctenophryne AMNH Guyana, Berbice DQ283383 DQ28436 DQ282846 DQ282993 DQ283698 4392 geayei A166444 River camp at ca. 18 mi (linear) SW Kwakwani (ca. 2 mi downriver from Kurundi River con¯uence), 200 ft, 5Њ5Ј6ЉN, 58Њ14Ј14ЉW Cycloramphus CFBH 5757 Brazil, SaÄo Paulo, DQ283097 DQ284147 DQ283813 DQ282675 DQ282924 DQ283498 4740 boraceiensis Picinguaba, Ubatuba Cyclorana SAMA Australia, no other DQ284124 DQ284124 AY844553 3062 australis R16906 data Dasypops schirchi CFBH-T 71 Brazil, EspõÂrito Santo, DQ283095 DQ284145 DQ283811 DQ282922 DQ283497 4302 Linhares, Reserva da Vale Dendrobates USNM Panama, Bocas del AY843581 DQ284072 AY844554 AY844781 AY844032 AY844211 4758 auratus 313818 Toro Dendrophryniscus MJH 7095 Peru, HuaÂnuco, RõÂo AY843582 DQ284096 AY844555 3056 minutus Llullapichis, Panguana Dendropsophus MJH 7116 Peru, HuaÂnuco, RõÂo AY843640 DQ284085 DQ283782 3071 marmoratus Llullapichis, Panguana Dendropsophus MACN Argentina, Misiones, AY549345 DQ284096 DQ283758 AY844089 DQ283456 4309 minutus 33799 GuaranõÂ, San Vicente, Campo Anexo INTA ``Cuartel RõÂo Victoria'' Dendropsophus MACN Argentina, Entre RõÂos, AY549346 DQ284051 AY844634 AY844852 AY844271 4200 nanus 37785 Dept Islas del Ibicuy Dendropsophus AMNH Brazil, Acre, Centro AY843652 AY844638 AY844856 AY844097 AY844274 4410 parviceps A139315 Experimental da Universidade do Acre at km 23 on Rio Branco±Porto Velho Rd Dendrotriton AF199232 516 rabbi* Dermatonotus AMNH No data (pet trade) DQ283329 2069 muelleri A168436 DQ283330 Dermophis UTA 56550 Mexico, , DQ283455 DQ284428 DQ282897 2710 oaxacae Tierra Colorada± Ayutla Hwy, 424 m Desmognathus UMMZ USA, North Carolina, DQ283253 DQ284278 DQ283923 DQ282775 DQ283614 3628 quadramaculatus 221202 Macon Co, Blue Ridge Parkway Desmognathus AY728225 2320 wrighti* Dicamptodon RAN 31288 USA, Idaho, Beneweh DQ283118 DQ284164 DQ283516 3394 aterrimus Co, Mannering Creek Dicamptodon TAT 1043 USA, Oregon, Lane DQ283261 DQ284284 2705 tenebrosus Co, Thompson Creek Didelphis AMNH Peru, Loreto, RõÂo DQ283321 DQ282811 1897 marsupialis A272836 Galvez, Nuevo San JuaÂn Didynamipus AY325991 2289 sjostedti* 302 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Dimorphognathus SAM 51540 Gabon, Loango DQ283175 DQ284212 DQ283867 DQ282725 DQ282943 DQ283564 4240 africanus National Park, near pitfall trap line #3, 02Њ20Ј27ЉS, 09Њ35Ј50ЉE Discodeles guppyi AMS Solomon Islands, DQ283200 DQ284232 DQ283883 DQ282743 DQ282947 3521 R137175 Malaita, Su'u Bay Discoglossus ZSM 725/ Portugal, Alamada DQ283243 DQ284270 DQ283915 DQ282770 DQ283609 3774 galganoi 2000 City Discoglossus AH Spain, Barcelona DQ283435 DQ284412 DQ284034 DQ282889 DQ283744 3796 pictus Duellmanohyla MVZ Costa Rica, AY549315 DQ284059 AY844556 AY844033 AY844212 4295 ru®oculis 207193 Guanacaste Prov, VolcaÂn Cacao RdS No data (pet trade) DQ283434 DQ284411 DQ282888 DQ283025 DQ283743 4419 Eburana AMNH Vietnam, Ha Giang DQ283394 DQ284008 DQ282854 DQ282999 DQ283707 4359 chloronota A163935 Prov, Vi Xuyen, Cao Bo Commune, Mt Tay Conn Linh II, Bac Trao River, near camp, just upstream, 600 m, 22Њ45Ј39ЉN, 104Њ52Ј23ЉE Ecnomiohyla SIUC 6998 Panama, Cocle Prov, AY843776 DQ284115 AY844629 AY844847 AY844088 AY844268 3846 miliaria El CopeÂ, Parque AY843777 Nacional ``Omar Torrijos'' Edalorhina perezi MJH 7082 Peru, HuaÂnuco, RõÂo AY843585 DQ284095 AY844558 AY844764 DQ283474 4200 Llullapichis, Panguana Elachistocleis AMNH Guyana, Dubulay DQ283405 DQ284386 2728 ovalis A141136 Ranch on Berbice River, 200 ft, 5Њ40Ј55ЉN, 57Њ51Ј32ЉW Eleutherodactylus JAC 21987 Mexico, Veracruz, DQ283318 DQ284318 DQ283649 3483 alfredi Municipio CoÂrdoba, Cruz de los Naranjos, 1100 m Eleutherodactylus CG (now Mexico, Sonora, DQ283271 DQ284291 DQ283935 DQ282786 DQ282963 DQ283627 4738 augusti UAZ Alamos unnumbered) Eleutherodactylus CFBH 5813 Brazil, SaÄo Paulo, DQ283092 DQ284142 DQ283807 DQ282918 DQ283493 4288 binotatus Parque Estadual da Serra do Mar, NuÂcleo Santa VirgõÂnia, SaÄo Luiz do Paraitinga Eleutherodactylus SIUC 7062 Panama, Cocle Prov, DQ283165 DQ284203 DQ282718 DQ282942 DQ283555 4404 bufoniformis Parque Nacional El Cope Eleutherodactylus CFBH 4450 Brazil, Minas Gerais, DQ283093 DQ284143 DQ283809 DQ282920 DQ283495 4271 juipoca PocËos de Caldas Eleutherodactylus USNM USA, Texas, Travis DQ283101 DQ284151 DQ283817 DQ282677 DQ283502 3324 marnockii 331345 Co, Austin, Univ of DQ283102 Texas Campus, near football stadium Eleutherodactylus UTA Mexico, Oaxaca, DQ283316 DQ284316 DQ283959 DQ282807 DQ283647 2855 nitidus A54771 Municipio CuicatlaÂn, Tutepetongo, 1619 m Eleutherodactylus USNM USA, Florida, Collier, DQ283107 DQ284155 DQ283821 DQ282680 DQ282929 DQ283506 4740 planirostris 547959; Naples, Parkshore DQ283108 DQ284294 DQ283937 DQ282788 DQ282964 DQ283629 P. Moler, subdivision, Parkview unnumbered Way, 26Њ11Ј24ЉN, 81Њ48Ј16ЉW; USA, Florida, Duval Co, 2826 Rosselle St 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 303

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Eleutherodactylus AMNH Bolivia, Santa Cruz, AY843586 DQ284372 AY844559 AY844785 AY844035 AY844213 4790 pluvicanorus A165195 Caballero, San JuaÂn Canton, Amboro National Park, 2050 m, 17Њ50Ј17ЉS, 64Њ23Ј30ЉW Eleutherodactylus SIUC 7066 Panama, CocleÂ, DQ283168 DQ284206 DQ283862 DQ283558 3804 punctariolus Parque Nacional El Cope Eleutherodactylus USNM-FS Panama, Bocas del DQ283105 DQ284154 DQ283820 DQ282928 DQ283505 4004 ranoides 195393 Toro, Isla Escudo de DQ283106 Veraguas, West Point Eleutherodactylus JAC 22721 Mexico, Oaxaca, El DQ283317 DQ284317 DQ283960 DQ282808 DQ282968 DQ283648 4702 rhodopis Mirador, Municipio Santa MarõÂa Chilchotla Ensatina AY728216 2311 eschscholtzii* Epicrionops sp. UMMZ Ecuador, Cotopaxi, DQ283130 DQ284171 DQ283523 3074 185825 San Francisco de Las Pampas Epipedobates UMMZ Pet trade, imported DQ283037 DQ284063 DQ283768 DQ282653 DQ282902 DQ283461 4761 boulengeri 227952 from Ecuador Euphlyctis AF249053 AF249111 AF249174 2069 cyanophlyctis* AF249015 Euproctus asper* U04694 840 U04695 Eupsophus MACN Argentina, NeuqueÂn, AY843587 DQ284120 AY844560 AY844786 AY844036 AY844214 4749 calcaratus 37980 Huiliches, Termas de Epulafquen Eurycea wilderae UMMZ USA, North Carolina, DQ283254 DQ283615 3040 221205 Macon Co, Deep Gap Exerodonta JAC 21736 Mexico, Chiapas, AY843619 DQ284099 AY844596 AY844815 AY844062 AY844240 4738 chimalapa Colonia Rodulfo Figueroa, El Carrizal, 1475 m Fejervarya AB070731 2391 cancrivorus* AF206473 AF206092 AF206137 Fejervarya AY014380 502 kirtisinghei* Fejervarya AMNH Vietnam, Nghe An AY843588 DQ284356 AY844561 AY844787 AY844037 3991 limnocharis A161230 Prov, Con Cuong Dist, Bong Khe Commune, 19Њ2Ј24ЉN, 104Њ54Ј24ЉE Fejervarya AY141843 AF249107 AF249170 2484 syhadrensis* AF249011 Flectonotus sp. CFBH 5720 Brazil, Santa Catarina, AY843589 AY844562 AY844788 AY844038 AY844215 4004 Santo Amaro da Imperatriz Gastrophryne RdS 726 Belize, Stann Creek DQ283426 DQ284404 DQ282883 DQ283019 DQ283735 4387 elegans Dist, Cockscomb Basin Wildlife Sanctuary Gastrophryne ATH 476 USA, Arizona, Santa DQ283268 DQ284288 DQ283932 DQ282784 DQ282961 DQ283624 4703 olivacea Cruz Co, Ruby Rd, vicinity Calabasas Canyon 304 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Gastrotheca NK A5286 Bolivia, Dept Santa AY843590 DQ284069 AY844563 AY844789 AY844039 3995 cf. marsupiata Cruz, Prov Caballero, San JuaÂn CantoÂn, AmboroÂ, National Park, near San JuaÂn del Potrero, on RõÂo Cerro Bravo, near 17Њ50Ј08ЉS, 64Њ23Ј23ЉW, 1800± 2100 m Gastrotheca JLG 09 Brazil, EspõÂrito Santo, AY843592 AY844564 AY844790 3130 ®ssipes Setiba, Guarapari Gazella thomsoni WCS 851199 No data M86501 DQ282812 DQ283652 3556 Gegeneophis AF461136 972 ramaswamii* AF461137 Genyophryne ABTC 49624 Papua New Guinea, DQ283209 DQ284241 DQ283890 DQ282752 DQ283585 4171 thomsoni Bolulo Geocrinia ABTC 7145 Australia, Victoria, DQ283294 DQ284306 DQ283947 DQ282799 DQ282965 DQ283637 3858 victoriana Tanjil Bren DQ283295 DQ283296 Geotrypetes FMNH Gabon, Prov de DQ283337 DQ284328 DQ283662 2816 seraphini 256782 Woleu-Ntem, 31 km ESE Minvoul, along IOBT trail (PK 29), 600 m, 2Њ4.8ЈN, 12Њ24.4ЈE Glandirana AF315127 882 minima* AF315153 Gyrinophilus UMMZ USA, North Carolina, DQ283255 DQ284279 DQ283924 DQ282776 DQ283616 3630 porphyriticus 221207 Macon Co, Deep Gap Hamptophryne RdS Peru (no other data) DQ283438 DQ284414 DQ282892 DQ283747 3891 boliviana Heleioporus ABTC 76692 Australia, New South DQ283306 DQ284311 DQ283953 DQ282804 DQ283642 3694 australiacus Wales, Mona Vale DQ283307 Heleophryne TMSA South Africa, Western AY843593 DQ284113 AY844565 AY844791 3458 purcelli 84157 Cape Prov, Cedarberg Range, head of Krom River Heleophryne regis AC 2544 South Africa, Western DQ283115 DQ284161 DQ283828 DQ282684 DQ283513 3758 Cape Prov, Montague Pass Hemidactylium UMFS USA, Michigan, Saint DQ283120 1587 scutatum 11564 Clair Co, Woodlot DQ283121 just S Marysville along Hwy 29/Busha Hwy, about 1 km NW jct Davis Rd, 42Њ53.3ЈN, 82Њ29.3ЈW Hemiphractus MJH 3689 Peru, Ucayali, 3 km AY843594 DQ284084 AY844566 AY844792 3431 helioi S, km 65 on Hwy Federico Basadre at Ivita Hemisus RdS 916 Tanzania, Arusha, AY326070 DQ284407 DQ284029 DQ282885 DQ283022 DQ283738 4619 marmoratus Masai Camp Herpele UTA Cameroon, DQ283359 DQ284346 DQ283980 DQ283679 3746 squalostoma A52349 Southwestern Prov, Nguti Heterixalus sp. UMMZ Madagascar, DQ283448 DQ284422 DQ283027 DQ283752 4004 219330 Mahajanga, Antsalova, Bemaraha Reserve Antranopasasy, 44.71635ЊN, 18.708016ЊE Heterixalus AY341630 AY341759 2392 tricolor* AY341697 AY341725 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 305

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Hoplobatrachus KU 290425 Ghana, Muni Lagoon, DQ283059 DQ284090 DQ283787 DQ282907 DQ283471 4303 occipitalis Winneba, 5Њ21Ј14ЉN, 0Њ42Ј12ЉW Hoplobatrachus FMNH Laos, Champasak DQ283141 DQ284181 DQ283842 DQ282697 DQ282934 DQ283533 4696 rugulosus 255191 Prov, Mounlapamok Dist, Dong Khanthung National Biodiversity Conservation Area, near Houay Khiem stream, 60 m, 14Њ08ЈN, 105Њ22ЈE Hoplophryne RdS 949 Tanzania, East DQ283419 DQ284398 DQ282876 DQ283015 DQ283730 3946 rogersi Usambara Mts, adjacent to Amanai Nature Reserve, 05Њ07Ј38.0ЉS, 38Њ37Ј22.6ЉE Huia nasica AMNH Vietnam, Ha Tinh DQ283345 DQ284333 DQ283971 DQ282821 DQ282970 DQ283667 4694 A161169 Prov, Huang Son Reserve, Rao An region, top of Pomu Mt, 900±1200 m, 18Њ20Ј53ЉN, 105Њ14Ј38ЉE Hyalinobatrachium JAC 21365 Mexico, Oaxaca, San DQ283453 DQ284 DQ284043 DQ283756 3800 ¯eischmanni Jose Pacõ®co± Candelaria Loxicha Hwy, 480 m Hydromantes CAS 206495 USA, California, Inyo DQ283227 DQ28425 2248 platycephalus Co, Elderberry Canyon, 37.37749ЊN, 118.63851ЊW Hyla arborea* ZFMK Germany (live AY843601 AY843822 AY844046 3270 specimen) Hyla cinerea MVZ USA, Texas, Travis AY549327 DQ284057 AY844597 AY844816 AY844063 AY844241 4736 145385 Co, Austin, municipal golf course Hylarana FMNH Cambodia, Siem Reap DQ283138 DQ283839 DQ282694 3121 erythraea 257285 Prov, Siem Reap Dist, Siem Reap town, Ͻ10 m, 13Њ22Ј29ЉN, 103Њ50Ј44ЉE Hylarana AMNH Vietnam, Ha Giang DQ283396 DQ284010 DQ282856 DQ283000 DQ283710 4358 taipehensis A163972 Prov, Yen Minh, Du Gia Commune, Khau Ria Village, rice paddy on edge of limestone forest S village, 934 m, 22Њ53Ј49ЉN, 105Њ14Ј48ЉE Hylodes phyllodes CFBH-T 249 Brazil, SaÄo Paulo, DQ283096 DQ284146 DQ283812 DQ282674 DQ282923 3989 Picinguaba, Ubatuba Hylorina sylvatica* AY389153 718 Hyloscirtus AMNH Bolivia, Dept Santa AY549321 DQ284070 AY844579 AY844050 AY844224 4367 armatus A165163 Cruz, Caballero, San JuaÂn Canton, Amboro National Park, near base camp on RõÂo Cerro Bravo, 17Њ50Ј17ЉS, 64Њ23Ј30ЉW Hyloscirtus SIUC 6924 Panama, Cocle Prov, AY843650 DQ284088 AY844636 AY844095 AY844273 4354 palmeri El CopeÂ, Parque Nacional ``Omar Torrijos'' Hymenochirus AY341634 AY341763 2339 boettgeri* AY341700 AY341726 306 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Hyperolius alticola CAS 202047 Uganda, Rukungiri DQ283225 DQ284255 DQ283902 DQ282762 DQ282952 DQ283595 3781 Dist, Bwindi Impenetrable National Park, Munyaga River, ca. 100 m downstream from Munyaga Falls Hyperolius AMNH Tanzania, Morogoro, DQ283389 DQ284375 DQ282851 DQ282997 DQ283704 3963 punticulatus A153299 Udzungwa Mts National Park, Njokamoni River drainage, 1100± 1200 m Hyperolius AMNH Tanzania, Morogoro, DQ283399 DQ284381 DQ282858 DQ283002 DQ283712 3492 tuberilinguis A153257 Udzungwa Mts DQ283400 National Park, Man'gula Camp Site 3 on Mwaya River, 350 m, 7Њ50Ј51ЉS, 36Њ53Ј0ЉE Hypogeophis UMMZ Seychelles, Silhouette DQ283131 DQ284172 DQ282687 DQ283524 3890 rostratus 181332 Island, Trail from La Passe to Jardin Marron Hypsiboas USNM Brazil, Pernambuco, AY549316 AY549369 AY844794 AY844218 3901 albomarginatus 284519 near CaraurucËu, on way to Serra dos Cavalos Hypsiboas boans RWM 17746 Venezuela, Amazonas, AY843610 DQ284086 AY844588 AY844809 AY844055 AY844231 4746 CanÄo Agua Blanca, 3.5 km SE Neblina base camp on RõÂo Baria Hypsiboas MAD 085 Guyana, Iwokrama, AY549336 DQ284076 AY844610 AY844828 DQ283466 4218 cinerascens Muri Scrub camp, 80 m Hypsiboas AMNH Guyana, Demerara, AY843648 AY844633 AY844851 AY844093 AY844270 4437 multifasciatus A141040 Ceiba Station, Madewini River, ca. 3 mi (linear) E Timehri airport Ichthyophis MW 375 India DQ283086 DQ284137 DQ282669 DQ283487 3847 cf. peninsularis (Univ of Kerala) Ichthyophis sp. FMNH Laos, Khammouan DQ283336 DQ284327 DQ283661 2988 256425 Nakai Dist, Nakai Nam Theun National Biodiversity Conservation Area, Annamite Mts, disturbed evergreen forest along Houay Ting Tou stream, 700 m, 17Њ58ЈN, 105Њ34ЈE Iguana iguana WCS No data NC002793 DQ284249 DQ283590 3416 Indirana sp.* AF249051 AF249122 AF249185 1399 Indirana sp.* AF249064 AF249123 AF249186 1406 Ingerana baluensis FMNH Malaysia, Sabah, DQ283142 DQ284182 DQ283843 DQ282698 DQ282935 DQ283534 4636 231085 Lahad Datu Dist, Danum Valley Research Center Ischnocnema MJH 7057 Peru, HuaÂnuco, DQ283061 DQ284091 DQ283788 DQ282661 2950 quixensis Panguana, RõÂo DQ283060 Llullapichis Isthmohyla MVZ Costa Rica, Heredia AY843659 DQ284058 AY844649 AY844117 3598 rivularis 149750 Prov, ``Chompipe'', vicinity VolcaÂn Barba Ixalotriton niger* AF451248 518 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 307

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Kalophrynus FMNH Malaysia, Sabah, DQ283146 DQ284186 DQ283846 DQ282702 DQ283537 4168 pleurostigma 230844 Lahad Datu Dist, Danum Valley Research Center Kaloula pulchra AMCC Vietnam, Ha Tinh DQ283397 DQ284379 DQ284011 DQ282857 DQ283001 DQ283711 4745 106697; RdS Prov, Huong Son DQ283398 DQ282874 DQ283012 DQ283727 02 Dist, Huong Son Reserve, Ngai Doi region, headquarters of Huong Son Reserve, 100 m, 18Њ27Ј08ЉN, 105Њ17Ј43ЉE; No data (pet trade) Kassina RdS 803 Tanzania, Iringa, DQ283437 DQ284413 DQ282891 DQ283026 DQ283746 4401 senegalensis Kibebe Farm, 07Њ48Ј12.4ЉS, 35Њ45Ј24.2ЉE Kurixalus eif®ngeri UMFS 5969 China, Taiwan, Nan- DQ283122 DQ284166 DQ283830 DQ282931 DQ283518 4272 Tou, Lu-Gu Chi-Tou, 900±1100 m Kurixalus UMFS 5702 China, Taiwan, Nan- DQ283054 DQ284087 DQ283783 DQ282905 DQ283468 4287 idiootocus Tou, Tung Fu, 750 m Kyarranus ABTC 25186 Australia, New South DQ283313 DQ283957 DQ283646 3442 sphagnicolus Wales, Dorrigo Mountain Laliostoma UMMZ Madagascar, Toliara, DQ283057 DQ284089 DQ283786 AF249169 DQ283470 4295 labrosum 213554 Sakaraha, Zombitsy Forest, 44.711666ЊS, 22.843333ЊE Lankanectes AF215393 AF249115 AF249178 2102 corrugatus* AF249019 AF249043 Latimeria AMNH Comoros, Grande NC࿞001804 DQ284319 AF131253 DQ283653 3865 chalumnae A59196 Comore Lechriodus ABTC 24921 Australia, New South DQ283282 DQ284299 DQ283942 DQ282793 DQ283632 3753 ¯etcheri Wales, Border Ranges DQ283283 National Park Leiopelma archeyi DMG 5123 New Zealand, North DQ283216 DQ284246 DQ283895 DQ283588 3395 Island, Coromandel DQ283215 Peninsula, Tapu Summit Leiopelma DMG 5135 New Zealand, DQ283217 DQ284247 DQ282755 DQ283589 3894 hochstetteri Waitakere Mts, Cowan Stream Lepidobatrachus AMNH No data (pet trade) DQ283152 DQ284191 DQ283851 DQ282707 DQ283543 4192 laevis A168407 Leptobrachium AMNH Vietnam, Ha Giang DQ283052 DQ284081 DQ283467 3053 chapaense A163791 Prov, Vi Xuyen, Cao Bo Commune, Mount Tay Conn Linh II, above Tham Ve Village, spring camp, 1420 m, 22Њ45Ј59ЉN, 104Њ49Ј56ЉE Leptobrachium CAS 222293 Myanmar, Rakhine DQ283239 DQ284265 DQ283911 DQ282767 DQ283605 4176 hasselti State, Gwa Township, Rakhine Yoma Elephant Sanctuary, Kyat stream camp, 17Њ42Ј14.0ЉN, 94Њ38Ј54.3ЉE Leptodactylodon UTA Cameroon, Southwest DQ283364 DQ284351 DQ283986 DQ282980 3587 bicolor A44492 Prov, plateau NW Ntale Village 308 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Leptodactylus AMNH Guyana, Southern DQ283404 DQ284385 DQ284015 DQ282862 AY341760 DQ283716 4744 fuscus A139088 Rupununi savanna, Aishalton (on Kubabawau Creek), 150 m, 2Њ28Ј31ЉN, 59Њ19Ј16ЉW Leptodactylus MACN Argentina, Buenos AY843688 DQ284104 AY844681 AY844302 3809 ocellatus 38648 Aires, Escobar, Loma Verde, Establecimiento ``Los Cipreses'' Leptolalax AMNH Vietnam, Ha Giang DQ283381 DQ284367 DQ284000 DQ282844 DQ283696 3246 bourretti A163810 Prov, Vi Xuyen, Cao Bo Commune, Mount Tay Conn Linh II, above Tham Ve village, stream 2 km SW base camp, 1420 m, 22Њ46Ј8ЉN, 104Њ49Ј51ЉE Leptopelis CAS 169938 Kenya, Kili® Dist, DQ283226 DQ284256 DQ283903 DQ283596 3760 argenteus 14.4 km W Kakayuni, towards Lake Jilore, on Kakayuni Rd roadside pond Leptopelis bocagei RdS 802 Tanzania, Iringa, DQ283418 DQ283014 DQ283729 3639 Kibebe Farm, 07Њ48Ј12.4ЉS, 35Њ45Ј24.2ЉE Leptopelis sp. AMNH No data (pet trade) DQ283161 DQ284199 DQ283856 DQ283552 3766 A168408 Leptopelis CAS 168661 Tanzania, Tanga DQ283242 DQ284268 DQ283608 3450 vermiculatus Region, Muheza Dist, East Usambara Mts, Amani-Muheza Rd 3± 5 km SE Amani Limnodynastes NTM Australia, Northern DQ283308 DQ284312 DQ283954 DQ282805 DQ283643 4174 depressus R26241 Territory, Elizabeth Downs, Daly River region Limnodynastes SAMA Australia, New South DQ283285 DQ284301 DQ283944 DQ282794 DQ283633 3743 dumerilli R34734 Wales, Langothlin DQ283286 Limnodynastes QMJ 57109 Australia, Queensland, DQ283280 DQ284298 DQ283941 DQ282792 DQ283631 3756 ornatus Heathlands DQ283281 Limnodynastes AH No data DQ283245 DQ284272 DQ283917 DQ282772 2962 peronii DQ283246 Limnodynastes AY326071 2408 salmini* Limnomedusa MACN Argentina, Misiones, AY843689 DQ284127 AY844682 AY844128 3594 macroglossa 38641 Aristobulo del Valle, Balneario CunÄapiru Limnonectes AY313724 2399 acanthi* Limnonectes ABTC 47812 Papua New Guinea, DQ283202 DQ284234 DQ283885 DQ282745 3443 grunniens Utai Limnonectes AY313749 2402 heinrichi* Limnonectes kuhlii AMNH Vietnam, Quang Binh AY313686 DQ284234 DQ283885 DQ282745 DQ282982 DQ283688 4686 A161202 Prov, Minh Hoa, Cha Lo Limnonectes AMNH Vietnam, Quang Nam DQ283378 DQ284364 DQ283997 DQ282841 DQ282989 3981 poilani A163717 Prov, TraÄMy,TraÄ TaÃp Commune, stream near Thon 2 Village, 920±1060 m, 15Њ9Ј37ЉN, 108Њ2Ј26ЉE Limnonectes AY313719 2358 visayanus* 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 309

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Lineatriton AF380808 516 lineolus* Liophryne ABTC 49566 Papua New Guinea, DQ283199 DQ284231 DQ283882 DQ282742 DQ283580 4171 rhododactyla Bulolo Lithobates AMNH Guyana, Magdalen's DQ283384 DQ284369 DQ284001 DQ282847 DQ282994 DQ283699 4686 palmipes A166454 Creek camp, Ϯ300 yd NW bank of Konawaruk River ca. 25 mi (linear) WSW Mabura Hill, 400 ft, 5Њ13Ј7ЉN, 59Њ2Ј43ЉW Lithodytes lineatus AMNH Guyana, Berebice AY843690 DQ284112 AY844683 AY844129 AY844303 4345 A166426 River camp at ca. 18 mi (linear) SW Kwakwani (ca. 2 mi downriver from Kurundi River con¯uence), 200 ft, 5Њ5Ј6ЉN, 58Њ14Ј14ЉW Litoria aurea AMS 52744 New Caledonia, Prov AY843691 DQ284098 AY844684 AY844130 AY844892 3989 Nord, Valle Phaaye, Nomac River, 8 km E Poum Litoria freycineti SAMA Australia, New South AY843693 DQ284122 AY844686 AY844894 3460 R12260 Wales, 16 km E Retreat Litoria SAMA Australia, Northern DQ283222 DQ284252 DQ283899 DQ282759 DQ283592 4144 genimaculata R41068 Territory, Mt Lewis Litoria inermis SAMA Australia, Western DQ283211 DQ284243 DQ283892 2718 R53945 Australia, 24 km N DQ283212 Tunnel Creek Gorge Litoria lesueurii SAMA Australia, New South DQ283204 DQ284236 DQ283887 DQ282747 3456 R35012 Wales, Murrumbidgee River Litoria meiriana SAMA Australia, Western AY843695 DQ284125 AY844688 AY844895 AY844132 3988 R17215 Australia, Black Rock, near Kununurra Litoria nannotis SAMA Australia, Queensland, DQ283218 DQ284248 DQ283896 DQ282756 3459 R40266 Paluma Lysapsus laevis AMCC Guyana, Southern AY843696 DQ284110 AY844689 AY844896 AY844133 AY844305 4746 10720 Rupununi Savannah, Aishalton (on Kubanawan Creek) Mannophryne MVZ Trinidad and Tobago, DQ283071 DQ284108 DQ283796 3052 trinitatis 199838 Nariva Parish, Tamana Cave, Charuma Ward Mantella UMMZ Madagascar, DQ283035 DQ284061 DQ283766 DQ282651 DQ282901 DQ283460 4666 aurantiaca 201411 Toamasina, Moramanga, Andasibe Region Mantella nigricans AMNH Madagascar, DQ283034 DQ284056 DQ283764 DQ283458 3730 A167477 Antsiranana, Ambanja, Antsaravy Ridge, Tsaratanana Reserve, 13Њ55Ј34ЉS, 48Њ54Ј21ЉE Mantidactylus AMNH Madagascar, AY843698 DQ284055 DQ283763 AY844898 DQ282900 3978 cf. femoralis A167581 Antsiranana, Ambanja, Ramena River Camp, Tsaratanana Reserve, 13Њ55Ј4ЉS, 48Њ53Ј16ЉE Mantidactylus UMMZ Madagascar, DQ283036 DQ284062 DQ283767 DQ282652 3445 peraccae 213278 Fianarantsoa, Ivohibe, Andringitra Volotsangana River (Camp 3), 47.016666ЊS, 22.277777ЊE 310 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Megaelosia goeldii MZUSP Brazil, Rio de Janeiro, DQ283072 DQ284109 DQ283797 DQ282911 3590 95879 TeresoÂpolis, Rio Beija Flor, 910 m, 22Њ24ЈS, 42Њ69ЈW Megistolotis SAMA Australia, Western DQ283289 DQ284303 DQ282796 DQ283634 3846 lignarius R37834 Australia, Kununurra Megophrys nasuta FMNH Malaysia, Sabah, DQ283342 DQ284331 DQ283969 DQ282818 3437 236525 Tenom Dist, Crocker Range National Park, Purulon camp, Sungai Kilampun Melanophryniscus MACN Argentina, Chaco, AY843699 DQ284060 DQ283765 AY844899 AY844306 4207 klappenbachi 38531 vicinity Resistencia Meristogenys FMNH Malaysia, Sabah, DQ283147 DQ284187 DQ283847 DQ282703 DQ283538 4176 orphnocnemis 230531 Lahad Datu Dist, Danum Valley Research Center, Sungai Palum Tambun Metacrinia WAM Australia, Western DQ283292 DQ284305 DQ283946 DQ282798 DQ283636 3077 nichollsi R106065 Australia, 9.5 km DQ283293 ENE Mt Frankland Micrixalus borealis CAS 205064 Myanmar, Rakhine DQ283235 DQ283909 DQ282766 DQ283603 2517 State, Gwa Township, DQ283236 ca. 0.5 mi S Pleasant Beach Resort, 17Њ43Ј3.7ЉN, 94Њ31Ј55.6ЉE Micrixalus fuscus* AF249024 AF249120 AF249183 2105 AF249056 Micrixalus AF249025 AF249121 AF249184 2103 kottigeharensis* AF249041 Microhyla AMNH Vietnam, Ha Giang DQ283382 DQ282845 DQ282992 DQ283697 4052 heymonsi A163850 Prov, Yen Minh, Du Gia Commune, Khau Ria Village, stream 1, below cascade, 22Њ54Ј27ЉN, 105Њ13Ј52ЉE Microhyla sp. RdS 05 No data (pet trade) DQ283422 DQ284400 DQ284025 DQ282879 DQ283017 DQ283732 4678 Micryletta AF285207 502 inornata* Minyobates USNM-FS Panama, Bocas del DQ283042 DQ284071 DQ283772 DQ282654 DQ283462 4224 claudiae 59980 Toro, S end of Isla Popa, 1 km E Sumwood Channel Mixophyes ABTC 25115 Australia, Queensland, DQ283314 DQ284315 DQ283958 2148 carbinensis Mt Lewis area DQ283315 Myobatachus WAM Australia, Western DQ283309 DQ284313 DQ283955 DQ283644 3337 gouldii R116075 Australia, Spalding DQ283310 Park Geraldton Nannophrys AF249016 AF249112 AF249175 2112 ceylonensis* AF249047 Nanorana pleskei* AF206111 1979 AF206156 AF206492 Nasikabatrachidae AY425725 902 sp.* AY425726 Nasikabatrachus AY364360 AY364406 1532 sahyadrensis* AY364381 Natalobatrachus AF215396 784 bonebergi* AF215198 Nectophryne afra UTA Cameroon, Southwest DQ283360 DQ284347 DQ283981 DQ282831 DQ282977 DQ283680 4705 A44673 Prov, vicinity Edienosa 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 311

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Nectophryne batesi RABI 031 Gabon, Rabi (Shell DQ283169 DQ284207 DQ282721 DQ283559 3863 Gabon), near Toucan Well Head, 01.47ЊS, 09.53ЊE Nectophrynoides RdS 951 Tanzania, East DQ283413 DQ284394 DQ284018 DQ282870 DQ283723 3774 tornieri Usambara Mts, adjacent to Amanai Nature Reserve, 05Њ07Ј38.0ЉS, 38Њ37Ј22.6ЉE Necturus cf. beyeri AMCC USA, Florida, Jackson DQ283151 DQ284190 DQ283542 2952 125608 Co, Econ®na Creek, 30Њ34.17ЈN, 85Њ21.72ЈW Necturus AMCC USA, New York, DQ283412 DQ283722 2622 maculosus 105652 Suffolk Co, Cold Spring Harbor, Cold Spring Harbor Fish Hatchery Nelsonophryne AY326067 2414 aequatorialis* Neobatrachus SAMA Australia, South DQ283290 DQ284304 DQ282797 DQ283635 3430 pictus R50636 Australia, 10 km S DQ283291 Robe Neobatrachus SAMA Australia, New South AY843700 DQ284123 AY844691 AY844307 3779 sudelli R12391 Wales, 30 km N Kenmore Nesionixalus CAS 218925 SaÄo Tome and DQ284123 DQ284261 DQ283906 DQ282764 DQ282953 DQ283600 4213 thomensis Principe, SaÄo Tome I, forest at radio tower S Bom Sucesso, 00Њ16Ј64.0ЉN, 06Њ36Ј20.0ЉE Nesomantis RAN 25162 Seychelles, Mahe, DQ283452 DQ284425 DQ284042 AY341761 DQ283755 3776 thomasseti junction of Foret Noir Rd with Grand Bois River and Congo Rouge Trail Neurergus AY147246 719 crocatus* AY147247 Nidirana UMMZ China, Taiwan, Taipei, DQ283117 DQ284163 DQ283829 DQ282685 DQ282930 DQ283515 4684 adenopleura 189963 Wu-Lai, Shao-Yi, trail along Tung-Ho Creek Nidirana AMNH Vietnam, Ha Tinh DQ283365 DQ284352 DQ283987 DQ282833 DQ282981 DQ283685 3791 chapaensis A161183 Prov, Huang Son DQ283366 Reserve, Rao An region, top of Pomu Mountain, 900±1200 m, 18Њ20Ј53ЉN, 105Њ14Ј38ЉE Notaden QMJ 57130 Australia, Queensland, DQ283287 DQ284302 DQ283945 DQ282795 3026 melanoscaphus Hervey Range DQ283288 Notophthalmus AMCC No data DQ283421 DQ284399 DQ284024 DQ282878 3001 viridiscens 106084 Nototriton AF199199 514 abscondens* Nyctibates UTA Cameroon, DQ283361 DQ284349 DQ283983 DQ282979 DQ283682 3876 corrugatus A44461 Southwestern Prov, vicinity Ediensoa Nyctibatrachus AF249018 AF249114 1578 cf. aliciae* AF249063 Nyctibatrachus AF249017 AF249113 AF249176 2688 major* AF249052 AY341687 Nyctimistes SAMA Papua New Guinea, AY843701 DQ284126 AY844692 AY844134 3588 pulchra R45336 Magidobo, SHP 312 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Nyctimystes dayi SAMA Australia, Queensland, DQ283220 DQ284250 DQ283921 DQ282757 DQ283591 4145 R41010; Pilgrim Sands; DQ284276 DQ283897 ZSM 748/ Australia, Queensland, 2000 Tully River tributary, 100 m, 50 km (by rd) NW Tully Nyctixalus pictus FMNH Malaysia, Sabah, DQ283133 DQ284174 DQ283834 DQ282689 DQ283526 4150 231095 Lahad Datu Dist, Danum Valley Research Center Nyctixalus spinosus ACD 1043 Philippines, Mindanao DQ283114 DQ284160 DQ283827 DQ283512 3768 Occidozyga FMNH Malaysia, Sabah, DQ283143 DQ284183 DQ283844 DQ282699 DQ282936 DQ283535 4262 baluensis 242747 Sipitang Dist, Mendolong camp, km 6.8 Occidozyga lima CAS 213254 Myanmar, Yangon AF161027 DQ284254 DQ283901 DQ282761 DQ282951 DQ283594 4143 Division, Hlaw Ga Park, Mingalardon Township, 17Њ02Ј36.5ЉN, 96Њ06Ј41.5ЉE Occidozyga AMNH Vietnam, Ha Tinh DQ283357 DQ284344 DQ283978 DQ282829 DQ282976 3560 martensii A161171 Prov, Huang Son Reserve, Rao An region, 160 m, 18Њ29Ј54ЉN, 105Њ13Ј51ЉE Odontophrynus ZSM 733/ Argentina, Prov DQ283247 DQ284273 DQ283918 DQ282773 DQ283611 4243 achalensis 2000; BB CoÂrdoba, Pampa de DQ283248 1324 Achala; Argentina, Prov CoÂrdoba, proximity of Pampilla, near Parador El CoÂndor Odontophrynus JF 1946 Argentina, Buenos AY843704 AY844695 AY844901 AY844309 3913 americanus Aires, Escobar, Loma Verde, Ea. ``Los Cipreses'' Odorrana grahami CAS 207504 China, Yunnan, DQ283241 DQ284267 DQ283913 DQ282769 DQ283607 4163 Baoshan Pref, Qushi, ca. 25Њ17ЈN, 98Њ36ЈE Oedipina AF199230 515 uniformis* Ophryophryne AMNH Vietnam, Quang Nam DQ283377 DQ284363 DQ283996 DQ282840 DQ282988 3995 hansi A163669 Prov, TraÄMy,TraÄ Don Commune, near Camp 1, 980±1020 m, 15Њ11Ј41ЉN, 108Њ2Ј25ЉE Ophryophryne AMNH Vietnam, Ha Giang DQ283391 DQ284376 DQ284006 DQ282852 DQ283705 4190 microstoma A163859 Prov, Yen Minh, Du Gia Commune, Khau Ria Village, stream 1, above cascade, 900 m, 22Њ54Ј8ЉN, 105Њ13Ј52ЉE Opisthothylax MB 5513 Gabon, Rabi (Shell DQ283174 DQ284211 DQ283866 DQ282971 DQ283563 4274 immaculatus (SAM); DPL Gabon), at Rabi 059, 3968 trap lines 1±3, 01.5633ЊS, 09.5109ЊE; Cameroon, Southwest Prov, Kumba±Mamfe rd, within 15 km N and S of Nguti Oreophryne AMS Papua New Guinea, 8 DQ283194 DQ284227 DQ282738 DQ283577 3446 brachypus R129618 km NNE Amelei Osornophryne AY326036 2412 guacamayo* 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 313

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Osteocephalus AMNH Venezuela, Amazonas, AY843709 DQ284075 AY844700 AY844905 AY844140 AY844313 4731 taurinus A131254 Neblina base camp on RõÂo Mawarinuma (ϭRõÂo Baria), 140 m Osteopilus USNM Cuba, Guantanamo, AY843712 DQ284049 AY844906 AY844316 3886 septentrionalis 317830 Guantanamo Bay Pachytriton AMNH No data (pet trade) DQ283446 DQ284421 1749 brevipes A168416 DQ283447 Pantherana AY115111 856 berlandieri* Pantherana capito AMCC USA, Florida, DQ283187 DQ284221 DQ283874 DQ282732 DQ283572 4158 125632 Hernando Co, Croom Wildlife Management Area, Seed Orchard Pantherana ASU 33310 USA, Arizona, DQ283270 DQ284290 DQ283934 DQ282962 DQ283626 4291 chiricahuensis Greenlee Co, Coleman Creek Pantherana forreri USNM Honduras, El Paraiso, DQ283103 DQ284152 DQ283818 DQ282678 DQ283503 4156 534222 12 km NNW Ojo de Agua Pantherana pipiens UMMZ USA, Michigan, Kent DQ283123 DQ284167 DQ283831 DQ282686 DQ283519 4161 227023 Co, Grand Rapids Ada Township Park Pantherana CG USA, Arizona, Pima DQ283272 DQ284292 DQ283936 DQ282787 DQ283628 4158 yavapaiensis Co, Cienega Creek Papurana daemeli SAMA Australia, Northern DQ283201 DQ284233 DQ283884 DQ282744 DQ282948 DQ283581 4682 R40355 Territory, Cape Tribulation Paracrinia SAMA Australia, Victoria, DQ283304 DQ284310 DQ283952 DQ283641 3327 haswelli R40951 near Marlo DQ283305 Paramesotriton sp. RdS Vietnam (no other DQ283428 1950 data) Paratelmatobius CFBH-T 240 Brazil, ParanaÂ, DQ283098 DQ284148 DQ283814 DQ282676 DQ282925 DQ283499 4726 sp. Piraquara Parvimolge AF451247 512 townsendi* Pedostibes hosei FMNH Malaysia, Sabah, DQ283164 DQ284202 DQ283859 DQ282717 3463 231190 Lahad Datu Dist, Danum Valley Research Center, Sungai Palum Tambun Pelobates AY236801 AY364386 1586 cultripes* AY364341 AY364363 Pelobates fuscus AH Germany, ThuÈringen, DQ283113 DQ284159 DQ283826 DQ283511 3788 Geroda (Triptis) Pelodytes AH Spain, Barcelona DQ283111 DQ284157 DQ283824 DQ282682 DQ283509 4175 punctatus Pelomedusa RAX 2055 Ghana, Muni Lagoon, NC001947 DQ282782 DQ283622 3545 subrufa (now KU) Winneba, 5Њ21Ј14ЉN, 0Њ42Ј12ЉW Pelophryne AF375503 1093 brevipes* AF375530 Pelophylax FMNH China, Sichuan, DQ283137 DQ284178 DQ283838 DQ282693 DQ282932 DQ283530 4694 nigromaculata 232879 Hongya Xian, Bin Ling Pelophylax AB023397 918 ridibunda* AY147983 Petropedetes UTA Cameroon, Southwest DQ283075 DQ284130 DQ283800 DQ282665 DQ283481 3671 cameronensis A44399 Prov, vicinity DQ283076 Ediensoa Petropedetes RABI 033 Gabon, Rabi (Shell DQ283177 DQ284214 DQ283869 DQ282727 3045 newtoni Gabon), Toucan Well DQ283178 Head, 01.4750ЊS, 09.5335ЊE 314 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Petropedetes UTA Cameroon, South DQ283074 DQ284129 DQ283799 DQ282664 DQ283480 3747 palmipes A52407 Prov, near Bipindi, vicinity 0658198, 0342287 (UTM 32 N) Petropedetes AY341694 AY364394 AY341757 2830 parkeri* AY364348 AY364369 Phaeognathus AY728233 2363 hubrichti* Phasmahyla CFBH 5756 Brazil, SaÄo Paulo, AY843716 AY844703 AY844909 AY844145 3661 guttata Ubatuba, Picinguaba Philautus AMNH Vietnam, Ha Giang DQ283392 DQ284007 DQ282853 DQ282998 DQ283706 3945 rhododiscus A163892 Prov, Vi Xuyen, Cao DQ283393 Bo Commune, Mount Tay Conn Linh II, above Tham Ve Village, base camp, 1420 m, 22Њ46Ј8ЉN, 104Њ49Ј51ЉE Phlyctimantis DPL 4057 Cameroon, DQ283355 DQ284343 DQ282828 DQ283677 3446 leonardi Southwestern Prov, DQ283356 Kumba±Mamfe Phobobates RG No data (Atlanta DQ283073 DQ284116 DQ283798 DQ282663 DQ283479 4223 silverstonei Botanical Garden, captive bred) Phrynobatrachus UTA Cameroon, Southwest DQ283084 DQ284135 DQ282668 DQ282916 DQ283485 3901 auritus A44704 Prov, vicinity Edienosa Phrynobatrachus CAS 199268 Cameroon, East Prov, DQ283240 DQ28426 DQ283912 DQ282768 DQ283606 4157 calcaratus Dja Reserve, Boumir Camp, 3Њ11Ј26ЉN, 12Њ48Ј42ЉE, 665 m Phrynobatrachus CAS 202048 Uganda, Rukunguri DQ283228 DQ2842 DQ283904 DQ283598 3312 dendrobates Prov Dist, Bwindi DQ283229 Impenetrable National Park, Munyaga River, ca. 100 m downstream Munyaga Falls Phrynobatrachus CAS 218995 SaÄo Tome and DQ283223 DQ284253 DQ283900 DQ282760 DQ283593 3742 dispar Principe, SaÄo Tome Id., Java, 00Њ15Ј39.9ЉN, 06Њ39Ј03.2ЉE Phrynobatrachus RdS 805 Tanzania, Iringa, DQ283424 DQ284402 DQ284026 DQ282881 DQ283733 4158 mababiensis Kibebe Farm, Netting Pond, 07Њ48Ј12.4ЉS, 35Њ45Ј24.2ЉE Phrynobatrachus RdS 881 Tanzania, Tatanda DQ283414 DQ284395 DQ284019 DQ282871 DQ283009 DQ283724 4628 natalensis Village, 08Њ29Ј27.2ЉS, 31Њ30Ј18.3ЉE Phrynodon UTA Cameroon, Southwest DQ283082 DQ284339 DQ283804 DQ282825 DQ282915 DQ283673 4662 sandersoni A44599; Prov, plateau NW DQ283083 DQ284134 DQ283975 DQ282667 DQ282973 DQ283484 UTA Ntale Village. ca. A44600 567 m Phrynomantis RdS No data (pet trade) DQ283154 DQ28419 DQ282709 DQ282940 DQ283545 3945 bifasciatus Phrynopus sp. AMNH Bolivia, La Paz, AY843720 DQ284371 DQ282995 AY844323 3901 A165108 Bautista Saavedra, Canton Charazani, ca. 4 km E Chullina, 3590 m, 15Њ10Ј12ЉS, 68Њ53Ј12ЉW Phrynopus sp.* KU 202652 AY326010 3446 Phyllobates USNM-FS Panama, Bocas del DQ283043 DQ282655 DQ283463 4157 lugubris 195116 Toro, S end of Isla Popa, 1 km E Sumwood Channel Phyllodytes JLG17 Brazil, EspõÂrito Santo, AY843721 AY844708 AY844913 AY844150 3312 luteolus Setiba, Guarapari 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 315

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Phyllomedusa AMNH Guyana, Berbice AY549363 AY844716 AY844921 AY844158 AY844329 3742 vaillanti A166288 River camp at ca. 18 mi (linear) SW Kwakwani, ca. 2 mi downriver from Kurundi River con¯uence, 200 ft, 5Њ5Ј6ЉN, 58Њ14Ј14ЉW Physalaemus RdS 788 Uruguay, Flores DQ283417 DQ284022 DQ282875 DQ283728 3885 gracilis Pipa carvalhoi AH No data DQ283251 DQ284277 DQ283922 DQ282774 DQ283613 4009 Pipa pipa USNM Venezuela, Amazonas, DQ283053 DQ283781 DQ282660 3168 562560 Dept RõÂo Negro, Neblina Base Camp on the RõÂo Baria, 00Њ49Ј50ЉN, 66Њ09Ј40ЉW, 140 m Platymantis USNM Palau Islands, DQ283104 DQ2841 DQ283819 DQ282679 DQ283504 4134 pelewensis 546385 Ngerekebesang I, Echang village, rd to Japanese bunkers just N Image Restaurant and Palau Sunrise Hotel, 7Њ21ЈN, 134Њ27ЈE Platymantis weberi AMS Solomon Islands, New DQ283196 DQ284229 DQ283880 DQ282740 3436 R134894 Georgia, Patutiva Platypelis grandis AMNH Madagascar, DQ283410 DQ284392 DQ282868 DQ283007 DQ283721 4336 A167214 Antsiranana, Vohemar, Bezavona Mountain, 13Њ31Ј58ЉS, 49Њ51Ј57ЉE Plectrohyla UTA Guatemala, AY843731 AY844719 AY844924 AY844160 3663 guatemalensis A-55140 Guatemala, Don Justo, Santa Rosalia, km 12.5 on the hwy to El Salvador Plethodon dunni TAT 1040 USA, Oregon, Lane DQ283262 DQ284285 DQ283930 DQ283620 3226 Co, Richardson Creek Rd Plethodon jordani UMMZ North Carolina, DQ283125 DQ284169 DQ283521 2431 210798 Macon Co, Coweeta DQ283126 Middle Elevation Plethodontohyla AMNH Madagascar, DQ283409 DQ284390 DQ282866 DQ283006 DQ283719 3914 sp. A167315 Antsiranana, Vohemar, Camp 1, Sorata Mountain, 13Њ41Ј9ЉS, 49Њ26Ј31ЉE Pleurodeles waltl AMNH No data (pet trade) DQ283445 DQ284420 DQ283751 3442 A168418 Pleurodema AMNH Guyana, Southern AY843733 DQ28411 AY844721 AY844926 3466 brachyops A139118 Rupununi Savanna, Aishalton (on Kubabawau Creek), 150 m, 2Њ28Ј31ЉN, 59Њ19Ј16ЉW Polypedates AF249028 AF249124 AF249187 2167 cruciger* AY341685 Polypedates AMNH Vietnam, Ha Tinh, DQ283048 DQ284079 DQ283777 3040 leucomystax A161395 Huong Son, Huong Son Reserve, Rao An region, top of Pomu Mountain, 900±1200 m, 18Њ20Ј53ЉN, 105Њ14Ј38ЉE Probreviceps RdS 942 Tanzania, East DQ283420 DQ282877 DQ283016 DQ283731 3149 macrodactylus Usambara Mts, adjacent to Amanai Nature Reserve, 05Њ07Ј38.0ЉS, 38Њ37Ј22.6ЉE 316 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Proceratophrys JF 1947 Argentina, Misiones, DQ283038 DQ2840 DQ283769 DQ282903 3255 avelinoi Guarani, San Vicente, DQ283039 Campo Anexo INTA ``Cuartel RõÂo Victoria'' Pseudacris crucifer JF No data (pet trade) AY843735 DQ284114 AY844723 AY844927 AY844163 DQ283478 4695 Pseudacris AMNH USA, Florida, AY843736 AY844724 AY844164 4416 ocularis A168472 Columbia Co, SR 250 2 mi W I-10 MACN Argentina, Corrientes, AY843740 DQ284128 AY844727 AY844167 AY844337 4357 37786 Dept Bellavista, San Roque±Bellavista rd Pseudoamolops UMMZ China, Taiwan, Tai- DQ283124 DQ284168 DQ283832 DQ283520 3762 sauteri 189938 Chung, Ha-Pin trout pond near Wu-Lin Pseudobranchus AMCC USA, Georgia, Long DQ283182 DQ284217 DQ283569 2935 striatus 125629 Co, 15.5 km E DQ283183 Glennville Pseudoeurycea JAC 21252 Mexico, Oaxaca, DQ283454 DQ284427 DQ283757 2917 conanti Sierra Madre del Sur, San Jose Pacõ®co± Portillo del Rayo Hwy, 1850 m, 16Њ1.701ЈN, 96Њ31.176ЈW Pseudopaludicola MACN Argentina, Corrientes, AY843741 DQ284117 AY844728 AY844930 AY844168 3989 falcipes 38647 Yapeyu Pseudophryne SAMA Australia, New South AY843988 AY844338 3118 bibroni R73293 Wales, S Para Reservoir Reserve Pseudophryne ABTC 25573 Australia, New South DQ283311 DQ284314 DQ283956 DQ282806 DQ283645 3276 coriacea Wales, Sheepstation DQ283312 Creek, Border Ranges Pseudorana johnsi AMNH Vietnam, Nghe An DQ283214 DQ284245 DQ283894 DQ282754 DQ283587 4148 A161191 Prov, Con Cuong, Chau Khe Commune, Ngun Stream, 300 m, 19Њ2Ј17ЉN, 104Њ42Ј6ЉE Ptychadena AF261249 1650 anchietae* AF261267 Ptychadena AMNH Ethiopia, Bale Prov, DQ283066 DQ283792 DQ283475 2497 cooperi A158394 creek just E Dinsho DQ283067 Ptychadena AMNH Madagascar, DQ283031 DQ284052 DQ283760 DQ282899 3670 mascareniensis A167415 Antsiranana, Ambanja, Mandrizavona Village, Ramena Valley, 13Њ48Ј3ЉS, 48Њ44Ј47ЉE Ptychohyla UTA Mexico, Oaxaca, AY843746 AY844733 AY844934 AY844171 3660 leonhardlschultei A-54782 Sierra Madre del Sur, Pochutla Hwy, 681 m Pyxicephalus AMNH No data (pet trade) DQ283157 DQ284195 DQ283853 DQ282711 DQ282941 DQ283548 4685 edulis A168412 Quasipaa AMNH Vietnam, Quang Nam DQ283379 DQ284365 DQ283998 DQ282842 DQ282990 DQ283694 4263 verrucospinosa A163740 Prov, TraÄMy,TraÄ Don Commune, near Camp 1, 980±1020 m, 15Њ11Ј41ЉN, 108Њ2Ј25ЉE Quasipaa ZSM 759/ China, Hong Kong, DQ283244 DQ284271 DQ283916 DQ282771 DQ282957 DQ283610 4698 exilispinosa 2000 Lantau Island, Sunset Peak Ramanella AF215382 499 obscura* Rana japonica FMNH China, Sichuan, DQ283136 DQ284177 DQ283837 DQ282692 DQ283529 4153 232896 Hongya Xian, Bin Ling 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 317

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Rana sylvatica AMCC USA, New Jersey, DQ283387 DQ284373 DQ284004 DQ282849 DQ282996 DQ283702 4687 108286 Monmouth Co, 2 mi N Manalapan, 40.27910ЊN, 74.38136ЊW Rana temporaria ZSM 762/ Germany, Thuringia, DQ283127 DQ284170 DQ283914 DQ282956 DQ283522 4284 2000; UMFS Schnellbach, 725 m DQ283128 DQ284269 8005 elevation; Ireland, DQ283129 Kells, Meath Ranodon sibiricus* NC004021 2428 Rhacophorus AMNH Vietnam, Quang Binh DQ283047 DQ283776 2729 annamensis A161414 Prov, Minh Hoa, Cha Lo Rhacophorus AMNH Vietnam, Ha Tinh AY843750 DQ284078 AY844737 3756 bipunctatus A161418 Prov, Huong Son Dist, Huon Son Reserve, Rao An region, top of Pomu Mountain Rhacophorus AMNH Vietnam, Quang Nam DQ283380 DQ284366 DQ283999 DQ282843 DQ282991 DQ283695 4692 calcaneus A163749 Prov, TraÄMy,TraÄ Don Commune, near Camp 1, 980±1020 m, 15Њ11Ј41ЉN, 108Њ2Ј25ЉE Rhacophorus AMNH Vietnam, Ha Tinh DQ283049 DQ283778 2716 orlovi A161405 Prov, Huong Son Dist, Huon Son Reserve, Nga Doi region, tributary of Nga Doi River, 240± 350 m, 18Њ29Ј50ЉN, 105Њ13Ј49ЉE Rhamphophryne AF375504 1583 festae* AF375531 Rheobatrachus ABTC 7317 Australia, Queensland, DQ283275 DQ284295 DQ283938 DQ282789 3007 silus Connondale Ranges DQ283276 Rhinatrema AMNH Guyana, Magdalen's DQ283385 DQ284370 DQ284002 DQ283700 2897 bivittatum A166059 Creek camp, Ϯ300 yd NW bank of Konawaruk River ca. 25 mi (linear) WSW Mabura Hill, 400 ft, 5Њ13Ј7ЉN, 59Њ2Ј43ЉW Rhinoderma IZUA 3504 Chile, X RegioÂn, DQ283324 DQ284320 DQ283963 DQ282813 DQ283654 4202 darwinii Valdivia, Reserva Forestal de Oncol Rhinophrynus WR USA, Texas, Starr Co, DQ283109 DQ284156 DQ283822 DQ282681 DQ283507 4198 dorsalis near McAllen (living specimen) Rhyacotriton UMFS USA, Oregon, DQ283110 DQ283823 DQ283508 2924 cascadae 11729 Multnomah Co, 122.114ЊW, 45.569ЊN Salamandra AMNH No data (pet trade) DQ283440 DQ284416 DQ284037 2604 salamandra A168419 Scaphiophryne AMNH Madagascar, AY843751 DQ284391 AY364390 DQ282867 AY844175 DQ283720 4706 marmorata A-167395 Antsiranana, Vohemnar, Sorata Mtn (1320 m) Scaphiopus couchii AMNH No data (pet trade) DQ283150 DQ283850 DQ282706 DQ283541 3860 A168413 Scaphiopus AMNH USA, Florida, DQ283156 DQ284194 DQ283852 DQ282710 DQ283547 4101 holbrooki A168434 Alachua Co, Rd 346, 1.2 mi W jct Florida Hwy 121 Scarthyla KU 215427 Peru, Madre de Dios, AY843752 AY844738 AY844938 3124 goinorum Cusco AmazoÂnico, 15 km E Puerto Maldonado 318 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Schismaderma RdS 796 Tanzania, Iringa, DQ283425 DQ284403 DQ284027 DQ282882 DQ283734 4218 carens Kibebe Farm, 07Њ48Ј12.4ЉS, 35Њ45Ј24.2ЉE Schistometopum BMNH Tanzania DQ283089 DQ284140 DQ283805 DQ283490 3749 gregorii 2002.98 Schoutedenella CAS 201752 Uganda, Rukungiri DQ283237 DQ284264 DQ283910 DQ283604 2964 schubotzi Dist, Bwindi DQ283238 Impenetrable National Park, Buhoma Rd (E side), ca. 25 m S Bizenga River, 00Њ59Ј33.9ЉS, 29Њ36Ј56.6ЉE Schoutedenella UTA Cameroon, Southwest DQ283077 DQ284131 DQ283801 DQ282666 DQ282912 DQ283482 4028 sylvatica A44685 Prov, vicinity Babong DQ283078 Schoutedenella CAS 207926 , DQ283232 DQ284262 DQ283907 DQ282954 DQ283601 3892 taeniata Bioko I, vicinity Moka Malabo, along road cut of Moka Rd, 03Њ21Ј39.5ЉN, 08Њ40Ј02.7ЉE Schoutedenella RdS 864 Tanzania, Uluguru DQ283431 DQ284408 DQ284030 DQ283023 DQ283739 3911 xenodactyloides Mts, Tegetero Village, 6Њ56Ј30ЉS, 37Њ43Ј10ЉE Scinax garbei MHNSM Peru, Madre de Dios, DQ283030 DQ283759 DQ282650 DQ282898 DQ283457 4446 7311 Prov Tambopata, Cusco AmazoÂnico, ca. 15 km E Puerto Maldonado, 200 m IWK 109 Guyana, Iwokrama, AY549365 DQ284045 AY844746 AY844944 AY844181 3993 Muri Scrub camp Scolecomorphus FMNH Tanzania, Tanga DQ283338 DQ284329 DQ282816 DQ283663 3792 vittatus 251843 Region, Muheza Dist, western edge Kwangumi Forest Reserve, 4.4 km W Mt Mhinduro, 2 km S Kwamtili Estate of®ces, 230 m, 4Њ56Ј30ЉS, 38Њ44ЈE Scotobleps UTA Cameroon, SW Prov, DQ283367 DQ284353 DQ283988 DQ282834 DQ283686 3743 gabonicus A44772 vicinity Ediensoa Scythrophrys CFBH 6072 Brazil, ParanaÂ, DQ283099 DQ284149 DQ283815 DQ282926 DQ283500 4334 sawayae Piraquara Sierrana maculata USNM Honduras, Atlantida, DQ283303 DQ284309 DQ283951 DQ282803 DQ282967 3975 559483 Parque Nacional Pico Bonito, Quebrada de Oro (tributary of RõÂo Viejo), 15Њ38ЈN, 86Њ48ЈW, 945 m Silurana tropicalis UTA Cameroon, East Prov, DQ283363 DQ284350 DQ283985 DQ283684 3834 A47158 vicinity Lipondji Village Siphonops hardyi MW 1032 Brazil DQ283088 DQ284139 DQ283489 2535 (BM) Siren intermedia* Y10946 2410 Siren lacertina AMCC USA, Florida, Putnam DQ283181 DQ284216 DQ282729 DQ283568 3825 125630 Co, Rodman Reservoir at FL Hwy 310, 29Њ32.5ЈN, 81Њ50.2ЈW Smilisca phaeota RdS 786 No data (Baltimore AY843764 DQ284083 AY844751 AY844948 AY844185 AY844351 4733 Natl ; captive bred) Sooglossus UMMZ No data DQ283449 DQ284423 DQ284040 DQ282895 DQ283028 DQ283753 4337 sechellensis (#15) DQ283450 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 319

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Spea hammondii RNF 3221 USA, California, San DQ283179 DQ283870 DQ282728 DQ283566 3857 Diego Co, Del Mar Mesa, 32.94738333ЊN, 117.15688333ЊW Speleomantes AY728215 2069 italicus* Sphaenorhynchus USNM Peru, Madre de Dios, AY549367 DQ28404 AY844754 AY844188 AY844352 4344 lacteus 152136 30 km (by air) SSW Puerto Maldonado, Tambopata Reserve Sphaerotheca USNM Myanmar, Sagaing, DQ283100 DQ28415 DQ283816 DQ282927 DQ283501 3883 breviceps 524017 Kanbular Township, Chatthin, ca. 2 km WNW Chatthin Wildlife Sancturary, San Myaung Camp, 23Њ34Ј46ЉN, 95Њ44Ј26ЉE, 200 m Sphaerotheca AF249014 AF249110 AF249173 2110 pluvialis* AF249042 Sphenophryne sp. AMS Papua New Guinea, DQ283205 DQ284237 DQ282748 3134 R122221 Namosado Spicospina WAM Australia, Western DQ283301 DQ284308 DQ283950 DQ282802 DQ282966 DQ283640 4267 ¯ammocaerulea R119457 Australia, 30 km NE DQ283302 Walpole Staurois FMNH Malaysia, Sabah, DQ283140 DQ284180 DQ283841 DQ282696 DQ283532 4150 tuberlinguis 243096 Sipitang Dist, Mendolong camp, Sungai Mendolong Stefania evansi AMNH Guyana, Iwokrama, AY843767 AY844755 AY844950 AY844189 AY844353 4032 A164211 Pakatau Creek, 85 m, 4Њ45ЈN, 59Њ01ЈW Stephopaedes AF220910 511 anotis* Strongylopus grayii AMNH South Africa, Western DQ283068 DQ283793 2719 A144979 Cape Prov, Bainskloof, at settlement at crest of pass in stream Stumpf®a AMNH Madagascar, DQ283411 DQ284393 DQ282869 DQ283008 3580 cf. psologlossa A167359 Antsiranana, Vohemar, Bezavona Mountain, 13Њ31Ј58ЉS, 49Њ51Ј57ЉE Sylvirana guentheri AMNH Vietnam, Ha Tinh DQ283265 DQ284287 DQ283931 DQ282783 DQ283623 4155 A161190; Prov, Ke Go Natural DQ283266 DQ284377 DQ284009 DQ282855 DQ283708 AMNH Reserve, Rao Cai; DQ283267 A163940 Vietnam, Ha Tinh, Yen Minh, Du Gia Commune, Khau Ria Village, rice paddy on edge of limestone forest south of village, 934 m, 22Њ53Ј49ЉN, 105Њ14Ј48ЉE Sylvirana AMNH Vietnam, Vinh Phu DQ283373 DQ284359 DQ283993 DQ282838 DQ282985 DQ283691 4687 maosonensis A161487 Prov, ca. 17 km NW Tam Dao Hill Station near Buddhist temple (E Tinh Sinh) Sylvirana AMNH Vietnam, Ha Tinh DQ283371 DQ284357 DQ283991 DQ282836 DQ282983 DQ283689 4284 nigrovittata A161280 Prov, Ke Go Natural Reserve, Rao Cai Sylvirana AF249022 AF249118 AF249181 2163 temporalis* AF249054 Synapturanus MJH 3986 No data DQ283064 DQ284094 DQ282908 DQ283473 2032 mirandaribeiroi 320 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Tachycnemis UMMZ Seychelles, Praslin, DQ283451 DQ284424 DQ284041 DQ282896 DQ283029 DQ283754 4719 seychellensis 189382 near entrance Vallee de Mai Taricha sp. AMNH No data (pet trade) DQ283444 DQ284419 DQ284039 2607 A168420 Taudactylus SAMA Australia, Queensland, DQ283277 DQ284296 DQ283939 DQ282790 2987 acutirostris R41092 Mt Lewis DQ283278 Taylorana AF261251 490 limborgi* Telmatobius AMNH Bolivia, La Paz, DQ283040 DQ283770 2740 jahuira A165110 Bautista Saavedra, Charazani Canton, stream 4, 15Њ7Ј49ЉS, 68Њ53Ј17ЉW Telmatobius AMNH Bolivia, La Paz, AY843769 DQ284068 AY844757 AY844952 AY844355 4192 marmoratus A165114 Bautista Saavedra, Charazani Canton, stream, 2700±2750 m, 15Њ7Ј49ЉS, 68Њ53Ј17ЉW Telmatobius sp. AMNH Bolivia, La Paz, DQ283041 DQ284067 DQ283771 3066 A165130 Bautista Saavedra, Charazani Canton, stream 4, 15Њ7Ј49ЉS, 68Њ53Ј17ЉW Telmatobufo IZUA 3054 Chile, VII RegioÂn, DQ283325 DQ284321 DQ283964 DQ282814 DQ283655 3216 venustus Altos de Vilches, RõÂo Licay Theloderma AMNH Vietnam, Vinh Phu DQ283050 DQ284080 DQ283779 DQ282659 DQ282904 3969 corticale A161499 Prov, ca. 500 m W Institute of Ecology and Biological Resources Station on SW outskirts of Tam Dao Thorius sp. JAC 21291 Mexico, Oaxaca, DQ283334 DQ284325 DQ283659 2925 Sierra MiahuatlaÂn, 2943 m Thoropa miliaris CFBH 3239 Brazil, SaÄo Paulo, DQ283331 2752 Picinguaba, Ubatuba Tlalocohyla picta RdS 606 Belize, Stann Creek AY843654 DQ284121 AY844640 AY844858 AY844099 AY844276 4739 Dist, Cockscomb Basin Wildlife Sanctuary Tomopterna AMNH South Africa, Western DQ283403 DQ284384 DQ284014 DQ282861 DQ283005 DQ283715 4694 delalandii A144981 Cape Prov, Stellenbosch air ®eld Torrentophryne AF160770 897 aspinia* AF160787 Trachycephalus UMMZ No data AY843771 DQ284097 AY844758 AY844953 AY844190 AY844356 4735 jordani 218914 Trachycephalus AMNH Guyana, Dubulay AY549362 AY844707 AY844912 AY844149 AY844322 4735 venulosus A141142 Ranch on Berbice River, 200 ft, 5Њ40Ј55ЉN, 57Њ51Ј32ЉW Trichobatrachus DPL 3932 Cameroon, Southwest AY843773 DQ284335 AY844760 AY844954 AY844192 DQ283669 4684 robustus Prov, Kumba±Mamfe Triprion petasatus RdS Belize, Hummingbird AY843774 DQ284082 AY844761 AY844955 AY844193 AY844357 4729 Hwy, 9.5 km from Western Hwy turnpoint Triturus cristatus AMNH No data (pet trade) DQ283441 DQ284417 DQ284038 DQ282894 DQ283749 3695 A168421 Trypheropsis KRL 823 Panama, Cocle Prov, DQ283256 DQ284280 DQ283925 DQ282777 DQ282958 DQ283617 4682 warszewitschii (voucher at Parque Nacional El Univ of Cope Panama) 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 321

Locus/partition Total Species ID number Locality mtDNA Histone H3 Rhodopsin SIA Tyrosinase 28S bp

Tylerana arfaki AMS Papua New Guinea, DQ283203 DQ284235 DQ283886 DQ282746 3982 R114913 near Haia Tylototriton AMCC No data DQ283395 DQ284378 DQ283709 3433 shanjing 105494 Typhlonectes BMNH Venezuela (no other DQ283085 DQ284136 DQ283486 2992 natans 2000.218 data) Uperoleia SAMA Australia, New South DQ283221 DQ284251 DQ283898 DQ282758 3445 laevigata R42629 Wales, Ourimbah State Forest Uraeotyphlus MW 1418 India (no other data) DQ283090 DQ284141 DQ282671 DQ283491 3822 narayani (Univ of Kerala) Vanzolinius RdS Ecuador (no other DQ283433 DQ284410 DQ284033 DQ282887 DQ283742 4204 discodactylus data) Werneria mertensi DPL 5107 Cameroon (no other DQ283348 DQ284338 DQ283974 DQ282824 DQ283672 4217 data) Wolterstorf®na DPL 5101 Cameroon (no other DQ283346 DQ284334 DQ283972 DQ282822 DQ283668 3778 parvipalmata data) Xenophrys AY236800 553 lateralis*(ϭX. major) Xenophrys major AMNH Vietnam, Vinh Phu DQ283374 DQ284360 DQ282986 DQ283692 3572 A161506 Prov, ca. 17 km NW Tam Dao Hill Station near Buddhist temple (E Tinh Sinh) Xenopus gilli AMNH South Africa, Western DQ283442 DQ284418 DQ283750 3161 A153027 Cape Prov, 5 km E of DQ283443 Betty's Bay, 34Њ22ЈS, 19Њ7ЈE Xenopus laevis* NC001573 BC054145 AY341764 X59734 4044 Y10943 322 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

APPENDIX 2 ACCESSION NUMBERS AND PUBLICATION REFERENCES FOR GENBANK SEQUENCES USED Accession numbers and publication references are provided for all 199 GenBank sequences included in this study. The locus mtDNA refers to 12S, tRNAVal, and 16S sequences.

Species Locus GenBank accession number Reference Acanthixalus spinosus mtDNA AJ437002, AF215214, AF465438 Vences, unpubl. data; RoÈdel et al., 2003; Vences et al., 2003c Afrixalus fornasini mtDNA U22071 Richards and Moore, 1996 Alligator sinensis mtDNA NC004448 Wu et al., unpubl. data Allophryne ruthveni mtDNA AF364511, AF364512 Austin et al., 2002 Alytes obstetricans Rhodopsin AY364385 Biju and Bossuyt, 2003 Ambystoma tigrinum Rhodopsin U36574 N. Chen et al., 1996 Amnirana galamensis Rhodopsin AY341808 Vences et al., 2003d Amolops hongkongensis mtDNA AF206072, AF206453, AF206117 L.Q. Chen et al., 2005 Andrias davidianus mtDNA AJ492192 Zhang et al., 2003a Aneides hardii mtDNA AY728226 Mueller et al., 2004 Anhydrophryne rattrayi mtDNA AF215504 Vences, unpubl. data Anodonthyla montana mtDNA AJ314812 Odierna et al., unpubl. data Ansonia muelleri mtDNA U52740, U52784 Graybeal, 1997 Ascaphus truei mtDNA AJ440760 Hertwig et al., 2004 Batrachoseps attenuatus mtDNA AY728228 Mueller et al., 2004 Batrachoseps wrightorum mtDNA AY728221 Mueller et al., 2004 Tyrosinase AF249168 Bossuyt and Milinkovitch, 2000 Brachytarsophrys feae mtDNA AY236799 GarcõÂa-ParõÂs et al., 2003 Bufo angusticeps mtDNA AF220852, AF220899 Cunningham and Cherry, 2000 Bufo biporcatus mtDNA AY325987 Darst and Cannatella, 2004 Bufo bufo mtDNA AY325988 Darst and Cannatella, 2004 Bufo bufo Rhodopsin U59921 Fyhrquist et al., unpubl. data Bufo celebensis mtDNA AF375513, AY180245 Darst and Cannatella, 2004; B.J. Evans et al., 2004 Bufo margaritifer mtDNA AF375514, AF375489 A. Gluesenkamp, unpubl. data Bufo mazatlanensis mtDNA U52755, U52723 Graybeal, 1997 Bufo nebulifer mtDNA AY325985 Darst and Cannatella, 2004 Callulina kreffti mtDNA AY326068 Darst and Cannatella, 2004 Callulops slateri mtDNA AF095339 Emerson et al., 2000b Capensibufo rosei mtDNA AF220864, AF220911 Cunningham and Cherry, 2000 Capensibufo tradouwi mtDNA AF220865, AF220912 Cunningham and Cherry, 2000 Centrolene geckoideum mtDNA X86230, X86264, X86298 Hay et al., 1995 Centrolene prosoblepon mtDNA AY364358, AY364379 Biju and Bossuyt, 2003 Centrolene prosoblepon Rhodopsin AY364404 Biju and Bossuyt, 2003 Chiromantis xerampelina mtDNA AF215348, AF458132 Vences, unpubl. data; J.A. Wilkinson et al., 2002 Clinotarsus curtipes mtDNA AF249058, AF249021 Bossuyt and Milinkovitch, 2000 Clinotarsus curtipes Rhodopsin AF249117 Bossuyt and Milinkovitch, 2000 Clinotarsus curtipes Tyrosinase AF249180 Bossuyt and Milinkovitch, 2000 Cryptobatrachus sp. mtDNA AY326050 Darst and Cannatella, 2004 Cryptotriton alvarezdeltoroi mtDNA AF199196 GarcõÂa-ParõÂs and Wake, 2000 Dendrotriton rabbi mtDNA AF199232 GarcõÂa-ParõÂs and Wake, 2000 Desmognathus wrighti mtDNA AY728225 Mueller et al., 2004 Didynamipus sjostedti mtDNA AY325991 Darst and Cannatella, 2004 Ensatina eschscholtzii mtDNA AY728216 Mueller et al., 2004 Euphlyctis cyanophlyctis mtDNA AF249053, AF249015 Bossuyt and Milinkovitch, 2000; Biju and Bossuyt, 2003 Euphlyctis cyanophlyctis Rhodopsin AF249111 Bossuyt and Milinkovitch, 2000 Euphlyctis cyanophlyctis Tyrosinase AF249174 Bossuyt and Milinkovitch, 2000 Euproctus asper mtDNA U04694, U04695 Caccone et al., 1994 Fejervarya cancrivorus mtDNA AB070731, AF206473, AF206092, AF206137 Chen et al., unpubl. data; Sumida et al., 2002 Fejervarya kirtisinghei mtDNA AY014380 Kosuch et al., 2001 Fejervarya syhadrensis mtDNA AY141843, AF249011 Bossuyt and Milinkovitch, 2000; Meegaskum- bura et al., 2002 Fejervarya syhadrensis Rhodopsin AF249107 Bossuyt and Milinkovitch, 2000 Fejervarya syhadrensis Tyrosinase AF249170 Bossuyt and Milinkovitch, 2000 Gazella thomsoni mtDNA M86501 Allard et al., 1992 mtDNA AF461136, AF461137 M. Wilkinson et al., 2002 Glandirana minima mtDNA AF315127, AF315153 Jiang and Zhou, 2001 Hemisus marmoratus mtDNA AY326070 Darst and Cannatella, 2004 mtDNA AY341630, AY341697, AY341725 Vences et al., 2003d Heterixalus tricolor Tyrosinase AY341759 Vences et al., 2003d Hydrophylax galamensis Tyrosinase AY341749 Vences et al., 2003d Hylorina sylvatica mtDNA AY389153 NunÄez, unpubl. data mtDNA AY341634, AY341700, AY341726 Vences et al., 2003d Hymenochirus boettgeri Tyrosinase AY341763 Vences et al., 2003d Iguana iguana mtDNA NC࿞002793 Janke et al., 2001 Indirana sp. 1 mtDNA AF249051 Bossuyt and Milinkovitch, 2000 Indirana sp. 1 Rhodopsin AF249122 Bossuyt and Milinkovitch, 2000 Indirana sp. 1 Tyrosinase AF249185 Bossuyt and Milinkovitch, 2000 Indirana sp. 2 mtDNA AF249064 Bossuyt and Milinkovitch, 2000 Indirana sp. 2 Rhodopsin AF249123 Bossuyt and Milinkovitch, 2000 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 323

Species Locus GenBank accession number Reference Indirana sp. 2 Tyrosinase AF249186 Bossuyt and Milinkovitch, 2000 Ixalotriton niger mtDNA AF451248 Parra-Olea, 2002 Laliostoma labrosum Tyrosinase AF249169 Bossuyt and Milinkovitch, 2000 Lankanectes corrugatus mtDNA AF215393, AF249019, AF249043 Vences, unpubl. data; Bossuyt and Milinkov- itch, 2000 Lankanectes corrugatus Rhodopsin AF249115 Bossuyt and Milinkovitch, 2000 Lankanectes corrugatus Tyrosinase AF249178 Bossuyt and Milinkovitch, 2000 Latimeria chalumnae mtDNA NC࿞001804 Zardoya and Meyer, 1996 Latimeria chalumnae Rhodopsin AF131253 Yokoyama et al., 1999 Tyrosinase AY341760 Vences et al., 2003d Limnodynastes salmini mtDNA AY326071 Darst and Cannatella, 2004 Limnonectes acanthi mtDNA AY313724 B.J. Evans et al., 2004 Limnonectes heinrichi mtDNA AY313749 B.J. Evans et al., 2004 Limnonectes kuhlii mtDNA AY313686 B.J. Evans et al., 2004 Limnonectes visayanus mtDNA AY313719 B.J. Evans et al., 2004 Lineatriton lineolus mtDNA AF380808 Parra-Olea and Wake, 2001 Micrixalus fuscus mtDNA AF249024 Bossuyt and Milinkovitch, 2000 Micrixalus fuscus mtDNA AF249056 Bossuyt and Milinkovitch, 2000 Micrixalus fuscus Rhodopsin AF249120 Bossuyt and Milinkovitch, 2000 Micrixalus fuscus Tyrosinase AF249183 Bossuyt and Milinkovitch, 2000 Micrixalus kottigeharensis mtDNA AF249025, AF249041 Bossuyt and Milinkovitch, 2000 Micrixalus kottigeharensis Rhodopsin AF249121 Bossuyt and Milinkovitch, 2000 Micrixalus kottigeharensis Tyrosinase AF249184 Bossuyt and Milinkovitch, 2000 mtDNA AF285207 Ziegler, unpubl. data Nannophrys ceylonensis mtDNA AF249016, AF249047 Bossuyt and Milinkovitch, 2000 Nannophrys ceylonensis Rhodopsin AF249112 Bossuyt and Milinkovitch, 2000 Nannophrys ceylonensis Tyrosinase AF249175 Bossuyt and Milinkovitch, 2000 Nanorana pleskei mtDNA AF206111, AF206156, AF206492 L.Q. Chen et al., 2005 Nasikabatrachus sahyadrensis mtDNA AY364360, AY364381 Biju and Bossuyt, 2003 Nasikabatrachus sahyadrensis Rhodopsin AY364406 Biju and Bossuyt, 2003 Nasikobatrachidae sp. mtDNA AY425725, AY425726 Dutta et al., 2004 Natalobatrachus bonebergi mtDNA AF215396, AF215198 Vences, unpubl. data Nelsonophryne aequatorialis mtDNA AY326067 Darst and Cannatella, 2004 Nesomantis thomasseti Tyrosinase AY341761 Vences et al., 2003d mtDNA AY147246, AY147247 Steinfartz et al., 2002 Nototriton abscondens mtDNA AF199199 GarcõÂa-ParõÂs and Wake, 2000 Nyctibatrachus cf. aliciae mtDNA AF249018, AF249063 Bossuyt and Milinkovitch, 2000 Nyctibatrachus cf. aliciae Rhodopsin AF249114 Bossuyt and Milinkovitch, 2000 mtDNA AF249017, AF249052, AY341687 Bossuyt and Milinkovitch, 2000; Vences et al., 2003d Nyctibatrachus major Rhodopsin AF249113 Bossuyt and Milinkovitch, 2000 Nyctibatrachus major Tyrosinase AF249176 Bossuyt and Milinkovitch, 2000 Occidozyga lima mtDNA AF161027 Marmayou et al., 2000 mtDNA AF199230 GarcõÂa-ParõÂs and Wake, 2000 Osornophryne guacamayo mtDNA AY326036 Darst and Cannatella, 2004 Parvimolge townsendi mtDNA AF451247 Parra-Olea, 2002 mtDNA AY236801, AY364341, AY364363 GarcõÂa-ParõÂs et al., 2003; Biju and Bossuyt, 2003 Pelobates cultripes Rhodopsin AY364386 Biju and Bossuyt, 2003 Pelomedusa subrufa mtDNA NC࿞001947 Zardoya and Meyer, 1998 Pelophryne brevipes mtDNA AF375503, AF375530 Gluesenkamp, unpubl. data Pelophylax ridibunda mtDNA AB023397, AY147983 Sumida et al., 2000b Petropedetes parkeri mtDNA AY341694, AY364348, AY364369 Biju and Bossuyt, 2003 Petropedetes parkeri Rhodopsin AY364394 Biju and Bossuyt, 2003 Petropedetes parkeri Tyrosinase AY341757 Vences et al., 2003d Phaeognathus hubrichti mtDNA AY728233 Mueller et al., 2004 Phrynopus sp. KU 202652 mtDNA AY326010 Darst and Cannatella, 2004 Polypedates cruciger mtDNA AF249028, AY341685 Bossuyt and Milinkovitch, 2000; Vences et al., 2003d Polypedates cruciger Rhodopsin AF249124 Bossuyt and Milinkovitch, 2000 Polypedates cruciger Tyrosinase AF249187 Bossuyt and Milinkovitch, 2000 Ptychadena anchietae mtDNA AF261249, AF261267 Richards et al., 2000 Ramanella obscura mtDNA AF215382 Vences, unpubl. data Rana berlandieri mtDNA AY115111 ZaldõÂvar-RiveroÂn et al., 2004 Ranodon sibiricus mtDNA NC࿞004021 Zhang et al., 2003b Rhamphophryne festae mtDNA AF375504, AF375531 Gluesenkamp, unpubl. data Rhodopsin AY364390 Biju and Bossuyt, 2003 Siren intermedia mtDNA Y10946 Feller and Hedges, 1998 Speleomantes italicus mtDNA AY728215 Mueller et al., 2004 Sphaerotheca pluvialis mtDNA AF249014, AF249042 Bossuyt and Milinkovitch, 2000 Sphaerotheca pluvialis Rhodopsin AF249110 Bossuyt and Milinkovitch, 2000 Sphaerotheca pluvialis Tyrosinase AF249173 Bossuyt and Milinkovitch, 2000 Stephopaedes anotis mtDNA AF220910 Cunningham and Cherry, 2000 Sylvirana temporalis mtDNA AF249022, AF249054 Bossuyt and Milinkovitch, 2000 Sylvirana temporalis Rhodopsin AF249118 Bossuyt and Milinkovitch, 2000 Sylvirana temporalis Tyrosinase AF249181 Bossuyt and Milinkovitch, 2000 Taylorana limborgi mtDNA AF261251 Richards et al., 2000 Torrentophryne aspinia mtDNA AF160770, AF160787 W. Liu et al., 2000 Xenophrys major mtDNA AY236800 GarcõÂa-ParõÂs et al., 2003 Xenopus laevis Rhodopsin BC054145 Klein et al., 2002 Xenopus laevis Tyrosinase AY341764 Vences et al., 2003d 324 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

APPENDIX 3 BASE-PAIR LENGTH OF 28S FRAGMENT

Length Length Higher taxon and family Species (bp) Higher taxon and family Species (bp)

Marsupialia Anura (continued) Didelphidae Didelphis marsupialis 1092 Bufonidae Bufo viridis 751 Bufonidae Bufo woodhousii 751 Diapsida Bufonidae Dendrophryniscus minutus 752 Alligatoridae Alligator sinensis 696 Bufonidae Melanophryniscus klappenbachi 740 Iguanidae Iguana iguana 699 Bufonidae Nectophryne afra 752 Testudines Bufonidae Nectophryne batesi 752 Chelydridae Chelydra serpentina 694 Bufonidae Nectophrynoides tornieri 753 Pelomedusidae Pelomedusa subrufa 694 Bufonidae Schismaderma carens 754 Bufonidae Werneria mertensi 751 Coelocantha Bufonidae Wolterstorf®na parvipalmata 750 Latimeriidae Latimeria chalumnae 691 Centrolenidae Centrolene prosoblepon 732 Anura Centrolenidae Cochranella bejaranoi 732 Alytidae Alytes obstetricans 706 Centrolenidae Hyalinobatrachium ¯eischmanni 732 Alytidae Discoglossus galganoi 706 Ceratobatrachidae Batrachylodes vertebralis 714 Alytidae Discoglossus pictus 706 Ceratobatrachidae Ceratobatrachus guentheri 720 Amphignathodontidae Flectonotus sp. 762 Ceratobatrachidae Playmantis pelewensis 713 Arthroleptidae Arthroleptis tanneri 721 Ceratophryidae Atelognatus patagonicus 732 Arthroleptidae Arthroleptis variabilis 722 Ceratophryidae 732 Arthroleptidae Astylosternus schioetzi 716 Ceratophryidae Ceratophrys cranwelli 728 Arthroleptidae Cardioglossa gratiosa 717 Ceratophryidae Telmatobius sp. 728 Arthroleptidae Cardioglossa leucomystax 719 Cryptobatrachidae Stefania evansi 786 Arthroleptidae Leptopelis argenteus 717 Cycloramphidae Alsodes gargola 757 Arthroleptidae Leptopelis bocagei 717 Cycloramphidae Cycloramphus boraceiensis 742 Arthroleptidae Leptopelis sp. 717 Cycloramphidae 757 Arthroleptidae Nyctibates corrugatus 717 Cycloramphidae Odontophrynus achalensis 780 Arthroleptidae Schoutedenella schubotzi 744 Cycloramphidae 778 Arthroleptidae Schoutedenella xenodactyloides 762 Cycloramphidae Rhinoderma darwinii 744 Arthroleptidae Scotobleps gabonicus 718 Dendrobatidae Allobates boulengeri 774 Arthroleptidae Trichobatrachus robustus 714 Dendrobatidae Ameerega femoralis 782 Batrachophrynidae Caudiverbera caudiverbera 709 Dendrobatidae Colostethus undulatus 775 Batrachophrynidae Telmatobufo venustus 710 Dendrobatidae Dendrobates auratus 759 Bombinatoridae Bombina microdeladigitora 710 Dendrobatidae Minyobates claudiae 760 Bombinatoridae Bombina orientalis 710 Dendrobatidae Phobobates silverstonei 771 Bombinatoridae Bombina variegata 710 Dendrobatidae Phyllobates lugubris 769 Brachycephalidae Barycholos ternetzi 744 Dicroglossidae Hoplobatrachus occipitalis 708 Brachycephalidae Brachycephalus ephippium 740 Dicroglossidae Hoplobatrachus rugulosus 708 Brachycephalidae Craugastor alfredi 759 Dicroglossidae Limnonectes kuhlii 709 Brachycephalidae 760 Dicroglossidae Occidozyga lima 708 Brachycephalidae Craugastor bufoniformis 744 Dicroglossidae Paa exilispinosa 714 Brachycephalidae Craugastor pluvicanorus 830 Dicroglossidae Quasipaa verrucospinosa 708 Brachycephalidae Craugastor punctariolus 756 Dicroglossidae Phrynoglossus baluensis 708 Brachycephalidae 756 Dicroglossidae Phrynoglossus borealis 708 Brachycephalidae Eleutherodactylus binotatus 747 Dicroglossidae Sphaerotheca breviceps 709 Brachycephalidae Eleutherodactylus juipoca 738 Heleophrynidae Heleophryne regis 719 Brachycephalidae Eleutherodactylus rugulosus 757 Hemisotidae Hemisus marmoratus 709 Brachycephalidae Euhyas planirostris 768 Hylidae Anotheca spinosa 743 Brachycephalidae Phrynopus sp. 743 Hylidae Hypsiboas albomarginatus 764 Brachycephalidae Syrrhophus marnockii 769 Hylidae 757 Brachycephalidae Syrrhophus nitidus 769 Hylidae Argenteohyla siemersi pederseni 740 Brevicipitidae Breviceps mossambicus 712 Hylidae Charadrahyla nephila 741 Brevicipitidae Callulina kisiwamsitu 710 Hylidae 789 Brevicipitidae Probreviceps macrodactylus 710 Hylidae 713 Bufonidae Atelopus spumarius 766 Hylidae 745 Bufonidae Bufo alvarius 751 Hylidae Duellmanohyla ru®oculis 738 Bufonidae Bufo amboroensis 751 Hylidae Ecnomiohyla miliaria 744 Bufonidae Bufo andrewsi 752 Hylidae Exerodonta chimalapa 743 Bufonidae Bufo arenarum 752 Hylidae Hyla cinerea 742 Bufonidae Bufo asper 751 Hylidae Hyloscirtus armatus 764 Bufonidae Bufo boreas 751 Hylidae Hyloscirtus palmeri 767 Bufonidae Bufo brauni 751 Hylidae Hypsiboas boans 757 Bufonidae Bufo camerunensis 732 Hylidae Hypsiboas granosus 767 Bufonidae Bufo cf. chilensis 752 Hylidae Hypsiboas multifasciatus 762 Bufonidae Bufo cognatus 751 Hylidae Litoria genimaculata 690 Bufonidae Bufo divergens 750 Hylidae Lysapsus laevis 757 Bufonidae Bufo granulosus 742 Hylidae Nyctimistes dayi 694 Bufonidae Bufo guttatus 752 Hylidae Osteocephalus taurinus 740 Bufonidae Bufo gutturalis 732 Hylidae Osteopilus septentrionalis 742 Bufonidae Bufo haematiticus 752 Hylidae Phrynohyas venulosa 744 Bufonidae Bufo latifrons 751 Hylidae Phyllomedusa vaillanti 795 Bufonidae Bufo maculatus 751 Hylidae Plectrohyla guatemalensis 744 Bufonidae Bufo punctatus 700 Hylidae Pseudacris crucifer 743 Bufonidae Bufo quercicus 751 Hylidae Pseudacris triseriata 747 Bufonidae Bufo terrestris 751 Hylidae Pseudis paradoxa 758 Bufonidae Bufo tuberosus 721 Hylidae Scinax garbei 778 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 325

Length Length Higher taxon and family Species (bp) Higher taxon and family Species (bp)

Anura (continued) Anura (continued) Hylidae Smilisca phaeota 745 Pelodytidae Pelodytes puntatus 703 Hylidae Sphaenorhynchus lacteus 753 Petropedetidae Arthroleptides sp. 747 Hylidae Tlalacohyla picta 747 Petropedetidae Conraua goliath 708 Hylidae Trachycephalus jordani 742 Petropedetidae Conraua robusta 708 Hylidae Triprion petasatus 743 Petropedetidae Petropedetes cameronensis 718 Hyperoliidae Afrixalus fornasinii 714 Petropedetidae Petropedetes palmipes 726 Hyperoliidae Afrixalus pygmaeus 714 Phrynobatrachidae Dimorphognathus africanus 708 Hyperoliidae Alexteroon obstetricans 716 Phrynobatrachidae Phrynobatrachus auritus 707 Hyperoliidae Cryptothylax gresshof® 714 Phrynobatrachidae Phrynobatrachus calcaratus 719 Hyperoliidae Heterixalus sp. 735 Phrynobatrachidae Phrynobatrachus dendrobates 708 Hyperoliidae Hyperolius alticola 714 Phrynobatrachidae Phrynobatrachus dispar 719 Hyperoliidae Hyperolius punticulatus 713 Phrynobatrachidae Phrynobatrachus mababiensis 708 Hyperoliidae Hyperolius tuberilinguis 713 Phrynobatrachidae Phrynobatrachus natalensis 708 Hyperoliidae Kassina senegalensis 714 Phrynobatrachidae Phrynodon sandersoni 710 Hyperoliidae Nesionixalus thomensis 714 Pipidae Silurana tropicalis 713 Hyperoliidae Opisthothylax immaculatus 714 Pipidae Xenopus gilli 713 Hyperoliidae Phlyctimantis leonardi 714 Ptychadenidae Ptychadena cooperi 714 Hyperoliidae Tachycnemis seychellensis 733 Pyxicephalidae Amietia angolensis 718 Leiopelmatidae Ascaphus truei 703 Pyxicephalidae Amietia fuscigula 721 Leiopelmatidae Leiopelma archeyi 703 Pyxicephalidae Amietia vertebralis 718 Leiopelmatidae Leiopelma hochstetteri 703 Pyxicephalidae Arthroleptella bicolor 731 Leptodactylidae Edalorhina perezi 756 Pyxicephalidae Aubria subsigillata 708 Leptodactylidae Leptodactylus fuscus 750 Pyxicephalidae Aubria subsigillata 708 Leptodactylidae Leptodactylus ocellatus 742 Pyxicephalidae Pyxicephalus edulis 708 Leptodactylidae Lithodytes lineatus 746 Pyxicephalidae Tomopterna delalandii 713 Leptodactylidae Paratelmatobius sp. 730 Ranidae Amerana mucosa 708 Leptodactylidae 761 Leptodactylidae Scythrophrys sawayae 728 Ranidae Amnirana albilabris 708 Leptodactylidae Vanzolinius discodactylus 745 Ranidae Amolops chapaensis 709 Limnodynastidae Adelotus brevis 721 Ranidae Aquarana catesbeiana 708 Limnodynastidae Heleioporus australiacus 720 Ranidae Aquarana clamitans 709 Limnodynastidae Lechriodus ¯etcheri 727 Ranidae Aquarana grylio 708 Limnodynastidae Limnodynastes depressus 724 Ranidae Aquarana heckscheri 708 Limnodynastidae Limnodynastes dumerilli 719 Ranidae Aquarana aurora 708 Limnodynastidae Limnodynastes lignarius 720 Ranidae Chalcorana chalconota 708 Limnodynastidae Limnodynastes ornatus 730 Ranidae Huia nasica 709 Limnodynastidae Neobatrachus pictus 721 Ranidae Hylarana taipehensis 708 Limnodynastidae Neobatrachus sudelli 721 Ranidae Lithobates palmipes 708 Limnodynastidae Philoria sphagnicola 724 Ranidae Meristogenys orphnocnemis 708 Mantellidae Aglyptodactylus madagascariensis 710 Ranidae Nidirana adenopleura 714 Mantellidae Laliostoma labrosum 712 Ranidae Nidirana chapaensis 708 Mantellidae Mantella aurantiaca 685 Ranidae Odorrana grahami 709 Mantellidae Mantella nigricans 685 Ranidae 709 Megophryidae 728 Ranidae Pantherana capito 708 Megophryidae Leptobrachium hasselti 726 Ranidae Pantherana chiricahuensis 708 Megophryidae Leptolalax pelodytoides 726 Ranidae Pantherana forreri 708 Megophryidae Ophryophryne microstoma 725 Ranidae Pantherana pipiens 708 Megophryidae Xenophrys major 726 Ranidae Pantherana yavapaiensis 708 Microhylidae Aphantophryne pansa 719 Ranidae Papurana daemeli 708 Microhylidae Calluella guttulata 725 Ranidae Pelophylax nigromaculata 708 Microhylidae Choerophryne sp. 719 Ranidae Pseudoamalops sauteri 713 Microhylidae Cophixalus sphagnicola 718 Ranidae Pseudorana johnsi 708 Microhylidae Ctenophryne geayei 727 Ranidae Rana japonica 709 Microhylidae Dasypops schirchi 719 Ranidae Rana sylvatica 707 Microhylidae Dyscophus guineti 716 Ranidae Rana temporaria 707 Microhylidae 720 Ranidae Staurois tuberlinguis 710 Microhylidae 721 Ranidae Sylvirana guentheri 708 Microhylidae Genyophryne thomsoni 728 Ranidae Sylvirana maosonensis 708 Microhylidae Hoplophryne rogersi 718 Ranidae Sylvirana nigrovittata 708 Microhylidae Kalophrynus pleurostigma 716 Ranidae Trypheropsis warszewitschii 708 Microhylidae Kaloula pulchra 732 Rhacophoridae Chirixalus doriae 709 Microhylidae Liophryne rhododactyla 718 Rhacophoridae Chirixalus vittatus 710 Microhylidae 725 Rhacophoridae Kurixalus eif®ngeri 709 Microhylidae Microhyla sp. 698 Rhacophoridae Kurixalus idiooticus 709 Microhylidae Oreophryne brachypus 719 Rhacophoridae Nyctixalus pictus 709 Microhylidae Phrynomantis bifasciatus 726 Rhacophoridae Nyctixalus spinosus 709 Microhylidae Platyelis sp. 716 Rhacophoridae Philautus rhododiscus 709 Microhylidae Plethodontohyla sp. 717 Rhacophoridae 709 Microhylidae Scaphiophryne marmorata 716 Rhacophoridae 709 Microhylidae Synapturanus mirandaribeiroi 718 Rhinophrynidae Rhinophrynus dorsalis 705 Myobatrachidae Arenophryne rotunda 726 Scaphiopodidae Scaphiopus couchii 703 Myobatrachidae Crinia nimba 724 Scaphiopodidae Scaphiopus holbrooki 703 Myobatrachidae Metacrinia nichollsi 726 Scaphiopodidae Spea hammondii 703 Myobatrachidae Myobatachus gouldii 726 Sooglossidae Nesomantis thomasseti 737 Myobatrachidae Pseudophryne bibroni 726 Sooglossidae Sooglossus seychellensis 741 Myobatrachidae Pseudophryne coriacea 726 Myobatrachidae Spicospina ¯ammocaerulea 726 Artiodactyla Pelobatidae Pelobates fuscus 713 Bovidae Gazella thomsoni 748 326 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Length Length Higher taxon and family Species (bp) Higher taxon and family Species (bp)

Caudata Caudata (continued) Ambystomatidae Dicamptodon ensatus 694 Sirenidae Pseudobranchus striatus 694 Amphiumidae Amphiuma tridactylum 694 Sirenidae Siren lacertina 694 Cryptobranchidae Cryptobranchus alleganiensis 694 Hynobiidae Batrachuperus pinchoni 694 Gymnophiona Plethodontidae Bolitoglossa rufescens 694 Caeciliidae Boulengerula uluguruensis 701 Plethodontidae Desmognathus quadramaculatus 695 Caeciliidae 709 Plethodontidae Eurycea wilderae 694 Caeciliidae Crotaphatrema tchabalmboensis 727 Plethodontidae Gyrinophilus porphyriticus 694 Caeciliidae 710 Plethodontidae Plethodon dunni 694 Caeciliidae 700 Plethodontidae Plethodon jordani 694 Caeciliidae Hypogeophis rostratus 702 Plethodontidae Pseudoeurycea conanti 695 Caeciliidae Schistometopum gregorii 701 Plethodontidae Thorius sp. 694 Caeciliidae Siphonops hardyi 700 Proteidae Necturus cf. beyeri 694 Caeciliidae 684 Proteidae Necturus maculosus 694 Ichthyophiidae Ichthyophis peninsularis 683 Rhyacotritonidae Rhyacotriton cascadae 694 Ichthyophiidae Ichthyophis sp. 697 Salamandridae Pleurodeles waltl 694 Ichthyophiidae 683 Salamandridae Triturus sp. 695 Rhinatrematidae Epicrionops sp. 714 Salamandridae Tylototriton shanjing 694 Rhinatrematidae 695

APPENDIX 4 BRANCH LENGTHS,BREMER SUPPORT, AND JACKKNIFE VALUES Values given correspond to branches numbered in ®gures 50, 52, 53, 54, 56, 58, 59, 60, 61, 62, 63, and 65.

Branch Bremer Branch Bremer Branch Taxon length support Jackknife Branch Taxon length support Jackknife

1 Amniota 117 96 100 43 Unnamed 21 9 100 2 Mammalia 81 67 100 44 Unnamed 20 6 92 3 103 83 100 45 Unnamed 22 9 99 4 Testudines 105 83 100 46 Unnamed 26 21 100 5 Diapsida 94 75 100 47 Unnamed 11 9 99 6 Amphibia 78 49 100 48 Unnamed 7 2 78 7 Gymnophiona 89 78 100 49 Plethosalamandroidei 50 41 99 8 Rhinatrematidae 49 30 100 50 Xenosalamandroidei 48 31 99 9 Stegokrotaphia 58 47 100 51 Plethodontidae 94 85 99 10 Ichthyophiidae 89 82 100 52 Plethodontinae 36 24 100 11 Unnamed 56 33 100 53 Unnamed 26 15 100 12 Caeciliidae 60 44 100 54 Unnamed 45 38 100 13 Unnamed 63 41 100 55 Unnamed 29 19 99 14 Unnamed 34 21 100 56 Unnamed 39 26 100 15 Unnamed 98 90 100 57 Desmognathus 37 26 100 16 Unnamed 44 27 100 58 Unnamed 26 13 100 17 Scolecomorphinae 47 40 100 59 Unnamed 77 66 100 18 Unnamed 73 32 100 60 Unnamed 34 13 97 19 Unnamed 31 23 100 61 Batrachoseps 85 43 100 20 Unnamed 55 39 100 62 Unnamed 15 6 88 21 Unnamed 24 19 100 63 Unnamed 25 15 99 22 Unnamed 25 17 100 64 Unnamed 48 34 100 23 Batrachia 72 109 100 65 Unnamed 24 21 99 24 Caudata 114 107 99 66 Unnamed 9 4 97 25 Cryptobranchoidei 58 40 100 67 Unnamed 7 1 51 26 Hynobiidae 91 80 100 68 Unnamed 10 4 87 27 Cryptobranchidae 122 115 100 69 Unnamed 9 6 94 28 Andrias 71 64 100 70 Unnamed 60 11 99 29 Diadectosalamandroidei 43 30 99 71 Unnamed 5 2 62 30 Hydatinosalamandroidei 38 29 100 72 Pseudoeurycea 7376 31 Perennibranchia 43 35 100 73 Unnamed 12 9 99 32 Proteidae 166 163 100 74 Anura 125 109 99 33 Sirenidae 108 98 100 75 Leiopelmatidae 55 41 100 34 Siren 57 49 100 76 Leiopelma 72 65 100 35 Treptobranchia 43 29 100 77 Lalagobatrachia 82 57 99 36 Ambystomatidae 78 69 100 78 Xenoanura 68 55 100 37 Dicamptodon 168 165 100 79 Pipidae 45 48 100 38 Ambystoma 135 128 100 80 Unnamed 36 130 100 39 Unnamed 54 52 100 81 Pipa 85 198 100 40 Salamandridae 72 63 100 82 Unnamed 145 17 100 41 Pleurodelinae 35 28 100 83 Xenopus 24 24 100 42 Unnamed 37 24 100 84 Sokolanura 36 20 99 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 327

Branch Bremer Branch Bremer Branch Taxon length support Jackknife Branch Taxon length support Jackknife

85 Costata 69 55 100 164 Arthroleptidae 57 37 100 86 Alytidae 68 48 100 165 Leptopelinae 112 100 100 87 Discoglossus 138 130 100 166 Unnamed 53 37 100 88 Bombinatoridae 200 198 100 167 Unnamed 72 72 100 89 Unnamed 27 17 100 168 Arthroleptinae 56 44 100 90 Unnamed 28 24 100 169 Astylosternini 54 39 100 91 Acosmanura 81 65 99 170 Unnamed 45 21 99 92 Anomocoela 85 65 100 171 Unnamed 78 60 100 93 Pelodytoidea 57 33 100 172 Arthroleptini 38 28 100 94 Scaphiopodidae 101 87 100 173 Unnamed 109 92 100 95 Scaphiopus 82 72 100 174 Cardioglossa 79 70 100 96 Pelobatoidea 74±75 53 100 175 Arthroleptis 55 42 100 97 Pelobatidae 69 66 100 176 Unnamed 45 32 100 98 Megophryidae 100 39 100 177 Unnamed 26 14 100 99 Unnamed 65±66 27 100 178 Unnamed 46 32 100 100 Leptobrachium 61±62 37 100 179 Unnamed 68 62 100 101 Unnamed 18±92 16 100 180 Natatanura 65 34 99 102 Xenophrys 31±35 31 100 181 Ptychadenidae 135 100 100 103 Unnamed 39±42 10 100 182 Unnamed 37 85 100 104 Ophryophryne 99 81 100 183 Victoranura 39 20 99 105 Neobatrachia 127 108 100 184 Ceratobatrachidae 114 43 100 106 Heleophrynidae 187 186 100 185 Unnamed 129 120 100 107 Phthanobatrachia 66 31 99 186 Unnamed 41 22 100 108 Ranoides 110 31 99 187 Unnamed 56 35 100 109 Allodapanura 45 31 100 188 Platymantis 100 82 100 110 Microhylidae 72 34 100 189 Telmatobatrachia 19 10 99 111 Unnamed 7 2 71 190 Micrixalidae 105 42 100 112 Unnamed 17 3 90 191 Ametrobatrachia 17 12 99 113 Unnamed 4 3 85 192 Africanura 32 21 99 114 Unnamed 36 33 97 193 Phrynobatrachidae 87 51 100 115 Unnamed 28 13 99 194 Unnamed 45 26 100 116 Unnamed 50 9 98 195 Unnamed 85 70 100 117 Unnamed 8 5 93 196 Unnamed 74 103 100 118 Cophylinae 118 24 100 197 Unnamed 62 35 100 119 Unnamed 71 6 99 198 Unnamed 115 61 100 120 Unnamed 8 11 98 199 Unnamed 52 31 100 121 Gastrophryninae 36 13 99 200 Pyxicephaloidea 25 14 99 122 Unnamed 67 58 100 201 Petropedetidae 33 15 99 123 Unnamed 44 15 99 202 Conraua 89 24 100 124 Unnamed 54 34 100 203 Unnamed 17 13 99 125 Unnamed 39 23 100 204 Indirana 30 22 100 126 Unnamed 29 18 99 205 Unnamed 51 39 100 127 Gastrophryne 58 47 100 206 Petropedetes 77 54 100 128 Unnamed 27 16 99 207 Unnamed 29 19 100 129 Unnamed 29 18 100 208 Unnamed 10 5 89 130 Microhylinae 54 42 100 209 Pyxicephalidae 36 21 100 131 Unnamed 24 45 98 210 Pyxicephalinae 117 105 100 132 Unnamed 43 50 100 211 Aubria 189 187 100 133 Unnamed 115 92 100 212 Cacosterninae 56 44 99 134 Unnamed 45 31 99 213 Unnamed 43 6 94 135 Asterophryinae 100 24 100 214 Unnamed 8 5 92 136 Unnamed 44 7 100 215 Unnamed 28 9 99 137 Unnamed 22 13 98 216 Unnamed 33 10 98 138 Unnamed 21 40 91 217 Unnamed 8 2 85 139 Unnamed 54 22 100 218 Amietia 33 6 99 140 Unnamed 32 14 100 219 Unnamed 4 3 85 141 Unnamed 38 7 98 220 Saukrobatrachia 26 17 99 142 Unnamed 10 7 98 221 Dicroglossidae 39 31 100 143 Afrobatrachia 52 37 100 222 Occidozyginae 42 36 100 144 Xenosyneunitanura 96 81 100 223 Unnamed 24 15 99 145 Brevicipitidae 79 49 100 224 Unnamed 57 21 99 146 Unnamed 77 53 100 225 Dicroglossinae 37 27 100 147 Callulina 103 103 100 226 Limnonectini 84 20 100 148 Laurentobatrachia 61 42 100 227 Unnamed 45 14 100 149 Hyperoliidae 114 64 100 228 Unnamed 34 14 99 150 Unnamed 14 9 98 229 Unnamed 9 6 98 151 Unnamed 31 21 100 230 Unnamed 13 9 99 152 Unnamed 76 29 100 231 Unnamed 30 10 99 153 Unnamed 52 27 100 232 Dicroglossini 43 32 100 154 Unnamed 43 32 100 233 Quasipaa 56 46 100 155 Afrixalus 36 16 99 234 Unnamed 26 20 100 156 Unnamed 98 68 100 235 Unnamed 71 65 100 157 Heterixalus 20 8 97 236 Fejervarya 1 60 39 100 158 Unnamed 42 23 100 237 Unnamed 43 32 100 159 Unnamed 82 37 100 238 Sphaerotheca 57 53 100 160 Alexteroon 19 19 100 239 Unnamed 26 6 86 161 Hyperolius 21 9 98 240 Fejervarya 2 16 15 100 162 Unnamed 14 8 98 241 Unnamed 29 19 100 163 Unnamed 28 10 97 242 Unnamed 28 16 100 328 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Branch Bremer Branch Bremer Branch Taxon length support Jackknife Branch Taxon length support Jackknife

243 Hoplobatrachus 27 14 99 322 Limnodynastidae 72 61 100 244 Aglaioanura 33 19 99 323 Unnamed 47 30 100 245 Rhacophoroidea 36 24 100 324 Unnamed 22 16 100 246 Mantellidae 52 40 100 325 Unnamed 26 18 100 247 Boophinae 81 67 100 326 Neobatrachus 80 75 100 248 Mantellinae 38 21 100 327 Unnamed 33 20 100 249 Laliostomini 50 30 100 328 Unnamed 79 67 100 250 Mantellini 52 44 100 329 Unnamed 35 17 99 251 Mantidactylus 52 35 100 330 Limnodynastes 57 47 100 252 Mantella 83 68 100 331 Unnamed 20 13 99 253 Rhacophoridae 57 46 100 332 Unnamed 31 22 100 254 Rhacophorinae 61 42 100 333 Unnamed 48 41 100 255 Unnamed 33 16 57 334 Myobatrachidae 38 24 100 256 Kurixalus 137 127 100 335 Unnamed 30 12 98 257 Unnamed 15 2 100 336 Unnamed 49 36 100 258 Unnamed 54 42 100 337 Unnamed 63 53 100 259 Unnamed 74 54 100 338 Unnamed 27 20 100 260 Nyctixalus 47 38 100 339 Unnamed 58 32 100 261 Unnamed 38 27 100 340 Unnamed 45 47 100 262 Rhacophorus 44 32 100 341 Unnamed 23 16 100 263 Unnamed 30 10 96 342 Unnamed 36 27 100 264 Unnamed 40 25 100 343 Unnamed 37 24 100 265 Unnamed 41 29 100 344 Unnamed 55 48 100 266 Polypedates 66 60 100 345 Pseudophryne 35 30 100 267 Chiromantis 55 37 100 346 Unnamed 56 27 100 268 Unnamed 57 39 100 347 Unnamed 5 3 85 269 Ranoidea 23 17 99 348 Nobleobatrachia 96 88 100 270 Nyctibatrachidae 18 8 97 349 Meridianura 51 35 100 271 Nyctibatrachus 75 64 100 350 Brachycephalidae 49 43 100 272 Ranidae 57 37 99 351 Unnamed 52 38 100 273 Unnamed 53 36 99 352 Unnamed 54 38 100 274 Hylarana 37 21 100 353 Unnamed 35 27 100 275 Unnamed 96 85 100 354 Unnamed 44 26 100 276 Unnamed 36 13 99 355 Unnamed 38 26 100 277 Unnamed 28 22 99 356 Unnamed 77 51 100 278 Hydrophylax 45 10 99 357 Unnamed 110 96 100 279 Unnamed 42 26 100 358 Syrrhophus 78 72 100 280 Sylvirana 27 13 99 359 Unnamed 32 25 100 281 Unnamed 37 22 100 360 Phrynopus 61 42 100 282 Unnamed 17 12 99 361 Craugastor 51 34 100 283 Unnamed 39 31 100 362 Unnamed 102 85 100 284 Unnamed 26±27 3 87 363 Unnamed 95 77 100 285 Unnamed 32±34 15 100 364 Unnamed 75 50 100 286 Unnamed 7±23 13 69 365 Unnamed 119 107 100 287 Unnamed 25±26 23 52 366 Cladophrynia 58 45 100 288 Pelophylax 12 2 58 367 Cryptobatrachidae 34 22 100 289 Unnamed 10±19 7 98 368 Tinctanura 43 29 100 290 Unnamed 32 10 99 369 Amphignathodontidae 44 33 100 291 Babina 56 32 100 370 Gastrotheca 67 47 100 292 Huia 70 55 100 371 Athesphatanura 41 37 100 293 Unnamed 28 19 99 372 Hylidae 35 37 100 294 Unnamed 43 16 99 373 Unnamed (Phyllomedu- 76 65 100 295 Unnamed 39 30 100 sinae ϩ Pelodryadinae) 296 Rana 50 39 100 374 Phyllomedusinae 87 45 100 297 Unnamed 52 39 100 375 Unnamed 42 24 99 298 Unnamed 34 26 100 376 Unnamed 34 16 99 299 Unnamed 31 16 100 377 Pelodryadinae 45 33 100 300 Unnamed 18 8 98 378 Unnamed 49 37 100 301 Lithobates 33 27 100 379 Unnamed 74 24 100 302 Unnamed 59 45 100 380 Unnamed 40 25 100 303 Unnamed 18 6 88 381 Unnamed 39 27 100 304 Unnamed 8 2 56 382 Unnamed 58 40 100 305 Unnamed 24 1 52 383 Unnamed 16 10 98 306 Unnamed 42 28 100 384 Unnamed 20 11 99 307 Unnamed 28 12 99 385 Unnamed 22 12 98 308 Unnamed 32 9 90 386 Hylinae 32 24 100 309 Unnamed 71 35 100 387 Unnamed 28 20 100 310 Unnamed 19 5 78 388 Unnamed 49 29 100 311 Unnamed 44 34 100 389 Scinax 115 97 100 312 Unnamed 9 7 99 390 Cophomantini 61 52 100 313 Unnamed 4 1 66 391 Hyloscirtus 61 40 100 314 Hyloides 60 35 100 392 Unnamed 61 33 100 315 Sooglossidae 50 42 100 393 Unnamed 44 30 100 316 Nasikabatrachus 122 117 100 394 Unnamed 48 24 100 317 Sooglossus 62 62 100 395 Unnamed 37 30 100 318 Notogaeanura 51 24 100 396 Unnamed 23 14 99 319 Australobatrachia 62 54 100 397 Unnamed 32 24 100 320 Batrachophrynidae 93 86 100 398 Unnamed 82 69 100 321 Myobatrachoidea 48 31 100 399 Unnamed 64 48 100 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 329

Branch Bremer Branch Bremer Branch Taxon length support Jackknife Branch Taxon length support Jackknife

400 Dendropsophus 69 55 100 464 Unnamed 56 32 100 401 Unnamed 49 25 100 465 Unnamed 39 24 100 402 Unnamed 53 37 100 466 Unnamed 53 29 100 403 Unnamed 29 22 100 467 Unnamed 47 35 100 404 Lophiohylini 65 42 100 468 Unnamed 68 45 100 405 Unnamed 23 14 90 469 Bufonidae 51 33 99 406 Osteocephalus 18 8 87 470 Unnamed 83 38 99 407 Unnamed 22 12 91 471 Unnamed 39 22 100 408 Trachycephalus 45 25 100 472 Atelopus 54 43 100 409 Hylini 85 78 100 473 Unnamed 114 114 100 410 Unnamed 29 15 100 474 Unnamed 40 25 100 411 Unnamed 43 32 100 475 Unnamed 58 20 100 412 Unnamed 27 19 99 476 Rhaebo 51 42 100 413 Unnamed 16 4 75 477 Unnamed 43 20 100 414 Unnamed 24 9 96 478 Unnamed 30 25 100 415 Unnamed 23 5 89 479 Unnamed 30 14 98 416 Unnamed 42 32 100 480 Nectophryne 134 107 100 417 Pseudacris 54 37 100 481 Unnamed 15 9 97 418 Unnamed 54 5 100 482 Unnamed 37 18 100 419 Unnamed 34 19 99 483 Unnamed 12 8 97 420 Unnamed 15 5 85 484 Unnamed 20 9 95 421 Unnamed 44 30 100 485 Unnamed 27 10 94 422 Unnamed 23 14 99 486 Unnamed 24 9 94 423 Unnamed 45 28 100 487 Ansonia 25 9 90 424 Leptodactyliformes 24 17 100 488 Unnamed 13 8 93 425 Diphyabatrachia 35 29 98 489 Unnamed 9 6 94 426 Centrolenidae 41 12 99 490 Unnamed 14 6 95 427 Centroleninae 67 22 100 491 Ingerophrynus 16 10 99 428 Unnamed 23 13 99 492 Unnamed 15 11 100 429 Unnamed 20 11 99 493 Unnamed 30 14 99 430 Leptodactylidae 30 23 99 494 Unnamed 12 8 92 431 Unnamed 50 41 98 495 Unnamed 18 12 99 432 Unnamed 70 47 100 496 Unnamed 31 16 99 433 Unnamed 26 10 97 497 Unnamed 35 15 99 434 Unnamed 30 15 99 498 Unnamed 15 10 98 435 Unnamed 65 40 100 499 Bufo (sensu stricto) 59 28 100 436 Leptodactylus 61 47 100 500 Unnamed 6 3 90 437 Unnamed 64 42 100 501 Unnamed 18 5 71 438 Unnamed 46 34 100 502 Unnamed 12 9 94 439 Unnamed 64 31 100 503 Capensibufo 11 10 99 440 Chthonobatrachia 18 13 100 504 Unnamed 10 7 94 441 Ceratophryidae 34 18 98 505 Unnamed 7 2 61 442 Telmatobiinae 70 62 100 506 Amietophrynus 9265 443 Unnamed 26 18 100 507 Unnamed 36 2 65 444 Ceratophryinae 22 2 62 508 Unnamed 22 15 99 445 Batrachylini 54 37 100 509 Unnamed 22 17 99 446 Ceratophryini 46 38 100 510 Unnamed 30 22 99 447 Unnamed 24 8 94 511 Unnamed 100 98 100 448 Hesticobatrachia 34 26 98 512 Unnamed 25 27 94 449 Cycloramphidae 21 9 98 513 Anaxyrus 22 15 99 450 Hylodinae 70 5 91 514 Unnamed 44 33 100 451 Unnamed 71 43 100 515 Unnamed 23 8 90 452 Cycloramphinae 19 9 82 516 Unnamed 28 21 100 453 Cycloramphini 42 30 96 517 Unnamed 30 26 100 454 Alsodini 28 4 80 518 Unnamed 13 6 81 455 Unnamed 8 4 81 519 Cranopsis 25 11 99 456 Unnamed 19 14 99 520 Unnamed 14 1 58 457 Unnamed 44 10 100 521 Unnamed 16 11 99 458 Unnamed 76 52 100 522 Chaunus 22 16 98 459 Odontophrynus 48 38 100 523 Unnamed 13 7 78 460 Agastorophrynia 39 30 98 524 Unnamed 14 9 80 461 Dendrobatoidea 51 39 100 525 Unnamed 13 6 75 462 Dendrobatidae 74 61 100 526 Unnamed 55 51 100 463 Unnamed 66 51 100 527 Unnamed 21 17 99 330 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

APPENDIX 5 DNA SEQUENCE TRANSFORMATIONS FOR SELECTED BRANCHES/TAXA Evidence is presented for taxa recognized solely on the basis of DNA sequence transformations, or taxa whose molecular evidence was speci®cally noted in the text. The table is organized by branch number, with those taxa lacking branch numbers following in alphabetical order. Optimization ambiguous transformations are excluded. Locus abbreviations are 28S (large nuclear ribosomal subunit), H1 (mitochondrial transcription unit H1), H3 (histone H3), rhod (rhodopsin exon 1), SIA (seven in absentia), and tyr (tyrosinase). Other abbreviations are Br/ Taxon/Frag (branch, taxon, and DNA fragment), Pos (position in aligned sequence), Anc (ancestral character), Der (derived character), A (adenine), C (cytosine), G (guanine), T (thymine), and ``Ð'' (gap).

Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der

29 (Diadectosalamandroidei) H1 frag. 23 743 Ð T H1 frag. 11 264 C T H1 frag. 23 1130 Ð G 28S frag. 4 71 T C H1 frag. 23 956 Ð A H1 frag. 11 822 A Ð H1 frag. 23 1726 A C H1 frag. 10 7 G C H1 frag. 23 1051 Ð A H1 frag. 11 1101 Ð C H1 frag. 6 89 Ð C H1 frag. 10 23 T A H1 frag. 23 1090 Ð C H1 frag. 12 41 A T H1 frag. 8 57 C T H1 frag. 11 315 A Ð H1 frag. 23 1169 G A H1 frag. 13 71 C A H1 frag. 8 316 T C H1 frag. 11 409 T Ð H1 frag. 23 1328 G A H1 frag. 13 130 C A H1 frag. 8 335 T C H1 frag. 13 198 C A H1 frag. 23 1750 T Ð H1 frag. 14 31 T A H1 frag. 8 628 T C H1 frag. 14 143 C T H1 frag. 4 205 Ð T H1 frag. 14 89 T A H1 frag. 8 790 T C H1 frag. 14 217 T A H1 frag. 4 640 Ð A H1 frag. 14 106 A C H1 frag. 9 89 A G H1 frag. 16 94 A G H1 frag. 6 23 T C H1 frag. 15 25 G A H1 frag. 9 409 T C H1 frag. 16 313 A Ð H1 frag. 6 75 A Ð H1 frag. 16 152 A T H3 frag. 1 66 C G H1 frag. 17 47 C A H1 frag. 7 35 C A H1 frag. 16 429 C A H3 frag. 1 81 C A H1 frag. 17 210 C T H1 frag. 8 156 C Ð H1 frag. 17 47 A T 46 (unnamed taxon) H1 frag. 18 276 A C H1 frag. 8 628 A T H1 frag. 17 182 T A H1 frag. 18 479 C A H1 frag. 18 349 C A H1 frag. 8 796 T A H1 frag. 18 79 A G H1 frag. 18 536 Ð A H1 frag. 19 136 G A SIA frag. 1 9 A T H1 frag. 18 116 T C H1 frag. 18 732 A T H1 frag. 19 195 C A SIA frag. 1 12 T C H1 frag. 18 366 Ð C H1 frag. 19 42 A C H1 frag. 19 331 G T SIA frag. 2 20 C T H1 frag. 18 756 T C H1 frag. 19 254 G Ð H1 frag. 19 376 G Ð 31 (Perennibranchia) H1 frag. 19 259 A Ð H1 frag. 19 278 C A H1 frag. 19 509 A Ð 28S frag. 2 312 C T H1 frag. 19 749 A T H1 frag. 19 331 T A H1 frag. 19 531 A Ð H1 frag. 10 199 C T H1 frag. 2 342 A T H1 frag. 19 431 C A H1 frag. 2 218 C A H1 frag. 11 36 T A H1 frag. 2 407 A C H1 frag. 19 612 T C H1 frag. 2 277 A Ð H1 frag. 11 72 C T H1 frag. 20 146 T A H1 frag. 19 796 C T H1 frag. 2 285 A Ð H1 frag. 11 249 Ð G H1 frag. 21 93 C T H1 frag. 19 808 C T H1 frag. 20 78 G A H1 frag. 11 819 T C H1 frag. 23 67 G A H1 frag. 20 54 A T H1 frag. 21 124 C Ð H1 frag. 11 1327 T Ð H1 frag. 23 670 Ð T H1 frag. 20 176 A G H1 frag. 22 70 A T H1 frag. 14 93 A T H1 frag. 23 1676 A Ð H1 frag. 20 182 T C H1 frag. 23 33 C Ð H1 frag. 14 166 C A H1 frag. 23 1707 A T H1 frag. 21 57 A C H1 frag. 23 260 C T H1 frag. 14 244 G A H1 frag. 3 169 T A H1 frag. 21 239 Ð T H1 frag. 23 283 C Ð H1 frag. 15 60 T A H1 frag. 4 64 T C H1 frag. 23 22 T C H1 frag. 23 981 T A H1 frag. 16 94 G Ð H1 frag. 4 262 A G H1 frag. 23 293 C T H1 frag. 23 1114 A T H1 frag. 16 127 T Ð H1 frag. 4 263 G A H1 frag. 23 668 C A H1 frag. 23 1338 C A H1 frag. 16 170 T Ð H1 frag. 4 458 A C H1 frag. 23 1346 A C H1 frag. 23 1405 A Ð H1 frag. 16 191 T Ð H1 frag. 4 487 Ð T H1 frag. 6 31 C T H1 frag. 23 1684 T Ð H1 frag. 16 221 A C H1 frag. 9 288 C T H1 frag. 7 11 G A H1 frag. 23 1687 T Ð H1 frag. 16 563 T G H1 frag. 9 763 T A H1 frag. 8 250 A T H1 frag. 23 1940 T Ð H1 frag. 18 488 C T H3 frag. 1 192 C G H1 frag. 8 628 C T H1 frag. 3 415 A T H1 frag. 18 628 T Ð SIA frag. 1 3 T C H1 frag. 8 675 Ð T H1 frag. 4 169 C A H1 frag. 19 131 A T SIA frag. 3 66 A G H1 frag. 8 735 T C H1 frag. 8 159 G C H1 frag. 20 25 A T SIA frag. 3 72 C T 49 (Plethosalamandroidei) H1 frag. 8 250 T A H1 frag. 21 76 T C SIA frag. 3 147 A G H1 frag. 10 24 A G H1 frag. 9 84 C A H1 frag. 23 43 T A 41 (Pleurodelinae) H1 frag. 10 101 G A H1 frag. 9 85 C A H1 frag. 23 1015 A T H1 frag. 10 101 G A H1 frag. 11 57 A T H3 frag. 2 26 T C H1 frag. 23 1118 Ð A H1 frag. 11 89 C T H1 frag. 11 89 C T 30 (Hydatinosalamandroidei) H1 frag. 23 1303 A T H1 frag. 11 264 T Ð H1 frag. 11 1191 T A H1 frag. 11 31 T C H1 frag. 23 1759 A T H1 frag. 11 1191 T A H1 frag. 12 52 T A H1 frag. 11 595 A C H1 frag. 23 1763 T A H1 frag. 11 1327 T Ð H1 frag. 13 172 C T H1 frag. 11 694 C Ð H1 frag. 23 1962 A T H1 frag. 12 6 A G H1 frag. 14 124 T C H1 frag. 11 1294 T A H1 frag. 8 28 A C H1 frag. 12 90 T A H1 frag. 15 58 A G H1 frag. 13 91 A Ð H1 frag. 8 57 C T H1 frag. 14 208 A C H1 frag. 16 4 C T H1 frag. 14 45 C T H1 frag. 8 69 A T H1 frag. 16 4 C T H1 frag. 16 681 T A H1 frag. 14 144 G A H1 frag. 8 110 C T H1 frag. 17 22 T C H1 frag. 17 46 A T H1 frag. 14 182 Ð T H1 frag. 8 589 C T H1 frag. 17 38 T C H1 frag. 17 160 T A H1 frag. 15 22 C T H1 frag. 8 634 T A H1 frag. 17 136 A T H1 frag. 18 479 A T H1 frag. 16 466 Ð T H1 frag. 8 714 A G H1 frag. 18 45 A Ð H1 frag. 18 750 G A H1 frag. 17 38 A T H1 frag. 9 54 T C H1 frag. 18 479 A C H1 frag. 18 854 T Ð H1 frag. 17 133 Ð A H1 frag. 9 73 C A H1 frag. 18 741 C Ð H1 frag. 19 417 T A H1 frag. 18 838 A T H1 frag. 9 333 A C H1 frag. 18 750 G A H1 frag. 19 432 Ð C H1 frag. 18 878 A T H3 frag. 1 48 G A H1 frag. 18 759 A T H1 frag. 20 140 A T H1 frag. 19 294 A Ð H3 frag. 1 103 C A H1 frag. 18 854 T A H1 frag. 21 134 G Ð H1 frag. 19 369 A T H3 frag. 1 135 C G H1 frag. 19 729 T C H1 frag. 21 177 C A H1 frag. 19 453 A Ð H3 frag. 1 204 C A H1 frag. 21 50 Ð A H1 frag. 21 260 T C H1 frag. 19 496 C Ð 35 (Treptobranchia) H1 frag. 21 260 T A H1 frag. 23 49 T A H1 frag. 19 668 A Ð 28S frag. 4 150 T C H1 frag. 22 11 T C H1 frag. 23 102 A T H1 frag. 2 73 T A H1 frag. 11 141 A T H1 frag. 23 293 T C H1 frag. 23 182 T Ð 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 331

Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der

H1 frag. 23 521 A Ð H1 frag. 23 1816 A G H1 frag. 1 40 T A H1 frag. 2 133 C T H1 frag. 23 623 A Ð H1 frag. 23 1833 Ð A H1 frag. 1 41 A G H1 frag. 2 210 G A H1 frag. 23 789 A Ð 86 (Alytidae) H1 frag. 1 64 A G H1 frag. 2 301 C A H1 frag. 23 902 C Ð 28S frag. 2 753 C Ð H1 frag. 10 72 A T H1 frag. 2 407 A C H1 frag. 23 951 G Ð 28S frag. 2 764 A T H1 frag. 10 261 T A H1 frag. 20 61 A G H1 frag. 23 953 G Ð 28S frag. 3 217 C G H1 frag. 11 10 A Ð H1 frag. 21 44 Ð A H1 frag. 23 1108 A Ð 28S frag. 3 424 G C H1 frag. 11 17 Ð G H1 frag. 21 177 C A H1 frag. 23 1124 A Ð 28S frag. 3 582 G C H1 frag. 11 31 T C H1 frag. 21 178 C A H1 frag. 23 1158 T Ð H1 frag. 11 409 T A H1 frag. 11 47 T C H1 frag. 21 251 A G H1 frag. 23 1173 A Ð H1 frag. 11 457 C A H1 frag. 11 67 C A H1 frag. 23 49 T C H1 frag. 23 1657 T A H1 frag. 11 565 C A H1 frag. 11 79 A G H1 frag. 23 100 T A H1 frag. 23 1766 T A H1 frag. 11 694 C Ð H1 frag. 11 88 T C H1 frag. 23 102 T A H1 frag. 6 77 Ð T H1 frag. 11 983 T A H1 frag. 11 141 A T H1 frag. 23 105 A C H1 frag. 8 184 T C H1 frag. 11 1161 A C H1 frag. 11 213 C T H1 frag. 23 236 A G H1 frag. 8 331 Ð C H1 frag. 11 1217 A C H1 frag. 11 230 C T H1 frag. 23 283 C T H1 frag. 8 345 G A H1 frag. 13 121 A T H1 frag. 11 778 A Ð H1 frag. 23 981 T A H1 frag. 8 369 A T H1 frag. 14 35 A C H1 frag. 11 910 C A H1 frag. 23 1097 G Ð H1 frag. 8 562 G A H1 frag. 14 166 C Ð H1 frag. 11 953 C G H1 frag. 23 1150 Ð T H1 frag. 8 634 T C H1 frag. 16 7 A C H1 frag. 11 1135 C Ð H1 frag. 23 1169 G A H1 frag. 9 209 T Ð H1 frag. 16 46 Ð T H1 frag. 11 1316 A T H1 frag. 23 1181 C G H1 frag. 9 520 A Ð H1 frag. 16 429 C Ð H1 frag. 12 4 T A H1 frag. 23 1270 T C H1 frag. 9 672 A T H1 frag. 17 56 Ð T H1 frag. 12 29 A C H1 frag. 23 1607 C A rhod frag. 1 94 G A H1 frag. 17 231 T A H1 frag. 12 52 G A H1 frag. 23 1695 A G rhod frag. 1 151 C T H1 frag. 17 320 Ð G H1 frag. 12 122 G A H1 frag. 23 1722 A C rhod frag. 2 93 C G H1 frag. 18 306 A C H1 frag. 14 9 G A H1 frag. 23 1745 C T 50 (Xenosalamandroidei) H1 frag. 18 447 T C H1 frag. 14 93 A C H1 frag. 23 1759 T C H1 frag. 11 40 G Ð H1 frag. 18 746 T A H1 frag. 14 100 T C H1 frag. 23 1824 T Ð H1 frag. 11 77 C T H1 frag. 19 91 C Ð H1 frag. 14 144 G T H1 frag. 23 1940 T A H1 frag. 11 213 C Ð H1 frag. 19 203 A C H1 frag. 14 146 A G H1 frag. 24 1 C T H1 frag. 11 577 Ð C H1 frag. 19 415 A C H1 frag. 14 149 G A H1 frag. 24 10 A C H1 frag. 11 1217 A C H1 frag. 19 439 A G H1 frag. 14 208 A C H1 frag. 24 17 C T H1 frag. 11 1311 C Ð H1 frag. 19 771 A C H1 frag. 15 19 C T H1 frag. 24 35 G A H1 frag. 12 59 A T H1 frag. 20 20 T Ð H1 frag. 15 22 C A H1 frag. 25 16 A C H1 frag. 12 74 A T H1 frag. 21 10 T C H1 frag. 15 40 C T H1 frag. 25 24 T C H1 frag. 13 70 T C H1 frag. 21 57 A T H1 frag. 16 5 A G H1 frag. 25 38 A G H1 frag. 14 250 C A H1 frag. 21 218 C T H1 frag. 16 18 T C H1 frag. 3 48 A T H1 frag. 15 15 T A H1 frag. 23 52 C T H1 frag. 16 23 G A H1 frag. 3 58 C T H1 frag. 15 23 T A H1 frag. 23 199 A C H1 frag. 16 94 T Ð H1 frag. 3 160 T C H1 frag. 15 25 G A H1 frag. 23 452 C Ð H1 frag. 16 127 T Ð H1 frag. 3 214 T C H1 frag. 15 60 T A H1 frag. 23 942 A T H1 frag. 16 241 A G H1 frag. 3 251 C T H1 frag. 16 5 A G H1 frag. 23 1154 A G H1 frag. 16 509 A G H1 frag. 3 283 A G H1 frag. 16 39 T A H1 frag. 23 1186 Ð A H1 frag. 16 535 A C H1 frag. 3 384 T C H1 frag. 16 201 T A H1 frag. 23 1256 T C H1 frag. 16 576 A C H1 frag. 3 391 A G H1 frag. 16 359 T C H1 frag. 23 1376 T C H1 frag. 16 590 A C H1 frag. 3 402 C A H1 frag. 16 672 A T H1 frag. 23 1496 Ð C H1 frag. 16 648 A G H1 frag. 3 407 C A H1 frag. 17 221 Ð A H1 frag. 23 1919 C Ð H1 frag. 16 696 G A H1 frag. 4 8 C A H1 frag. 18 322 A C H1 frag. 23 1962 A T H1 frag. 17 2 C T H1 frag. 4 26 A G H1 frag. 18 488 C T H1 frag. 25 15 C T H1 frag. 17 60 G A H1 frag. 4 169 C A H1 frag. 18 748 T C H1 frag. 25 86 G A H1 frag. 17 125 T C H1 frag. 4 260 T C H1 frag. 18 802 Ð A H1 frag. 6 49 T A H1 frag. 17 187 A T H1 frag. 4 268 T C H1 frag. 19 15 A G H1 frag. 6 81 A C H1 frag. 17 372 T C H1 frag. 4 283 T A H1 frag. 19 97 T C H1 frag. 6 183 Ð T H1 frag. 18 164 T Ð H1 frag. 4 286 A G H1 frag. 19 99 T C H1 frag. 8 132 T A H1 frag. 18 185 C Ð H1 frag. 4 298 A T H1 frag. 19 460 T G H1 frag. 8 634 T C H1 frag. 18 380 Ð A H1 frag. 4 363 T A H1 frag. 19 573 G T H1 frag. 9 520 A Ð H1 frag. 18 433 A C H1 frag. 4 365 T C H1 frag. 19 635 T Ð H1 frag. 9 768 Ð A H1 frag. 18 501 A T H1 frag. 4 386 T A H1 frag. 20 60 A G H3 frag. 1 51 T A H1 frag. 18 579 Ð T H1 frag. 4 404 A G H1 frag. 20 146 T A H3 frag. 1 81 A C H1 frag. 18 604 A T H1 frag. 4 408 A C H1 frag. 20 176 A G H3 frag. 1 114 T C H1 frag. 18 656 A Ð H1 frag. 4 434 A G H1 frag. 21 17 Ð A H3 frag. 1 126 G T H1 frag. 18 717 A C H1 frag. 4 439 C A H1 frag. 21 243 T C H3 frag. 2 39 G T H1 frag. 18 748 T C H1 frag. 4 452 A Ð H1 frag. 23 167 A C H3 frag. 2 42 C G H1 frag. 18 766 C T H1 frag. 4 467 G T H1 frag. 23 293 T C rhod frag. 1 90 T C H1 frag. 18 830 C A H1 frag. 4 481 A G H1 frag. 23 965 Ð T rhod frag. 1 99 T C H1 frag. 18 863 G A H1 frag. 4 489 T C H1 frag. 23 1081 A G rhod frag. 1 101 G A H1 frag. 18 866 C A H1 frag. 4 517 A G H1 frag. 23 1703 T C rhod frag. 2 3 A G H1 frag. 18 883 C T H1 frag. 4 592 T C H1 frag. 23 1752 A Ð rhod frag. 2 69 T C H1 frag. 19 109 C T H1 frag. 4 637 T C H1 frag. 23 1796 T Ð SIA frag. 4 7 T C H1 frag. 19 244 A C H1 frag. 4 649 A C H1 frag. 25 34 C A SIA frag. 4 61 T A H1 frag. 19 250 Ð G H1 frag. 6 35 T C H1 frag. 6 27 G A 88 (Bombinatoridae/Bombina) H1 frag. 19 259 A C H1 frag. 6 52 T A H1 frag. 6 91 T A 28S frag. 2 132 T C H1 frag. 19 283 Ð T H1 frag. 6 75 A C H1 frag. 8 316 T A 28S frag. 2 467 Ð T H1 frag. 19 313 Ð C H1 frag. 6 85 T A H1 frag. 8 598 A Ð 28S frag. 2 711 Ð G H1 frag. 19 350 A G H1 frag. 6 137 C A H1 frag. 8 667 C T 28S frag. 2 763 Ð A H1 frag. 19 449 T A H1 frag. 6 176 Ð C 72 (Pseudoeurycea) 28S frag. 3 370 G C H1 frag. 19 531 A T H1 frag. 7 83 A Ð H1 frag. 21 57 A C 28S frag. 3 420 Ð A H1 frag. 19 823 C A H1 frag. 7 84 A Ð H1 frag. 23 49 A G 28S frag. 3 535 C T H1 frag. 19 826 A T H1 frag. 8 316 T C H1 frag. 23 1376 T C 28S frag. 4 83 T C H1 frag. 2 79 Ð G H1 frag. 8 582 T C H1 frag. 23 1776 C Ð H1 frag. 1 18 T C H1 frag. 2 88 A C H1 frag. 8 628 A C H1 frag. 23 1798 T Ð H1 frag. 1 20 A T H1 frag. 2 108 C T H1 frag. 8 644 A G 332 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der

H1 frag. 8 647 A C H1 frag. 3 248 G A H1 frag. 4 120 T C H1 frag. 9 226 A T H1 frag. 8 711 G A H1 frag. 3 266 C T H1 frag. 4 232 C T H1 frag. 9 788 A C H1 frag. 8 735 C Ð H1 frag. 3 366 C T H1 frag. 4 268 T C H3 frag. 1 108 C A H1 frag. 8 787 C Ð H1 frag. 3 391 A G H1 frag. 4 286 A G H3 frag. 1 147 A G H1 frag. 8 792 C T H1 frag. 3 402 C T H1 frag. 4 481 A G rhod frag. 1 33 G A H1 frag. 8 807 Ð G H1 frag. 3 410 A T H1 frag. 6 164 Ð A rhod frag. 1 168 C T H1 frag. 8 828 T A H1 frag. 4 280 C T H1 frag. 8 316 T C rhod frag. 1 169 A G H1 frag. 9 42 G T H1 frag. 4 375 Ð C H1 frag. 8 488 C T rhod frag. 1 174 G A H1 frag. 9 281 T C H1 frag. 4 391 A C H1 frag. 9 652 T C rhod frag. 2 106 A T H1 frag. 9 453 A T H1 frag. 6 137 C A H1 frag. 9 775 A T 97 (Pelobatidae) H3 frag. 1 84 G C H1 frag. 7 35 C G rhod frag. 1 69 C T H1 frag. 10 174 Ð C H3 frag. 1 222 C T H1 frag. 8 306 C T SIA frag. 3 48 G T H1 frag. 11 53 T G H3 frag. 2 5 T G H1 frag. 8 629 Ð T SIA frag. 3 66 G A H1 frag. 11 134 A C H3 frag. 2 33 G A H1 frag. 9 453 A T SIA frag. 3 153 A G H1 frag. 11 141 A T H3 frag. 2 63 C T H3 frag. 1 18 G C 96 (Pelobatoidea) H1 frag. 11 409 T C H3 frag. 2 66 C T H3 frag. 1 72 C T 28S frag. 2 589 Ð C H1 frag. 11 517 Ð C rhod frag. 1 78 G A H3 frag. 1 99 C T 28S frag. 2 590 Ð G H1 frag. 11 795 Ð C rhod frag. 1 104 T A H3 frag. 1 138 C T 28S frag. 2 721 Ð C H1 frag. 11 901 Ð A rhod frag. 1 110 C T H3 frag. 1 195 G T H1 frag. 1 2 T C H1 frag. 11 991 Ð T rhod frag. 1 122 C T H3 frag. 1 204 C T H1 frag. 10 72 A Ð H1 frag. 11 1081 Ð A rhod frag. 1 131 T C H3 frag. 2 29 T C H1 frag. 10 76 A Ð H1 frag. 11 1161 A T rhod frag. 1 134 G C rhod frag. 1 78 G A H1 frag. 11 116 Ð C H1 frag. 11 1266 A T rhod frag. 1 135 T C rhod frag. 1 120 C T H1 frag. 11 138 A C H1 frag. 11 1311 C A rhod frag. 1 137 C T rhod frag. 1 122 C A H1 frag. 11 565 C T H1 frag. 11 1333 T C rhod frag. 1 150 C T rhod frag. 1 153 G A H1 frag. 11 988 Ð C H1 frag. 12 41 A G rhod frag. 2 21 C T rhod frag. 2 9 A G H1 frag. 11 1217 A Ð H1 frag. 22 70 A G rhod frag. 2 31 C T rhod frag. 2 33 G C H1 frag. 11 1248 C Ð H1 frag. 23 18 Ð A rhod frag. 2 54 C T rhod frag. 2 73 G T H1 frag. 13 168 C A H1 frag. 23 25 A Ð rhod frag. 2 67 T G SIA frag. 1 12 T C H1 frag. 14 51 A G H1 frag. 23 50 C T rhod frag. 2 112 C T SIA frag. 1 36 T C H1 frag. 14 102 T C H1 frag. 23 72 C T 92 (Anomocoela) SIA frag. 2 14 A G H1 frag. 14 129 C T H1 frag. 23 89 G A H1 frag. 11 622 C Ð SIA frag. 3 43 T A H1 frag. 14 217 A T H1 frag. 23 490 T C H1 frag. 11 897 Ð T SIA frag. 3 44 C G H1 frag. 15 54 G A H1 frag. 23 559 T A H1 frag. 11 898 Ð C SIA frag. 3 171 G C H1 frag. 16 32 C T H1 frag. 23 602 T A H1 frag. 11 1076 Ð T SIA frag. 4 49 T G H1 frag. 16 319 Ð A H1 frag. 23 762 A T H1 frag. 11 1294 T C SIA frag. 4 67 T C H1 frag. 16 429 C T H1 frag. 23 1015 A C H1 frag. 13 69 C A 93 (Pelodytoidea) H1 frag. 17 52 A T H1 frag. 23 1161 Ð A H1 frag. 14 154 G Ð H1 frag. 17 231 C Ð 28S frag. 2 442 C Ð H1 frag. 23 1181 C A H1 frag. 14 166 C A 28S frag. 2 613 C Ð H1 frag. 18 209 A Ð H1 frag. 23 1316 C T H1 frag. 14 208 A T 28S frag. 2 714 G Ð H1 frag. 18 615 C T H1 frag. 23 1711 A C H1 frag. 14 250 C T 28S frag. 2 753 C Ð H1 frag. 18 727 C T H1 frag. 23 1793 A C H1 frag. 16 59 Ð G 28S frag. 2 764 A Ð H1 frag. 19 66 A T H1 frag. 23 1799 Ð C H1 frag. 16 292 A G H1 frag. 11 16 A T H1 frag. 19 376 G C H1 frag. 23 1908 Ð A H1 frag. 16 485 G C H1 frag. 11 230 C T H1 frag. 19 531 A C H1 frag. 25 15 C T H1 frag. 16 590 A C H1 frag. 12 116 G A H1 frag. 2 108 C T H1 frag. 25 86 G A H1 frag. 16 668 Ð C H1 frag. 14 106 A C H1 frag. 2 360 T Ð H1 frag. 6 84 Ð C H1 frag. 17 5 T A H1 frag. 14 142 C T H1 frag. 21 133 A T H1 frag. 6 199 C T H1 frag. 17 46 A C H1 frag. 14 146 A G H1 frag. 21 219 Ð T H1 frag. 6 213 A C H1 frag. 17 407 T A H1 frag. 14 260 G T H1 frag. 23 105 A T H1 frag. 8 28 A C H1 frag. 18 7 C T H1 frag. 15 21 T C H1 frag. 23 789 A T H1 frag. 18 388 C T H1 frag. 16 31 A G H1 frag. 23 1221 C T H1 frag. 8 41 A T H1 frag. 18 784 Ð C H1 frag. 16 94 T Ð H1 frag. 23 1478 G Ð H1 frag. 8 69 C T H1 frag. 18 838 A Ð H1 frag. 16 127 T A H1 frag. 23 1518 A Ð H1 frag. 8 179 C T H1 frag. 19 5 Ð C H1 frag. 16 201 T A H1 frag. 23 1607 C Ð H1 frag. 8 192 G A H1 frag. 19 109 C T H1 frag. 16 382 A C H1 frag. 23 1657 C Ð H1 frag. 8 565 A T H1 frag. 19 249 Ð A H1 frag. 16 509 A Ð H1 frag. 23 1676 C Ð H1 frag. 8 667 C T H1 frag. 19 278 C T H1 frag. 16 690 A T H1 frag. 23 1787 T Ð H1 frag. 8 828 T A H1 frag. 19 668 C T H1 frag. 17 437 C A H1 frag. 23 1962 A Ð H1 frag. 9 46 T C H1 frag. 19 808 C T H1 frag. 18 95 A C H1 frag. 3 58 C A H1 frag. 9 202 C Ð H1 frag. 2 73 T C H1 frag. 18 138 A Ð H1 frag. 3 108 A C H1 frag. 9 256 C T H1 frag. 2 256 A C H1 frag. 18 315 Ð A H1 frag. 3 214 T A H1 frag. 9 367 A T H1 frag. 2 301 C Ð H1 frag. 18 545 Ð C H1 frag. 4 26 A T H1 frag. 9 618 A C H1 frag. 20 68 G A H1 frag. 18 830 C T H1 frag. 4 148 A T H1 frag. 9 781 Ð C H1 frag. 20 146 T A H1 frag. 19 273 Ð T H1 frag. 4 211 A T H1 frag. 9 782 Ð C H1 frag. 20 176 A G H1 frag. 19 274 Ð T H1 frag. 4 263 G A rhod frag. 1 6 T C H1 frag. 21 113 A T H1 frag. 19 600 C T H1 frag. 4 271 T A rhod frag. 1 94 G A H1 frag. 23 250 G T H1 frag. 2 100 A C H1 frag. 4 441 A C rhod frag. 1 134 G C H1 frag. 23 606 Ð C H1 frag. 2 203 A G H1 frag. 4 493 C T rhod frag. 1 140 A T H1 frag. 23 737 Ð T H1 frag. 20 16 T A H1 frag. 6 66 C Ð rhod frag. 1 159 G A H1 frag. 23 762 C A H1 frag. 20 77 G A H1 frag. 6 167 A T rhod frag. 2 6 C T H1 frag. 23 1097 G C H1 frag. 23 25 A T H1 frag. 6 181 A C rhod frag. 2 24 C T H1 frag. 23 1160 Ð G H1 frag. 23 38 C T H1 frag. 7 81 C T rhod frag. 2 37 C T H1 frag. 23 1256 T A H1 frag. 23 96 A T H1 frag. 7 84 A C rhod frag. 2 41 C T H1 frag. 23 1338 C A H1 frag. 23 902 C T H1 frag. 8 232 A C rhod frag. 2 54 C T H1 frag. 23 1358 A T H1 frag. 23 963 A Ð H1 frag. 8 628 A T rhod frag. 2 66 G C H1 frag. 23 1444 G Ð H1 frag. 23 1019 Ð T H1 frag. 8 696 A G rhod frag. 2 100 G T H1 frag. 23 1460 G T H1 frag. 23 1169 G T H1 frag. 8 711 G A rhod frag. 2 101 C A H1 frag. 23 1669 C T H1 frag. 23 1334 A T H1 frag. 8 804 A G rhod frag. 2 115 C T H1 frag. 23 1951 A T H1 frag. 3 92 C T H1 frag. 8 818 C T rhod frag. 2 136 T C H1 frag. 3 120 T A H1 frag. 3 117 G A H1 frag. 9 89 G A rhod frag. 2 139 G A 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 333

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99 (unnamed taxon) H1 frag. 12 125 A T 102 (unnamed taxon) H1 frag. 23 1962 A T H1 frag. 10 93 A G H1 frag. 13 18 G T H1 frag. 21 57 A T H1 frag. 3 124 T A H1 frag. 11 16 A T H1 frag. 13 172 C A H1 frag. 21 113 T C H1 frag. 3 342 G T H1 frag. 11 77 C T H1 frag. 14 83 A T H1 frag. 21 133 T C H1 frag. 4 308 Ð G H1 frag. 11 86 A G H1 frag. 14 108 C T H1 frag. 21 190 A G H1 frag. 4 391 A C H1 frag. 11 95 T C H1 frag. 14 193 T C H1 frag. 21 277 C A H1 frag. 4 673 T A H1 frag. 11 116 C A H1 frag. 14 250 T A H1 frag. 22 55 T C H1 frag. 6 46 C A H1 frag. 11 124 C Ð H1 frag. 16 54 Ð G H1 frag. 23 5 T C H1 frag. 6 81 A C H1 frag. 11 264 C T H1 frag. 16 292 G T H1 frag. 23 17 G A H1 frag. 7 76 C T H1 frag. 11 315 A G H1 frag. 16 319 A T H1 frag. 23 25 A G H1 frag. 7 83 A Ð H1 frag. 11 392 C Ð H1 frag. 16 333 A T H1 frag. 23 40 T C H1 frag. 7 84 A Ð H1 frag. 11 852 C T H1 frag. 16 648 A C H1 frag. 23 67 G A H1 frag. 8 162 C A H1 frag. 12 112 G A H1 frag. 17 46 C A H1 frag. 23 114 A T H1 frag. 8 179 C T H1 frag. 13 39 A Ð H1 frag. 17 52 T C H1 frag. 23 195 Ð C H1 frag. 8 192 G A H1 frag. 13 73 A Ð H1 frag. 17 120 T Ð H1 frag. 23 224 A T H1 frag. 8 352 Ð A H1 frag. 13 107 G A H1 frag. 17 122 A Ð H1 frag. 23 260 C T H1 frag. 8 369 G T H1 frag. 14 13 T A H1 frag. 17 136 A Ð H1 frag. 23 283 C T H1 frag. 8 488 C A H1 frag. 14 64 A T H1 frag. 17 160 A Ð H1 frag. 23 606 C T H1 frag. 8 544 A C H1 frag. 16 1 A G H1 frag. 17 187 A Ð H1 frag. 23 762 A C H1 frag. 8 580 C A H1 frag. 16 7 A G H1 frag. 17 210 T C H1 frag. 23 854 C T H1 frag. 8 647 A C H1 frag. 16 127 T C H1 frag. 17 251 T C H1 frag. 23 1027 A C H1 frag. 8 726 A T H1 frag. 16 359 C T H1 frag. 18 138 A Ð H1 frag. 23 1074 T C H1 frag. 8 822 Ð A H1 frag. 16 563 C Ð H1 frag. 18 218 C T H1 frag. 23 1081 A T H1 frag. 9 73 C A H1 frag. 16 576 C T H1 frag. 18 433 A C H1 frag. 23 1114 A T H1 frag. 9 281 T A H1 frag. 16 658 C T H1 frag. 18 501 A C H1 frag. 23 1135 T Ð H1 frag. 9 633 C Ð H1 frag. 17 47 C T H1 frag. 18 539 A Ð H1 frag. 23 1160 T Ð rhod frag. 1 A C H1 frag. 17 182 T A H1 frag. 18 656 A G H1 frag. 23 1169 G Ð rhod frag. 1 3 T C H1 frag. 17 416 Ð G H1 frag. 18 698 A G H1 frag. 23 1201 G A rhod frag. 1 12 T C H1 frag. 18 52 A C H1 frag. 18 707 T G H1 frag. 23 1226 T A rhod frag. 1 21 C T H1 frag. 18 164 T C H1 frag. 19 109 T Ð H1 frag. 23 1256 A T rhod frag. 1 107 T C H1 frag. 18 276 A C H1 frag. 19 123 A C H1 frag. 23 1458 Ð T rhod frag. 1 117 T A H1 frag. 18 581 C T H1 frag. 19 286 C A H1 frag. 23 1687 A T rhod frag. 1 157 T C H1 frag. 18 746 T C H1 frag. 19 715 A T H1 frag. 23 1825 T C rhod frag. 2 3 A G H1 frag. 18 756 T C H1 frag. 19 749 C T H1 frag. 24 8 C T rhod frag. 2 51 T C H1 frag. 18 830 C A H1 frag. 20 25 A G H1 frag. 24 17 C T rhod frag. 2 73 G A H1 frag. 19 278 T C H1 frag. 20 66 T C H1 frag. 25 20 A T rhod frag. 2 82 G C H1 frag. 20 20 C G H1 frag. 21 171 A T 108 (Ranoides) rhod frag. 2 126 C A H1 frag. 20 140 T C H1 frag. 21 179 C T 28S frag. 2 720 G Ð SIA frag. 2 41 A G H1 frag. 21 8 C T H1 frag. 22 20 T C H1 frag. 10 14 C T SIA frag. 2 59 C T H1 frag. 23 789 T Ð H1 frag. 23 48 A G H1 frag. 10 19 G A SIA frag. 3 12 C A H1 frag. 23 944 G A H1 frag. 23 299 G Ð H1 frag. 11 27 G A SIA frag. 3 108 T C H1 frag. 23 963 A Ð H1 frag. 23 898 Ð T H1 frag. 12 14 C T SIA frag. 3 117 C T H1 frag. 23 968 C Ð H1 frag. 23 922 Ð T H1 frag. 12 21 G A SIA frag. 3 129 C T H1 frag. 23 1027 A Ð H1 frag. 23 1160 G T H1 frag. 12 127 A T tyr frag. 1 40 C T H1 frag. 23 1097 C Ð H1 frag. 23 1181 C T H1 frag. 13 151 C T tyr frag. 2 14 G A H1 frag. 23 1320 C T H1 frag. 23 1303 C A H1 frag. 14 72 Ð A tyr frag. 2 93 A C H1 frag. 23 1376 T A H1 frag. 23 1686 Ð C H1 frag. 14 142 C T tyr frag. 2 96 A C H1 frag. 23 1695 A G H1 frag. 23 1781 A T H1 frag. 14 147 G A tyr frag. 2 100 G C H1 frag. 23 1704 C T H1 frag. 23 1824 T C H1 frag. 14 149 G A tyr frag. 2 108 G A H1 frag. 25 14 G A H1 frag. 23 1882 A T H1 frag. 15 19 C T tyr frag. 2 128 G A H1 frag. 25 87 C T H1 frag. 23 1968 T A H1 frag. 15 20 C T tyr frag. 2 138 T G H3 frag. 1 72 T C H1 frag. 25 32 G A H1 frag. 15 60 T A tyr frag. 2 172 C T H3 frag. 1 120 T C H1 frag. 6 62 A Ð H1 frag. 16 72 Ð G tyr frag. 2 207 C T H3 frag. 2 39 G C H1 frag. 6 67 T Ð H1 frag. 16 152 A T tyr frag. 2 270 G C rhod frag. 1 33 A C H1 frag. 6 75 A Ð H1 frag. 16 249 Ð T tyr frag. 3 63 C A rhod frag. 1 78 A G H1 frag. 6 85 T Ð H1 frag. 16 467 G Ð tyr frag. 3 85 A G rhod frag. 1 135 T C H1 frag. 6 98 A Ð H1 frag. 16 665 T C tyr frag. 3 88 C T rhod frag. 2 61 G A H1 frag. 6 104 C Ð H1 frag. 17 160 C Ð tyr frag. 3 91 A T rhod frag. 2 85 C G H1 frag. 6 115 A Ð H1 frag. 17 306 Ð A tyr frag. 3 103 C T rhod frag. 2 108 A T H1 frag. 6 181 C Ð H1 frag. 18 93 C T tyr frag. 3 119 C T SIA frag. 2 32 G A H1 frag. 6 213 A Ð H1 frag. 18 185 C T tyr frag. 3 127 A T SIA frag. 2 41 A G H1 frag. 8 40 T C H1 frag. 19 155 T C tyr frag. 3 128 C T SIA frag. 2 59 C T H1 frag. 8 173 T G H1 frag. 19 215 A G tyr frag. 3 140 G A SIA frag. 3 48 G C H1 frag. 8 184 T C H1 frag. 19 216 T C tyr frag. 3 164 G A SIA frag. 3 54 C T H1 frag. 8 232 C Ð H1 frag. 19 286 C A tyr frag. 3 173 C T SIA frag. 3 78 A G H1 frag. 8 525 C A H1 frag. 19 826 A T 109 (Allodapanura) SIA frag. 4 88 T G H1 frag. 8 619 Ð C H1 frag. 2 16 C A H1 frag. 1 68 C T 101 (unnamed taxon) H1 frag. 9 28 A G H1 frag. 2 196 T C H1 frag. 11 536 C T H1 frag. 10 7 A C H1 frag. 9 48 T C H1 frag. 2 200 G A H1 frag. 11 622 C T H1 frag. 10 9 T C H1 frag. 9 281 T C H1 frag. 2 301 C A H1 frag. 11 868 Ð A H1 frag. 10 23 T G H1 frag. 9 309 Ð C H1 frag. 2 419 A C H1 frag. 11 872 Ð A H1 frag. 10 192 Ð T H1 frag. 9 558 A C H1 frag. 22 10 T C H1 frag. 11 938 Ð C H1 frag. 10 199 C A H1 frag. 9 618 A Ð H1 frag. 23 250 G A H1 frag. 11 1089 C A H1 frag. 10 261 T C H1 frag. 9 739 A G H1 frag. 23 274 C A H1 frag. 12 65 Ð A H1 frag. 11 19 A G H1 frag. 9 818 T C H1 frag. 23 729 C T H1 frag. 16 221 C Ð H1 frag. 11 130 A C H3 frag. 1 0 C T H1 frag. 23 1386 Ð A H1 frag. 16 313 C Ð H1 frag. 11 469 Ð T H3 frag. 1 27 C T H1 frag. 23 1607 C A H1 frag. 16 509 A T H1 frag. 11 887 A T H3 frag. 1 28 C A H1 frag. 23 1695 A T H1 frag. 18 371 A T H1 frag. 11 1336 T C H3 frag. 1 81 A C H1 frag. 23 1759 T A H1 frag. 19 54 T A H1 frag. 12 14 A T H3 frag. 1 216 G A H1 frag. 23 1828 C A H1 frag. 19 185 G A 334 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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H1 frag. 19 208 C T H1 frag. 9 226 A G H1 frag. 22 36 G A H1 frag. 17 118 G T H1 frag. 19 244 A C H1 frag. 9 652 T C H1 frag. 23 9 A C H1 frag. 17 300 Ð G H1 frag. 19 796 A G H1 frag. 9 775 C T H1 frag. 23 22 T C H1 frag. 18 322 A G H1 frag. 21 277 C T H3 frag. 1 114 T C H1 frag. 23 102 A T H1 frag. 18 393 Ð C H1 frag. 22 37 T C H3 frag. 1 117 G C H1 frag. 23 190 C Ð H1 frag. 18 447 C Ð H1 frag. 23 762 C A H3 frag. 1 193 C A H1 frag. 23 250 T Ð H1 frag. 19 119 A T H1 frag. 23 1041 T Ð H3 frag. 2 29 T C H1 frag. 23 265 A C H1 frag. 2 154 T C H1 frag. 23 1905 T Ð H3 frag. 2 39 G C H1 frag. 23 559 C T H1 frag. 21 57 A Ð H1 frag. 3 407 A Ð rhod frag. 1 36 A G H1 frag. 23 1015 A C H1 frag. 3 398 C A H1 frag. 4 578 Ð T rhod frag. 1 66 T C H1 frag. 23 1181 C A H1 frag. 4 672 T A H1 frag. 4 639 Ð C rhod frag. 1 117 A C H1 frag. 23 1292 Ð C H1 frag. 8 41 A C H1 frag. 6 51 Ð A rhod frag. 1 135 C T H1 frag. 23 1310 T C H1 frag. 8 428 Ð T H1 frag. 6 181 A T rhod frag. 1 162 T C H1 frag. 23 1316 T C H1 frag. 8 545 C T H1 frag. 8 241 Ð C rhod frag. 2 43 T A H1 frag. 23 1532 C Ð H1 frag. 9 46 A T H1 frag. 8 316 T A rhod frag. 2 53 A T H1 frag. 23 1739 A T H1 frag. 9 349 Ð T H1 frag. 8 335 T A rhod frag. 2 130 G Ð H1 frag. 23 1750 A C H1 frag. 9 383 C A H1 frag. 8 423 C T SIA frag. 3 9 T C H1 frag. 23 1846 T C H1 frag. 9 440 Ð C H1 frag. 9 580 C A SIA frag. 3 120 G C H1 frag. 24 20 T C H3 frag. 1 198 G A rhod frag. 1 99 T C SIA frag. 3 168 T C H1 frag. 6 25 T C SIA frag. 2 38 C T SIA frag. 3 33 C T SIA frag. 3 177 A G H1 frag. 6 27 G A SIA frag. 3 9 C T SIA frag. 3 64 T C 111 (unnamed taxon) H1 frag. 8 23 T C tyr frag. 1 46 T C tyr frag. 2 210 C T 28S frag. 2 369 Ð C H1 frag. 8 24 T A tyr frag. 2 47 T G tyr frag. 3 35 G T H1 frag. 2 66 Ð C H1 frag. 8 41 A T tyr frag. 2 83 T C tyr frag. 3 64 A G H1 frag. 2 438 T A H1 frag. 8 90 Ð C tyr frag. 3 47 G A tyr frag. 3 79 C T H1 frag. 3 355 A T H1 frag. 8 139 C T 129 (unnamed taxon) tyr frag. 3 122 G A H1 frag. 4 337 C T H1 frag. 8 568 A C H1 frag. 11 872 C A tyr frag. 3 181 G A H1 frag. 4 351 G A H1 frag. 8 611 A T H1 frag. 11 1161 A Ð 110 (Microhylidae) H3 frag. 1 55 C A H1 frag. 8 667 A C H1 frag. 13 159 T C 28S frag. 2 473 T C 118 (Cophylinae) H1 frag. 8 828 T C H1 frag. 16 201 T Ð 28S frag. 2 644 Ð C 28S frag. 2 567 C Ð H1 frag. 9 132 T Ð H1 frag. 17 33 A T 28S frag. 2 719 C G H1 frag. 11 16 A G H1 frag. 9 138 C Ð H1 frag. 17 176 Ð A 28S frag. 2 790 C A H1 frag. 11 47 T Ð H1 frag. 9 173 A Ð H1 frag. 18 267 Ð T 28S frag. 3 424 G C H1 frag. 11 67 C Ð H1 frag. 9 185 A Ð H1 frag. 18 276 A T H1 frag. 10 261 C T H1 frag. 11 409 T Ð H1 frag. 9 202 A Ð H1 frag. 18 349 C T H1 frag. 11 144 T C H1 frag. 11 663 A C H1 frag. 9 226 G Ð H1 frag. 18 821 T C H1 frag. 11 726 Ð G H1 frag. 11 726 G C H1 frag. 9 315 C Ð H1 frag. 19 600 C T H1 frag. 11 1045 A T H1 frag. 11 778 C Ð H1 frag. 9 333 C Ð H1 frag. 2 152 C Ð H1 frag. 11 1217 A C H1 frag. 11 910 C T H1 frag. 9 343 A Ð H1 frag. 20 20 C A H1 frag. 12 52 C Ð H1 frag. 11 938 C Ð H1 frag. 9 367 A Ð H1 frag. 21 90 C A H1 frag. 13 8 G A H1 frag. 11 983 T Ð H1 frag. 9 506 C Ð H1 frag. 23 942 C T H1 frag. 13 125 A C H1 frag. 11 1342 T A H1 frag. 9 520 A Ð H1 frag. 23 981 A Ð H1 frag. 14 44 T C H1 frag. 12 6 A C H1 frag. 9 533 A Ð H1 frag. 23 997 A T H1 frag. 15 50 T C H1 frag. 13 6 A Ð H1 frag. 9 558 A Ð H1 frag. 23 1103 Ð A H1 frag. 16 16 A G H1 frag. 13 33 T Ð H1 frag. 9 580 A Ð H1 frag. 3 2 A G H1 frag. 16 25 T C H1 frag. 13 75 T Ð H1 frag. 9 693 A Ð H1 frag. 3 398 C A H1 frag. 16 414 A T H1 frag. 13 127 A T H1 frag. 9 798 C Ð H1 frag. 3 402 C T H1 frag. 16 556 Ð T H1 frag. 13 132 G A H1 frag. 9 818 T C H1 frag. 4 191 T C H1 frag. 16 557 Ð T H1 frag. 14 13 T C H3 frag. 1 45 C T H1 frag. 8 84 Ð C H1 frag. 17 231 C T H1 frag. 14 91 T A H3 frag. 1 69 G A H1 frag. 8 241 C T H1 frag. 17 423 A G H1 frag. 14 193 T C H3 frag. 1 192 C G H1 frag. 8 568 A C H1 frag. 18 116 A T H1 frag. 14 243 G A H3 frag. 1 195 A C H1 frag. 9 216 T C H1 frag. 18 397 Ð A H1 frag. 14 263 T C SIA frag. 1 36 T C rhod frag. 1 51 T C H1 frag. 18 607 Ð C H1 frag. 15 23 C A SIA frag. 2 44 C T SIA frag. 2 59 T C H1 frag. 18 727 C T H1 frag. 16 33 C T SIA frag. 3 33 T C SIA frag. 3 42 T C H1 frag. 18 872 A C H1 frag. 16 86 Ð T SIA frag. 3 42 T G 130 (Microhylinae) H1 frag. 19 376 T A H1 frag. 16 355 Ð C SIA frag. 3 60 A G 28S frag. 2 434 Ð G H1 frag. 19 668 A T H1 frag. 16 547 C Ð SIA frag. 3 64 C T 28S frag. 2 682 Ð C H1 frag. 2 7 C A H1 frag. 16 557 A C SIA frag. 3 159 C G 28S frag. 3 491 Ð G H1 frag. 2 407 A T H1 frag. 17 38 A T SIA frag. 4 76 T G 28S frag. 3 603 A G H1 frag. 20 129 T A H1 frag. 18 45 T C tyr frag. 1 50 A G H1 frag. 1 50 A G H1 frag. 23 190 Ð C H1 frag. 18 93 T C tyr frag. 2 92 G A H1 frag. 1 57 A G H1 frag. 23 213 A G H1 frag. 18 94 A C tyr frag. 2 191 G A H1 frag. 10 52 C T H1 frag. 23 1532 Ð C H1 frag. 18 408 Ð C tyr frag. 2 194 A G H1 frag. 11 368 A C H1 frag. 23 1607 A T H1 frag. 18 581 C T tyr frag. 3 91 T A H1 frag. 11 595 A T H1 frag. 23 1846 C T H1 frag. 18 707 T C tyr frag. 3 113 T A H1 frag. 11 868 A C H1 frag. 23 1946 Ð A H1 frag. 18 821 T C tyr frag. 3 126 C A H1 frag. 11 1089 A T H1 frag. 4 120 T Ð H1 frag. 18 872 C A tyr frag. 3 153 G A H1 frag. 11 1327 T A H1 frag. 4 201 Ð A H1 frag. 19 83 C A 121 (Gastrophryninae) H1 frag. 12 65 A G H1 frag. 4 308 G C H1 frag. 19 331 G Ð H1 frag. 1 71 A T H1 frag. 13 125 C A H1 frag. 4 330 T Ð H1 frag. 19 338 A Ð H1 frag. 11 67 C A H1 frag. 16 170 T A H1 frag. 4 637 T C H1 frag. 19 350 G Ð H1 frag. 11 595 A T H1 frag. 16 556 T C H1 frag. 5 12 A G H1 frag. 19 356 C Ð H1 frag. 11 778 C A H1 frag. 16 614 T C H1 frag. 6 167 A Ð H1 frag. 19 376 A T H1 frag. 11 1000 A T H1 frag. 17 31 T Ð H1 frag. 8 24 A T H1 frag. 19 439 G Ð H1 frag. 13 69 C A H1 frag. 17 182 T A H1 frag. 8 28 A T H1 frag. 19 715 T C H1 frag. 13 151 T C H1 frag. 17 296 T A H1 frag. 8 122 A C H1 frag. 20 66 A G H1 frag. 14 124 T C H1 frag. 17 423 G A H1 frag. 8 260 C A H1 frag. 20 176 A G H1 frag. 15 58 A G H1 frag. 18 185 C T H1 frag. 8 313 C T H1 frag. 21 10 C T H1 frag. 16 191 C T H1 frag. 19 278 C A H1 frag. 8 523 A C H1 frag. 21 171 G A H1 frag. 16 414 T C H1 frag. 20 1 C T H1 frag. 9 216 Ð T H1 frag. 21 238 Ð T H1 frag. 16 590 A Ð H1 frag. 20 68 G A 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 335

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H1 frag. 20 73 A G H1 frag. 10 217 T Ð H1 frag. 8 824 Ð C H1 frag. 12 148 T Ð H1 frag. 21 10 C T H1 frag. 10 269 C T H1 frag. 9 226 G A H1 frag. 12 186 A C H1 frag. 21 171 G A H1 frag. 11 7 G A H1 frag. 9 281 C A H1 frag. 14 72 A Ð H1 frag. 23 52 C T H1 frag. 11 88 T C H1 frag. 9 725 C T H1 frag. 16 39 T A H1 frag. 23 602 T Ð H1 frag. 11 778 C Ð H3 frag. 1 12 C T H1 frag. 16 535 A Ð H1 frag. 23 1221 C T H1 frag. 11 870 Ð C H3 frag. 1 84 C G H1 frag. 16 672 A T H1 frag. 23 1331 C T H1 frag. 11 887 A T H3 frag. 1 189 G C H1 frag. 17 274 A Ð H1 frag. 23 1518 A C H1 frag. 11 1314 Ð T SIA frag. 3 6 A G H1 frag. 18 276 A C H1 frag. 23 1616 C A H1 frag. 11 1315 Ð T SIA frag. 3 51 T C H1 frag. 18 306 A T H1 frag. 4 13 A T H1 frag. 11 1336 T Ð SIA frag. 3 60 A C H1 frag. 18 349 C T H1 frag. 6 181 C T H1 frag. 11 1342 T Ð SIA frag. 3 84 G C H1 frag. 18 508 Ð T H1 frag. 6 199 A T H1 frag. 12 41 A G SIA frag. 3 123 T C H1 frag. 18 648 C A H1 frag. 8 41 A T H1 frag. 12 143 A C SIA frag. 4 22 G T H1 frag. 19 123 A C H1 frag. 8 313 T A H1 frag. 13 130 C T SIA frag. 4 23 T C H1 frag. 19 216 C A H1 frag. 8 523 C T H1 frag. 14 9 G A SIA frag. 4 46 T C H1 frag. 19 439 G T H1 frag. 9 609 C A H1 frag. 14 238 A T SIA frag. 4 76 T C H1 frag. 19 509 C T H3 frag. 1 T C H1 frag. 16 33 C T 143 (Afrobatrachia) H1 frag. 2 32 G A H3 frag. 1 124 A C H1 frag. 16 72 G A 28S frag. 2 386 C G H1 frag. 2 67 Ð C rhod frag. 1 57 T C H1 frag. 16 115 Ð C 28S frag. 2 639 G Ð H1 frag. 2 133 C T rhod frag. 1 66 C T H1 frag. 16 241 A Ð 28S frag. 2 655 G Ð H1 frag. 2 277 A Ð rhod frag. 1 125 T C H1 frag. 16 573 Ð T 28S frag. 2 768 C A H1 frag. 20 60 G A SIA frag. 1 4 T C H1 frag. 16 665 C T H1 frag. 1 59 T A H1 frag. 20 115 T Ð SIA frag. 2 32 G C H1 frag. 16 677 G A H1 frag. 10 43 A Ð H1 frag. 21 218 C Ð SIA frag. 3 3 A G H1 frag. 16 680 G T H1 frag. 11 264 C T H1 frag. 23 52 C T SIA frag. 3 111 A G H1 frag. 17 54 C T H1 frag. 11 429 Ð A H1 frag. 23 1074 T Ð SIA frag. 3 153 A G H1 frag. 17 210 A C H1 frag. 11 565 C A H1 frag. 23 1131 G C tyr frag. 1 21 T G H1 frag. 18 1 G A H1 frag. 11 819 C A H1 frag. 23 1338 C Ð tyr frag. 2 130 T C H1 frag. 18 116 C T H1 frag. 11 1327 T G H1 frag. 23 1444 G Ð tyr frag. 3 50 T A H1 frag. 18 306 A C H1 frag. 12 80 C Ð H1 frag. 23 1825 A T 134 (unnamed taxon) H1 frag. 18 530 C Ð H1 frag. 12 112 G A H1 frag. 3 58 C T 28S frag. 2 567 C T H1 frag. 18 563 A C H1 frag. 12 116 G A H1 frag. 3 184 A T H1 frag. 10 199 C A H1 frag. 18 597 Ð A H1 frag. 13 91 T A H1 frag. 3 297 C A H1 frag. 11 53 T G H1 frag. 18 717 A C H1 frag. 16 333 A C H1 frag. 3 351 Ð C H1 frag. 11 439 C A H1 frag. 18 741 C Ð H1 frag. 17 118 G Ð H1 frag. 3 365 T C H1 frag. 11 852 C A H1 frag. 18 792 A T H1 frag. 17 333 C T H1 frag. 4 94 A T H1 frag. 11 938 C T H1 frag. 18 883 C T H1 frag. 18 388 C T H1 frag. 4 292 T Ð H1 frag. 11 1294 T C H1 frag. 19 24 G Ð H1 frag. 19 3 C T H1 frag. 4 351 G A H1 frag. 11 1333 T A H1 frag. 19 66 A T H1 frag. 2 152 C T H1 frag. 6 181 T C H1 frag. 13 69 C A H1 frag. 19 369 C A H1 frag. 2 171 A C H1 frag. 7 96 A T H1 frag. 16 19 C A H1 frag. 19 439 G A H1 frag. 2 437 C A H1 frag. 8 46 T C H1 frag. 16 27 T A H1 frag. 2 7 A C H1 frag. 20 176 A G H1 frag. 8 51 C T H1 frag. 16 648 A G H1 frag. 2 285 A Ð H1 frag. 21 133 A T H1 frag. 8 52 A G H1 frag. 16 681 C T H1 frag. 20 9 A T H1 frag. 23 22 T C H1 frag. 8 74 A T H1 frag. 17 333 C T H1 frag. 20 60 G Ð H1 frag. 23 59 G A H1 frag. 8 181 A G H1 frag. 18 13 A G H1 frag. 21 8 C T H1 frag. 23 822 Ð T H1 frag. 8 369 T Ð H1 frag. 18 539 T Ð H1 frag. 21 172 G A H1 frag. 23 981 T A H1 frag. 8 458 Ð T H1 frag. 18 724 Ð C H1 frag. 21 270 G C H1 frag. 23 1169 C A H1 frag. 8 556 T C H1 frag. 19 42 A T H1 frag. 22 10 C T H1 frag. 3 22 C A H1 frag. 8 558 G A H1 frag. 19 61 G Ð H1 frag. 22 11 C T H1 frag. 3 169 A T H1 frag. 8 565 A C H1 frag. 19 119 A T H1 frag. 22 13 C T H1 frag. 3 176 A T H1 frag. 8 611 A T H1 frag. 2 65 T Ð H1 frag. 22 23 G A H1 frag. 3 338 Ð C H1 frag. 8 816 T C H1 frag. 2 154 T C H1 frag. 22 61 G A H1 frag. 3 396 Ð T H1 frag. 8 828 T A H1 frag. 2 180 A T H1 frag. 23 52 C T H1 frag. 4 592 T A H1 frag. 9 5 C A H1 frag. 2 439 C T H1 frag. 23 67 G A H1 frag. 6 23 C T H1 frag. 9 457 Ð G H1 frag. 21 57 A Ð H1 frag. 23 103 A C H1 frag. 8 69 C T H1 frag. 9 538 Ð C H1 frag. 21 163 Ð A H1 frag. 23 190 C Ð H1 frag. 8 352 A G H1 frag. 9 539 Ð C H1 frag. 23 452 C T H1 frag. 23 274 A C H1 frag. 8 550 G A H1 frag. 9 693 A Ð H1 frag. 23 623 A C H1 frag. 23 283 C T H1 frag. 8 626 C A H1 frag. 9 840 C A H1 frag. 23 1169 C T H1 frag. 23 440 Ð A H1 frag. 9 397 Ð A rhod frag. 1 2 A C H1 frag. 3 58 C T H1 frag. 23 693 A Ð H1 frag. 9 506 A T rhod frag. 2 3 G A H1 frag. 3 362 G A H1 frag. 23 1131 G A H1 frag. 9 558 A Ð rhod frag. 2 42 C G H1 frag. 4 283 T C H1 frag. 23 1303 C T H1 frag. 9 818 T C rhod frag. 2 80 G A H1 frag. 4 316 C A H1 frag. 23 1816 A C rhod frag. 2 126 A G rhod frag. 2 127 C G H1 frag. 5 17 C A H1 frag. 23 1946 A Ð tyr frag. 1 G A rhod frag. 2 129 A T H1 frag. 6 91 T C H1 frag. 3 2 A T tyr frag. 1 12 A C SIA frag. 1 36 T C H1 frag. 8 40 A T H1 frag. 3 8 T C tyr frag. 2 157 A C SIA frag. 1 39 C T H1 frag. 8 74 A T H1 frag. 3 251 C T tyr frag. 2 195 G A SIA frag. 2 53 A G H1 frag. 8 260 A C H1 frag. 3 252 C T tyr frag. 2 258 A G SIA frag. 3 24 C T H1 frag. 8 441 A C H1 frag. 3 321 G A tyr frag. 3 53 T C SIA frag. 3 54 C T H1 frag. 8 696 A G H1 frag. 3 398 C T 144 (Xenosyneunitanura) SIA frag. 3 78 G A H1 frag. 8 735 A G H1 frag. 4 391 A C 28S frag. 2 330 G Ð SIA frag. 3 114 G T H1 frag. 9 343 A C H1 frag. 4 441 C A 28S frag. 2 719 C Ð SIA frag. 4 22 G A H1 frag. 9 645 Ð T H1 frag. 4 592 C A H1 frag. 11 67 C T SIA frag. 4 73 G T H1 frag. 9 818 T C H1 frag. 4 670 A C H1 frag. 11 139 A C tyr frag. 1 7 C G SIA frag. 1 39 C T H1 frag. 5 12 G A H1 frag. 11 368 A C tyr frag. 1 46 T C 135 (Asterophryinae) H1 frag. 5 35 G A H1 frag. 11 852 C T tyr frag. 1 82 C T 28S frag. 2 453 C T H1 frag. 8 41 A C H1 frag. 11 937 Ð C tyr frag. 2 11 C T 28S frag. 2 650 Ð G H1 frag. 8 86 Ð A H1 frag. 11 1071 C A tyr frag. 2 39 A C H1 frag. 1 22 G A H1 frag. 8 313 T C H1 frag. 11 1217 A T tyr frag. 2 138 G A H1 frag. 10 52 C A H1 frag. 8 488 T A H1 frag. 11 1259 T Ð tyr frag. 2 177 C T 336 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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tyr frag. 2 204 C A H1 frag. 19 749 T C H1 frag. 11 1226 Ð T H1 frag. 4 346 C A tyr frag. 2 213 T A H1 frag. 23 693 A T H1 frag. 12 6 A G H1 frag. 4 349 C A tyr frag. 3 44 G A H1 frag. 23 1074 T A H1 frag. 12 7 G T H1 frag. 4 351 G T tyr frag. 3 51 G A H1 frag. 23 1221 C Ð H1 frag. 12 12 T C H1 frag. 4 577 Ð C tyr frag. 3 52 A C H1 frag. 23 1911 Ð T H1 frag. 12 84 Ð C H1 frag. 4 637 T Ð tyr frag. 3 123 C T tyr frag. 2 23 C T H1 frag. 12 186 A C H1 frag. 5 9 A G 148 (Laurentobatrachia) tyr frag. 2 65 C T H1 frag. 13 18 G T H1 frag. 5 33 A T 28S frag. 2 714 G C tyr frag. 2 67 C G H1 frag. 13 91 A T H1 frag. 6 25 T C 28S frag. 2 764 A C tyr frag. 2 222 T C H1 frag. 13 130 C T H1 frag. 7 42 A C 28S frag. 3 306 Ð C tyr frag. 3 79 C G H1 frag. 13 156 T A H1 frag. 7 97 C A H1 frag. 11 392 C A 164 (Arthroleptidae) H1 frag. 14 28 C A H1 frag. 8 69 T C H1 frag. 11 868 A C 28S frag. 2 319 Ð G H1 frag. 14 60 Ð T H1 frag. 8 177 A C H1 frag. 11 1089 A Ð 28S frag. 3 370 G C H1 frag. 14 143 C A H1 frag. 8 181 A G H1 frag. 12 103 A Ð H1 frag. 1 G A H1 frag. 14 238 A T H1 frag. 8 298 G A H1 frag. 12 127 T A H1 frag. 1 30 C T H1 frag. 14 250 C A H1 frag. 8 488 A C H1 frag. 14 91 T A H1 frag. 10 54 Ð T H1 frag. 14 260 G T H1 frag. 8 544 C T H1 frag. 16 31 A G H1 frag. 10 101 G A H1 frag. 15 42 A T H1 frag. 8 550 A G H1 frag. 16 359 C A H1 frag. 10 266 C T H1 frag. 16 141 Ð A H1 frag. 8 694 Ð A H1 frag. 16 547 C A H1 frag. 11 12 G A H1 frag. 16 170 T A H1 frag. 8 716 Ð T H1 frag. 16 590 A T H1 frag. 11 89 C T H1 frag. 16 333 C T H1 frag. 8 735 A G H1 frag. 17 11 A C H1 frag. 11 910 T Ð H1 frag. 16 450 A C H1 frag. 9 67 T C H1 frag. 17 47 C T H1 frag. 11 927 A C H1 frag. 16 648 A T H1 frag. 9 126 C T H1 frag. 17 182 T A H1 frag. 13 168 A C H1 frag. 16 691 A G 168 (Arthroleptinae) H1 frag. 17 318 Ð C H1 frag. 14 65 G Ð H1 frag. 17 5 T C H1 frag. 11 67 C A H1 frag. 18 93 T C H1 frag. 14 129 T C H1 frag. 17 12 A T H1 frag. 11 264 T C H1 frag. 18 371 T Ð H1 frag. 15 54 A G H1 frag. 17 296 C T H1 frag. 11 368 A C H1 frag. 18 766 C T H1 frag. 16 127 T C H1 frag. 17 372 T Ð H1 frag. 11 409 T Ð H1 frag. 18 782 A Ð H1 frag. 16 614 T A H1 frag. 18 164 T C H1 frag. 11 938 C A H1 frag. 18 863 G A H1 frag. 17 31 T A H1 frag. 18 306 A C H1 frag. 11 1191 T Ð H1 frag. 19 24 G T H1 frag. 17 429 A C H1 frag. 18 530 C A H1 frag. 12 113 Ð G H1 frag. 19 91 C T H1 frag. 18 52 A C H1 frag. 18 648 C T H1 frag. 12 116 A G H1 frag. 19 247 C T H1 frag. 18 95 A C H1 frag. 18 673 A C H1 frag. 14 93 A C H1 frag. 19 403 C A H1 frag. 18 388 T A H1 frag. 18 717 A T H1 frag. 14 208 A T H1 frag. 19 816 G A H1 frag. 18 503 Ð A H1 frag. 18 825 Ð A H1 frag. 16 94 T Ð H1 frag. 2 16 A C H1 frag. 19 119 A T H1 frag. 19 14 Ð A H1 frag. 16 382 A C H1 frag. 2 118 C A H1 frag. 19 338 A Ð H1 frag. 19 54 A T H1 frag. 16 436 Ð A H1 frag. 2 210 A G H1 frag. 19 560 A T H1 frag. 19 66 A T H1 frag. 16 629 A C H1 frag. 2 238 C T H1 frag. 2 15 C A H1 frag. 19 172 A C H1 frag. 17 182 A G H1 frag. 2 345 Ð C H1 frag. 20 60 G T H1 frag. 19 318 T Ð H1 frag. 17 383 A T H1 frag. 20 73 A G H1 frag. 20 180 C T H1 frag. 19 439 G A H1 frag. 17 424 Ð C H1 frag. 20 182 C T H1 frag. 23 17 G A H1 frag. 19 826 T A H1 frag. 18 155 Ð T H1 frag. 21 76 C T H1 frag. 23 693 A C H1 frag. 2 44 G A H1 frag. 18 628 T A H1 frag. 22 55 T C H1 frag. 23 1214 T Ð H1 frag. 2 176 Ð A H1 frag. 18 720 A C H1 frag. 23 250 A T H1 frag. 23 1221 C A H1 frag. 2 207 G A H1 frag. 18 866 A G H1 frag. 23 1238 Ð G H1 frag. 3 262 T C H1 frag. 2 237 C T H1 frag. 19 3 T C H1 frag. 23 1359 G A H1 frag. 3 355 A G H1 frag. 2 240 G A H1 frag. 19 449 T C H1 frag. 23 1427 A C H1 frag. 3 362 G A H1 frag. 2 425 C T H1 frag. 19 485 Ð A H1 frag. 23 1556 Ð A H1 frag. 4 120 T A H1 frag. 2 437 A C H1 frag. 19 531 A C H1 frag. 3 124 A T H1 frag. 4 630 T A H1 frag. 20 23 C A H1 frag. 19 715 A T H1 frag. 3 165 Ð A H1 frag. 7 15 T C H1 frag. 4 370 Ð A H1 frag. 8 139 C T H1 frag. 21 76 T Ð H1 frag. 2 328 C T H1 frag. 4 673 A C H1 frag. 8 263 Ð A H1 frag. 21 113 A T H1 frag. 23 25 C T H1 frag. 6 199 C A H1 frag. 8 545 A T H1 frag. 21 243 T Ð H1 frag. 23 59 A G H1 frag. 7 55 C T H1 frag. 9 73 A C H1 frag. 22 37 C T H1 frag. 23 216 Ð G H1 frag. 8 249 A Ð H1 frag. 9 202 C A H1 frag. 22 47 Ð T H1 frag. 23 602 T Ð H1 frag. 8 735 T A H1 frag. 9 492 A C H1 frag. 22 55 C Ð H1 frag. 23 997 A T H1 frag. 9 729 Ð A H3 frag. 1 42 C T H1 frag. 23 2 G C H1 frag. 23 1097 G A rhod frag. 2 6 C G H3 frag. 1 57 G C H1 frag. 23 52 C A H1 frag. 23 1169 A C rhod frag. 2 27 C T H3 frag. 1 66 C T H1 frag. 23 100 A T H1 frag. 23 1518 A C rhod frag. 2 82 C G H3 frag. 1 126 A C H1 frag. 23 182 T G H1 frag. 23 1951 A C rhod frag. 2 118 T C rhod frag. 2 21 C T H1 frag. 23 1131 G C H1 frag. 3 58 C A SIA frag. 3 51 T C rhod frag. 2 61 G A H1 frag. 23 1154 A T H1 frag. 3 169 T Ð SIA frag. 3 156 T A tyr frag. 1 55 T C H1 frag. 23 1181 C A H1 frag. 3 176 T G tyr frag. 1 14 T G tyr frag. 2 17 C T H1 frag. 23 1321 A G H1 frag. 3 338 C T tyr frag. 1 29 C G tyr frag. 2 20 G A H1 frag. 23 1460 G T H1 frag. 4 292 T Ð tyr frag. 2 53 C T tyr frag. 2 56 G A H1 frag. 23 1554 T C H1 frag. 4 325 A T tyr frag. 3 47 T A tyr frag. 2 92 A G H1 frag. 23 1704 C A H1 frag. 4 386 T C tyr frag. 3 134 A G tyr frag. 2 207 T A H1 frag. 24 8 C T H1 frag. 6 55 T G 161 (Hyperolius) 165 (Leptopelinae) H1 frag. 25 44 A T H1 frag. 6 62 A T 28S frag. 2 567 G A 28S frag. 2 606 Ð C H1 frag. 3 8 A C H1 frag. 8 74 A T H1 frag. 16 401 Ð C 28S frag. 3 603 A G H1 frag. 3 44 A T H1 frag. 8 441 A C H1 frag. 18 24 C T H1 frag. 1 22 G A H1 frag. 3 249 A T H1 frag. 8 523 A C H1 frag. 18 254 T A H1 frag. 10 6 C T H1 frag. 3 266 C A H1 frag. 8 667 A T H1 frag. 18 509 A T H1 frag. 10 24 G A H1 frag. 4 211 A T H1 frag. 9 27 A G H1 frag. 19 54 A T H1 frag. 10 261 C T H1 frag. 4 254 T C H1 frag. 9 343 A Ð H1 frag. 19 178 C T H1 frag. 11 16 A G H1 frag. 4 280 C T H1 frag. 9 409 A C H1 frag. 19 331 G A H1 frag. 11 72 C A H1 frag. 4 298 A T tyr frag. 2 23 C T H1 frag. 19 396 A T H1 frag. 11 130 A C H1 frag. 4 316 C T tyr frag. 2 35 C T H1 frag. 19 651 A C H1 frag. 11 146 C T H1 frag. 4 337 C A tyr frag. 3 50 T C H1 frag. 19 729 C A H1 frag. 11 663 A G H1 frag. 4 343 G T tyr frag. 3 52 A C 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 337

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169 (Astylosternini) H1 frag. 23 40 C T H1 frag. 16 590 A C H1 frag. 18 530 C T H1 frag. 10 55 T A H1 frag. 23 1338 C T H1 frag. 17 350 A T H1 frag. 18 563 A T H1 frag. 11 40 G Ð H1 frag. 23 1427 C A H1 frag. 18 654 A T H1 frag. 18 727 C T H1 frag. 11 656 Ð G H1 frag. 8 711 G A H1 frag. 18 746 T C H1 frag. 18 766 C T H1 frag. 11 657 Ð C H1 frag. 8 796 C A H1 frag. 18 838 A T H1 frag. 18 863 G A H1 frag. 11 694 T C H1 frag. 8 804 A T H1 frag. 18 866 A G H1 frag. 19 148 A T H1 frag. 11 778 A C H1 frag. 9 17 A G H1 frag. 19 24 G A H1 frag. 19 197 A T H1 frag. 11 1113 A C H1 frag. 9 22 A G H1 frag. 19 91 C T H1 frag. 19 201 A T H1 frag. 11 1311 A C H1 frag. 9 52 T C H1 frag. 19 278 C A H1 frag. 19 213 T C H1 frag. 12 148 T A H1 frag. 9 54 T C H1 frag. 19 415 A T H1 frag. 19 461 Ð C H1 frag. 13 159 A C H1 frag. 9 256 C T H1 frag. 19 560 A T H1 frag. 2 118 C A H1 frag. 14 64 T C H1 frag. 9 432 T Ð H1 frag. 19 635 C Ð H1 frag. 2 140 C T H1 frag. 14 72 A C SIA frag. 2 32 C A H1 frag. 2 133 C A H1 frag. 2 215 A T H1 frag. 14 214 Ð C SIA frag. 3 39 C T H1 frag. 2 371 Ð C H1 frag. 2 277 A G H1 frag. 15 50 T C 175 (Arthroleptis) H1 frag. 2 432 C T H1 frag. 2 389 T C H1 frag. 16 333 C Ð 28S frag. 2 790 C A H1 frag. 2 437 C T H1 frag. 2 420 A G H1 frag. 16 590 T C 28S frag. 3 187 G C H1 frag. 2 443 G A H1 frag. 20 68 G A H1 frag. 16 626 Ð C 28S frag. 3 263 Ð C H1 frag. 20 62 Ð G H1 frag. 20 146 T G H1 frag. 16 660 Ð G 28S frag. 3 264 Ð C H1 frag. 20 77 G Ð H1 frag. 22 72 T A H1 frag. 17 11 C T H1 frag. 10 54 T A H1 frag. 23 2 G C H1 frag. 23 265 A T H1 frag. 17 251 T Ð H1 frag. 11 368 C Ð H1 frag. 23 25 C T H1 frag. 23 883 C Ð H1 frag. 17 350 A C H1 frag. 11 505 A Ð H1 frag. 23 184 Ð C H1 frag. 23 983 Ð G H1 frag. 17 407 T C H1 frag. 11 536 T Ð H1 frag. 23 299 G C H1 frag. 23 997 A T H1 frag. 17 435 Ð A H1 frag. 11 1245 A C H1 frag. 23 1108 C Ð H1 frag. 23 1201 G A H1 frag. 18 816 Ð C H1 frag. 12 41 T C H1 frag. 23 1840 T A H1 frag. 23 1245 C T H1 frag. 19 24 T A H1 frag. 12 113 G A H1 frag. 23 1940 T Ð H1 frag. 23 1346 A T H1 frag. 19 217 A C H1 frag. 15 50 T C H1 frag. 3 8 A C H1 frag. 23 1356 A G H1 frag. 19 364 Ð A H1 frag. 15 58 A G H1 frag. 3 160 T A H1 frag. 23 1444 G A H1 frag. 2 65 T C H1 frag. 16 681 T C H1 frag. 4 64 T Ð H1 frag. 23 1555 Ð A H1 frag. 20 129 T C H1 frag. 17 54 C T H1 frag. 4 223 A C H1 frag. 23 1704 C T H1 frag. 21 133 T A H1 frag. 17 306 A T H1 frag. 4 280 C Ð H1 frag. 23 1951 A C H1 frag. 23 299 G A H1 frag. 17 333 T A H1 frag. 4 439 C A H1 frag. 23 1977 C T H1 frag. 23 785 Ð T H1 frag. 18 155 T C H1 frag. 6 229 C A H1 frag. 24 5 T C H1 frag. 23 1074 T Ð H1 frag. 18 539 A C H1 frag. 7 24 C Ð H1 frag. 24 16 C T H1 frag. 23 1088 A C H1 frag. 18 821 T A H1 frag. 8 74 A T H1 frag. 24 33 A G H1 frag. 23 1124 A T H1 frag. 18 830 A C H1 frag. 8 407 C T H1 frag. 25 16 A G H1 frag. 23 1226 T C H1 frag. 18 865 A T H1 frag. 8 554 G A H1 frag. 3 41 G A H1 frag. 23 1346 A C H1 frag. 19 460 C Ð H1 frag. 8 735 T G H1 frag. 3 129 Ð T H1 frag. 23 1750 T A H1 frag. 19 796 A Ð H1 frag. 9 693 A C H1 frag. 3 138 Ð G H1 frag. 23 1962 T A H1 frag. 20 115 T C H3 frag. 1 57 G C H1 frag. 3 142 Ð A H1 frag. 3 342 T C H1 frag. 20 129 T C H3 frag. 1 162 G C H1 frag. 3 170 Ð G H1 frag. 3 391 A G H1 frag. 23 133 G A H3 frag. 1 189 G C H1 frag. 3 190 Ð T H1 frag. 4 434 A G H1 frag. 23 1074 T C H3 frag. 2 39 G A H1 frag. 3 193 Ð C H1 frag. 5 35 A G H1 frag. 23 1303 C A rhod frag. 2 42 C G H1 frag. 3 194 Ð T H1 frag. 8 28 A C H1 frag. 23 1607 A C rhod frag. 2 97 A C H1 frag. 3 195 Ð T H1 frag. 8 407 C Ð H1 frag. 23 1732 T A tyr frag. 1 18 T C H1 frag. 3 200 Ð C H1 frag. 8 611 A C H1 frag. 23 1992 C T tyr frag. 1 46 T G H1 frag. 3 206 Ð T H1 frag. 8 626 A C H1 frag. 6 59 Ð C tyr frag. 2 23 C T H1 frag. 3 207 Ð T H1 frag. 9 511 A T H1 frag. 7 24 C T tyr frag. 2 101 A G H1 frag. 3 226 Ð T H1 frag. 9 703 Ð C H1 frag. 7 40 A C tyr frag. 2 159 T C H1 frag. 3 227 Ð T tyr frag. 1 29 G A H1 frag. 7 43 G A tyr frag. 2 183 C T H1 frag. 3 241 Ð T tyr frag. 2 83 C A H1 frag. 7 78 C T tyr frag. 2 208 G T H1 frag. 3 243 Ð T tyr frag. 2 240 C T H1 frag. 7 95 C T tyr frag. 2 249 T G H1 frag. 3 249 A G tyr frag. 2 243 C A H1 frag. 8 28 A T tyr frag. 3 52 A G H1 frag. 3 258 C A tyr frag. 3 153 A G H1 frag. 8 41 A C tyr frag. 3 137 T G H1 frag. 3 291 Ð T 172 (Arthroleptini) H1 frag. 8 710 A T 181 (Ptychadenidae/Ptychadena) H1 frag. 3 292 Ð A H1 frag. 10 204 Ð C H1 frag. 9 14 A G H1 frag. 1 20 A G H1 frag. 3 293 Ð A H1 frag. 11 15 A T H1 frag. 9 57 T C H1 frag. 1 54 T A H1 frag. 3 294 Ð A H1 frag. 11 149 T C H1 frag. 9 79 G A H1 frag. 1 76 C T H1 frag. 3 295 Ð C H1 frag. 11 762 A C H1 frag. 9 85 C A H1 frag. 10 43 A Ð H1 frag. 3 298 Ð A H1 frag. 11 1327 G A H1 frag. 9 90 G A H1 frag. 14 89 T C H1 frag. 3 306 Ð C H1 frag. 11 1336 T C tyr frag. 2 19 A G H1 frag. 14 106 A C H1 frag. 3 343 Ð C H1 frag. 12 21 A G tyr frag. 2 41 C T H1 frag. 14 208 A T H1 frag. 3 344 Ð G H1 frag. 12 41 A T tyr frag. 2 126 C G H1 frag. 15 42 A T H1 frag. 3 346 Ð G H1 frag. 13 117 C A tyr frag. 2 219 T C H1 frag. 16 7 A G H1 frag. 3 347 Ð G H1 frag. 14 35 A T tyr frag. 2 273 T C H1 frag. 16 201 T C H1 frag. 3 348 Ð G H1 frag. 14 44 T A tyr frag. 3 61 G C H1 frag. 16 249 T C H1 frag. 4 13 A G H1 frag. 14 75 Ð T tyr frag. 3 70 T C H1 frag. 16 292 A G H1 frag. 4 214 Ð T H1 frag. 14 106 A T tyr frag. 3 114 T A H1 frag. 16 359 C A H1 frag. 4 335 A G H1 frag. 14 243 G A tyr frag. 3 134 G A H1 frag. 16 548 Ð A H1 frag. 4 338 T A H1 frag. 17 18 A T 180 (Natatanura) H1 frag. 16 549 Ð A H1 frag. 4 447 A G H1 frag. 17 437 C T 28S frag. 2 768 C Ð H1 frag. 17 356 Ð C H1 frag. 5 20 A C H1 frag. 18 276 A T H1 frag. 1 38 C T H1 frag. 17 446 C A H1 frag. 5 29 G A H1 frag. 19 216 C T H1 frag. 1 64 A T H1 frag. 18 38 T A H1 frag. 6 12 C T H1 frag. 19 278 C T H1 frag. 10 23 T C H1 frag. 18 52 A G H1 frag. 6 49 T Ð H1 frag. 19 460 A C H1 frag. 11 368 A C H1 frag. 18 209 A G H1 frag. 6 104 C Ð H1 frag. 19 788 A C H1 frag. 11 778 A Ð H1 frag. 18 232 C T H1 frag. 6 137 C T H1 frag. 19 796 G A H1 frag. 13 127 A T H1 frag. 18 282 Ð G H1 frag. 6 163 C A H1 frag. 20 10 T C H1 frag. 14 93 A T H1 frag. 18 411 A T H1 frag. 7 11 G A H1 frag. 20 73 G A H1 frag. 16 450 A Ð H1 frag. 18 514 A T H1 frag. 7 54 C T 338 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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H1 frag. 8 47 C A H1 frag. 13 121 A Ð H1 frag. 9 467 G A H1 frag. 23 293 T A H1 frag. 8 148 C T H1 frag. 13 125 A T H1 frag. 9 522 Ð T H1 frag. 23 727 Ð C H1 frag. 8 300 A G H1 frag. 13 132 G A H1 frag. 9 618 A Ð H1 frag. 23 729 T A H1 frag. 8 352 A T H1 frag. 14 7 T G rhod frag. 1 3 C T H1 frag. 23 940 Ð A H1 frag. 8 488 A T H1 frag. 14 13 T C rhod frag. 1 9 T C H1 frag. 23 1014 Ð T H1 frag. 8 557 Ð G H1 frag. 14 69 Ð T rhod frag. 1 161 A T H1 frag. 23 1124 A Ð H1 frag. 8 568 A Ð H1 frag. 14 126 A G rhod frag. 2 42 G T H1 frag. 23 1214 T C H1 frag. 8 569 A G H1 frag. 14 166 C T rhod frag. 2 61 G A H1 frag. 23 1288 T C H1 frag. 8 714 A C H1 frag. 14 244 G A SIA frag. 2 20 C T H1 frag. 23 1295 T C H1 frag. 8 752 Ð A H1 frag. 14 263 T C SIA frag. 2 71 A C H1 frag. 23 1378 G A H1 frag. 8 818 C A H1 frag. 15 6 A G SIA frag. 3 109 C A H1 frag. 23 1676 C T H1 frag. 9 71 T Ð H1 frag. 15 33 A C SIA frag. 3 123 T C H1 frag. 23 1766 C A H1 frag. 9 202 C T H1 frag. 15 34 G A SIA frag. 3 126 C T H1 frag. 23 1770 A G H1 frag. 9 227 Ð G H1 frag. 15 57 T C SIA frag. 4 52 C T H1 frag. 23 1802 T C H1 frag. 9 258 Ð A H1 frag. 16 337 Ð T SIA frag. 4 55 G A H1 frag. 23 1803 G A H1 frag. 9 383 C Ð H1 frag. 17 372 T C tyr frag. 1 52 T C H1 frag. 23 1811 C T H1 frag. 9 432 T Ð H1 frag. 18 164 T C tyr frag. 2 26 C T H1 frag. 23 1977 C T H1 frag. 9 618 A T H1 frag. 18 185 T A tyr frag. 2 66 A G H1 frag. 23 1995 G A H1 frag. 9 739 A T H1 frag. 18 488 T A tyr frag. 2 67 C T H1 frag. 24 10 A T H1 frag. 9 798 A G H1 frag. 18 530 C A tyr frag. 2 80 C T H1 frag. 24 24 G A rhod frag. 1 C T H1 frag. 18 655 Ð C tyr frag. 2 285 G A H1 frag. 6 85 T C rhod frag. 1 27 G A H1 frag. 18 761 T C tyr frag. 3 13 C T H1 frag. 8 49 Ð C rhod frag. 1 107 C G H1 frag. 18 878 T C tyr frag. 3 14 G T H1 frag. 8 59 T Ð rhod frag. 1 128 C T H1 frag. 18 885 A C tyr frag. 3 53 T A H1 frag. 8 62 T C rhod frag. 1 129 C T H1 frag. 19 278 A T tyr frag. 3 56 G A H1 frag. 8 469 T A rhod frag. 1 135 C T H1 frag. 19 427 T C tyr frag. 3 67 C T H1 frag. 8 553 A G rhod frag. 1 165 C T H1 frag. 19 596 C A 189 (Telmatobatrachia) H1 frag. 8 554 A T rhod frag. 1 171 C T H1 frag. 19 622 A G H1 frag. 11 598 Ð T H1 frag. 8 714 A G rhod frag. 2 41 C T H1 frag. 2 438 T C H1 frag. 11 600 Ð G H1 frag. 8 816 T C rhod frag. 2 54 C T H1 frag. 20 61 A G H1 frag. 11 1093 Ð A H1 frag. 8 828 T Ð rhod frag. 2 86 A T H1 frag. 21 57 A C H1 frag. 21 251 T A H1 frag. 9 28 A G 183 (Victoranura) H1 frag. 21 270 G Ð H1 frag. 22 62 A G H1 frag. 9 33 T C 28S frag. 2 764 A Ð H1 frag. 23 69 T C H1 frag. 23 644 Ð C H1 frag. 9 43 A C H1 frag. 11 392 C A H1 frag. 23 83 A G H1 frag. 23 762 C A H1 frag. 9 48 T C H1 frag. 11 663 A T H1 frag. 23 96 A T H1 frag. 23 1041 T A H1 frag. 9 71 T A H1 frag. 12 120 A C H1 frag. 23 213 A G H1 frag. 23 1233 A G H1 frag. 9 170 Ð G H1 frag. 13 168 A C H1 frag. 23 236 A Ð H1 frag. 23 1411 Ð C H1 frag. 9 171 Ð A H1 frag. 17 118 G A H1 frag. 23 260 C Ð H1 frag. 23 1518 A C H1 frag. 9 453 A G H1 frag. 17 231 C T H1 frag. 23 1097 G A H1 frag. 23 1657 T C H1 frag. 9 467 G C H1 frag. 17 333 C A H1 frag. 23 1131 G A H1 frag. 23 1865 A G H1 frag. 9 495 T C H1 frag. 18 276 A T H1 frag. 23 1338 C A H1 frag. 23 1970 T Ð H1 frag. 9 672 T A H1 frag. 18 388 C T H1 frag. 23 1346 A C H1 frag. 6 50 Ð A H1 frag. 9 755 C Ð H1 frag. 19 737 Ð C H1 frag. 23 1364 A C H1 frag. 8 249 A T rhod frag. 1 12 C T H1 frag. 2 88 A Ð H1 frag. 23 1427 A T H1 frag. 9 98 C T rhod frag. 1 159 G C H1 frag. 2 100 A T H1 frag. 23 1478 A Ð rhod frag. 2 86 A C rhod frag. 1 162 T C H1 frag. 2 162 Ð A H1 frag. 23 1743 A C tyr frag. 3 181 G T rhod frag. 2 57 C T H1 frag. 2 342 A Ð H1 frag. 23 1776 A G 190 (Micrixalidae) rhod frag. 2 112 C T H1 frag. 21 76 C T H1 frag. 23 1798 T C H1 frag. 10 159 A C rhod frag. 2 132 A T H1 frag. 23 182 T C H1 frag. 23 1840 A C H1 frag. 11 16 A G tyr frag. 1 19 C A H1 frag. 23 1376 T C H1 frag. 23 1905 T Ð H1 frag. 11 27 A G tyr frag. 1 21 T G H1 frag. 23 1816 A G H1 frag. 24 6 T C H1 frag. 11 36 T C tyr frag. 1 43 C T H1 frag. 23 1968 T C H1 frag. 24 29 A G H1 frag. 11 67 C A tyr frag. 1 46 G A H1 frag. 4 106 Ð C H1 frag. 3 44 A T H1 frag. 11 79 A G tyr frag. 2 36 T C H1 frag. 4 211 A Ð H1 frag. 3 303 C T H1 frag. 11 112 Ð A tyr frag. 2 41 T C H1 frag. 4 308 G A H1 frag. 3 315 Ð A H1 frag. 11 113 Ð C tyr frag. 2 46 G A H1 frag. 6 91 T C H1 frag. 3 362 G A H1 frag. 11 138 A T tyr frag. 2 47 C T H1 frag. 8 41 A C H1 frag. 4 306 A G H1 frag. 11 312 Ð A tyr frag. 2 92 A G H1 frag. 8 260 C Ð H1 frag. 4 330 T C H1 frag. 11 910 C A tyr frag. 2 98 C T H1 frag. 8 313 C T H1 frag. 4 475 Ð A H1 frag. 11 1190 Ð C tyr frag. 2 100 C G H1 frag. 8 423 C Ð H1 frag. 5 17 A C H1 frag. 11 1248 A T tyr frag. 2 174 G A H1 frag. 9 256 C T H1 frag. 7 42 A C H1 frag. 11 1316 A T tyr frag. 2 189 C G H1 frag. 9 495 Ð T H1 frag. 7 76 T C H1 frag. 11 1327 T C tyr frag. 2 201 T C H1 frag. 9 517 T A H1 frag. 8 62 T A H1 frag. 12 74 T A tyr frag. 2 208 T G H1 frag. 9 755 A C H1 frag. 8 122 A T H1 frag. 21 124 A C tyr frag. 2 216 C T H3 frag. 1 3 C T H1 frag. 8 221 Ð C H1 frag. 21 133 A C tyr frag. 2 222 T C rhod frag. 2 100 G C H1 frag. 8 223 Ð C H1 frag. 21 155 T C tyr frag. 2 233 A G tyr frag. 1 G A H1 frag. 8 345 G A H1 frag. 22 8 T A tyr frag. 2 266 A G tyr frag. 1 28 T C H1 frag. 8 488 A Ð H1 frag. 22 20 T C tyr frag. 3 4 G A tyr frag. 2 258 A G H1 frag. 8 514 C T H1 frag. 22 53 C T tyr frag. 3 56 G T tyr frag. 2 276 C T H1 frag. 8 551 A T H1 frag. 22 54 T C tyr frag. 3 94 C T tyr frag. 3 98 G C H1 frag. 8 568 A C H1 frag. 22 55 T C tyr frag. 3 128 T C 184 (Ceratobatrachidae) H1 frag. 8 696 A Ð H1 frag. 22 63 G A tyr frag. 3 173 T C 28S frag. 3 134 T G H1 frag. 8 732 Ð C H1 frag. 23 25 T A 191 (Ametrobatrachia) H1 frag. 10 269 C T H1 frag. 8 735 G A H1 frag. 23 38 T C H1 frag. 10 261 C T H1 frag. 11 7 G A H1 frag. 8 796 C G H1 frag. 23 103 A C H1 frag. 11 467 Ð A H1 frag. 11 16 A T H1 frag. 8 804 A T H1 frag. 23 129 C T H1 frag. 11 963 Ð T H1 frag. 11 505 C T H1 frag. 8 831 G A H1 frag. 23 149 A G H1 frag. 23 963 A T H1 frag. 11 852 C T H1 frag. 9 104 C A H1 frag. 23 152 A G H1 frag. 23 1062 Ð T H1 frag. 12 128 T C H1 frag. 9 326 Ð C H1 frag. 23 184 C A H1 frag. 23 1077 Ð A H1 frag. 13 91 T C H1 frag. 9 453 A T H1 frag. 23 258 Ð G H1 frag. 23 1781 C T 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 339

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H1 frag. 7 36 Ð T SIA frag. 3 78 G A 209 (Pyxicephalidae) H1 frag. 23 623 A C H1 frag. 8 232 A C SIA frag. 3 99 G A 28S frag. 3 424 G C H1 frag. 23 729 T C H1 frag. 8 352 A G SIA frag. 3 105 A G H1 frag. 10 52 T A H1 frag. 23 883 C Ð H1 frag. 8 387 Ð C SIA frag. 3 120 G T H1 frag. 10 101 G A H1 frag. 23 1077 A T H1 frag. 9 202 C T SIA frag. 3 153 A G H1 frag. 11 89 C T H1 frag. 23 1097 G A H1 frag. 9 315 A T SIA frag. 4 61 T C H1 frag. 11 264 C T H1 frag. 23 1154 C T H1 frag. 9 492 A T tyr frag. 1 4 A C H1 frag. 13 127 T A H1 frag. 23 1181 C A H1 frag. 9 788 A T tyr frag. 1 13 C T H1 frag. 14 85 C A H1 frag. 23 1214 T C H1 frag. 9 798 A T tyr frag. 1 28 C T H1 frag. 14 142 T C H1 frag. 23 1334 T C tyr frag. 3 155 A G tyr frag. 1 46 G T H1 frag. 15 25 A G H1 frag. 23 1376 C T 193 (Phrynobatrachidae) tyr frag. 2 80 C T H1 frag. 16 513 Ð T H1 frag. 23 1386 A T H1 frag. 1 50 A G tyr frag. 2 157 A C H1 frag. 17 388 Ð G H1 frag. 23 1554 T Ð H1 frag. 1 66 C T tyr frag. 2 159 C T H1 frag. 17 437 C T H1 frag. 23 1569 A Ð H1 frag. 10 55 T A tyr frag. 2 171 C T H1 frag. 18 260 Ð C H1 frag. 23 1722 G A H1 frag. 10 260 C A tyr frag. 2 193 C A H1 frag. 19 369 T Ð H1 frag. 23 1798 T A H1 frag. 11 27 A G tyr frag. 2 213 C T H1 frag. 19 771 C T H1 frag. 23 1803 G Ð H1 frag. 11 57 A Ð tyr frag. 2 241 T A H1 frag. 2 162 A T H1 frag. 23 1816 G A H1 frag. 11 457 C T tyr frag. 3 39 A G H1 frag. 21 133 A C H1 frag. 23 1824 T C H1 frag. 11 694 C Ð tyr frag. 3 134 A G H1 frag. 23 152 A G H1 frag. 24 17 C T H1 frag. 11 983 C A 200 (Pyxicephaloidea) H1 frag. 23 644 C A H1 frag. 24 19 C A H1 frag. 11 1045 A T 28S frag. 2 473 T C H1 frag. 23 1088 G C H1 frag. 3 58 T A H1 frag. 11 1093 A C H1 frag. 1 38 T C H1 frag. 23 1288 T C H1 frag. 3 262 C T H1 frag. 12 120 C A H1 frag. 11 887 A T H1 frag. 23 1776 A T H1 frag. 3 268 C T H1 frag. 13 110 Ð C H1 frag. 11 953 C A H1 frag. 3 209 Ð T H1 frag. 3 362 G A H1 frag. 13 127 T Ð H1 frag. 11 1232 A C H1 frag. 3 313 Ð A H1 frag. 3 391 A G H1 frag. 14 31 T A H1 frag. 16 359 C T H1 frag. 3 342 T C H1 frag. 4 13 A G H1 frag. 14 104 T C H1 frag. 17 81 A Ð H1 frag. 6 26 A T H1 frag. 4 137 Ð A H1 frag. 16 72 G A H1 frag. 18 411 A T H1 frag. 8 272 A Ð H1 frag. 4 325 T C H1 frag. 16 576 A T H1 frag. 18 628 T C H3 frag. 1 193 C A H1 frag. 4 383 Ð C H1 frag. 17 210 A C H1 frag. 18 741 C A rhod frag. 1 21 T C H1 frag. 4 404 T A H1 frag. 17 333 A Ð H1 frag. 19 147 C T rhod frag. 1 104 C T H1 frag. 4 414 A Ð H1 frag. 17 392 Ð G H1 frag. 20 68 G A rhod frag. 1 165 C G H1 frag. 4 452 A C H1 frag. 18 254 T A H1 frag. 23 789 A T tyr frag. 2 64 A T H1 frag. 4 585 Ð C H1 frag. 18 447 T A H1 frag. 23 1346 A T tyr frag. 2 67 C G H1 frag. 4 649 A C H1 frag. 18 766 C T H1 frag. 23 1951 A T tyr frag. 2 97 A T H1 frag. 4 673 A C H1 frag. 18 767 C T H1 frag. 3 26 T C tyr frag. 3 51 G A H1 frag. 5 3 C A H1 frag. 18 861 G A H1 frag. 4 100 T A tyr frag. 3 120 C T H1 frag. 6 34 G A 210 (Pyxicephalinae) H1 frag. 6 167 T C H1 frag. 18 863 G A H1 frag. 4 191 C A H1 frag. 1 20 A G H3 frag. 1 6 G A H1 frag. 19 172 A C H1 frag. 6 62 A T H1 frag. 11 67 C Ð H3 frag. 1 45 C T H1 frag. 19 542 A C H1 frag. 6 91 C T H1 frag. 11 141 A T H3 frag. 1 57 C T H1 frag. 19 651 T C H1 frag. 8 568 A C H1 frag. 11 467 A Ð H3 frag. 1 99 C T H1 frag. 19 822 A C H1 frag. 8 628 A T H1 frag. 11 983 C Ð H3 frag. 1 189 C G H1 frag. 2 118 C A H1 frag. 9 188 Ð C H1 frag. 11 1327 T A H3 frag. 2 42 C G H1 frag. 2 162 A C H1 frag. 9 202 T A H1 frag. 12 153 T A rhod frag. 1 9 T C H1 frag. 21 76 T A H1 frag. 9 609 C T H1 frag. 13 69 C A rhod frag. 1 36 A G H1 frag. 23 40 T C 201 (Petropedetidae) H1 frag. 13 117 C T rhod frag. 1 125 C T H1 frag. 23 182 C T H1 frag. 1 52 C T H1 frag. 14 72 A Ð rhod frag. 2 21 C T H1 frag. 23 299 C T H1 frag. 11 57 A T H1 frag. 14 78 A Ð rhod frag. 2 28 C T H1 frag. 23 559 C T H1 frag. 11 130 T C H1 frag. 14 149 A G rhod frag. 2 33 G A H1 frag. 23 729 T C H1 frag. 11 392 A C H1 frag. 14 217 T A rhod frag. 2 112 C T H1 frag. 23 1074 T A H1 frag. 11 483 Ð T H1 frag. 15 5 G T SIA frag. 3 82 G A H1 frag. 23 1097 G A H1 frag. 12 80 C A H1 frag. 15 19 T C SIA frag. 3 102 C T H1 frag. 23 1427 A Ð H1 frag. 13 87 Ð A H1 frag. 15 23 C A SIA frag. 3 153 A G H1 frag. 23 1554 T Ð H1 frag. 13 172 C A H1 frag. 16 2 T C SIA frag. 4 22 A T H1 frag. 3 155 G Ð H1 frag. 16 333 A C H1 frag. 16 221 C T SIA frag. 4 55 G A H1 frag. 3 266 C A H1 frag. 16 535 A C H1 frag. 16 266 T G tyr frag. 1 4 A C H1 frag. 3 339 Ð C H1 frag. 17 118 A T H1 frag. 16 292 A C tyr frag. 1 18 C T H1 frag. 4 673 A C H1 frag. 18 153 Ð T H1 frag. 16 563 A C tyr frag. 1 28 C T H1 frag. 6 49 T A H1 frag. 18 276 T A H1 frag. 17 231 T A tyr frag. 1 29 A G H1 frag. 6 137 C Ð H1 frag. 18 322 C T H1 frag. 17 274 A T tyr frag. 1 46 G T H1 frag. 6 213 A C H1 frag. 18 539 A C H1 frag. 17 372 T A tyr frag. 1 55 C T H1 frag. 8 10 T C H1 frag. 23 1427 A G H1 frag. 18 164 T C tyr frag. 2 20 G A H1 frag. 8 93 C Ð H1 frag. 23 1607 A T H1 frag. 18 254 T C tyr frag. 2 47 C T H1 frag. 8 122 A C H1 frag. 23 1811 C T H1 frag. 18 349 T Ð tyr frag. 2 83 C T H1 frag. 8 249 T Ð H1 frag. 3 8 C A H1 frag. 18 411 T C tyr frag. 2 144 G A H1 frag. 8 554 A G H1 frag. 3 370 G A H1 frag. 18 870 C T tyr frag. 2 153 G A H1 frag. 8 569 A G H1 frag. 3 415 T C H1 frag. 19 42 T C tyr frag. 2 159 C T H1 frag. 8 714 A Ð H1 frag. 4 4 T C H1 frag. 19 160 A T tyr frag. 2 194 C A H1 frag. 8 816 T Ð H1 frag. 4 292 T A H1 frag. 19 175 A C tyr frag. 2 213 C T H1 frag. 8 838 C T H1 frag. 4 505 Ð A H1 frag. 19 213 T A tyr frag. 2 234 C T H1 frag. 9 432 T A H1 frag. 8 29 T C H1 frag. 19 216 T A tyr frag. 2 237 C T H1 frag. 9 453 A T H1 frag. 9 226 A T H1 frag. 19 278 A C tyr frag. 3 31 C T H1 frag. 9 708 A C H1 frag. 9 564 Ð C H1 frag. 19 496 A T tyr frag. 3 70 T A H1 frag. 9 739 A G H3 frag. 2 39 A C H1 frag. 20 176 A G tyr frag. 3 167 C T H3 frag. 1 168 C A SIA frag. 3 42 T A H1 frag. 21 57 A C 212 (Cacosterninae) rhod frag. 1 54 A G SIA frag. 3 120 G A H1 frag. 23 22 T C 28S frag. 2 315 Ð G rhod frag. 1 171 C T tyr frag. 1 52 T A H1 frag. 23 140 T C 28S frag. 2 496 Ð T rhod frag. 2 79 C T tyr frag. 2 104 T C H1 frag. 23 224 C G 28S frag. 2 613 C G SIA frag. 3 51 T C tyr frag. 2 165 T C H1 frag. 23 299 C Ð 28S frag. 2 724 Ð C 340 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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28S frag. 2 727 Ð C H1 frag. 4 649 A G H1 frag. 11 622 A T H1 frag. 11 1316 A T 28S frag. 2 729 Ð A H1 frag. 5 7 T C H1 frag. 11 1035 Ð C H1 frag. 11 1327 T G H1 frag. 1 50 A G H1 frag. 7 96 A C H1 frag. 12 52 C A H1 frag. 12 136 Ð C H1 frag. 1 70 A G H1 frag. 8 511 A G H1 frag. 12 143 A C H1 frag. 12 148 T Ð H1 frag. 11 762 A Ð tyr frag. 2 183 C T H1 frag. 13 8 G T H1 frag. 12 153 A C H1 frag. 11 819 C T tyr frag. 3 2 C A H1 frag. 13 130 C T H1 frag. 12 187 G A H1 frag. 12 21 A G 220 (Saukrobatrachia) H1 frag. 14 13 T A H1 frag. 13 125 A T H1 frag. 16 301 Ð T H1 frag. 11 663 T A H1 frag. 14 127 A G H1 frag. 14 93 T C H1 frag. 18 116 A C H1 frag. 11 1161 A C H1 frag. 14 208 A T H1 frag. 16 382 T C H1 frag. 18 488 T A H1 frag. 16 313 C T H1 frag. 15 34 G A H1 frag. 16 585 Ð C H1 frag. 18 604 A C H1 frag. 18 254 T Ð H1 frag. 15 56 T C H1 frag. 17 47 T A H1 frag. 18 648 C A H1 frag. 18 411 A C H1 frag. 16 94 T C H1 frag. 18 110 Ð C H1 frag. 19 145 A T H1 frag. 18 877 T C H1 frag. 16 305 Ð C H1 frag. 18 322 C A H1 frag. 19 148 T G H1 frag. 19 560 T A H1 frag. 16 658 A G H1 frag. 18 488 T A H1 frag. 19 406 Ð T H1 frag. 19 749 C A H1 frag. 22 6 G T H1 frag. 18 530 C A H1 frag. 19 695 A T H1 frag. 2 210 A G H1 frag. 22 8 T G H1 frag. 18 717 C T H1 frag. 2 171 C T H1 frag. 2 328 C A H1 frag. 22 17 C G H1 frag. 18 767 C T H1 frag. 2 312 Ð T H1 frag. 2 437 T C H1 frag. 22 20 T C H1 frag. 18 861 G A H1 frag. 2 407 T A H1 frag. 23 452 C T H1 frag. 23 67 G A H1 frag. 19 771 C T H1 frag. 21 190 A G H1 frag. 23 1181 C A H1 frag. 23 81 G A H1 frag. 2 360 T C H1 frag. 23 25 T C H1 frag. 23 1245 C T H1 frag. 23 1041 A T H1 frag. 20 73 A G H1 frag. 23 182 C T H1 frag. 23 1358 C T H1 frag. 23 1221 C T H1 frag. 22 23 G A H1 frag. 23 602 T A H1 frag. 3 214 T G H1 frag. 23 1811 C T H1 frag. 23 38 T C H1 frag. 23 1245 C T H1 frag. 4 223 C A H1 frag. 23 1825 A T H1 frag. 23 52 C T H1 frag. 23 1732 T C H1 frag. 4 306 A C H1 frag. 23 1919 A T H1 frag. 23 69 T C H1 frag. 23 1743 A G H1 frag. 8 211 Ð A H1 frag. 24 19 C A H1 frag. 23 83 A G H1 frag. 3 8 C T H1 frag. 8 711 G T rhod frag. 1 78 G A H1 frag. 23 113 T C H1 frag. 3 47 C T H1 frag. 9 228 Ð G rhod frag. 1 134 G C H1 frag. 23 224 C T H1 frag. 4 283 C T H1 frag. 9 383 C Ð rhod frag. 2 80 G A H1 frag. 23 602 T A H1 frag. 4 343 A G H1 frag. 9 672 T C SIA frag. 3 9 T C H1 frag. 23 896 Ð A H1 frag. 6 213 A C H3 frag. 1 114 T C SIA frag. 3 42 T C H1 frag. 23 1233 G A H1 frag. 7 13 A T H3 frag. 2 33 G C SIA frag. 3 54 T C H1 frag. 23 1478 A C H1 frag. 7 39 C T H3 frag. 2 60 G C SIA frag. 3 138 T C H1 frag. 23 1882 A G H1 frag. 8 69 C T 221 (Dicroglossidae) SIA frag. 3 160 C T H1 frag. 23 1962 T C H1 frag. 8 122 A C H1 frag. 10 14 T C SIA frag. 4 76 T C H1 frag. 3 22 A C H1 frag. 8 313 T C H1 frag. 10 19 A G SIA frag. 4 79 T G H1 frag. 3 150 A T H1 frag. 8 628 T G H1 frag. 11 320 Ð T 225 (Dicroglossinae) H1 frag. 3 160 A G H1 frag. 9 73 C A H1 frag. 11 409 T A H1 frag. 11 53 T G H1 frag. 3 252 C T H1 frag. 9 212 Ð G H1 frag. 11 983 C T H1 frag. 12 80 C A H1 frag. 3 342 T C H1 frag. 9 226 A C H1 frag. 12 103 A T H1 frag. 13 16 A T H1 frag. 3 361 G A H1 frag. 9 716 Ð G H1 frag. 13 172 C A H1 frag. 13 43 Ð G H1 frag. 3 365 T C H1 frag. 9 739 A G H1 frag. 14 25 Ð C H1 frag. 13 55 A C H1 frag. 4 673 A C rhod frag. 1 78 G A H1 frag. 14 63 A G H1 frag. 14 6 C T H1 frag. 5 6 C T rhod frag. 1 85 A C H1 frag. 14 89 T C H1 frag. 14 90 T A H1 frag. 6 25 T C rhod frag. 1 131 C T H1 frag. 16 382 A T H1 frag. 14 177 C T H1 frag. 6 27 G A rhod frag. 2 147 T G H1 frag. 17 210 A C H1 frag. 15 15 A T H1 frag. 8 29 T C SIA frag. 1 12 A G H1 frag. 17 407 T Ð H1 frag. 17 333 A C H1 frag. 8 37 C A SIA frag. 2 14 A G H1 frag. 18 116 A T H1 frag. 18 138 A T H1 frag. 8 52 A C tyr frag. 1 75 T C H1 frag. 18 490 Ð C H1 frag. 18 550 A C H1 frag. 8 69 C T tyr frag. 2 66 A G H1 frag. 18 868 A C H1 frag. 18 886 G A tyr frag. 2 68 T C H1 frag. 19 449 T C H1 frag. 19 42 A T H1 frag. 8 556 T G tyr frag. 2 183 T C H1 frag. 19 729 C Ð H1 frag. 19 212 C T H1 frag. 8 667 A T 218 (Amietia) H1 frag. 21 251 A T H1 frag. 19 757 Ð C H1 frag. 8 816 T A H1 frag. 1 70 G A H1 frag. 21 270 G A H1 frag. 21 277 C A H1 frag. 9 42 G A H1 frag. 11 642 Ð C H1 frag. 23 1060 C A H1 frag. 22 15 A G H1 frag. 9 71 T A H1 frag. 11 819 T C H1 frag. 23 1169 C T H1 frag. 23 293 T Ð H1 frag. 9 460 Ð T H1 frag. 11 963 T C H1 frag. 23 1427 A G H1 frag. 23 623 A C H3 frag. 2 33 C G H1 frag. 14 144 A G H1 frag. 23 1518 C Ð H1 frag. 23 1154 A G H3 frag. 2 36 A C H1 frag. 15 42 A T H1 frag. 25 53 C T H1 frag. 23 1183 Ð C H3 frag. 2 42 C G H1 frag. 16 359 T C H1 frag. 25 69 G A H1 frag. 23 1569 A T H3 frag. 2 60 C G H1 frag. 16 614 T A H1 frag. 7 40 A Ð H1 frag. 23 1732 T C rhod frag. 1 142 A G H1 frag. 17 167 Ð T H1 frag. 8 116 T A H1 frag. 23 1846 C T rhod frag. 2 91 C T H1 frag. 18 260 C A H1 frag. 9 343 A T H1 frag. 6 213 A Ð SIA frag. 1 42 G A H1 frag. 18 411 T C H1 frag. 9 521 Ð C H1 frag. 7 13 A T SIA frag. 3 48 G A H1 frag. 18 488 A T H1 frag. 9 693 C A H1 frag. 8 47 C A SIA frag. 3 78 G A H1 frag. 18 891 T A rhod frag. 1 10 A G H1 frag. 8 306 C T SIA frag. 3 108 C T H1 frag. 19 201 A G rhod frag. 1 72 G A H1 frag. 8 553 A G SIA frag. 3 168 A C H1 frag. 19 449 T C rhod frag. 2 57 C T H1 frag. 8 568 A C SIA frag. 4 31 G A H1 frag. 2 215 C Ð SIA frag. 3 120 G A H1 frag. 9 432 T C tyr frag. 1 4 A C H1 frag. 21 133 C G SIA frag. 3 156 T A H1 frag. 9 545 C Ð tyr frag. 1 11 A G H1 frag. 23 114 A T tyr frag. 1 85 C T SIA frag. 3 3 G A tyr frag. 1 44 T A H1 frag. 23 294 Ð C tyr frag. 3 53 T C SIA frag. 3 141 T C tyr frag. 1 73 C T H1 frag. 23 559 C T tyr frag. 3 167 C T tyr frag. 2 31 T C tyr frag. 2 87 A C H1 frag. 23 1464 Ð T 222 (Occidozyginae) tyr frag. 3 98 C T tyr frag. 2 100 C T H1 frag. 23 1828 A T H1 frag. 10 55 T C 226 (Limnonectini) tyr frag. 2 124 T C H1 frag. 24 10 A T H1 frag. 10 205 Ð A H1 frag. 1 64 T C tyr frag. 2 285 G T H1 frag. 3 20 T C H1 frag. 11 57 A T H1 frag. 10 101 G A tyr frag. 3 64 A C H1 frag. 3 51 G A H1 frag. 11 92 T C H1 frag. 11 89 C T tyr frag. 3 88 T G H1 frag. 3 126 C T H1 frag. 11 392 A Ð H1 frag. 11 146 C T tyr frag. 3 115 C T H1 frag. 4 298 A G H1 frag. 11 595 A Ð H1 frag. 11 320 T A tyr frag. 3 128 T C 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 341

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228 (unnamed taxon) H1 frag. 23 644 C T H1 frag. 21 57 A C H1 frag. 4 292 T A H1 frag. 10 55 T A H1 frag. 24 5 T C H1 frag. 21 90 A T H1 frag. 4 325 T A H1 frag. 10 217 T C H1 frag. 24 33 A G H1 frag. 22 61 A G H1 frag. 4 365 T C H1 frag. 11 561 Ð T H1 frag. 4 94 A T H1 frag. 23 105 A T H1 frag. 4 434 A G H1 frag. 11 1093 A C H1 frag. 4 169 C A H1 frag. 23 963 T A H1 frag. 4 458 A C H1 frag. 12 127 T A H1 frag. 4 561 Ð T H1 frag. 23 1181 A C H1 frag. 7 11 G A H1 frag. 13 82 T C H1 frag. 6 49 T A H1 frag. 23 1233 G C H1 frag. 8 94 Ð T H1 frag. 13 172 A C H1 frag. 8 122 A T H1 frag. 23 1386 A Ð H1 frag. 8 174 A G H1 frag. 14 127 A G H1 frag. 8 369 T Ð H1 frag. 23 1743 A C H1 frag. 8 201 T A H1 frag. 14 128 G A H1 frag. 8 488 A C H1 frag. 23 1816 G A H1 frag. 8 249 A C H1 frag. 16 39 T C H1 frag. 8 735 G A H1 frag. 23 1846 C T H1 frag. 8 294 T C H1 frag. 16 672 A G H1 frag. 9 233 Ð C H1 frag. 25 34 C A H1 frag. 8 300 G A H1 frag. 17 26 A G H1 frag. 9 533 A C H1 frag. 4 223 A T H1 frag. 8 313 T C H1 frag. 17 38 A T H1 frag. 9 708 A G H1 frag. 4 592 T C H1 frag. 8 545 T C H1 frag. 18 322 A T H3 frag. 1 15 C A H1 frag. 7 54 C T H1 frag. 8 726 T C H1 frag. 18 514 A C H3 frag. 2 18 A T H1 frag. 8 544 T C H1 frag. 8 788 Ð T H1 frag. 18 746 C T H3 frag. 2 19 G C H1 frag. 9 392 Ð C H1 frag. 8 804 A G H1 frag. 19 695 A C 245 (Rhacophoroidea) H1 frag. 9 493 Ð A H1 frag. 8 838 C T H1 frag. 2 218 T C H1 frag. 11 467 A T H3 frag. 1 78 C A H1 frag. 9 42 G T H1 frag. 2 371 C T H1 frag. 11 1139 Ð T rhod frag. 1 104 C T H1 frag. 9 54 T C H1 frag. 22 43 C T H1 frag. 13 35 Ð A rhod frag. 1 144 C T H1 frag. 9 67 C T H1 frag. 22 62 G A H1 frag. 16 127 T C rhod frag. 2 80 G A H1 frag. 9 506 A Ð H1 frag. 23 559 C Ð H1 frag. 16 313 T A rhod frag. 2 134 C G H3 frag. 1 81 A G H1 frag. 23 1077 A T H1 frag. 16 429 C A tyr frag. 1 28 C T H3 frag. 1 126 A G H1 frag. 23 1338 C A H1 frag. 16 509 A C tyr frag. 1 40 T A H3 frag. 1 195 C A H1 frag. 23 1478 C T H1 frag. 18 322 C T tyr frag. 1 55 T C H3 frag. 1 243 T C H1 frag. 23 1834 A C H1 frag. 18 563 A T tyr frag. 2 100 C T H3 frag. 2 5 T G H1 frag. 3 214 G A H1 frag. 18 830 A C 247 (Boophinae/Boophis) H3 frag. 2 18 T G H1 frag. 4 100 T C H1 frag. 18 838 T C H1 frag. 1 36 A G rhod frag. 1 165 C T H1 frag. 7 39 C T H1 frag. 23 452 T A H1 frag. 1 48 A T 248 (Mantellinae) H1 frag. 8 816 A C H1 frag. 23 623 A Ð H1 frag. 10 261 T C H1 frag. 11 315 A T H1 frag. 9 281 A T H1 frag. 23 1015 A T H1 frag. 11 134 A T H1 frag. 11 368 C T H1 frag. 9 609 C T H1 frag. 23 1776 A G H1 frag. 11 317 Ð C H1 frag. 11 422 Ð A H1 frag. 9 618 A T H1 frag. 3 108 A T H1 frag. 11 887 A T H1 frag. 11 457 T C H1 frag. 9 693 A T H1 frag. 3 124 C A H1 frag. 11 1071 C T H1 frag. 11 983 C Ð 232 (Dicroglossini) H1 frag. 3 331 Ð T H1 frag. 13 82 T C H1 frag. 11 1135 C Ð H1 frag. 11 409 T G H1 frag. 4 120 T A H1 frag. 13 126 A G H1 frag. 12 9 C A H1 frag. 11 513 Ð A H1 frag. 4 191 C T H1 frag. 14 193 T Ð H1 frag. 13 156 T A H1 frag. 11 598 T C H1 frag. 4 404 T C H1 frag. 15 40 C T H1 frag. 14 144 A G H1 frag. 11 606 Ð C H1 frag. 4 408 A T H1 frag. 16 36 T C H1 frag. 14 208 C T H1 frag. 11 831 Ð C H1 frag. 4 439 A T H1 frag. 16 94 T C H1 frag. 15 22 T C H1 frag. 11 983 T C H1 frag. 8 201 Ð T H1 frag. 16 361 Ð A H1 frag. 18 145 Ð T H1 frag. 11 1146 Ð A H1 frag. 8 249 T A H1 frag. 16 676 A G H1 frag. 18 411 C Ð H1 frag. 12 21 A G H1 frag. 8 568 A C H1 frag. 17 38 A G H1 frag. 18 530 C A H1 frag. 12 52 A C H1 frag. 9 104 C A H1 frag. 18 93 T C H1 frag. 18 654 T A H1 frag. 21 133 C A H3 frag. 1 168 C A H1 frag. 18 185 T C H1 frag. 19 42 A T H1 frag. 23 2 C G H3 frag. 1 184 C A H1 frag. 18 283 Ð A H1 frag. 19 449 T C H1 frag. 23 644 C T H3 frag. 1 186 C A H1 frag. 18 868 A C H1 frag. 2 240 G A H1 frag. 23 981 C T tyr frag. 1 29 A G H1 frag. 19 531 A T H1 frag. 2 425 C T H1 frag. 23 1411 C T tyr frag. 2 53 C T H1 frag. 19 542 A T H1 frag. 23 420 Ð T H1 frag. 6 22 C T tyr frag. 2 98 C A H1 frag. 2 16 A C H1 frag. 23 710 Ð T H1 frag. 8 233 A T tyr frag. 2 198 C T H1 frag. 2 118 T C H1 frag. 23 789 A T H1 frag. 8 281 T Ð tyr frag. 3 32 T G H1 frag. 2 241 C T H1 frag. 23 1334 A C H1 frag. 8 298 T Ð tyr frag. 3 98 C T H1 frag. 20 66 A G H1 frag. 23 1704 C T H1 frag. 9 43 T C 246 (Mantellidae) H1 frag. 20 68 A G H1 frag. 4 106 C Ð H1 frag. 9 54 C T H1 frag. 11 602 Ð T H1 frag. 20 73 A G H1 frag. 4 149 Ð A H1 frag. 9 202 A C H1 frag. 12 80 C A H1 frag. 20 151 A G H1 frag. 5 10 Ð G H1 frag. 9 367 A T H1 frag. 13 40 Ð C H1 frag. 20 180 C T H1 frag. 5 16 A T H1 frag. 9 798 G A H1 frag. 14 72 A Ð H1 frag. 23 21 G A H1 frag. 7 36 T G rhod frag. 2 139 T C H1 frag. 16 2 T C H1 frag. 23 23 T C H1 frag. 8 33 C T tyr frag. 1 4 A G H1 frag. 16 170 T C H1 frag. 23 38 T C H1 frag. 8 387 C A tyr frag. 1 67 C T H1 frag. 16 221 C A H1 frag. 23 40 T C H1 frag. 8 425 Ð C tyr frag. 2 84 G A H1 frag. 16 547 C T H1 frag. 23 43 T C H1 frag. 9 71 T A tyr frag. 2 261 T C H1 frag. 18 144 Ð A H1 frag. 23 52 C T H3 frag. 1 18 G C tyr frag. 3 119 T C H1 frag. 18 752 A G H1 frag. 23 224 C T H3 frag. 1 117 G C 214 (Aglaioanura) H1 frag. 18 758 G A H1 frag. 23 293 T C H3 frag. 1 204 C A H1 frag. 11 457 C T H1 frag. 18 761 T C H1 frag. 23 764 Ð T H3 frag. 2 20 C A H1 frag. 11 762 A C H1 frag. 18 870 C T H1 frag. 23 1041 A T tyr frag. 1 21 T C H1 frag. 12 4 A G H1 frag. 18 877 C T H1 frag. 23 1062 T G 249 (Laliostomini) H1 frag. 12 14 T C H1 frag. 19 54 T A H1 frag. 23 1124 A G 28S frag. 2 714 G C H1 frag. 14 208 A C H1 frag. 19 148 A T H1 frag. 23 1226 T A H1 frag. 10 159 A G H1 frag. 14 217 T A H1 frag. 19 154 A G H1 frag. 23 1722 G A H1 frag. 11 23 T C H1 frag. 16 535 A T H1 frag. 19 164 A C H1 frag. 23 1811 C T H1 frag. 11 79 A G H1 frag. 17 333 A T H1 frag. 19 181 G A H1 frag. 23 1828 A T H1 frag. 11 1217 A C H1 frag. 18 143 Ð C H1 frag. 19 209 C T H1 frag. 23 1951 A T H1 frag. 13 127 T A H1 frag. 18 371 A Ð H1 frag. 19 216 T C H1 frag. 3 58 T A H1 frag. 14 89 T C H1 frag. 2 118 C T H1 frag. 2 20 C A H1 frag. 3 372 T A H1 frag. 14 177 C T H1 frag. 2 162 A T H1 frag. 2 90 Ð A H1 frag. 3 402 T C H1 frag. 16 1 A G H1 frag. 2 238 C T H1 frag. 2 362 Ð A H1 frag. 4 215 Ð A H1 frag. 16 201 T C H1 frag. 20 68 G A H1 frag. 20 23 C A H1 frag. 4 271 T C H1 frag. 16 566 Ð T 342 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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H1 frag. 17 33 A T H1 frag. 23 1342 A T 256 (Kurixalus) H1 frag. 24 33 G A H1 frag. 17 103 A T H1 frag. 23 1676 C T 28S frag. 2 753 C T H1 frag. 25 30 C T H1 frag. 17 306 A C H1 frag. 4 100 T A H1 frag. 1 68 T C H1 frag. 25 34 C A H1 frag. 18 125 Ð G H1 frag. 4 306 C A H1 frag. 10 43 A C H1 frag. 25 41 A G H1 frag. 18 126 Ð T H1 frag. 5 9 A T H1 frag. 10 143 A Ð H1 frag. 3 26 T C H1 frag. 18 209 A G H1 frag. 5 28 G A H1 frag. 11 23 T C H1 frag. 3 141 T C H1 frag. 18 322 T Ð H1 frag. 6 25 T A H1 frag. 11 368 C A H1 frag. 3 150 A G H1 frag. 18 494 C T H1 frag. 6 213 A C H1 frag. 11 439 T A H1 frag. 3 266 C T H1 frag. 18 581 C A H1 frag. 8 47 C A H1 frag. 11 536 G T H1 frag. 3 268 C T H1 frag. 18 717 C T H1 frag. 9 27 A G H1 frag. 11 663 C Ð H1 frag. 3 321 G A H1 frag. 19 83 C A H1 frag. 9 46 T A H1 frag. 11 762 C Ð H1 frag. 4 316 C T H1 frag. 19 181 A T H1 frag. 9 49 T C H1 frag. 11 852 C Ð H1 frag. 4 373 G A H1 frag. 19 278 A T H1 frag. 9 59 T A H1 frag. 11 958 Ð T H1 frag. 4 380 C T H1 frag. 19 286 A C H1 frag. 9 73 A T H1 frag. 11 959 Ð T H1 frag. 4 463 A T H1 frag. 19 424 A G H1 frag. 9 256 T C H1 frag. 11 1138 Ð T H1 frag. 5 32 T C H1 frag. 19 749 A C H1 frag. 9 281 A C H1 frag. 11 1248 A T H1 frag. 7 35 C T H1 frag. 19 771 C T H1 frag. 9 498 Ð C H1 frag. 11 1327 T G H1 frag. 7 43 G A H1 frag. 2 171 C A 254 (Rhacophorinae) H1 frag. 12 80 C T H1 frag. 7 79 Ð T H1 frag. 2 360 T C H1 frag. 11 600 G Ð H1 frag. 12 103 A T H1 frag. 7 80 Ð A H1 frag. 21 260 A T H1 frag. 11 963 T C H1 frag. 12 174 A G H1 frag. 7 87 A C H1 frag. 23 1427 A T H1 frag. 11 1056 Ð T H1 frag. 13 198 C T H1 frag. 7 96 A T H1 frag. 23 1478 A G H1 frag. 11 1113 A T H1 frag. 14 68 Ð A H1 frag. 8 37 C T H1 frag. 23 1919 A T H1 frag. 11 1217 A Ð H1 frag. 14 208 C Ð H1 frag. 8 59 T C H1 frag. 3 26 T C H1 frag. 11 1324 A T H1 frag. 14 217 A T H1 frag. 8 62 T C H1 frag. 4 316 C T H1 frag. 12 148 T A H1 frag. 14 260 G A H1 frag. 8 122 T A H1 frag. 4 373 G A H1 frag. 13 69 C T H1 frag. 16 191 C A H1 frag. 8 352 G A H1 frag. 6 25 T C H1 frag. 13 123 A T H1 frag. 16 464 A C H1 frag. 8 553 A G H1 frag. 8 177 A G H1 frag. 13 168 C A H1 frag. 16 672 A T H1 frag. 8 568 C T H1 frag. 8 211 A C H1 frag. 14 106 A C H1 frag. 16 700 C A H1 frag. 8 626 C A H1 frag. 8 441 A T H1 frag. 15 3 A T H1 frag. 17 16 C A H1 frag. 8 711 T A H1 frag. 8 469 C A H1 frag. 16 7 A G H1 frag. 17 306 T A H1 frag. 8 816 T A H1 frag. 8 696 A G H1 frag. 16 243 Ð G H1 frag. 17 350 T A H1 frag. 8 838 C T H1 frag. 9 84 C T H1 frag. 16 400 Ð C H1 frag. 18 13 A G H1 frag. 9 52 C T H1 frag. 9 229 Ð T H1 frag. 16 535 T C H1 frag. 18 143 C T H1 frag. 9 409 C Ð H1 frag. 9 546 Ð T H1 frag. 16 614 T Ð H1 frag. 18 433 T Ð H1 frag. 9 498 C A H1 frag. 9 609 C Ð H1 frag. 17 47 T A H1 frag. 18 488 T A H1 frag. 9 558 C T H1 frag. 9 739 A G H1 frag. 18 140 Ð C H1 frag. 18 539 T A H1 frag. 9 693 C T tyr frag. 2 273 C T H1 frag. 18 767 C T H1 frag. 18 185 T A H1 frag. 9 755 C A tyr frag. 3 155 G A H1 frag. 18 861 G A H1 frag. 9 798 T C H1 frag. 18 838 C A 253 (Rhacophoridae) H1 frag. 19 197 A T H1 frag. 9 840 T C H1 frag. 18 866 G A H1 frag. 10 52 T A H1 frag. 19 203 A T rhod frag. 1 161 A T H1 frag. 19 91 T A H1 frag. 11 663 A C H1 frag. 19 212 C T rhod frag. 2 134 C G H1 frag. 19 376 T A H1 frag. 11 694 C A H1 frag. 19 224 A T tyr frag. 1 29 G C H1 frag. 19 427 T C H1 frag. 11 819 C T H1 frag. 19 350 A G tyr frag. 2 8 T A H1 frag. 19 622 A G H1 frag. 11 1093 A Ð H1 frag. 19 369 C A tyr frag. 2 36 T C H1 frag. 2 16 A C H1 frag. 13 163 A T H1 frag. 19 379 Ð T tyr frag. 2 50 G T H1 frag. 2 180 A C H1 frag. 15 34 G A H1 frag. 19 396 A T tyr frag. 2 77 C T H1 frag. 2 287 Ð A H1 frag. 16 31 A G H1 frag. 19 823 C A tyr frag. 2 168 C T H1 frag. 20 146 T C H1 frag. 16 72 G A H1 frag. 2 7 C T tyr frag. 2 213 C T H1 frag. 21 133 A C H1 frag. 16 242 Ð T H1 frag. 2 20 C A tyr frag. 2 266 A G H1 frag. 16 333 A T H1 frag. 23 25 T C H1 frag. 2 154 C T tyr frag. 2 277 C T H1 frag. 16 590 C Ð H1 frag. 23 529 Ð C H1 frag. 2 187 C A tyr frag. 3 2 C T H1 frag. 16 665 C T H1 frag. 23 602 T A H1 frag. 2 192 Ð A tyr frag. 3 28 T A H1 frag. 17 306 A T H1 frag. 23 644 T C H1 frag. 2 207 G A tyr frag. 3 31 C T H1 frag. 18 52 A T H1 frag. 23 854 A C H1 frag. 2 237 C T tyr frag. 3 63 A C H1 frag. 18 232 C T H1 frag. 23 1015 T Ð H1 frag. 2 301 A C tyr frag. 3 70 T C H1 frag. 18 306 A T H1 frag. 23 1781 T C H1 frag. 2 430 Ð T tyr frag. 3 158 G C H1 frag. 18 349 C A H1 frag. 23 1793 A T H1 frag. 2 439 C Ð tyr frag. 3 170 C T H1 frag. 18 457 Ð A H1 frag. 24 1 C T H1 frag. 21 57 A T 267 (Chiromantis) H1 frag. 18 604 A T H1 frag. 24 35 G A H1 frag. 22 5 G A H1 frag. 11 264 T C H1 frag. 18 615 C T H1 frag. 24 38 T C H1 frag. 22 18 C T H1 frag. 11 409 G Ð H1 frag. 18 650 Ð A H1 frag. 3 407 T A H1 frag. 22 20 T C H1 frag. 11 648 Ð C H1 frag. 18 732 C T H1 frag. 4 106 C T H1 frag. 23 48 C A H1 frag. 11 963 C Ð H1 frag. 18 878 T C H1 frag. 4 191 T A H1 frag. 23 81 G A H1 frag. 11 1089 T Ð H1 frag. 19 24 A T H1 frag. 4 439 T C H1 frag. 23 902 C T H1 frag. 12 74 T A H1 frag. 19 271 T A H1 frag. 4 562 Ð C H1 frag. 23 942 C A H1 frag. 12 120 C A H1 frag. 19 356 C T H1 frag. 7 36 T Ð H1 frag. 23 997 A C H1 frag. 14 64 C T H1 frag. 19 796 A T H1 frag. 7 97 C A H1 frag. 23 1154 C A H1 frag. 14 102 T C H1 frag. 19 826 T A H1 frag. 8 201 T C H1 frag. 23 1256 C T H1 frag. 16 27 T C H1 frag. 2 140 C A H1 frag. 8 313 T C H1 frag. 23 1359 G A H1 frag. 16 31 G C H1 frag. 2 154 T C H1 frag. 8 335 T G H1 frag. 23 1386 A C H1 frag. 16 266 T C H1 frag. 2 191 Ð T H1 frag. 8 469 C A H1 frag. 23 1478 A T H1 frag. 17 200 Ð A H1 frag. 2 196 C A H1 frag. 8 488 C A H1 frag. 23 1758 G A H1 frag. 17 306 T Ð H1 frag. 2 218 T A H1 frag. 8 523 T A H1 frag. 23 1811 C T H1 frag. 17 350 T A H1 frag. 23 163 C Ð H1 frag. 9 16 Ð T H1 frag. 23 1840 A C H1 frag. 17 446 C T H1 frag. 23 199 A T H1 frag. 9 28 A Ð H1 frag. 23 1974 C T H1 frag. 18 1 G A H1 frag. 23 250 A G H1 frag. 9 48 T Ð H1 frag. 23 1977 C T H1 frag. 18 140 C T H1 frag. 23 521 T A H1 frag. 9 55 Ð A H1 frag. 24 5 C T H1 frag. 18 185 A Ð H1 frag. 23 1214 T C H1 frag. 9 432 T C H1 frag. 24 16 C T H1 frag. 18 209 A T H1 frag. 23 1256 T C H1 frag. 9 558 A C H1 frag. 24 17 C T H1 frag. 18 581 C A 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 343

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H1 frag. 18 628 C A H1 frag. 19 560 A T H1 frag. 11 1266 A C H1 frag. 19 749 A C H1 frag. 18 868 A T H1 frag. 19 771 C T H1 frag. 21 83 Ð C H1 frag. 2 389 T C H1 frag. 19 651 C T H1 frag. 23 182 C A H1 frag. 23 942 C T H1 frag. 2 397 C T H1 frag. 2 7 C A H1 frag. 23 199 A T H1 frag. 23 1169 C T H1 frag. 20 62 G A H1 frag. 2 16 C A H1 frag. 23 224 C T H1 frag. 23 1338 C A H1 frag. 22 61 A G H1 frag. 2 44 G A H1 frag. 23 238 Ð T H1 frag. 23 1607 C T H1 frag. 22 62 G A H1 frag. 2 154 C T H1 frag. 23 789 A C H1 frag. 24 10 A G H1 frag. 23 11 C T H1 frag. 2 237 C T H1 frag. 23 1062 T Ð H1 frag. 6 81 C T H1 frag. 23 96 T C H1 frag. 2 328 T Ð H1 frag. 23 1181 A Ð H1 frag. 6 199 C T H1 frag. 23 584 Ð G H1 frag. 2 419 T C H1 frag. 23 1221 C A H1 frag. 7 13 A T H1 frag. 23 1834 A Ð H1 frag. 20 7 G A H1 frag. 23 1233 G A H1 frag. 8 249 T C H1 frag. 24 19 C A H1 frag. 20 25 A T H1 frag. 23 1245 T C H1 frag. 8 457 Ð A H1 frag. 3 47 T C H1 frag. 22 15 A G H1 frag. 23 1378 G Ð H1 frag. 8 542 G A H1 frag. 3 266 C A H1 frag. 22 45 T C H1 frag. 23 1607 A C H1 frag. 8 548 C T H1 frag. 3 342 G Ð H1 frag. 22 70 C T H1 frag. 23 1867 Ð A H1 frag. 8 792 A C H1 frag. 4 575 A C H1 frag. 23 103 C A H1 frag. 6 25 T C H1 frag. 9 247 Ð T H1 frag. 6 229 G Ð H1 frag. 23 1131 G A H1 frag. 6 46 A C H1 frag. 9 533 C Ð H1 frag. 8 40 A T H1 frag. 23 1169 C A H1 frag. 6 91 C T H1 frag. 9 681 Ð C H1 frag. 8 792 A C H1 frag. 23 1359 G A H1 frag. 7 35 C Ð H1 frag. 9 739 A G H1 frag. 9 315 T Ð H1 frag. 3 55 C A H1 frag. 7 39 C A rhod frag. 1 94 G A H1 frag. 9 400 A T H1 frag. 4 263 G A H1 frag. 8 139 C T rhod frag. 2 9 A G H1 frag. 9 638 T C H1 frag. 4 408 T A H1 frag. 8 161 C A rhod frag. 2 61 G A H3 frag. 1 117 G C H1 frag. 4 493 C T H1 frag. 8 748 A G 287 (unnamed taxon) rhod frag. 2 86 A T H1 frag. 4 670 A C H1 frag. 9 71 T C H1 frag. 11 27 A G SIA frag. 2 53 A G H1 frag. 6 62 A Ð H1 frag. 9 84 C T H1 frag. 11 72 C T SIA frag. 2 59 T C H1 frag. 8 10 T C H1 frag. 9 233 C A H1 frag. 11 88 T Ð SIA frag. 3 45 T C H1 frag. 8 569 A G rhod frag. 1 3 C T H1 frag. 11 392 A T tyr frag. 1 25 G A H1 frag. 8 626 C A rhod frag. 1 9 T C H1 frag. 11 1071 C Ð tyr frag. 1 58 C T H1 frag. 9 17 A G rhod frag. 1 134 G T H1 frag. 13 151 T C tyr frag. 2 41 C T H1 frag. 9 263 Ð A rhod frag. 2 33 G A H1 frag. 15 40 T A tyr frag. 2 85 T A H1 frag. 9 570 Ð A rhod frag. 2 112 C T H1 frag. 17 210 A T tyr frag. 3 22 A G rhod frag. 2 132 A C 274 (Hylarana) H1 frag. 17 306 A C tyr frag. 3 109 C T rhod frag. 2 134 C A H1 frag. 10 73 Ð T H1 frag. 18 328 Ð T tyr frag. 3 155 A C rhod frag. 2 136 T A H1 frag. 11 368 C T H1 frag. 18 433 A T 314 (Hyloides) 270 (Nyctibatrachidae) H1 frag. 11 409 C A H1 frag. 18 615 C A 28S frag. 2 354 Ð C H1 frag. 12 112 A G H1 frag. 11 457 T C H1 frag. 18 648 A T 28S frag. 2 483 Ð C H1 frag. 21 113 A C H1 frag. 11 663 A C H1 frag. 18 870 T A 28S frag. 3 379 Ð C H1 frag. 21 124 A C H1 frag. 13 69 C G H1 frag. 19 369 A T 28S frag. 3 385 Ð C H1 frag. 23 105 A C H1 frag. 14 177 C T H1 frag. 19 376 A T 28S frag. 3 389 Ð C H1 frag. 23 641 Ð T H1 frag. 16 152 T A H1 frag. 21 251 A T 28S frag. 3 487 Ð G H1 frag. 23 1676 C T H1 frag. 16 259 Ð G H1 frag. 23 105 A C H1 frag. 11 1294 T C H1 frag. 23 1977 C T H1 frag. 16 629 A T H1 frag. 23 587 Ð G H1 frag. 12 103 A T H1 frag. 25 14 G A H1 frag. 18 411 C A H1 frag. 23 693 C T H1 frag. 14 208 A T H1 frag. 25 87 C T H1 frag. 18 504 Ð T H1 frag. 23 789 C A H1 frag. 16 94 T A H1 frag. 7 13 A T H1 frag. 18 838 T C H1 frag. 23 1015 A G H1 frag. 16 414 A C H1 frag. 8 172 T C H1 frag. 19 61 G Ð H1 frag. 8 773 T C H1 frag. 17 54 C A H1 frag. 9 492 T C H1 frag. 19 338 A C H1 frag. 9 409 T C H1 frag. 17 372 T C H1 frag. 9 506 A T H1 frag. 19 449 T C H1 frag. 9 455 Ð G H1 frag. 18 232 C Ð rhod frag. 2 139 T A H1 frag. 2 301 A C H1 frag. 9 708 A G H1 frag. 18 322 A C tyr frag. 2 250 A C H1 frag. 21 90 A C 288 (Pelophylax) H1 frag. 18 632 Ð A tyr frag. 3 13 C T H1 frag. 23 152 A G H1 frag. 16 201 T A H1 frag. 18 727 C A tyr frag. 3 28 T C H1 frag. 23 250 A T H1 frag. 16 241 A G H1 frag. 18 782 A T tyr frag. 3 120 C A H1 frag. 23 439 Ð T H1 frag. 16 313 A C H1 frag. 19 109 C T 272 (Ranidae) H1 frag. 23 613 Ð C H1 frag. 16 547 C A H1 frag. 19 376 T C H1 frag. 10 35 Ð T H1 frag. 23 729 T Ð H1 frag. 17 350 T C H1 frag. 2 154 T C H1 frag. 11 409 T C H1 frag. 23 902 C A H1 frag. 17 372 T C H1 frag. 2 333 Ð A H1 frag. 11 467 A Ð H1 frag. 23 1041 A T H1 frag. 18 164 T C H1 frag. 20 68 G A H1 frag. 11 852 C T H1 frag. 23 1288 T C H1 frag. 18 232 C Ð H1 frag. 21 251 T C H1 frag. 11 953 C Ð H1 frag. 23 1303 A T H1 frag. 18 330 A T H1 frag. 23 52 C T H1 frag. 11 1000 A T H1 frag. 23 1356 A G H1 frag. 19 350 A C H1 frag. 23 283 C A H1 frag. 12 160 C A H1 frag. 23 1834 A T H1 frag. 19 523 Ð C H1 frag. 23 559 C A H1 frag. 14 109 C A H1 frag. 3 268 C T H1 frag. 20 5 A C H1 frag. 23 981 T Ð H1 frag. 14 238 A G H1 frag. 4 289 Ð A 296 (Rana) H1 frag. 23 1131 G Ð H1 frag. 16 7 A G H1 frag. 4 541 Ð G H1 frag. 10 60 Ð C H1 frag. 23 1569 A Ð H1 frag. 16 17 A G H1 frag. 4 548 A T H1 frag. 11 7 G T H1 frag. 3 258 C A H1 frag. 16 24 T C H1 frag. 5 18 C A H1 frag. 11 1179 Ð A H1 frag. 3 327 Ð C H1 frag. 16 563 A T H1 frag. 6 229 A G H1 frag. 11 1217 T C H1 frag. 4 94 A C H1 frag. 17 33 A C H1 frag. 9 409 T C H1 frag. 13 168 C A H1 frag. 4 529 Ð C H1 frag. 17 182 T A SIA frag. 4 79 T G H1 frag. 14 93 T C H1 frag. 4 548 A C H1 frag. 17 251 T G 285 (unnamed taxon) H1 frag. 17 33 C A H1 frag. 4 603 Ð A H1 frag. 17 274 A G H1 frag. 10 3 A G H1 frag. 17 58 C A H1 frag. 4 637 T A H1 frag. 17 383 A T H1 frag. 10 24 A C H1 frag. 17 118 T C H1 frag. 6 137 C Ð H1 frag. 18 433 T A H1 frag. 10 26 T C H1 frag. 18 276 A T H1 frag. 8 741 C A H1 frag. 18 758 G T H1 frag. 10 261 C T H1 frag. 18 654 T C H1 frag. 8 828 T A H1 frag. 18 766 C T H1 frag. 11 12 G A H1 frag. 18 758 T A H1 frag. 9 609 C A H1 frag. 18 767 C T H1 frag. 11 505 C T H1 frag. 19 146 G A H1 frag. 9 755 A C H1 frag. 18 861 G A H1 frag. 11 565 C T H1 frag. 19 181 G A H1 frag. 9 788 A C H1 frag. 18 863 G A H1 frag. 11 600 G A H1 frag. 19 217 A T H3 frag. 1 69 G A H1 frag. 19 178 T C H1 frag. 11 973 Ð T H1 frag. 19 531 A C H3 frag. 1 117 G C H1 frag. 19 518 Ð T H1 frag. 11 1217 T A H1 frag. 19 596 C T H3 frag. 2 5 T C 344 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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rhod frag. 2 103 T C H1 frag. 18 717 A C H1 frag. 5 9 A G H1 frag. 22 23 G A SIA frag. 2 66 T C H1 frag. 18 741 C G H1 frag. 5 28 G A H1 frag. 23 59 G A SIA frag. 3 48 G A H1 frag. 18 775 T Ð H1 frag. 6 13 C T H1 frag. 23 224 C T SIA frag. 3 75 G A H1 frag. 19 201 A C H1 frag. 6 43 Ð C H1 frag. 23 452 C T tyr frag. 1 7 C G H1 frag. 19 729 C A H1 frag. 6 62 A T H1 frag. 23 784 Ð A tyr frag. 1 18 T A H1 frag. 2 118 C A H1 frag. 8 24 A G H1 frag. 23 872 Ð C tyr frag. 1 55 T C H1 frag. 20 129 T A H1 frag. 8 177 A G H1 frag. 23 874 Ð C tyr frag. 1 58 C T H1 frag. 21 90 A Ð H1 frag. 8 232 A Ð H1 frag. 23 875 Ð C tyr frag. 2 8 T G H1 frag. 23 67 G A H1 frag. 8 354 Ð C H1 frag. 23 1097 G A tyr frag. 2 195 G A H1 frag. 23 963 A Ð H1 frag. 8 580 C T H1 frag. 23 1154 C T tyr frag. 2 261 T C H1 frag. 23 1154 A C H1 frag. 9 28 A G H1 frag. 23 1245 C T tyr frag. 3 25 T C H1 frag. 23 1607 C T H1 frag. 9 87 T A H1 frag. 23 1316 C T tyr frag. 3 49 G A H1 frag. 3 321 G A H1 frag. 9 124 A G H1 frag. 23 1321 A G tyr frag. 3 98 G C H1 frag. 4 120 T A H1 frag. 9 580 C T H1 frag. 23 1328 G A 315 (Sooglossidae) H1 frag. 4 148 A C H1 frag. 9 659 Ð A H1 frag. 23 1351 C T H1 frag. 10 3 A G H1 frag. 4 619 Ð C H3 frag. 1 81 A C H1 frag. 23 1508 Ð G H1 frag. 10 20 G A H1 frag. 4 624 Ð C H3 frag. 1 117 C T H1 frag. 23 1704 C T H1 frag. 10 26 T C H1 frag. 6 169 Ð C rhod frag. 2 79 C G H1 frag. 23 1776 A G H1 frag. 10 141 Ð A H1 frag. 7 84 A T SIA frag. 1 21 G A H1 frag. 25 17 T C H1 frag. 11 13 A T H1 frag. 8 10 T C SIA frag. 4 19 A T H1 frag. 25 20 C A H1 frag. 11 144 T C H1 frag. 8 369 G C 320 (Batrachophrynidae) H3 frag. 1 42 C G H1 frag. 11 409 T Ð H1 frag. 8 569 A G 28S frag. 2 354 C Ð H3 frag. 2 8 G T H1 frag. 11 595 A Ð H1 frag. 8 626 C T 28S frag. 2 386 C Ð H3 frag. 2 31 C T H1 frag. 11 663 A C H1 frag. 9 71 T A 28S frag. 2 505 C Ð rhod frag. 2 15 C T H1 frag. 11 887 A Ð H1 frag. 9 558 A Ð 28S frag. 2 655 G Ð rhod frag. 2 118 T C H1 frag. 11 1000 A Ð rhod frag. 1 97 A T 28S frag. 2 671 G Ð SIA frag. 1 18 T C H1 frag. 12 74 T C rhod frag. 2 42 C G 28S frag. 2 692 G Ð SIA frag. 3 60 A G H1 frag. 12 153 A C SIA frag. 1 39 C T 28S frag. 3 332 C Ð SIA frag. 3 90 C T H1 frag. 21 155 T C SIA frag. 3 156 T G 28S frag. 3 370 C Ð SIA frag. 3 153 A G H1 frag. 22 13 C T SIA frag. 4 55 G A 28S frag. 3 385 C G SIA frag. 3 165 T C H1 frag. 23 149 A G tyr frag. 1 31 C T 28S frag. 3 424 G Ð SIA frag. 3 177 A G H1 frag. 23 236 A G tyr frag. 1 43 C T 28S frag. 3 487 G Ð SIA frag. 3 183 C G H1 frag. 23 452 C T tyr frag. 2 68 T C H1 frag. 11 67 C A SIA frag. 4 13 C G H1 frag. 23 762 C T tyr frag. 3 10 C T H1 frag. 11 92 T C SIA frag. 4 49 T G H1 frag. 23 1201 G Ð tyr frag. 3 50 T C H1 frag. 11 144 T C 321 (Myobatrachoidea) H1 frag. 23 1225 Ð A tyr frag. 3 51 G C H1 frag. 11 409 T C 28S frag. 3 93 Ð C H1 frag. 23 1268 Ð C tyr frag. 3 61 G A H1 frag. 11 536 C A 28S frag. 3 297 Ð C H1 frag. 23 1270 A C 319 (Australobatrachia) H1 frag. 11 684 Ð C 28S frag. 3 575 G C H1 frag. 23 1295 T C 28S frag. 2 567 C T H1 frag. 11 983 T C H1 frag. 10 115 C T H1 frag. 23 1348 A C 28S frag. 2 639 G Ð H1 frag. 11 1129 Ð C H1 frag. 10 269 C T H1 frag. 23 1359 G Ð 28S frag. 2 719 C Ð H1 frag. 12 14 C T H1 frag. 11 7 G A H1 frag. 23 1444 G A H1 frag. 1 52 T G H1 frag. 12 74 T C H1 frag. 11 457 A Ð H1 frag. 23 1726 C A H1 frag. 10 250 G A H1 frag. 12 143 A T H1 frag. 11 1017 Ð A H1 frag. 23 1940 T C H1 frag. 11 53 T G H1 frag. 13 168 A C H1 frag. 11 1217 A C H1 frag. 8 30 C A H1 frag. 11 146 C T H1 frag. 14 141 T C H1 frag. 12 160 T C H1 frag. 8 59 T C H1 frag. 11 671 Ð T H1 frag. 14 144 A G H1 frag. 14 217 A T H1 frag. 8 115 Ð A H1 frag. 11 677 Ð G H1 frag. 14 243 A C H1 frag. 14 252 A C H1 frag. 8 441 A Ð H1 frag. 11 685 Ð C H1 frag. 15 22 T C H1 frag. 17 437 C A H1 frag. 8 523 A T H1 frag. 12 153 A Ð H1 frag. 15 27 A G H1 frag. 18 349 C A H1 frag. 8 551 A T H1 frag. 12 186 A C H1 frag. 16 7 A G H1 frag. 18 447 T A H1 frag. 8 565 A C H1 frag. 12 187 G A H1 frag. 16 19 T A H1 frag. 18 514 A C H1 frag. 8 583 Ð C H1 frag. 13 74 Ð A H1 frag. 16 191 C T H1 frag. 2 140 C T H1 frag. 8 584 Ð C H1 frag. 13 156 T C H1 frag. 16 230 Ð T H1 frag. 23 789 A Ð H1 frag. 8 593 T C H1 frag. 14 268 T G H1 frag. 16 414 C A H1 frag. 23 883 C Ð H1 frag. 8 598 A C H1 frag. 16 266 C Ð H1 frag. 16 505 Ð G H1 frag. 23 902 C Ð H1 frag. 8 643 A G H1 frag. 16 398 A C H1 frag. 16 614 T Ð H1 frag. 23 942 C T H1 frag. 8 687 G A H1 frag. 16 629 A Ð H1 frag. 16 691 A G H1 frag. 23 1074 T A H1 frag. 8 726 A C H1 frag. 17 333 C A H1 frag. 16 696 A G H1 frag. 23 1334 A C H1 frag. 9 432 T C H1 frag. 17 383 A G H1 frag. 17 5 T C H1 frag. 23 1364 A C H1 frag. 9 467 G A H1 frag. 18 254 T C H1 frag. 17 38 A G H1 frag. 23 1444 G A H1 frag. 9 579 Ð T H1 frag. 18 378 Ð C H1 frag. 17 54 A C H1 frag. 23 1554 T C H1 frag. 9 739 A C H1 frag. 18 563 A C H1 frag. 17 160 C A H1 frag. 3 22 C A H1 frag. 9 840 T A H1 frag. 18 842 Ð C H1 frag. 17 289 Ð G H1 frag. 4 191 T A rhod frag. 1 60 T C H1 frag. 19 531 A C H1 frag. 17 296 C T H1 frag. 4 280 C T rhod frag. 1 110 C T H1 frag. 2 20 C G H1 frag. 18 13 A G H1 frag. 6 23 C T 318 (Notogaeanura) H1 frag. 2 171 A T H1 frag. 18 604 A C H1 frag. 8 37 C T 28S frag. 3 370 G C H1 frag. 2 218 T A H1 frag. 18 648 C T H1 frag. 8 40 A T H1 frag. 1 68 C T H1 frag. 22 15 A G H1 frag. 18 746 T C H1 frag. 8 132 T C H1 frag. 10 43 A Ð H1 frag. 23 40 T C H1 frag. 18 821 T C H1 frag. 8 173 T C H1 frag. 11 96 T C H1 frag. 23 602 T C H1 frag. 19 3 C T H1 frag. 8 274 Ð A H1 frag. 11 457 C A H1 frag. 23 1027 A G H1 frag. 19 42 A C H1 frag. 8 275 Ð A H1 frag. 11 1327 T A H1 frag. 23 1612 Ð A H1 frag. 19 140 C T H1 frag. 8 316 T A H1 frag. 13 130 C T H1 frag. 23 1796 A C H1 frag. 19 155 T C H1 frag. 8 726 A C H1 frag. 13 159 A T H1 frag. 3 51 G A H1 frag. 19 197 A C H1 frag. 8 748 A G H1 frag. 14 177 C T H1 frag. 3 126 C T H1 frag. 19 215 A G H1 frag. 9 17 A G H1 frag. 16 16 A G H1 frag. 3 370 A T H1 frag. 19 350 A G H1 frag. 9 54 T C H1 frag. 16 25 T C H1 frag. 4 238 Ð T H1 frag. 19 788 A T H1 frag. 9 739 A T H1 frag. 17 118 G A H1 frag. 4 408 A C H1 frag. 20 61 A G H3 frag. 1 51 C T H1 frag. 18 358 Ð A H1 frag. 4 504 Ð A H1 frag. 20 68 A G H3 frag. 1 111 G A H1 frag. 18 628 T C H1 frag. 4 637 A C H1 frag. 21 121 Ð C rhod frag. 1 169 A G 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 345

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SIA frag. 2 62 C G H1 frag. 11 887 A T H1 frag. 23 1781 C T H1 frag. 23 568 Ð T SIA frag. 3 48 A C H1 frag. 12 112 G A H1 frag. 23 1951 A Ð H1 frag. 23 635 Ð C SIA frag. 3 120 G C H1 frag. 13 125 A C H1 frag. 23 1962 A T H1 frag. 23 908 Ð T 322 (Limnodynastidae) H1 frag. 14 208 T A H1 frag. 24 5 T C H1 frag. 23 1027 A T 28S frag. 2 129 A G H1 frag. 16 614 T A H1 frag. 24 33 A G H1 frag. 23 1627 A C 28S frag. 2 330 G C H1 frag. 17 107 Ð G H1 frag. 3 108 A T H1 frag. 23 1704 C A 28S frag. 2 354 C T H1 frag. 17 182 T Ð H1 frag. 3 355 A G H1 frag. 23 1754 A T 28S frag. 2 442 C G H1 frag. 17 251 T C H1 frag. 4 286 A T H1 frag. 24 17 C T 28S frag. 2 692 G A H1 frag. 18 322 C T H1 frag. 4 346 T C H1 frag. 4 169 C A 28S frag. 2 790 C A H1 frag. 18 378 C A H1 frag. 4 617 Ð C H1 frag. 5 12 A G 28S frag. 2 797 G A H1 frag. 18 838 A T H1 frag. 6 181 A C H1 frag. 6 167 A Ð 28S frag. 3 8 T C H1 frag. 2 32 A G H1 frag. 7 24 C Ð H1 frag. 7 96 A C 28S frag. 3 53 G A H1 frag. 2 218 A C H1 frag. 7 74 G A H1 frag. 8 523 A T 28S frag. 3 74 Ð A H1 frag. 2 301 C A H1 frag. 8 69 C T H1 frag. 8 722 Ð T 28S frag. 3 75 G C H1 frag. 2 407 A T H1 frag. 8 281 T C H1 frag. 9 93 C T 28S frag. 3 134 T G H1 frag. 21 155 T A H1 frag. 8 667 C T H1 frag. 9 333 C A 28S frag. 3 187 G Ð H1 frag. 23 40 C A H1 frag. 9 5 C T H1 frag. 9 788 C A 28S frag. 3 478 Ð G H1 frag. 23 236 A G H1 frag. 9 52 T C H3 frag. 1 3 C T 28S frag. 3 586 C G H1 frag. 23 283 A C H1 frag. 9 202 C A H3 frag. 1 48 A G 28S frag. 4 130 Ð T H1 frag. 23 623 A Ð H1 frag. 9 226 A G H3 frag. 1 193 C A H1 frag. 10 76 A C H1 frag. 23 1060 C T H1 frag. 9 506 A C H3 frag. 1 195 G A H1 frag. 11 15 A C H1 frag. 23 1226 T C H1 frag. 9 775 C T rhod frag. 1 36 A G H1 frag. 11 1045 A C H1 frag. 23 1342 A T H1 frag. 9 818 T C rhod frag. 1 104 T C H1 frag. 11 1135 C A H1 frag. 23 1348 A C H3 frag. 1 45 C T rhod frag. 2 118 T C H1 frag. 11 1336 T C H1 frag. 23 1627 A T H3 frag. 1 55 C A 350 (Brachycephalidae) H1 frag. 12 52 C T H1 frag. 23 1919 A Ð H3 frag. 1 111 G T 28S frag. 2 330 G C H1 frag. 13 160 Ð A H1 frag. 3 108 A C H3 frag. 1 114 T A 28S frag. 2 787 G A H1 frag. 14 13 T A H1 frag. 3 214 T C H3 frag. 1 147 A G H1 frag. 11 457 A Ð H1 frag. 14 64 A C H1 frag. 4 232 A C H3 frag. 1 225 G C H1 frag. 11 1268 C T H1 frag. 14 83 A C H1 frag. 6 167 A Ð H3 frag. 1 243 A G H1 frag. 12 8 G A H1 frag. 14 142 C T H3 frag. 1 12 C T H3 frag. 2 42 C G H1 frag. 12 163 A Ð H1 frag. 15 15 A C H3 frag. 1 105 G T rhod frag. 1 9 T C H1 frag. 13 39 A Ð H1 frag. 16 4 C T SIA frag. 2 44 C G rhod frag. 1 12 T A H1 frag. 14 109 C A H1 frag. 16 31 A C SIA frag. 2 48 T C rhod frag. 1 93 A C H1 frag. 14 129 C T H1 frag. 16 106 Ð T SIA frag. 4 88 T C rhod frag. 1 107 T G H1 frag. 14 146 A G H1 frag. 16 414 C T 348 (Nobleobatrachia) rhod frag. 1 128 C T H1 frag. 14 149 G A H1 frag. 16 429 C A H1 frag. 1 40 T A rhod frag. 2 24 C T H1 frag. 15 54 G A H1 frag. 16 683 T C H1 frag. 10 101 G A rhod frag. 2 73 G A H1 frag. 16 16 G A H1 frag. 17 33 A C H1 frag. 11 89 C T rhod frag. 2 82 G C H1 frag. 16 25 C T H1 frag. 17 274 A T H1 frag. 11 126 C T rhod frag. 2 126 C T H1 frag. 17 231 A Ð H1 frag. 17 350 A C H1 frag. 11 439 C Ð rhod frag. 2 127 C G H1 frag. 18 488 T A H1 frag. 17 446 C A H1 frag. 11 1017 Ð T rhod frag. 2 129 A T H1 frag. 18 604 A T H1 frag. 18 750 G A H1 frag. 11 1170 Ð T SIA frag. 1 27 A G H1 frag. 18 830 C T H1 frag. 18 756 T C H1 frag. 12 52 C T SIA frag. 2 14 A C H1 frag. 19 331 G Ð H1 frag. 18 821 T A H1 frag. 13 97 A T SIA frag. 2 44 C G H1 frag. 19 356 C Ð H1 frag. 18 872 A C H1 frag. 14 9 G A SIA frag. 3 24 C A H1 frag. 19 439 G Ð H1 frag. 2 198 Ð A H1 frag. 14 238 A T SIA frag. 3 39 C T H1 frag. 19 453 A Ð H1 frag. 21 124 A C H1 frag. 16 36 T Ð SIA frag. 3 42 T C H1 frag. 19 460 A Ð H1 frag. 23 274 C T H1 frag. 16 201 T Ð SIA frag. 3 135 G A H1 frag. 19 635 C T H1 frag. 23 1181 C T H1 frag. 16 313 C A SIA frag. 3 138 T A H1 frag. 2 407 A T H1 frag. 23 1607 T C H1 frag. 16 359 C T SIA frag. 4 1 C T H1 frag. 21 277 C T H1 frag. 23 1743 A C H1 frag. 16 386 Ð T SIA frag. 4 79 T G H1 frag. 22 53 C T H1 frag. 23 1763 T C H1 frag. 16 467 G A 349 (Meridianura) H1 frag. 22 63 G A H1 frag. 23 1940 T A H1 frag. 16 487 Ð A H1 frag. 10 199 C T H1 frag. 23 490 T C H1 frag. 4 75 Ð C H1 frag. 16 590 A T H1 frag. 11 622 C T H1 frag. 23 1256 T A H1 frag. 4 169 C A H1 frag. 16 672 A Ð H1 frag. 11 694 C Ð H1 frag. 23 1348 A T H1 frag. 4 408 C T H1 frag. 17 58 A T H1 frag. 11 927 A Ð H1 frag. 23 1444 G C H1 frag. 4 529 C Ð H1 frag. 17 196 Ð A H1 frag. 11 1089 C A H1 frag. 8 148 C T H1 frag. 7 42 A C H1 frag. 18 185 C T H1 frag. 11 1294 C T H1 frag. 8 369 C T H1 frag. 8 545 A C H1 frag. 18 514 A Ð H1 frag. 12 14 C T H1 frag. 8 536 G A H1 frag. 8 667 C A H1 frag. 18 868 A T H1 frag. 13 8 G A H1 frag. 9 545 C Ð H1 frag. 9 84 A C H1 frag. 18 872 A T H1 frag. 13 169 Ð C H1 frag. 9 725 C A H1 frag. 9 173 A G H1 frag. 18 883 C T H1 frag. 14 208 T A rhod frag. 1 21 C T rhod frag. 1 9 T C H1 frag. 19 403 C A H1 frag. 16 152 A T rhod frag. 1 51 T C rhod frag. 1 69 C T H1 frag. 19 651 T Ð H1 frag. 16 429 C A rhod frag. 1 162 T C rhod frag. 1 90 T C H1 frag. 19 749 C T H1 frag. 16 658 A C SIA frag. 3 84 A C rhod frag. 1 97 T C H1 frag. 19 814 Ð T H1 frag. 16 683 T A tyr frag. 1 7 G A rhod frag. 1 131 T G H1 frag. 2 196 T A H1 frag. 17 46 A C tyr frag. 1 12 A C rhod frag. 1 137 C T H1 frag. 2 301 C Ð H1 frag. 17 47 C T tyr frag. 2 128 G A rhod frag. 1 140 A C H1 frag. 23 173 Ð T H1 frag. 17 182 T A tyr frag. 2 222 T C rhod frag. 2 3 A G H1 frag. 23 250 G A H1 frag. 17 383 A Ð tyr frag. 2 234 C T rhod frag. 2 66 G C H1 frag. 23 283 A T H1 frag. 18 322 C T tyr frag. 2 266 A G rhod frag. 2 67 T G H1 frag. 23 623 A T H1 frag. 18 358 A T tyr frag. 3 98 C T rhod frag. 2 69 T A H1 frag. 23 997 A Ð H1 frag. 18 821 T C tyr frag. 3 153 A G rhod frag. 2 93 C G H1 frag. 23 1041 T C H1 frag. 19 376 C T 358 (Syrrhophus) rhod frag. 2 136 T C H1 frag. 23 1181 C A H1 frag. 19 668 A T 28S frag. 2 405 Ð G 334 (Myobatrachidae) H1 frag. 23 1221 C T H1 frag. 19 695 C T 28S frag. 2 406 Ð G H1 frag. 11 27 G A H1 frag. 23 1303 C A H1 frag. 19 771 A T 28S frag. 2 480 Ð C H1 frag. 11 622 C A H1 frag. 23 1342 A T H1 frag. 2 171 A C 28S frag. 2 573 Ð G H1 frag. 11 778 C T H1 frag. 23 1364 A T H1 frag. 23 224 C T 28S frag. 2 574 Ð G 346 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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28S frag. 2 575 Ð G 28S frag. 2 719 C Ð H1 frag. 11 1049 Ð A H1 frag. 8 828 A T 28S frag. 2 578 Ð G 28S frag. 2 762 Ð A H1 frag. 11 1071 C A H1 frag. 9 791 Ð T 28S frag. 2 630 Ð C H1 frag. 10 7 A G H1 frag. 13 69 C T 368 (Tinctanura) 28S frag. 3 97 Ð C H1 frag. 11 40 G Ð H1 frag. 13 82 C A 28S frag. 2 243 T G 28S frag. 3 204 Ð C H1 frag. 11 83 Ð A H1 frag. 13 151 C Ð 28S frag. 2 254 A C 28S frag. 3 208 Ð C H1 frag. 11 1113 A C H1 frag. 16 191 C T 28S frag. 2 339 Ð T 28S frag. 3 210 Ð C H1 frag. 12 80 A T H1 frag. 16 386 T A 28S frag. 2 692 G C 28S frag. 3 211 Ð C H1 frag. 12 126 Ð A H1 frag. 17 413 Ð T 28S frag. 3 370 C A 28S frag. 3 221 Ð T H1 frag. 13 69 C A H1 frag. 18 349 C A 28S frag. 3 385 T Ð 28S frag. 3 222 Ð T H1 frag. 13 127 A T H1 frag. 18 654 A Ð 28S frag. 3 424 G Ð 28S frag. 3 313 Ð C H1 frag. 13 156 T C H1 frag. 18 656 G Ð H1 frag. 1 36 A G 28S frag. 3 314 Ð C H1 frag. 14 208 A Ð H1 frag. 18 717 C T H1 frag. 1 72 C T 28S frag. 3 317 Ð C H1 frag. 15 33 A C H1 frag. 18 732 C T H1 frag. 11 421 Ð C 28S frag. 3 484 Ð G H1 frag. 16 1 A G H1 frag. 18 822 Ð A H1 frag. 11 541 Ð T 28S frag. 3 485 Ð G H1 frag. 16 450 A T H1 frag. 18 838 A T H1 frag. 11 910 C Ð 28S frag. 4 131 T C H1 frag. 17 4 A G H1 frag. 19 808 C A H1 frag. 11 1113 A T H1 frag. 11 57 A Ð H1 frag. 17 33 A C H1 frag. 2 438 T A H1 frag. 13 33 C A H1 frag. 11 96 C T H1 frag. 17 274 A T H1 frag. 22 61 A G H1 frag. 16 170 C Ð H1 frag. 11 1191 C T H1 frag. 18 254 C A H1 frag. 23 316 Ð C H1 frag. 16 298 Ð T H1 frag. 11 1259 T C H1 frag. 18 433 C A H1 frag. 23 350 Ð T H1 frag. 18 584 Ð T H1 frag. 13 33 C T H1 frag. 18 821 T A H1 frag. 23 362 Ð T H1 frag. 18 615 C T H1 frag. 13 93 Ð A H1 frag. 18 838 A C H1 frag. 23 379 Ð C H1 frag. 18 628 C T H1 frag. 13 97 T A H1 frag. 19 61 G C H1 frag. 23 387 Ð A H1 frag. 19 509 C T H1 frag. 14 51 A G H1 frag. 19 123 A T H1 frag. 23 397 Ð C H1 frag. 19 796 A T H1 frag. 14 102 T C H1 frag. 19 307 A Ð H1 frag. 23 762 C A H1 frag. 2 184 Ð T H1 frag. 15 28 G A H1 frag. 19 318 G Ð H1 frag. 23 807 Ð T H1 frag. 2 238 C T H1 frag. 16 191 C A H1 frag. 19 338 A Ð H1 frag. 23 811 Ð T H1 frag. 2 420 A G H1 frag. 16 266 C T H1 frag. 19 449 C A H1 frag. 23 1154 C T H1 frag. 23 48 C T H1 frag. 16 414 C A H1 frag. 2 171 C A H1 frag. 23 1329 Ð T H1 frag. 23 327 Ð T H1 frag. 17 52 A T H1 frag. 2 187 C T H1 frag. 23 1382 Ð A H1 frag. 23 404 Ð T H1 frag. 17 160 C A H1 frag. 2 196 A T H1 frag. 3 20 T A H1 frag. 23 416 Ð T H1 frag. 17 296 C A H1 frag. 20 20 T A H1 frag. 4 94 C T H1 frag. 23 707 Ð A H1 frag. 17 407 C T H1 frag. 20 54 A T H1 frag. 4 529 C A H1 frag. 23 729 C A H1 frag. 18 615 C A H1 frag. 20 77 A G H1 frag. 4 617 C A H1 frag. 23 1338 C A H1 frag. 18 648 C T H1 frag. 23 5 A T H1 frag. 4 619 C T H1 frag. 23 1518 A C H1 frag. 18 707 T A H1 frag. 23 24 A T H1 frag. 6 199 C A H1 frag. 23 1695 A G H1 frag. 19 15 A Ð H1 frag. 23 38 T C H1 frag. 8 37 C T H1 frag. 23 1759 T C H1 frag. 19 61 G T H1 frag. 23 559 A C H1 frag. 8 628 C T H1 frag. 23 1828 C A H1 frag. 19 91 C A H1 frag. 23 823 Ð T H1 frag. 8 727 Ð C H1 frag. 3 108 T A H1 frag. 19 245 Ð C H1 frag. 23 1154 C Ð H1 frag. 9 48 C T H1 frag. 3 214 T A H1 frag. 19 278 C T H1 frag. 23 1245 T C H1 frag. 9 506 C T H1 frag. 3 355 G T H1 frag. 19 695 T A H1 frag. 23 1322 C T SIA frag. 1 3 C T H1 frag. 8 741 A T H1 frag. 2 420 A G H1 frag. 23 1393 Ð C SIA frag. 1 21 G A H1 frag. 9 383 C A H1 frag. 21 161 Ð C H1 frag. 23 1684 T A tyr frag. 1 75 T C H1 frag. 9 580 C T H1 frag. 23 102 A T H1 frag. 3 26 A C tyr frag. 2 207 C A H1 frag. 9 714 Ð G H1 frag. 23 140 T A H1 frag. 4 526 Ð G tyr frag. 3 73 C T tyr frag. 3 128 C T H1 frag. 23 577 Ð A H1 frag. 5 17 C A tyr frag. 3 181 G T 369 (Amphignathodontidae) H1 frag. 23 623 T A H1 frag. 6 226 A C 367 (Cryptobatrachidae) H1 frag. 10 86 A T H1 frag. 23 1146 Ð A H1 frag. 7 9 G A H1 frag. 11 23 T C H1 frag. 10 97 T C H1 frag. 23 1338 C A H1 frag. 7 55 C T H1 frag. 11 264 C G H1 frag. 10 199 T C H1 frag. 23 1451 Ð G H1 frag. 7 74 A T H1 frag. 11 713 Ð T H1 frag. 11 126 T C H1 frag. 23 1824 T Ð H1 frag. 8 369 T C H1 frag. 11 887 A C H1 frag. 11 819 C A H1 frag. 23 1974 C T H1 frag. 8 714 A G H1 frag. 11 1333 T C H1 frag. 11 1000 A Ð H1 frag. 24 38 T C H1 frag. 9 28 A G H1 frag. 13 156 T C H1 frag. 11 1071 A T H1 frag. 25 90 T C H1 frag. 9 49 T C H1 frag. 13 159 T A H1 frag. 11 1170 T C H1 frag. 3 47 C T H1 frag. 9 98 T C H1 frag. 13 163 A C H1 frag. 11 1248 C T H1 frag. 4 64 T A H1 frag. 9 409 T C H1 frag. 14 28 C T H1 frag. 13 169 C A H1 frag. 4 343 G A H1 frag. 9 652 T C H1 frag. 15 43 A G H1 frag. 14 83 A T H1 frag. 4 346 C A H3 frag. 1 126 T G H1 frag. 16 398 A T H1 frag. 16 382 A C H1 frag. 8 37 C T rhod frag. 1 10 A G H1 frag. 18 164 T C H1 frag. 16 509 A C H1 frag. 8 62 T C rhod frag. 1 12 A C H1 frag. 18 254 T A H1 frag. 17 182 A C H1 frag. 8 722 T Ð tyr frag. 1 25 G A H1 frag. 18 325 Ð A H1 frag. 17 196 A C H1 frag. 9 47 G A tyr frag. 2 62 T G H1 frag. 18 494 C T H1 frag. 18 185 T C H1 frag. 9 51 C T tyr frag. 2 68 C T H1 frag. 19 203 A C H1 frag. 18 632 A Ð H1 frag. 9 151 Ð A tyr frag. 2 171 T C H1 frag. 19 286 C T H1 frag. 19 338 A Ð H1 frag. 9 226 G A tyr frag. 3 38 A G H1 frag. 19 788 A C H1 frag. 19 378 Ð C rhod frag. 1 9 C T tyr frag. 3 82 T C H1 frag. 20 20 T A H1 frag. 19 749 T C rhod frag. 2 136 T C tyr frag. 3 119 C T H1 frag. 22 15 A G H1 frag. 23 173 T G tyr frag. 2 105 G A 366 (Cladophrynia) H1 frag. 23 72 C T H1 frag. 23 224 T C tyr frag. 2 131 T C 28S frag. 2 172 Ð C H1 frag. 23 89 G A H1 frag. 23 356 Ð C tyr frag. 3 73 C T 28S frag. 2 518 Ð G H1 frag. 23 173 T A H1 frag. 23 815 Ð A tyr frag. 3 88 A T 28S frag. 2 570 Ð G H1 frag. 23 213 A G H1 frag. 23 1233 A G 361 (Craugastor) 28S frag. 2 768 C A H1 frag. 23 902 C A H1 frag. 23 1435 Ð G 28S frag. 2 240 C T 28S frag. 3 337 Ð G H1 frag. 23 908 T C H1 frag. 23 1657 T C 28S frag. 2 262 Ð T 28S frag. 3 344 Ð G H1 frag. 6 26 A Ð H1 frag. 8 177 A G 28S frag. 2 264 Ð G 28S frag. 3 345 Ð G H1 frag. 8 74 A T H1 frag. 8 249 A C 28S frag. 2 490 Ð G H1 frag. 11 27 G A H1 frag. 8 122 A C H1 frag. 8 470 Ð C 28S frag. 2 491 Ð T H1 frag. 11 53 T G H1 frag. 8 139 C T H1 frag. 8 667 T C 28S frag. 2 584 G Ð H1 frag. 11 279 Ð A H1 frag. 8 300 A G H1 frag. 9 22 A G 28S frag. 2 671 G C H1 frag. 11 505 A T H1 frag. 8 544 A T H1 frag. 9 281 T A 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 347

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H1 frag. 9 367 A T tyr frag. 2 77 C T tyr frag. 3 161 T C tyr frag. 3 62 Ð C H1 frag. 9 708 A C tyr frag. 3 64 A Ð tyr frag. 3 170 C T tyr frag. 3 155 A G rhod frag. 1 66 T C tyr frag. 3 114 T A 377 (Pelodryadinae) 442 (Telmatobiinae) rhod frag. 1 135 C T tyr frag. 3 127 A T H1 frag. 11 421 C A H1 frag. 10 199 T C rhod frag. 2 80 G A 372 (unnamed taxon) H1 frag. 11 480 Ð C H1 frag. 11 505 T C rhod frag. 2 134 C A 28S frag. 2 312 C G H1 frag. 12 103 T A H1 frag. 11 887 T C SIA frag. 3 39 T C 28S frag. 2 453 C T H1 frag. 14 31 T A H1 frag. 11 1012 C T SIA frag. 3 144 T C 28S frag. 2 537 Ð T H1 frag. 17 231 T C H1 frag. 11 1117 Ð G tyr frag. 1 73 T A 28S frag. 2 757 G A H1 frag. 17 337 Ð A H1 frag. 11 1266 A C tyr frag. 2 98 C T 28S frag. 3 281 Ð T H1 frag. 18 276 A Ð H1 frag. 12 9 C A tyr frag. 2 194 C A 28S frag. 3 370 A T H1 frag. 18 374 Ð G H1 frag. 12 90 T C 371 (Athesphatanura) 28S frag. 4 131 T C H1 frag. 18 766 C T H1 frag. 12 148 T C 28S frag. 2 719 C Ð 28S frag. 4 138 T G H1 frag. 18 861 A Ð H1 frag. 13 97 T A 28S frag. 2 788 Ð A H1 frag. 1 72 T C H1 frag. 19 376 T A H1 frag. 13 132 G A 28S frag. 3 54 A G H1 frag. 11 53 G T H1 frag. 19 695 T A H1 frag. 13 156 T C 28S frag. 3 379 C Ð H1 frag. 11 393 Ð T H1 frag. 19 796 T C H1 frag. 13 172 C T 28S frag. 3 389 C Ð H1 frag. 11 536 C Ð H1 frag. 2 7 C A H1 frag. 13 179 C T H1 frag. 11 392 C Ð H1 frag. 11 887 A T H1 frag. 2 133 T Ð H1 frag. 14 93 A C H1 frag. 11 565 C A H1 frag. 11 1050 Ð T H1 frag. 2 238 T C H1 frag. 15 33 A C H1 frag. 11 1161 A T H1 frag. 11 1071 A T H1 frag. 20 20 T A H1 frag. 15 40 C T H1 frag. 11 1191 C T H1 frag. 11 1248 C T H1 frag. 20 23 C A H1 frag. 16 21 T A H1 frag. 14 250 C A H1 frag. 12 139 A T H1 frag. 23 103 T C H1 frag. 16 31 A G H1 frag. 15 42 A G H1 frag. 16 4 C T H1 frag. 23 236 A T H1 frag. 16 57 A G H1 frag. 16 388 Ð T H1 frag. 17 38 A C H1 frag. 23 400 Ð A H1 frag. 16 152 T C H1 frag. 16 547 C A H1 frag. 17 47 T A H1 frag. 23 733 T A H1 frag. 16 333 A T H1 frag. 17 118 A Ð H1 frag. 17 161 Ð A H1 frag. 23 1245 C T H1 frag. 16 547 A C H1 frag. 17 296 C T H1 frag. 18 24 C T H1 frag. 23 1256 T A H1 frag. 16 590 T C H1 frag. 17 333 C T H1 frag. 18 322 T C H1 frag. 23 1356 A G H1 frag. 16 671 T C H1 frag. 17 407 C A H1 frag. 18 433 C Ð H1 frag. 23 1444 G A H1 frag. 17 54 A C H1 frag. 18 648 C G H1 frag. 18 530 C T H1 frag. 23 1687 A T H1 frag. 17 81 A T H1 frag. 18 830 C A H1 frag. 18 604 A T H1 frag. 23 1704 A T H1 frag. 17 85 A C H1 frag. 19 147 C T H1 frag. 18 648 G A H1 frag. 3 355 T A H1 frag. 17 182 A C H1 frag. 2 437 C T H1 frag. 18 717 T A H1 frag. 4 148 C A H1 frag. 17 255 Ð A H1 frag. 21 139 Ð A H1 frag. 18 750 G A H1 frag. 4 343 G A H1 frag. 17 279 T C H1 frag. 23 379 C A H1 frag. 19 140 C T H1 frag. 6 181 C A H1 frag. 18 45 T A H1 frag. 23 883 C A H1 frag. 2 180 A T H1 frag. 8 441 A T H1 frag. 18 615 T C H1 frag. 23 1359 G A H1 frag. 2 184 T C H1 frag. 8 511 G A H1 frag. 18 632 A T H1 frag. 18 838 T C H1 frag. 23 1627 C A H1 frag. 2 277 A Ð H1 frag. 8 647 A T H1 frag. 23 1754 T A H1 frag. 18 887 A T H1 frag. 2 439 C T H1 frag. 9 256 C Ð H1 frag. 4 169 A C H1 frag. 19 147 T C H1 frag. 20 7 G A H3 frag. 2 57 C T H1 frag. 4 286 T A H1 frag. 19 148 G A H1 frag. 23 417 Ð A rhod frag. 1 101 G A H1 frag. 4 548 C T H1 frag. 19 201 C A H1 frag. 23 419 Ð A tyr frag. 2 68 C T H1 frag. 8 29 T C H1 frag. 19 403 A T H1 frag. 23 490 T A tyr frag. 2 198 C G H1 frag. 8 281 C T H1 frag. 19 509 T A H1 frag. 23 902 C A tyr frag. 2 207 A C H1 frag. 9 52 C T H1 frag. 19 729 A C H1 frag. 23 1337 A T tyr frag. 3 63 G A H1 frag. 9 798 A C H1 frag. 19 823 C T H1 frag. 23 1410 Ð T tyr frag. 3 82 T C H3 frag. 1 168 T G H1 frag. 21 243 T C H1 frag. 23 1828 A Ð tyr frag. 3 93 C G rhod frag. 2 127 G C H1 frag. 21 251 C A H1 frag. 24 5 C T tyr frag. 3 99 T C rhod frag. 2 129 T A H1 frag. 23 54 A G H1 frag. 24 33 G A 386 (Hylinae) SIA frag. 3 84 A T H1 frag. 23 86 G T 28S frag. 3 187 G C tyr frag. 1 12 A C H1 frag. 3 22 A T H1 frag. 23 182 A G tyr frag. 2 222 T C H1 frag. 3 327 C A 28S frag. 3 344 G C H1 frag. 23 224 T A tyr frag. 3 88 A G H1 frag. 6 199 A C 28S frag. 3 487 G C H1 frag. 23 387 T C 372 (Hylidae) H1 frag. 8 37 T C H1 frag. 11 279 A T H1 frag. 23 903 A T 28S frag. 2 284 T C H1 frag. 8 667 T C H1 frag. 12 148 T A H1 frag. 23 1233 A T 28S frag. 2 358 Ð G H1 frag. 9 281 T C H1 frag. 14 93 A T H1 frag. 23 1256 T C 28S frag. 2 516 Ð G H1 frag. 9 409 A T H1 frag. 16 382 A T H1 frag. 23 1337 A T H1 frag. 12 21 G A H1 frag. 9 818 C T H1 frag. 16 563 A C H1 frag. 23 1434 Ð C H1 frag. 14 64 A T H3 frag. 1 T C H1 frag. 17 182 A Ð H1 frag. 23 1824 T C H1 frag. 17 160 C T H3 frag. 1 3 T C H1 frag. 19 635 C A H1 frag. 23 1825 T C H1 frag. 17 231 A T rhod frag. 1 90 T C H1 frag. 21 113 A T H1 frag. 4 394 Ð C H1 frag. 17 437 C T rhod frag. 1 168 C T H1 frag. 23 365 Ð C H1 frag. 8 249 A T H1 frag. 18 518 Ð A rhod frag. 1 169 A G H1 frag. 23 855 Ð C H1 frag. 8 727 C T H1 frag. 18 581 C T SIA frag. 2 59 C T H1 frag. 23 1687 A G H1 frag. 9 281 T C H1 frag. 18 616 Ð C SIA frag. 4 13 C T H1 frag. 23 1763 T C H1 frag. 9 367 A T H1 frag. 18 872 T A SIA frag. 4 52 C T H1 frag. 4 386 T A H3 frag. 1 6 A G H1 frag. 19 286 C T SIA frag. 4 67 T C H1 frag. 4 401 T C H3 frag. 1 84 G C H1 frag. 23 397 C T tyr frag. 1 22 T G H1 frag. 4 548 T Ð H3 frag. 1 102 C T H1 frag. 23 452 C T tyr frag. 1 73 T C H1 frag. 8 582 T C H3 frag. 1 124 C A H1 frag. 23 733 Ð T tyr frag. 2 14 A G H1 frag. 9 28 A G H3 frag. 1 126 T G H1 frag. 23 811 T A tyr frag. 2 111 C T H1 frag. 9 48 T C H3 frag. 1 192 G A H1 frag. 4 292 A G tyr frag. 2 130 A T H1 frag. 9 343 A C H3 frag. 2 66 C T H1 frag. 4 391 A T tyr frag. 2 141 G A H1 frag. 9 609 A C rhod frag. 2 139 T A H1 frag. 4 619 T A tyr frag. 2 193 C A H3 frag. 2 29 C T 424 (Leptodactyliformes) H1 frag. 8 514 C T tyr frag. 2 223 G A rhod frag. 2 16 T C 28S frag. 2 190 C Ð H1 frag. 8 727 C T tyr frag. 2 224 A G SIA frag. 3 48 A T 28S frag. 2 202 C Ð H1 frag. 9 46 T A tyr frag. 2 266 A G tyr frag. 1 76 T A 28S frag. 2 203 G Ð H1 frag. 9 580 T A tyr frag. 3 2 C A tyr frag. 2 67 T C 28S frag. 2 243 G Ð rhod frag. 2 118 C T tyr frag. 3 58 Ð G tyr frag. 2 225 C A 28S frag. 2 518 G Ð rhod frag. 2 147 T A tyr frag. 3 128 T A tyr frag. 2 273 T C 28S frag. 2 519 G Ð 348 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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28S frag. 2 521 G Ð H1 frag. 3 384 T C H1 frag. 9 506 T A H1 frag. 4 271 T C 28S frag. 2 570 G Ð H1 frag. 5 9 A G H1 frag. 9 511 T A H1 frag. 4 386 T A 28S frag. 2 584 G Ð H1 frag. 6 25 T C rhod frag. 1 9 C T H1 frag. 4 434 A G 28S frag. 2 675 C Ð H1 frag. 8 40 T A rhod frag. 1 169 A G H1 frag. 7 84 T A 28S frag. 3 345 G C H1 frag. 9 281 T C 430 (Leptodactylidae) H1 frag. 8 200 C Ð H1 frag. 1 73 T A H1 frag. 9 333 A C 28S frag. 2 187 A G H1 frag. 8 544 A T H1 frag. 11 421 C A rhod frag. 1 93 C T 28S frag. 2 284 T C H1 frag. 9 652 T A H1 frag. 11 626 Ð G rhod frag. 1 107 G A 28S frag. 2 671 G C H3 frag. 2 36 G C H1 frag. 11 957 Ð A rhod frag. 1 171 C A 28S frag. 3 115 T A rhod frag. 1 104 C G H1 frag. 17 279 Ð T rhod frag. 2 79 C T H1 frag. 11 421 A T rhod frag. 1 169 A G H1 frag. 18 371 A Ð rhod frag. 2 92 A T H1 frag. 11 762 A C rhod frag. 2 42 G A H1 frag. 18 637 Ð T rhod frag. 2 103 C T H1 frag. 11 957 A C tyr frag. 2 136 G T H1 frag. 2 210 A G rhod frag. 2 109 C T H1 frag. 12 21 G A tyr frag. 2 141 G A H1 frag. 2 389 T C rhod frag. 2 118 C T H1 frag. 13 82 A T tyr frag. 2 183 C T H1 frag. 2 407 A T SIA frag. 3 126 C T H1 frag. 15 50 A C tyr frag. 3 14 G T H1 frag. 23 404 T A SIA frag. 4 73 G A H1 frag. 16 563 A C tyr frag. 3 67 T C H1 frag. 4 672 C T 427 (Centroleninae) H1 frag. 18 433 C T tyr frag. 3 93 C G tyr frag. 3 22 A G H1 frag. 10 93 A G H1 frag. 18 542 Ð C 440 (Chthonobatrachia) 425 (Diphyabatrachia) H1 frag. 10 266 C T H1 frag. 18 615 T A H1 frag. 11 315 A C 28S frag. 2 339 T G H1 frag. 11 12 G A H1 frag. 19 201 C A H1 frag. 11 631 Ð A 28S frag. 2 535 G Ð H1 frag. 11 146 C T H1 frag. 19 403 A T H1 frag. 11 887 A T 28S frag. 3 344 G C H1 frag. 11 953 C A H1 frag. 19 496 T A H1 frag. 13 55 A C H1 frag. 1 57 T A H1 frag. 11 1116 Ð C H1 frag. 19 544 Ð T H1 frag. 14 64 A C H1 frag. 11 368 A C H1 frag. 12 132 Ð C H1 frag. 21 155 T Ð H1 frag. 18 530 C T H1 frag. 11 505 T A H1 frag. 12 187 G A H1 frag. 23 224 T C H1 frag. 23 96 A T H1 frag. 11 819 C A H1 frag. 14 64 A C H1 frag. 23 807 T Ð H1 frag. 23 173 T A H1 frag. 12 9 C A H1 frag. 14 83 A C H1 frag. 23 1321 G A H1 frag. 23 335 Ð A H1 frag. 15 33 A G H1 frag. 16 21 T A H1 frag. 23 1825 T C H1 frag. 23 903 Ð A H1 frag. 16 127 G A H1 frag. 16 25 C T H1 frag. 4 223 A T H1 frag. 23 1245 C T H1 frag. 16 292 A C H1 frag. 16 565 Ð C H1 frag. 5 17 C A H1 frag. 3 58 C T H1 frag. 16 388 T A H1 frag. 17 16 C A H1 frag. 6 81 A T H3 frag. 1 162 G C H1 frag. 16 487 A Ð H1 frag. 17 47 T C H1 frag. 9 739 A T H3 frag. 1 174 T G H1 frag. 16 658 C T H1 frag. 17 54 A C H1 frag. 9 798 C T H3 frag. 2 42 G C H1 frag. 17 182 A T H1 frag. 17 103 T C H3 frag. 1 108 A T SIA frag. 3 84 T C H1 frag. 17 413 T C H1 frag. 17 251 T Ð rhod frag. 2 126 T C SIA frag. 4 70 T C H1 frag. 18 324 Ð G H1 frag. 17 279 T C 436 (Leptodactylus) tyr frag. 2 273 T C H1 frag. 18 411 T C H1 frag. 17 333 T C 28S frag. 2 488 Ð G 441 (Ceratophryidae) H1 frag. 18 494 C Ð H1 frag. 17 407 A T 28S frag. 2 505 C G 28S frag. 2 172 C Ð H1 frag. 2 328 T A H1 frag. 17 446 T A 28S frag. 3 219 Ð T 28S frag. 2 567 C Ð H1 frag. 23 89 G A H1 frag. 18 52 A T 28S frag. 3 337 G C 28S frag. 3 344 G T H1 frag. 23 381 Ð C H1 frag. 18 411 C A H1 frag. 1 57 G T 28S frag. 3 487 G Ð H1 frag. 23 408 Ð C H1 frag. 18 553 Ð C H1 frag. 1 73 A T H1 frag. 1 68 T C H1 frag. 23 1461 Ð A H1 frag. 18 866 A G H1 frag. 10 125 Ð A H1 frag. 10 217 T A H1 frag. 23 1714 A G H1 frag. 18 868 T A H1 frag. 11 415 Ð A H1 frag. 11 16 A G H1 frag. 23 1748 T C H1 frag. 19 124 Ð C H1 frag. 11 595 A T H1 frag. 11 778 A T H1 frag. 4 603 A T H1 frag. 19 356 C T H1 frag. 11 626 G T H1 frag. 11 1049 A T H1 frag. 8 57 T C H1 frag. 19 496 T C H1 frag. 11 722 Ð T H1 frag. 17 103 T A H1 frag. 8 200 A C H1 frag. 19 673 Ð A H1 frag. 11 887 A T H1 frag. 17 210 T A H1 frag. 8 554 A G H1 frag. 19 788 A T H1 frag. 11 1017 T Ð H1 frag. 17 254 Ð A H1 frag. 9 633 C Ð H1 frag. 19 808 A G H1 frag. 11 1170 T A H1 frag. 17 335 Ð A H1 frag. 9 755 C A H1 frag. 2 238 T C H1 frag. 12 60 A T H1 frag. 18 13 A G H1 frag. 9 775 T C H1 frag. 2 308 Ð C H1 frag. 14 83 A C H1 frag. 18 256 Ð G SIA frag. 3 90 C T H1 frag. 2 360 T A H1 frag. 14 89 T C H1 frag. 18 276 A T tyr frag. 3 161 T C H1 frag. 20 1 A C H1 frag. 16 16 A G H1 frag. 18 349 A T 426 (Centrolenidae) H1 frag. 21 113 A T H1 frag. 16 691 A G H1 frag. 18 488 T A H1 frag. 1 22 G A H1 frag. 23 350 T C H1 frag. 17 4 A G H1 frag. 18 606 Ð C H1 frag. 11 92 T C H1 frag. 23 387 T A H1 frag. 17 5 T C H1 frag. 19 376 T A H1 frag. 11 634 Ð T H1 frag. 23 811 T C H1 frag. 17 52 A T H1 frag. 19 753 Ð A H1 frag. 11 1017 T Ð H1 frag. 23 854 A C H1 frag. 17 333 T C H1 frag. 23 100 A T H1 frag. 11 1049 A C H1 frag. 23 1027 T C H1 frag. 17 350 A G H1 frag. 23 102 A T H1 frag. 11 1191 T C H1 frag. 23 1074 T A H1 frag. 18 209 A G H1 frag. 23 409 Ð T H1 frag. 11 1327 A C H1 frag. 23 1627 A C H1 frag. 18 474 A T H1 frag. 23 464 Ð T H1 frag. 12 80 A C H1 frag. 23 1687 A T H1 frag. 18 530 C A H1 frag. 23 811 T A H1 frag. 13 55 A T H1 frag. 23 1743 A C H1 frag. 18 707 T A H1 frag. 3 214 A G H1 frag. 13 168 A C H1 frag. 23 1750 T C H1 frag. 18 822 A T H1 frag. 3 384 T C H1 frag. 18 209 A G H1 frag. 23 1754 A C H1 frag. 19 668 T Ð H1 frag. 6 115 A T H1 frag. 18 254 T A H1 frag. 23 1763 T C H1 frag. 19 771 T A H1 frag. 8 93 C T H1 frag. 18 322 T C H1 frag. 4 67 Ð C H1 frag. 2 389 C T H1 frag. 9 708 A C H1 frag. 18 629 Ð A H1 frag. 4 232 A T H1 frag. 21 139 C T H1 frag. 9 798 C T H1 frag. 18 637 T A H1 frag. 4 386 T G H1 frag. 23 312 Ð C H3 frag. 1 108 A T H1 frag. 19 203 A C H1 frag. 4 391 A C H1 frag. 23 379 A T H3 frag. 1 129 G A H1 frag. 19 729 A C H1 frag. 4 499 T C H1 frag. 23 416 T A 444 (Ceratophryinae) H1 frag. 23 103 T C H1 frag. 4 672 T C H1 frag. 23 795 C A 28S frag. 2 187 A Ð H1 frag. 23 792 Ð T H1 frag. 6 181 C A H1 frag. 23 854 A C H1 frag. 11 264 C A H1 frag. 23 908 T Ð H1 frag. 7 84 T C H1 frag. 23 927 Ð A H1 frag. 11 290 Ð T H1 frag. 23 1154 T A H1 frag. 8 28 A C H1 frag. 23 1627 A C H1 frag. 11 368 A T H1 frag. 23 1607 T A H1 frag. 8 120 Ð A H1 frag. 23 1717 A G H1 frag. 11 421 A T H1 frag. 23 1798 T A H1 frag. 9 31 C A H1 frag. 4 22 A T H1 frag. 13 163 A T H1 frag. 25 22 A G H1 frag. 9 343 A C H1 frag. 4 64 T A H1 frag. 15 56 T C H1 frag. 3 26 T C H1 frag. 9 432 T A H1 frag. 4 94 T G H1 frag. 16 388 T A 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 349

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H1 frag. 18 539 A C H1 frag. 11 36 T C H1 frag. 4 280 C T 452 (Cycloramphinae) H1 frag. 19 286 C T H1 frag. 11 778 T Ð H3 frag. 2 5 C T H1 frag. 11 368 A C H1 frag. 20 7 G A H1 frag. 11 1170 T A rhod frag. 1 60 T C H1 frag. 11 663 C Ð H1 frag. 23 452 C A H1 frag. 12 74 A T rhod frag. 2 6 C G H1 frag. 11 1095 Ð T H1 frag. 23 1154 T C H1 frag. 13 8 A G SIA frag. 3 141 C T H1 frag. 11 1248 C T H1 frag. 23 1338 A T H1 frag. 14 31 T C tyr frag. 2 195 A G H1 frag. 12 160 T A H1 frag. 23 1554 T A H1 frag. 14 250 A C tyr frag. 3 65 A G H1 frag. 17 198 Ð T H1 frag. 23 1754 A T H1 frag. 15 33 A G tyr frag. 3 181 T G H1 frag. 18 358 T A H1 frag. 4 202 Ð A H1 frag. 16 221 A T 450 (Hylodinae) H1 frag. 18 451 Ð A H1 frag. 6 181 C A H1 frag. 16 266 A T H1 frag. 11 16 A C H1 frag. 18 520 Ð T H1 frag. 9 580 T C H1 frag. 16 292 A T H1 frag. 11 457 A T H1 frag. 19 788 A T H3 frag. 1 3 T C H1 frag. 16 509 A C H1 frag. 11 565 A T H1 frag. 23 1754 A T H3 frag. 2 57 C T H1 frag. 16 576 C T H1 frag. 12 163 A T H1 frag. 4 126 C T SIA frag. 3 141 C T H1 frag. 17 47 T C H1 frag. 13 6 A Ð H1 frag. 7 6 A T 445 (Batrachylini) H1 frag. 18 494 C A H1 frag. 13 55 C Ð H1 frag. 9 633 C T 28S frag. 2 330 G A H1 frag. 18 628 T A H1 frag. 14 4 G A H3 frag. 1 15 A C 28S frag. 2 339 T C H1 frag. 18 717 T C H1 frag. 14 38 A C H3 frag. 1 108 A G 28S frag. 2 714 G Ð H1 frag. 18 761 T A H1 frag. 14 51 A G rhod frag. 1 169 A G 28S frag. 3 295 Ð C H1 frag. 19 271 C T H1 frag. 15 11 G A rhod frag. 2 93 C G 28S frag. 3 353 Ð A H1 frag. 19 376 A C H1 frag. 16 191 T C tyr frag. 3 82 T C 28S frag. 3 354 Ð A H1 frag. 2 277 A G H1 frag. 16 633 Ð G 453 (Cycloramphini) 28S frag. 3 416 C A H1 frag. 25 22 A G H1 frag. 16 691 A T 28S frag. 2 246 G T 28S frag. 3 453 C A H1 frag. 25 40 T C H1 frag. 17 296 T C 28S frag. 2 567 C G H1 frag. 1 57 T A H1 frag. 3 258 A T H1 frag. 17 429 A T 28S frag. 3 54 G A H1 frag. 11 72 C T H1 frag. 3 376 Ð C H1 frag. 18 139 Ð C 28S frag. 3 337 G Ð H1 frag. 11 953 C T H1 frag. 3 391 T C H1 frag. 18 191 Ð C 28S frag. 3 344 G Ð H1 frag. 11 1217 A Ð H1 frag. 4 169 C A H1 frag. 18 488 T A 28S frag. 3 345 C T H1 frag. 12 52 T C H1 frag. 4 415 Ð G H1 frag. 19 24 G T 28S frag. 3 347 A T H1 frag. 12 153 A T H1 frag. 4 499 T C H1 frag. 19 42 A T H1 frag. 11 53 T G H1 frag. 13 39 A T H1 frag. 6 81 A C H1 frag. 19 668 T C H1 frag. 11 541 T A H1 frag. 13 55 C T H1 frag. 6 213 A C H1 frag. 19 749 T C H1 frag. 14 89 T C H1 frag. 13 159 T A H1 frag. 8 116 T A H1 frag. 23 72 C T H1 frag. 16 386 A C H1 frag. 14 28 C A H1 frag. 9 432 T A H1 frag. 23 89 G A H1 frag. 17 140 C T H1 frag. 14 238 T A H1 frag. 9 633 C Ð H1 frag. 23 173 A Ð H1 frag. 17 160 T A H1 frag. 16 535 C A H1 frag. 9 710 Ð A H1 frag. 23 318 Ð T H1 frag. 17 251 T Ð H1 frag. 17 168 Ð T rhod frag. 1 125 T C H1 frag. 23 379 A T H1 frag. 17 372 T A H1 frag. 17 296 T A 448 (Hesticobatrachia) H1 frag. 23 404 A T H1 frag. 18 539 A C H1 frag. 17 413 T C H1 frag. 19 350 A G 28S frag. 2 339 T C H1 frag. 23 416 T A H1 frag. 18 433 C A 28S frag. 2 532 Ð G H1 frag. 2 342 A C H1 frag. 23 559 A C H1 frag. 18 491 Ð C 28S frag. 2 533 Ð G H1 frag. 20 182 T C H1 frag. 23 729 A T H1 frag. 18 530 T C 28S frag. 3 347 Ð A H1 frag. 21 243 T A H1 frag. 23 1256 T A H1 frag. 18 563 A C 28S frag. 3 376 Ð C H1 frag. 23 10 C T H1 frag. 23 1444 G Ð H1 frag. 18 637 T Ð H1 frag. 11 53 G T H1 frag. 23 48 T C H1 frag. 23 1627 A T H1 frag. 18 830 A G H1 frag. 11 72 C T H1 frag. 23 362 T A H1 frag. 25 30 C T H1 frag. 19 278 C A H1 frag. 11 663 A C H1 frag. 24 19 C A H1 frag. 4 191 T C H1 frag. 19 596 C T H1 frag. 13 82 A C H1 frag. 25 17 T C H1 frag. 4 262 A G H1 frag. 21 139 A T H1 frag. 16 292 A T H1 frag. 3 26 T C H1 frag. 4 441 C A H1 frag. 22 61 G A H1 frag. 16 576 C T H1 frag. 4 94 C T H1 frag. 4 493 C T H1 frag. 23 794 Ð C H1 frag. 17 160 C T H1 frag. 4 316 C T H1 frag. 4 499 T C H1 frag. 23 908 T Ð H1 frag. 17 350 A Ð H1 frag. 4 614 A C H1 frag. 23 1060 C T H1 frag. 18 822 A T H1 frag. 9 367 A T H1 frag. 6 27 G A H1 frag. 23 1329 T A H1 frag. 19 271 C T H1 frag. 9 409 A C H1 frag. 6 62 A T H1 frag. 4 191 T A H1 frag. 19 356 C T H1 frag. 9 708 A T H1 frag. 8 93 C T H1 frag. 4 211 A C H1 frag. 19 600 C T H1 frag. 9 775 T A H1 frag. 8 122 A T H1 frag. 4 403 Ð A H1 frag. 2 218 T A H3 frag. 1 69 C G H1 frag. 8 272 T A H1 frag. 4 672 T C H1 frag. 23 105 A T H3 frag. 1 96 C T H1 frag. 8 511 G A H1 frag. 6 26 A C H1 frag. 23 902 C T H3 frag. 1 111 T A H1 frag. 9 618 A C H1 frag. 6 62 A T H1 frag. 23 1669 C T H3 frag. 1 192 G C H1 frag. 9 798 C T H1 frag. 7 19 G A H1 frag. 4 94 T C rhod frag. 1 A G H3 frag. 1 42 C T H1 frag. 8 511 G A H1 frag. 4 211 A Ð rhod frag. 1 39 A C H3 frag. 1 99 C T H1 frag. 9 755 C T H1 frag. 6 25 T C rhod frag. 1 87 A G H3 frag. 1 168 G T rhod frag. 1 96 C T H1 frag. 9 580 T A rhod frag. 1 97 T C H3 frag. 1 240 G A rhod frag. 1 99 T A H1 frag. 9 755 C T rhod frag. 1 125 T C H3 frag. 2 42 C G rhod frag. 2 6 C G SIA frag. 3 39 T C rhod frag. 2 12 A G 454 (Alsodini) rhod frag. 2 93 C G SIA frag. 3 48 A G rhod frag. 2 16 T C 28S frag. 2 206 Ð G rhod frag. 2 126 T G SIA frag. 3 180 C T rhod frag. 2 113 G A 28S frag. 2 209 Ð C SIA frag. 1 3 T C tyr frag. 3 155 A C SIA frag. 3 126 C A 28S frag. 2 703 Ð C SIA frag. 3 126 C T 449 (Cycloramphidae) SIA frag. 3 129 C T 28S frag. 3 487 G C SIA frag. 3 168 A C H1 frag. 11 852 C T SIA frag. 4 76 T G H1 frag. 11 505 T C 446 (Ceratophyrini) H1 frag. 11 983 T C tyr frag. 1 76 T C H1 frag. 11 1012 C T 28S frag. 2 182 C T H1 frag. 12 103 T A tyr frag. 2 19 A G H1 frag. 11 1049 A T 28S frag. 2 283 Ð T H1 frag. 13 33 A Ð tyr frag. 2 29 C T H1 frag. 12 139 A T 28S frag. 2 671 G T H1 frag. 16 535 C A tyr frag. 2 44 T C H1 frag. 14 250 A C 28S frag. 2 788 A Ð H1 frag. 17 140 Ð C tyr frag. 2 50 T C H1 frag. 18 474 A T 28S frag. 2 790 T C H1 frag. 18 276 A Ð tyr frag. 2 225 C G H1 frag. 18 604 A Ð 28S frag. 3 54 G A H1 frag. 18 563 A T tyr frag. 2 266 A G H1 frag. 18 830 A C 28S frag. 3 344 T Ð H1 frag. 19 752 Ð C tyr frag. 3 47 T G H1 frag. 19 376 T G 28S frag. 3 370 A Ð H1 frag. 2 328 T Ð tyr frag. 3 63 G C H1 frag. 23 807 T C 28S frag. 3 577 Ð C H1 frag. 21 57 A Ð tyr frag. 3 73 T C H1 frag. 23 903 A C H1 frag. 1 41 A G H1 frag. 23 904 Ð T tyr frag. 3 98 C T H1 frag. 23 1519 Ð T 350 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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H1 frag. 5 9 A G H1 frag. 23 635 C Ð H1 frag. 11 7 G A H1 frag. 18 79 A T H1 frag. 9 315 A Ð H1 frag. 23 693 A Ð H1 frag. 11 126 T C H1 frag. 18 109 T A rhod frag. 2 69 T C H1 frag. 23 698 G Ð H1 frag. 11 1166 Ð G H1 frag. 18 237 T A rhod frag. 2 139 T A H1 frag. 23 942 T C H1 frag. 12 9 A C H1 frag. 18 254 C A SIA frag. 1 12 T G H1 frag. 23 1245 T C H1 frag. 12 160 T G H1 frag. 18 349 A G SIA frag. 1 39 T G H1 frag. 23 1554 T A H1 frag. 14 4 G A H1 frag. 18 571 T C SIA frag. 2 8 T C H1 frag. 24 19 C A H1 frag. 14 144 A G H1 frag. 18 868 A T SIA frag. 4 22 G A H1 frag. 4 94 C A H1 frag. 14 177 T C H1 frag. 19 145 A C tyr frag. 2 273 C T H1 frag. 4 265 G A H1 frag. 15 22 T C H1 frag. 19 181 G A tyr frag. 3 91 A G H1 frag. 4 392 Ð C H1 frag. 16 21 A T H1 frag. 19 203 A G tyr frag. 3 140 A G H1 frag. 4 484 C T H1 frag. 16 298 T A H1 frag. 19 209 C T tyr frag. 3 155 C A H1 frag. 5 8 Ð G H1 frag. 16 382 T A H1 frag. 19 771 C T 460 (Agastorophrynia) H1 frag. 5 16 A Ð H1 frag. 16 535 C T H1 frag. 19 808 A G 28S frag. 2 330 G Ð H1 frag. 6 23 C T H1 frag. 17 196 T C H1 frag. 2 88 T C 28S frag. 2 494 G C H1 frag. 8 122 A C H1 frag. 17 413 T G H1 frag. 2 171 T C 28S frag. 3 232 Ð G H1 frag. 8 260 C T H1 frag. 18 52 A T H1 frag. 21 133 C T 28S frag. 3 283 Ð T H1 frag. 8 369 C T H1 frag. 19 123 A C H1 frag. 22 55 T C 28S frag. 3 284 Ð T H1 frag. 8 545 A T H1 frag. 19 449 T A H1 frag. 23 224 T C 28S frag. 3 289 Ð C H1 frag. 9 432 T Ð H1 frag. 19 542 C T H1 frag. 23 274 C T 28S frag. 3 371 Ð C H1 frag. 9 672 A T H1 frag. 2 437 T A H1 frag. 23 283 T C H1 frag. 11 536 C T H1 frag. 9 755 T A H1 frag. 23 50 C T H1 frag. 11 626 G Ð H1 frag. 9 775 T C H1 frag. 23 776 Ð A H1 frag. 23 335 A C H1 frag. 11 1089 A C H3 frag. 1 162 C G H1 frag. 23 1711 A C H1 frag. 23 416 T C H1 frag. 13 69 T Ð H3 frag. 2 42 C G H1 frag. 23 1793 T C H1 frag. 23 421 A T H1 frag. 16 241 A C 469 (Bufonidae) H1 frag. 3 20 A T H1 frag. 23 789 C Ð H1 frag. 16 629 A Ð 28S frag. 2 172 C Ð H1 frag. 3 299 T C H1 frag. 23 811 T A H1 frag. 17 196 A T 28S frag. 2 312 C Ð H1 frag. 3 391 T Ð H1 frag. 23 902 T C H1 frag. 18 234 Ð T 28S frag. 2 613 G Ð H1 frag. 4 316 C T H1 frag. 23 1074 A G H1 frag. 18 411 T C 28S frag. 2 639 G Ð H1 frag. 4 373 G A H1 frag. 23 1154 A G H1 frag. 18 604 A C 28S frag. 2 655 G Ð H1 frag. 4 386 T A H1 frag. 23 1233 A G H1 frag. 18 615 T A 28S frag. 2 769 A C H1 frag. 4 461 A C H1 frag. 23 1245 T C H1 frag. 18 717 T A 28S frag. 3 76 Ð T H1 frag. 4 603 C T H1 frag. 23 1256 A G H1 frag. 19 42 A C 28S frag. 3 87 Ð C H1 frag. 6 34 G A H1 frag. 23 1364 T C H1 frag. 2 88 A T 28S frag. 3 347 A G H1 frag. 8 161 T C H1 frag. 23 1607 T A H1 frag. 21 76 A T H1 frag. 1 41 A G H1 frag. 8 488 T A H1 frag. 23 1940 T C H1 frag. 23 340 Ð T H1 frag. 10 101 A G H1 frag. 8 601 Ð C H1 frag. 24 12 G A H1 frag. 23 1634 Ð C H1 frag. 11 27 A G H1 frag. 8 628 T Ð H1 frag. 6 169 C Ð H1 frag. 23 1952 Ð A H1 frag. 11 88 T C H1 frag. 8 647 A G H1 frag. 8 461 Ð A H1 frag. 24 1 C T H1 frag. 11 89 T C H1 frag. 8 721 C T H1 frag. 9 46 T C H1 frag. 4 64 T C H1 frag. 11 827 Ð T H1 frag. 8 809 G A rhod frag. 1 94 G A H1 frag. 4 169 C T H1 frag. 11 830 Ð A H1 frag. 9 315 A Ð rhod frag. 1 128 T A H1 frag. 4 447 A G H1 frag. 11 1170 T C H1 frag. 9 321 T C rhod frag. 2 79 C G H1 frag. 6 163 C T H1 frag. 12 9 C A H1 frag. 9 714 A C rhod frag. 2 113 G A H1 frag. 8 272 T C H1 frag. 13 29 T A H3 frag. 1 111 T A rhod frag. 2 116 T C H1 frag. 8 511 G A H1 frag. 14 193 T A H3 frag. 1 162 C G rhod frag. 2 129 T C rhod frag. 2 126 T G H1 frag. 14 238 T C H3 frag. 1 213 C T rhod frag. 2 130 A C SIA frag. 1 39 T C H1 frag. 16 152 T Ð rhod frag. 1 102 G T 506 (Amietophrynus) SIA frag. 3 138 A C H1 frag. 16 382 A T rhod frag. 2 79 C T H1 frag. 23 17 A G SIA frag. 3 168 A C H1 frag. 16 485 T A 491 (Ingerophrynus) H1 frag. 23 1337 C T tyr frag. 1 23 T A H1 frag. 17 372 T A H1 frag. 2 20 T C H1 frag. 23 1554 T A tyr frag. 1 49 G C H1 frag. 17 410 Ð T H1 frag. 2 323 Ð C H1 frag. 23 1739 C A tyr frag. 1 75 C T H1 frag. 19 28 Ð A H1 frag. 23 387 A T H1 frag. 23 1781 T C 461 (Dendrobatoidea) H1 frag. 19 201 C A H1 frag. 23 1097 G A H1 frag. 23 1798 T C H1 frag. 11 279 A C H1 frag. 2 14 A C H1 frag. 23 1796 A C H1 frag. 23 1973 T G H1 frag. 11 413 Ð C H1 frag. 2 133 T A H1 frag. 23 1828 C A H1 frag. 6 199 A T H1 frag. 11 509 Ð A H1 frag. 2 218 A C H1 frag. 4 264 A G H1 frag. 8 562 A G H1 frag. 11 1000 A Ð H1 frag. 20 7 G A H1 frag. 4 401 T Ð 513 (Anaxyrus) H1 frag. 11 1071 A C H1 frag. 23 96 T A H1 frag. 4 489 T C H1 frag. 1 40 A G H1 frag. 12 21 G A H1 frag. 23 421 Ð A H1 frag. 8 232 C T H1 frag. 11 887 T C H1 frag. 12 74 A T H1 frag. 23 1342 T C H1 frag. 8 249 A T H1 frag. 11 1089 C T H1 frag. 14 93 A T H1 frag. 23 1714 A G H1 frag. 8 260 A C H1 frag. 11 1235 Ð A H1 frag. 14 208 A T H1 frag. 23 1748 T C H1 frag. 8 313 C T H1 frag. 14 193 A T H1 frag. 15 33 A C H1 frag. 3 394 Ð C H1 frag. 8 523 C T H1 frag. 14 208 A T H1 frag. 15 56 T C H1 frag. 4 191 T C H1 frag. 8 735 C T H1 frag. 16 535 C T H1 frag. 4 548 T C H1 frag. 9 321 T Ð H1 frag. 16 99 C T H1 frag. 16 683 A T H1 frag. 8 33 C A 499 (Bufo) H1 frag. 16 221 A T H1 frag. 18 641 Ð C H1 frag. 8 562 G A H1 frag. 1 71 A C H1 frag. 16 648 T C H1 frag. 18 830 A T H1 frag. 8 727 C A H1 frag. 11 31 T C H1 frag. 17 333 A C H1 frag. 19 146 G A H3 frag. 1 114 A C H1 frag. 11 1259 T C H1 frag. 18 741 A C H1 frag. 19 147 T C H3 frag. 1 129 G A H1 frag. 13 7 T Ð H1 frag. 19 376 A C H1 frag. 19 148 G T rhod frag. 1 85 A C H1 frag. 13 14 A Ð H1 frag. 19 668 T C H1 frag. 19 203 A C rhod frag. 2 3 A G H1 frag. 13 121 T C H1 frag. 23 103 A T H1 frag. 19 286 C T rhod frag. 2 9 A G H1 frag. 13 193 T C H1 frag. 23 1214 A T H1 frag. 2 223 A T rhod frag. 2 42 G C H1 frag. 14 208 A G H1 frag. 3 108 C A H1 frag. 23 25 T C rhod frag. 2 51 T C H1 frag. 15 33 C T H1 frag. 7 39 T C H1 frag. 23 199 A T SIA frag. 3 171 G C H1 frag. 16 19 C T H1 frag. 7 84 T C H1 frag. 23 213 A G 476 (Rhaebo) H1 frag. 16 96 Ð G H1 frag. 7 96 C A H1 frag. 23 452 C Ð 28S frag. 2 466 Ð T H1 frag. 16 308 Ð A H1 frag. 8 313 C T H1 frag. 23 521 A Ð 28S frag. 2 505 C G H1 frag. 16 648 T C H1 frag. 8 523 C T H1 frag. 23 568 T Ð H1 frag. 10 269 C T H1 frag. 18 24 T C rhod frag. 2 80 G A 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 351

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519 (Cranopsis) H1 frag. 11 963 C Ð H1 frag. 21 57 A T H1 frag. 11 852 C T H1 frag. 11 1161 T A H1 frag. 11 983 C Ð H1 frag. 21 90 T C H1 frag. 11 933 Ð C H1 frag. 12 148 A G H1 frag. 11 1000 C Ð H1 frag. 21 218 C T H1 frag. 11 1057 Ð C H1 frag. 12 186 C A H1 frag. 11 1113 G Ð H1 frag. 22 62 G A H1 frag. 11 1089 T G H1 frag. 14 93 A T H1 frag. 11 1135 C Ð H1 frag. 23 21 G A H1 frag. 11 1232 A C H1 frag. 16 221 A C H1 frag. 11 1161 C Ð H1 frag. 23 24 A G H1 frag. 11 1245 A C H1 frag. 16 648 T A H1 frag. 11 1191 C Ð H1 frag. 23 163 C Ð H1 frag. 11 1248 A G H1 frag. 16 678 C A H1 frag. 11 1232 A Ð H1 frag. 23 182 T G H1 frag. 11 1333 T C H1 frag. 17 333 A T H1 frag. 11 1327 T A H1 frag. 23 199 A Ð H1 frag. 12 74 T C H1 frag. 17 372 A T H1 frag. 11 1333 T C H1 frag. 23 238 T C H1 frag. 12 125 T A H1 frag. 17 411 Ð T H1 frag. 12 9 C T H1 frag. 23 274 C A H1 frag. 12 153 A G H1 frag. 18 322 C A H1 frag. 12 103 A T H1 frag. 23 942 C A H1 frag. 12 187 G A H1 frag. 18 539 T A H1 frag. 12 128 C T H1 frag. 23 1214 T C H1 frag. 13 4 T C H1 frag. 18 581 A T H1 frag. 12 148 T C H1 frag. 23 1226 T Ð H1 frag. 13 33 T C H1 frag. 18 707 C T H1 frag. 12 173 C T H1 frag. 23 1256 T C H1 frag. 13 125 A C H1 frag. 18 732 C T H1 frag. 12 177 G Ð H1 frag. 23 1270 A G H1 frag. 13 127 T C H1 frag. 2 88 T A H1 frag. 12 186 T C H1 frag. 23 1288 T C H1 frag. 14 9 G A H1 frag. 21 139 T C H1 frag. 13 82 T C H1 frag. 23 1342 T C H1 frag. 14 63 A G H1 frag. 23 1074 A T H1 frag. 13 116 C T H1 frag. 23 1518 T A H1 frag. 14 64 C T H1 frag. 23 1181 A T H1 frag. 13 141 A Ð H1 frag. 23 1711 C A H1 frag. 14 65 G A H1 frag. 3 130 T C H1 frag. 13 167 Ð T H1 frag. 23 1726 C T H1 frag. 14 80 Ð C H1 frag. 4 452 A G H1 frag. 13 168 C A H1 frag. 23 1732 T G H1 frag. 14 102 T C H1 frag. 4 672 C T H1 frag. 13 172 C T H1 frag. 24 24 G A H1 frag. 14 238 A G H1 frag. 8 69 T C H1 frag. 14 12 C T H1 frag. 25 50 T Ð H1 frag. 14 250 C T H1 frag. 8 735 T C H1 frag. 14 35 A Ð H1 frag. 6 81 C T H1 frag. 15 12 A G H1 frag. 9 708 A T H1 frag. 14 51 A G H1 frag. 6 163 C T H1 frag. 15 15 A T 522 (Chaunus) H1 frag. 14 63 A G H1 frag. 6 199 C T H1 frag. 15 45 C T 28S frag. 2 453 C T H1 frag. 14 86 C G H1 frag. 7 15 T C H1 frag. 15 50 C T H1 frag. 11 1170 C T H1 frag. 14 89 T C H1 frag. 7 91 Ð T H1 frag. 16 8 G A H1 frag. 13 8 T A H1 frag. 14 102 T C H1 frag. 7 92 C T H1 frag. 16 10 A G H1 frag. 14 193 A G H1 frag. 14 158 C Ð H1 frag. 8 29 T C H1 frag. 16 127 C T H1 frag. 16 127 T C H1 frag. 14 242 G A H1 frag. 8 109 Ð T H1 frag. 16 201 T C H1 frag. 16 590 A G H1 frag. 14 260 G T H1 frag. 8 116 C T H1 frag. 16 359 C T H1 frag. 17 310 Ð C H1 frag. 15 36 T C H1 frag. 8 139 T C H1 frag. 16 563 A C H1 frag. 20 23 C T H1 frag. 15 43 G A H1 frag. 8 159 G A H1 frag. 17 23 A T H1 frag. 21 76 A C H1 frag. 15 50 C T H1 frag. 8 161 A C H1 frag. 17 429 A G H1 frag. 23 224 T C H1 frag. 16 127 T A H1 frag. 8 249 T A H1 frag. 18 433 T C H1 frag. 23 335 A Ð H1 frag. 16 170 T A H1 frag. 8 344 Ð G H1 frag. 18 494 C T H1 frag. 23 362 C Ð H1 frag. 16 266 T A H1 frag. 8 350 Ð T H1 frag. 18 741 C T H1 frag. 23 397 C A H1 frag. 16 429 C T H1 frag. 8 351 Ð T H1 frag. 18 883 C T H1 frag. 23 888 Ð C H1 frag. 16 464 A T H1 frag. 8 387 T A H1 frag. 19 54 T A H1 frag. 23 1233 A G H1 frag. 16 535 A T H1 frag. 8 501 Ð A H1 frag. 19 119 A G H1 frag. 23 1321 G A H1 frag. 16 576 C G H1 frag. 8 503 C A H1 frag. 19 154 A G H1 frag. 23 1973 A C H1 frag. 16 614 T C H1 frag. 8 514 T C H1 frag. 19 197 A C H1 frag. 4 246 A T H1 frag. 17 20 G A H1 frag. 8 525 C T H1 frag. 19 203 A C H1 frag. 6 181 C T H1 frag. 17 22 T C H1 frag. 8 551 G A H1 frag. 19 211 A C H1 frag. 8 444 Ð A H1 frag. 17 81 A T H1 frag. 8 626 T C H1 frag. 19 213 T C H1 frag. 9 43 C A H1 frag. 17 231 G A H1 frag. 8 628 T G H1 frag. 19 216 T C rhod frag. 1 51 T C H1 frag. 17 383 T A H1 frag. 8 665 Ð T H1 frag. 19 356 T A Amolops/Amolops hongkongensis H1 frag. 17 446 A C H1 frag. 8 696 A G H1 frag. 19 514 Ð T H1 frag. 10 55 A Ð H1 frag. 18 79 A G H1 frag. 8 714 A G H1 frag. 19 542 A G H1 frag. 11 86 A Ð H1 frag. 18 106 Ð C H1 frag. 8 816 T Ð H1 frag. 19 600 C T H1 frag. 11 89 C Ð H1 frag. 18 107 Ð A H1 frag. 9 66 Ð T H1 frag. 19 612 T C H1 frag. 11 91 A Ð H1 frag. 18 108 Ð A H1 frag. 9 67 C G H1 frag. 19 749 A C H1 frag. 11 92 T Ð H1 frag. 18 109 T C H1 frag. 9 71 C T H1 frag. 2 42 C A H1 frag. 11 95 T Ð H1 frag. 18 411 C T H1 frag. 9 81 C T H1 frag. 2 328 A C H1 frag. 11 96 T Ð H1 frag. 18 447 T A H1 frag. 9 281 A Ð H1 frag. 2 420 A G H1 frag. 11 120 G Ð H1 frag. 18 514 T Ð H1 frag. 9 367 C A H1 frag. 2 424 A C H1 frag. 11 130 T Ð H1 frag. 18 530 T A H1 frag. 9 467 G A H1 frag. 20 1 T C H1 frag. 11 134 A Ð H1 frag. 18 628 C T H1 frag. 9 528 Ð T H1 frag. 20 16 T C H1 frag. 11 138 A Ð H1 frag. 18 667 Ð C H1 frag. 9 615 Ð T H1 frag. 21 57 A C H1 frag. 11 139 A Ð H1 frag. 18 866 G A H1 frag. 9 652 A Ð H1 frag. 21 76 T C H1 frag. 11 146 C Ð H1 frag. 19 24 A T H1 frag. 9 693 C T H1 frag. 21 124 A G H1 frag. 11 155 A Ð H1 frag. 19 54 T A H1 frag. 9 775 C T H1 frag. 21 251 A T H1 frag. 11 160 G Ð H1 frag. 19 91 T A Aquixalus/Aquixalus gracilipes H1 frag. 22 10 C A H1 frag. 11 213 C Ð H1 frag. 19 148 A T H1 frag. 1 36 A T H1 frag. 22 11 C A H1 frag. 11 230 C Ð H1 frag. 19 154 A G H1 frag. 1 38 T C H1 frag. 22 23 G A H1 frag. 11 264 C Ð H1 frag. 19 159 Ð A H1 frag. 1 72 C T H1 frag. 23 559 C A H1 frag. 11 315 A Ð H1 frag. 19 203 A C H1 frag. 10 19 A G H1 frag. 23 1060 C T H1 frag. 11 368 C Ð H1 frag. 19 211 A G H1 frag. 10 24 A G H1 frag. 23 1169 C A H1 frag. 11 409 C Ð H1 frag. 19 349 Ð A H1 frag. 10 43 A Ð H1 frag. 23 1181 A C H1 frag. 11 425 A Ð H1 frag. 19 453 A Ð H1 frag. 10 269 C T H1 frag. 23 1221 C A H1 frag. 11 439 C Ð H1 frag. 19 518 T A H1 frag. 11 7 G A H1 frag. 23 1226 T C H1 frag. 11 505 C Ð H1 frag. 19 531 A C H1 frag. 11 31 T C H1 frag. 23 1316 T C H1 frag. 11 565 C Ð H1 frag. 19 578 A T H1 frag. 11 36 T C H1 frag. 23 1342 T C H1 frag. 11 600 G Ð H1 frag. 19 729 C A H1 frag. 11 79 A G H1 frag. 23 1478 A G H1 frag. 11 622 A Ð H1 frag. 19 796 A G H1 frag. 11 86 A G H1 frag. 23 1607 A G H1 frag. 11 663 A Ð H1 frag. 19 819 T C H1 frag. 11 282 Ð G H1 frag. 23 1726 C T H1 frag. 11 668 C Ð H1 frag. 20 16 T C H1 frag. 11 505 C T H1 frag. 23 1750 T C H1 frag. 11 852 T Ð H1 frag. 20 146 T C H1 frag. 11 819 T A H1 frag. 23 1865 G A 352 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

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H1 frag. 23 1882 A T H1 frag. 13 33 A C H1 frag. 4 430 A G H1 frag. 8 748 G C H1 frag. 23 1968 C Ð H1 frag. 13 47 T A H1 frag. 4 447 A G H1 frag. 8 772 Ð C H1 frag. 25 17 T C H1 frag. 13 121 T C H1 frag. 4 484 C T H1 frag. 8 773 T C H1 frag. 25 32 G A H1 frag. 13 159 T A H1 frag. 4 529 A T H1 frag. 8 796 T C H1 frag. 3 16 G A H1 frag. 13 169 C A H1 frag. 7 74 A G H1 frag. 8 838 C A H1 frag. 3 20 T A H1 frag. 14 78 T A H1 frag. 8 40 T C H1 frag. 9 1 T C H1 frag. 3 176 A G H1 frag. 14 166 C T H1 frag. 8 59 T C H1 frag. 9 22 A G H1 frag. 3 184 A G H1 frag. 14 244 G A H1 frag. 9 367 A G H1 frag. 9 27 A G H1 frag. 3 342 T A H1 frag. 15 54 A G H1 frag. 9 490 Ð G H1 frag. 9 28 A G H1 frag. 3 402 T A H1 frag. 16 1 A T H3 frag. 1 75 T C H1 frag. 9 48 T C H1 frag. 4 24 A G H1 frag. 16 11 A G H3 frag. 1 138 C T H1 frag. 9 52 T C H1 frag. 4 26 A G H1 frag. 16 100 T Ð H3 frag. 1 213 C T H1 frag. 9 84 T A H1 frag. 4 94 T G H1 frag. 16 127 T Ð H3 frag. 2 81 G C H1 frag. 9 90 G A H1 frag. 4 169 A Ð H1 frag. 16 279 Ð A rhod frag. 1 42 A G H1 frag. 9 367 C G H1 frag. 4 274 A G H1 frag. 16 292 C A rhod frag. 1 94 G T H1 frag. 9 481 T A H1 frag. 4 283 T C H1 frag. 16 398 T A rhod frag. 1 96 T G H1 frag. 9 507 Ð T H1 frag. 4 292 T A H1 frag. 16 443 Ð T rhod frag. 2 69 T C H1 frag. 9 580 C A H1 frag. 4 298 A G H1 frag. 16 537 T C rhod frag. 2 80 A G H1 frag. 9 637 Ð T H1 frag. 4 404 C T H1 frag. 16 590 A T rhod frag. 2 112 C T H1 frag. 9 672 T A H1 frag. 4 408 T C H1 frag. 16 648 A G rhod frag. 2 118 C A H1 frag. 9 755 T C H1 frag. 4 461 A G H1 frag. 16 691 A T rhod frag. 2 130 A T H1 frag. 9 840 T C H1 frag. 4 477 T A H1 frag. 17 372 A G SIA frag. 2 32 A G rhod frag. 1 45 C T H1 frag. 4 565 Ð A H1 frag. 17 413 T C SIA frag. 2 53 A C Opisthodon/Limnodynastes ornatus H1 frag. 4 627 Ð T H1 frag. 18 1 G C SIA frag. 4 76 T G 28S frag. 2 448 Ð C H1 frag. 4 637 C A H1 frag. 18 83 T A Meristogenys/Meristogenys 28S frag. 2 473 T C H1 frag. 4 663 T C H1 frag. 18 138 A G orphnocnemis 28S frag. 2 492 Ð G H1 frag. 5 5 G A H1 frag. 18 254 C Ð H1 frag. 10 6 T G 28S frag. 2 493 Ð T H1 frag. 6 31 C T H1 frag. 18 357 Ð C H1 frag. 10 80 Ð C 28S frag. 3 389 C T H1 frag. 6 62 A T H1 frag. 18 727 C A H1 frag. 10 217 T A H1 frag. 1 22 G T H1 frag. 6 81 C T H1 frag. 18 822 T C H1 frag. 11 57 T A H1 frag. 1 33 C T H1 frag. 6 137 C Ð H1 frag. 19 54 A C H1 frag. 11 79 A G H1 frag. 11 368 C T H1 frag. 6 181 A C H1 frag. 19 63 Ð A H1 frag. 11 368 C T H1 frag. 11 392 C T H1 frag. 8 37 C G H1 frag. 19 140 T C H1 frag. 11 392 A T H1 frag. 11 409 T G H1 frag. 8 41 T C H1 frag. 19 195 C T H1 frag. 11 446 Ð T H1 frag. 11 536 C T H1 frag. 8 69 C T H1 frag. 19 203 A G H1 frag. 11 595 A Ð H1 frag. 11 681 Ð G H1 frag. 8 97 Ð T H1 frag. 19 244 A G H1 frag. 11 762 C T H1 frag. 11 682 A T H1 frag. 8 181 A G H1 frag. 19 403 A T H1 frag. 11 819 C T H1 frag. 11 849 Ð T H1 frag. 8 232 T C H1 frag. 19 668 T C H1 frag. 11 887 C T H1 frag. 11 1079 A Ð H1 frag. 8 321 A G H1 frag. 2 32 A G H1 frag. 11 963 C T H1 frag. 11 1161 A T H1 frag. 8 550 G A H1 frag. 2 63 Ð T H1 frag. 11 983 C T H1 frag. 11 1336 C T H1 frag. 8 551 A G H1 frag. 2 100 A T H1 frag. 11 1205 Ð T H1 frag. 12 96 Ð C H1 frag. 8 569 A Ð H1 frag. 2 133 C A H1 frag. 11 1333 T C H1 frag. 13 91 T C H1 frag. 8 611 A G H1 frag. 2 184 C A H1 frag. 12 106 Ð C H1 frag. 13 117 C T H1 frag. 8 711 T C H1 frag. 2 277 G A H1 frag. 21 133 A T H1 frag. 13 141 A C H1 frag. 9 5 T A H1 frag. 2 437 T C H1 frag. 21 178 C T H1 frag. 16 382 T A H1 frag. 9 14 A G H1 frag. 2 439 T C H1 frag. 22 62 G A H1 frag. 16 429 C T H1 frag. 9 22 A C H1 frag. 20 7 A G H1 frag. 23 299 C T H1 frag. 16 547 A G H1 frag. 9 46 A C H1 frag. 20 20 T A H1 frag. 23 506 Ð G H1 frag. 16 654 Ð G H1 frag. 9 57 T C H1 frag. 20 129 A C H1 frag. 23 963 T A H1 frag. 16 681 C T H1 frag. 9 73 T C H1 frag. 20 182 T C H1 frag. 23 1003 Ð T H1 frag. 17 12 C T H1 frag. 9 367 A T H1 frag. 21 155 T C H1 frag. 23 1015 A C H1 frag. 17 38 A G H1 frag. 9 506 A C H1 frag. 23 87 G A H1 frag. 23 1077 G Ð H1 frag. 17 52 A T H1 frag. 9 812 A G H1 frag. 23 199 C A H1 frag. 23 1303 A G H1 frag. 17 118 G Ð rhod frag. 1 21 T C H1 frag. 23 213 A G H1 frag. 23 1342 T C H1 frag. 17 160 C A rhod frag. 1 85 A C H1 frag. 23 359 Ð T H1 frag. 23 1427 C T H1 frag. 17 423 G A rhod frag. 1 87 G A H1 frag. 23 404 A Ð H1 frag. 23 1951 T A H1 frag. 18 185 C T rhod frag. 1 104 C T H1 frag. 23 416 T G H1 frag. 24 17 C T H1 frag. 18 377 Ð T rhod frag. 1 107 C T H1 frag. 23 623 T C H1 frag. 25 20 C A H1 frag. 18 378 A T rhod frag. 2 86 C A H1 frag. 23 811 T C H1 frag. 6 25 C T H1 frag. 18 388 C T rhod frag. 2 97 C A H1 frag. 23 854 A C H1 frag. 6 31 C A H1 frag. 18 411 C A rhod frag. 2 98 C A H1 frag. 23 912 A T H1 frag. 6 62 A T H1 frag. 18 512 Ð A rhod frag. 2 100 C G H1 frag. 23 1068 Ð C H1 frag. 6 126 Ð T H1 frag. 18 550 A T rhod frag. 2 113 G A H1 frag. 23 1181 A C H1 frag. 6 167 A Ð H1 frag. 18 563 C A rhod frag. 2 115 A C H1 frag. 23 1245 T C H1 frag. 6 181 A T H1 frag. 18 628 T Ð rhod frag. 2 126 A C H1 frag. 23 1270 A C H1 frag. 6 213 C T H1 frag. 18 727 A T rhod frag. 2 130 G A H1 frag. 23 1329 C T H1 frag. 7 6 A C H1 frag. 18 767 T C Duttaphrynus/Bufo melanostictus H1 frag. 23 1554 T G H1 frag. 7 78 C T H1 frag. 18 830 C T 28S frag. 4 40 G A H1 frag. 23 1607 T C H1 frag. 8 29 T C H1 frag. 18 838 A C H1 frag. 1 20 A G H1 frag. 23 1781 T A H1 frag. 8 69 C T H1 frag. 18 861 A G H1 frag. 10 86 T A H1 frag. 23 1793 T G H1 frag. 8 85 Ð C H1 frag. 18 872 C A H1 frag. 11 91 A G H1 frag. 23 1940 T A H1 frag. 8 161 A G H1 frag. 18 878 T C H1 frag. 11 536 T C H1 frag. 23 1962 C T H1 frag. 8 181 G A H1 frag. 19 119 C T H1 frag. 11 622 T Ð H1 frag. 3 22 A T H1 frag. 8 352 A G H1 frag. 19 197 A C H1 frag. 11 663 T G H1 frag. 3 26 T C H1 frag. 8 514 T C H1 frag. 19 415 C T H1 frag. 11 1009 Ð C H1 frag. 3 401 Ð A H1 frag. 8 550 G A H1 frag. 19 439 G T H1 frag. 11 1049 A C H1 frag. 4 126 T A H1 frag. 8 553 A G H1 frag. 19 542 C T H1 frag. 11 1316 A T H1 frag. 4 169 T G H1 frag. 8 568 T A H1 frag. 19 796 A G H1 frag. 12 59 T Ð H1 frag. 4 265 G A H1 frag. 8 626 T A H1 frag. 2 16 C T H1 frag. 12 153 A C H1 frag. 4 401 T Ð H1 frag. 8 714 A G H1 frag. 2 65 T C H1 frag. 12 160 T C H1 frag. 4 408 A G H1 frag. 8 735 A G H1 frag. 2 328 C T 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 353

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H1 frag. 2 438 A G H1 frag. 19 135 A G H1 frag. 11 1048 Ð T H1 frag. 20 54 A T H1 frag. 20 25 G A H1 frag. 19 137 T C H1 frag. 11 1049 A T H1 frag. 21 76 A G H1 frag. 20 115 T C H1 frag. 19 231 T C H1 frag. 11 1071 A C H1 frag. 21 133 C T H1 frag. 21 113 A G H1 frag. 19 254 A G H1 frag. 11 1170 C T H1 frag. 21 139 C T H1 frag. 23 224 C T H1 frag. 19 600 T A H1 frag. 11 1248 A T H1 frag. 23 11 C T H1 frag. 23 547 Ð A H1 frag. 19 662 Ð C H1 frag. 13 14 A T H1 frag. 23 25 T C H1 frag. 23 652 Ð C H1 frag. 19 729 A Ð H1 frag. 13 116 A G H1 frag. 23 48 T C H1 frag. 23 654 T C H1 frag. 19 826 A T H1 frag. 13 156 T C H1 frag. 23 70 C T H1 frag. 23 693 A G H1 frag. 2 73 T C H1 frag. 14 35 A G H1 frag. 23 103 A C H1 frag. 23 729 C T H1 frag. 2 180 T A H1 frag. 14 56 A G H1 frag. 23 335 A G H1 frag. 23 942 T A H1 frag. 2 184 C A H1 frag. 14 193 A G H1 frag. 23 693 T A H1 frag. 23 1233 G A H1 frag. 2 218 C A H1 frag. 15 60 T A H1 frag. 23 1154 A G H1 frag. 3 214 T C H1 frag. 2 277 A G H1 frag. 16 14 C T H1 frag. 23 1181 A G H1 frag. 4 169 A C H1 frag. 2 420 A G H1 frag. 16 40 G A H1 frag. 23 1245 T C H1 frag. 4 254 C T H1 frag. 20 7 A G H1 frag. 16 118 Ð C H1 frag. 23 1303 T C H1 frag. 4 502 Ð A H1 frag. 20 10 T C H1 frag. 16 119 Ð C H1 frag. 23 1322 C T H1 frag. 4 503 Ð A H1 frag. 21 113 T A H1 frag. 16 127 T C H1 frag. 23 1358 T C H1 frag. 6 163 C T H1 frag. 21 133 C A H1 frag. 16 216 Ð T H1 frag. 23 1359 G A H3 frag. 1 108 C T H1 frag. 23 48 T C H1 frag. 16 221 A G H1 frag. 23 1427 A C rhod frag. 1 3 T C H1 frag. 23 67 A G H1 frag. 16 311 Ð C H1 frag. 23 1714 G A rhod frag. 1 36 A G H1 frag. 23 100 A C H1 frag. 16 398 T C H1 frag. 23 1739 A T rhod frag. 2 85 G A H1 frag. 23 103 A C H1 frag. 16 590 A G H1 frag. 23 1748 C T rhod frag. 2 94 C T H1 frag. 23 274 C T H1 frag. 16 671 C T H1 frag. 23 1750 T A SIA frag. 1 4 T C H1 frag. 23 623 T C H1 frag. 17 16 C A H1 frag. 23 1828 C T SIA frag. 1 12 T C H1 frag. 23 693 T C H1 frag. 17 58 T C H1 frag. 8 10 C T SIA frag. 1 21 A G H1 frag. 23 799 T A H1 frag. 17 210 C A H1 frag. 8 93 T C SIA frag. 2 47 C G H1 frag. 23 902 T C H1 frag. 18 322 C A H1 frag. 8 172 T C SIA frag. 2 53 A C H1 frag. 23 908 T A H1 frag. 18 501 C T H1 frag. 8 232 C T SIA frag. 2 71 G C H1 frag. 23 1041 T C H1 frag. 18 604 T A H1 frag. 8 268 Ð C SIA frag. 3 3 A G H1 frag. 23 1329 C T H1 frag. 18 707 C T H1 frag. 8 272 T A SIA frag. 3 48 C T H1 frag. 23 1333 T C H1 frag. 18 741 A G H1 frag. 8 281 T C SIA frag. 3 69 C G H1 frag. 23 1356 A G H1 frag. 18 822 T A H1 frag. 8 298 G A SIA frag. 3 96 G A H1 frag. 23 1359 G A H1 frag. 19 153 A T H1 frag. 8 313 C T SIA frag. 3 144 T C H1 frag. 23 1595 Ð A H1 frag. 19 197 A T H1 frag. 8 520 A C SIA frag. 3 156 A G H1 frag. 23 1643 Ð A H1 frag. 19 376 T C H1 frag. 8 544 A T SIA frag. 4 19 T C H1 frag. 23 1793 T C H1 frag. 19 823 C T H1 frag. 8 556 T A SIA frag. 4 37 T C H1 frag. 23 1798 T C H1 frag. 2 171 T C H1 frag. 8 569 G A SIA frag. 4 67 T C H1 frag. 23 1828 C T H1 frag. 20 23 C A H1 frag. 8 611 T C Phrynoidis/Bufo asper H1 frag. 23 1919 A T H1 frag. 23 224 T C H1 frag. 8 628 T C H1 frag. 1 38 A C H1 frag. 23 1940 T C H1 frag. 23 387 A T H1 frag. 8 727 C Ð H1 frag. 1 50 A G H1 frag. 23 1962 C A H1 frag. 23 421 A T H1 frag. 8 787 C T H1 frag. 1 57 A T H1 frag. 24 10 A G H1 frag. 23 807 A T H1 frag. 9 5 C A H1 frag. 10 86 T A H1 frag. 24 17 C T H1 frag. 23 902 T C H1 frag. 9 281 T C H1 frag. 10 101 G A H1 frag. 3 22 A C H1 frag. 23 1088 G A H1 frag. 9 362 Ð C H1 frag. 11 23 T C H1 frag. 3 55 G A H1 frag. 23 1233 A G H1 frag. 9 545 C T H1 frag. 11 488 Ð T H1 frag. 3 58 C T H1 frag. 23 1256 A G H1 frag. 9 632 Ð T H1 frag. 11 565 A T H1 frag. 3 124 C A H1 frag. 23 1518 A C H1 frag. 9 633 C T H1 frag. 11 887 T C H1 frag. 3 327 C T H1 frag. 23 1793 T C H1 frag. 9 672 A T H1 frag. 11 1046 Ð A H1 frag. 3 368 A T H1 frag. 3 327 C T H1 frag. 9 693 A T H1 frag. 11 1217 T A H1 frag. 4 239 Ð C H1 frag. 4 191 C A H1 frag. 9 714 G A H1 frag. 11 1232 T C H1 frag. 4 417 Ð G H1 frag. 4 223 C A H1 frag. 9 775 A C H1 frag. 11 1294 T C H1 frag. 4 473 A G H1 frag. 4 254 C A H1 frag. 9 818 C T H1 frag. 11 1327 T C H1 frag. 6 27 G A H1 frag. 4 286 A G Salamandrinae/Salamandra H1 frag. 11 1336 C T H1 frag. 6 163 A G H1 frag. 4 452 A T salamandra H1 frag. 12 52 T C H1 frag. 8 139 C T H1 frag. 4 592 A T H1 frag. 10 23 A G H1 frag. 12 125 T C H1 frag. 8 177 A C H1 frag. 4 670 A C H1 frag. 10 250 G A H1 frag. 12 143 C T H1 frag. 8 316 T C H1 frag. 7 6 A T H1 frag. 11 86 G A H1 frag. 12 148 A G H1 frag. 8 582 T C H1 frag. 8 57 T C H1 frag. 11 210 Ð C H1 frag. 13 117 A T H1 frag. 9 47 G A H1 frag. 8 200 C T H1 frag. 11 211 Ð C H1 frag. 14 35 A C H1 frag. 9 321 T C H1 frag. 8 232 C A H1 frag. 11 212 Ð A H1 frag. 14 59 A G H1 frag. 9 511 T C H1 frag. 8 316 T C H1 frag. 11 499 Ð T H1 frag. 14 78 T A H1 frag. 9 672 A C H1 frag. 8 446 Ð A H1 frag. 11 593 Ð C H1 frag. 14 90 T C H1 frag. 9 775 A G H1 frag. 8 554 A G H1 frag. 11 818 Ð T H1 frag. 14 113 T C H3 frag. 1 15 C A H1 frag. 9 27 G A H1 frag. 11 1161 A T H1 frag. 16 3 T C H3 frag. 1 69 C G H1 frag. 9 281 T Ð H1 frag. 11 1266 C T H1 frag. 16 19 T A H3 frag. 1 114 C A H1 frag. 9 618 A G H1 frag. 11 1342 T A H1 frag. 16 388 A Ð H3 frag. 1 195 A G H1 frag. 9 693 A C H1 frag. 12 41 T C H1 frag. 16 439 Ð C rhod frag. 2 42 C T H3 frag. 1 6 A G H1 frag. 12 45 T A H1 frag. 16 547 A T SIA frag. 4 76 T A rhod frag. 1 3 C T H1 frag. 12 59 A C H1 frag. 16 652 Ð C Pseudepidalea/Bufo viridis rhod frag. 2 136 C T H1 frag. 13 97 A T H1 frag. 17 58 T C 28S frag. 3 133 C A SIA frag. 1 36 C T H1 frag. 13 116 A G H1 frag. 17 333 C G H1 frag. 10 93 A G SIA frag. 1 42 G C H1 frag. 13 168 A C H1 frag. 17 413 T C H1 frag. 10 199 C T SIA frag. 3 90 C T H1 frag. 13 172 C T H1 frag. 17 437 C T H1 frag. 10 217 C A Rhinella/Bufo margaritifer H1 frag. 14 146 A G H1 frag. 18 24 T C H1 frag. 11 31 T C H1 frag. 19 151 A T H1 frag. 14 238 A G H1 frag. 18 45 T C H1 frag. 11 36 C T H1 frag. 19 153 A G H1 frag. 15 21 T C H1 frag. 18 138 A G H1 frag. 11 79 A T H1 frag. 19 244 A C H1 frag. 15 33 A G H1 frag. 18 570 T C H1 frag. 11 86 A G H1 frag. 19 307 C T H1 frag. 16 8 G A H1 frag. 18 821 A C H1 frag. 11 95 T C H1 frag. 19 424 G A H1 frag. 16 127 T C H1 frag. 18 830 A G H1 frag. 11 1008 Ð T H1 frag. 19 623 C T H1 frag. 16 201 T A 354 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der Br/Taxon/Frag Pos Anc Der

H1 frag. 16 232 Ð T H1 frag. 23 70 C T H1 frag. 9 775 A G H1 frag. 23 1154 A T H1 frag. 16 382 A Ð H1 frag. 23 140 T A H1 frag. 9 788 A T H1 frag. 23 1303 T C H1 frag. 16 509 A T H1 frag. 23 556 Ð T H3 frag. 1 37 C T H1 frag. 23 1321 G A H1 frag. 17 120 T A H1 frag. 23 920 A G H3 frag. 1 63 C T H1 frag. 23 1329 C A H1 frag. 17 231 A G H1 frag. 23 1015 A G H3 frag. 1 99 C T H1 frag. 23 1333 T C H1 frag. 18 121 A Ð H1 frag. 23 1081 A T H3 frag. 1 105 G C H1 frag. 23 1358 T C H1 frag. 18 138 A T H1 frag. 23 1088 A T H3 frag. 1 219 C T H1 frag. 23 1364 T C H1 frag. 18 254 A C H1 frag. 23 1090 C T rhod frag. 1 10 G A H1 frag. 23 1695 G A H1 frag. 18 276 C T H1 frag. 23 1338 A G rhod frag. 1 78 G C H1 frag. 18 366 C A H1 frag. 23 1752 A G rhod frag. 1 129 C T H1 frag. 23 1732 T C H1 frag. 18 514 A C H1 frag. 23 1763 T C rhod frag. 1 140 C T H1 frag. 23 1759 C T H1 frag. 18 563 A T H1 frag. 23 1776 A G rhod frag. 1 153 G T H1 frag. 23 1824 T C H1 frag. 18 707 C Ð H1 frag. 24 34 T A rhod frag. 2 27 C T H1 frag. 24 17 T C H1 frag. 18 872 T C H1 frag. 25 16 A G rhod frag. 2 33 G C H1 frag. 24 19 C A H1 frag. 19 42 A G H1 frag. 25 20 A C rhod frag. 2 39 G T H1 frag. 25 20 C T H1 frag. 19 153 A T H1 frag. 7 35 A T rhod frag. 2 46 G T Vandijkophrynus H1 frag. 19 195 A T H1 frag. 7 78 C T Stephopaedes H1 frag. 22 55 T C H1 frag. 19 203 A C H1 frag. 8 545 T G H1 frag. 23 22 T C H1 frag. 23 5 T C H1 frag. 19 237 A G H1 frag. 8 554 A G H1 frag. 23 70 C T H1 frag. 23 73 C T H1 frag. 19 439 A C H1 frag. 8 562 G A H1 frag. 23 86 G A H1 frag. 23 199 T C H1 frag. 19 715 A C H1 frag. 8 710 A T H1 frag. 23 102 A T H1 frag. 23 557 Ð C H1 frag. 19 823 C T H1 frag. 8 714 A G H1 frag. 23 140 T C H1 frag. 23 789 C A H1 frag. 20 51 T A H1 frag. 8 792 A Ð H1 frag. 23 340 C T H1 frag. 23 1071 Ð T H1 frag. 20 107 A G H1 frag. 9 90 G A H1 frag. 23 362 T Ð H1 frag. 21 23 C T H1 frag. 9 283 Ð T H1 frag. 23 404 A Ð H1 frag. 23 1072 Ð T H1 frag. 21 57 A C H1 frag. 9 284 Ð T H1 frag. 23 568 T C H1 frag. 23 1108 T C H1 frag. 21 90 A T H1 frag. 9 294 Ð A H1 frag. 23 762 T C H1 frag. 23 1181 A C H1 frag. 21 218 C T H1 frag. 9 629 Ð G H1 frag. 23 799 A C H1 frag. 23 1342 T C H1 frag. 21 243 T C H1 frag. 9 633 A T H1 frag. 23 807 T C H1 frag. 23 1549 Ð C H1 frag. 23 54 A T H1 frag. 9 672 A C H1 frag. 23 917 C A H1 frag. 23 1627 A T H1 frag. 23 61 G A H1 frag. 9 708 A G H1 frag. 23 1124 A T H1 frag. 23 1745 G A 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 355

APPENDIX 6

NOMENCLATURAL NOTES

AMPHIBIA: Amphibia in the sense that we use it (1890: 245) as a subclass for all living amphibi- (Gray, 1825; de Queiroz and Gauthier, 1992; Can- ans. If one is unwilling to accept ``Amphibia'' as natella and Hillis, 1993) corresponds reasonably restricted to crown-group amphibians because of closely to Lissamphibia of recent authors, al- the rather large paleontological literature constru- though our concept excludes all fossil taxa outside ing this term to early tetrapods, then Neobatrachii, of the living crown group, and is identical to the unlike Lissamphibia, is the taxonomic name of meaning of the term as used by the vast majority choice because it is untroubled by variable appli- of scientists in day-to-day discourse. Lissamphi- cation (Dubois, 2004b). (It is, however, homony- bia was originally conceived of by Haeckel mous with Neobatrachia Reig, 1958, a taxon of (1866) to include salamanders and frogs, but spe- frogs [Dubois, 2004b], so should this become a ci®cally excluded caecilians, making it synony- communication problem in the future, a new name mous with Batrachii Latreille (1800) and Batra- must be selected to replace Neobatrachia Reig.) chia of Ra®nesque (1814) (which were Latiniza- GYMNOPHIONA AND APODA: The names Apoda tions of the French vernacular Batraciens Brong- and Gymnophiona have been used more-or-less niart, 1800a; Dubois, 2004b).35 Gadow (1901) interchangeably for the taxon of caecilians for a subsequently transferred caecilians into his Lis- long time. The ®rst use of the name Apoda above samphibia, and this concept of the taxon has per- the family group was by Linnaeus (1758) for a sisted (e.g., Parsons and Williams, 1963), even as group of ®shes, thereby making all subsequent the familiar name ``Amphibia'' has had its intend- uses of Apoda above the family group in Am- ed meaning concomitantly eroded through its var- phibia junior homonyms (although homonymy in iable use for very different concepts of living and above-family-group nomenclature does not have fossil groups (e.g., Huxley, 1863; Cope, 1880; agreed-upon procedures to address it, and hom- Romer, 1933; Milner, 1993; Laurin and Reisz, onymy above the family-group level is considered 1997; Laurin, 2002; Ruta et al., 2003). We think, by many to be a nonproblem). Oppel (1810) pro- as did de Queiroz and Gauthier (1992) and Can- posed the name Apoda explicitly as a family for natella and Hillis (1993), that by restricting the caecilians, rendering his use of this name unavail- name Amphibia to the best-known group (living able for above-family taxonomy under the provi- amphibians; the concept of Gray, 1825, not of sions of the current International Code of Zoolog- Linnaeus, 1758, the latter a heterogeneous taxon ical Nomenclature (Dubois, 1984a: 112; ICZN, containing various amphibians and reptiles) will 1999; but see Dubois, 2004b). Fischer von Wald- stabilize nomenclature without putting undue re- heim (1813) applied the name as a composite tax- straint on the formulation of systematic hypothe- on containing caecilians, amphisbaenians, and ses. Dubois (2004b) has suggested that the name . Merrem (1820) was the ®rst to use the Amphibia should be attributed to de Blainville name Apoda, as have many subsequent authors, (1816). This attribution would require that one ar- in the modern sense as an order for caecilians. bitrarily choose between two uses by de Blainville Dubois (2004b) regarded the homonymy of Apo- in the original paper. On page 107 of his Prod- da Linnaeus, 1758, and Apoda Merrem, 1820, as romus, he uses the term ``Amphybiens'' as a good reason to reject use of Apoda Merrem, 1820. French colloquial equivalent of his equally French Although there are no rules in unregulated no- NudipellifeÁres. On page 111, he uses the name menclature, we agree with Dubois (2004b) for a ``Amphibiens'' for an order containing solely slightly different reason: that the name Apoda has ``Protees et les Sirens'' (proteids and sirenids), rendering Amphibiens de Blainville a synonym of been usedÐrecentlyÐin confusing ways in in¯u- Perennibranchia Latreille (1825). We follow de ential publications. Trueb and Cloutier (1991) Blainville in his use of the term as a formal taxon considered the name Apoda Merrem, 1820, to ap- ply to the living crown group of caecilians and nameÐin other words, as a synonym of Peren- ϩ nibranchia Latreille (1825). the name Gymnophiona to apply to Apoda Another synonym of Amphibia, as we employ Eocaecilia (a fossil form). Cannatella and Hillis it, is Neobatrachii, coined by Sarasin and Sarasin (1993) and S.E. Evans and Sigogneau-Russell (2001) subsequently considered the name Gym- nophiona to apply to the living crown group and 35 Although in regulated nomenclature the coining of the name Apoda to apply to their Gymnophiona certain vernacular (i.e., non-Latinized) names can be re- ϩ garded as constituting the coining of a new scienti®c Eocaecilia, and Dubois (2005) treated Gym- names (e.g., Art. 11.7.2; ICZN, 1999), we see no reason nophiona as including both the living crown to extend that practice to unregulated nomenclature. group (his epifamily Caecilioidia) and Eocaecilia 356 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(his epifamily Eocaecilioidia). This is problemat- to Gymnophiona, and is therefore not synony- ic, and for this reason we provide the name Par- mous. abatrachia (etymology: para- [Greek: beside, re- CAUDATA: Caudata Scopoli (1777) originally sembling] ϩ batrachos [Greek: frog, i.e., with ref- included several taxa as well as salaman- erence to Batrachia]) for the taxon composed of ders and clearly referred to a taxon quite different living caecilians ϩ Eocaecilia Jenkins and Walsh, from that with which it is associated today. Du- 1993. The diagnosis of Parabatrachia is identical meÂril (1806: 94) used the name Caudati as the to Gymnophiona, except that limbs are retained Latin equivalent of his UrodeÁles (but as an ex- (Trueb and Cloutier, 1991). plicit family and therefore unavailable for use in Gymnophia Ra®nesque (1814: 104) is the old- unregulated nomenclatureÐcontra Dubois, est name available for the living crown-group tax- 2004b). This was likely a subsequent use of Sco- on, and this was assumed (Dubois, 1984a, 2004b) poli's Caudata, but rede®ned, excluding the reptile to have been emended to Gymnophiona by J. taxa, and with a new content. Oppel (1810) used MuÈller (1832: 198), although there is no evidence Caudata in the modern sense of content, but fol- in MuÈller's paper that he was aware of the pub- lowed earlier authors in its application as a fam- lication of Ra®nesque (1814). It appears that J. ily-group taxon, rendering it unavailable for use MuÈller (1831) published the name Gymnophidia above the family-group level (contra Dubois, as alternative name for his Coeciliae (presumably 2004b). Fischer von Waldheim (1813: 58) treated a subsequent usage of Caeciliae Wagler, 1830) in Caudati as an unranked taxon, above the family ignorance of Ra®nesque's (1814) earlier paper, group, for salamanders and as a synonym of his and provided the name Gymnophiona J. MuÈller, Urodeli, and it is this author to whom should be 1832, as a replacement name for his earlier Gym- attributed Caudata in the sense that we now use nophidia. Gymnophia Ra®nesque, 1814, is an ear- it. Stejneger (1907) used the name Caudata in its lier name, but predominant usage favors Gym- modern usage, but attributed the name/concept to nophiona J. MuÈller, 1832. Other names that are Scopoli. available for this taxon are: (1) Nuda Fitzinger, UrodeÁles DumeÂril (1806), unfortunately, was 1826; (2) Caeciliae Wagler, 1830; (3) Gymnophi- also coined as a family for salamanders and is dia J. MuÈller, 1831; and (4) Pseudo-ophidia de therefore unavailable for above-family-group no- Blainville, 1835 (a Latinization of Pseudophidiens menclature. It also was not Latinized, although de Blainville, 1816). Gymnodermia Ra®nesque, some such family-group and genus-group names 1815, is not available for this taxon, having orig- are protected in regulated nomenclature. Fischer inally been formed as a family-group taxon com- von Waldheim (1813) was the ®rst user of the posed of caecilians and amphisbaenians. name Caudati as an order, but he applied the name BATRACHIA: As a concept, Batrachia extends as a synonym of his Urodeli. from Batraciens Brongniart (1800a), a French ver- The nomenclatural question here is not one of nacular name for salamanders plus frogs, but spe- priority. Clearly, Caudata Scopoli does not apply ci®cally excludes caecilians. This was subse- to the taxon of salamanders, and Fischer von quently Latinized (brought into scienti®c nomen- Waldheim (1813) named Urodeli and Caudati as clature in our view) by Latreille (1800) as Batra- synonyms, with all uses of these names prior to chii and as Batrachia of Ra®nesque (1814). Trueb Fischer von Waldheim (1813) being applied as and Cloutier (1991) applied the name Batrachia families and therefore unavailable under the cur- to the clade composed of salamanders plus frogs, rent Code. We therefore accept Caudati Fischer which is, in fact, the original content of the taxon von Waldheim (1813), as emended to Caudata, as so named (Dubois, 2004b). As early as Merrem the name for crown-group salamanders, because (1820) the content of the group called Batrachia this is the usage preferred by most working am- was expanded to include all living amphibians. phibian systematists (e.g., Duellman and Trueb, Other available names for this taxon (in terms of 1986). implied content), although all junior to Batrachia Trueb and Cloutier (1991) and Cannatella and Latreille, 1800, are Achelata Fischer von Wald- Hillis (1993) applied the name Urodela to the heim, 1813; Dipnoa Leuckart, 1821; Amphibia larger group of fossil and living salamanders. This Latreille, 1825; Caducibranchia Berthold, 1827; was a novel application and the ®rst time that Astatodipnoa Wagler, 1828; Lissamphibia Haeck- Urodela and Caudata (named originally as objec- el, 1866; and Paratoidia Gardiner, 1982. Parato- tive synonyms) were explicitly construed to apply idea de Queiroz and Gauthier, 1992 (an apparently to different taxa. Should this usage be accepted, incorrect subsequent spelling of Paratoidia Gar- the authorship of Urodela in this sense should be diner, 1982), was de®ned as Batrachia plus all fos- attributed to Trueb and Cloutier (1991), who pro- sil relatives more closely related to Batrachia than vided both the concept and the underyling diag- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 357 nosis, although the name would be a homonym of therefore unavailable for above-family-group no- all earlier uses. menclature. Similarly, Ra®nesque (1815) coined Another name of note in this discussion is Gra- the name Meantia as an explicit family-group dientia, which has had some use for the taxon of name (and therefore unavailable) for Larvarius (ϭ salamanders, but as originally conceived (Lauren- Proteus), Proteus, Exobranchia (ϭ Necturus), and ti, 1768) the taxon was heterogeneous, containing Sirena (ϭ Siren). The next oldest name that can salamanders, , at least one frog, and be legitimately construed to attach to this taxon is several (Dubois, 2004b). It was not until Perennibranchia Latreille (1825), which was Merrem (1820) that Gradientia was used as an coined to contain Siren and Proteus. Other avail- order for salamanders. But, importantly in our able names that are synonyms are Branchiurom- view, Gradientia does not enjoy any current us- algaei Ritgen, 1828; Dysmolgae Ritgen, 1828; Di- age. plopneumena Hogg, 1838; Manentibranchia Names that are synonymous with Caudata in Hogg, 1838; Externibranchia Hogg, 1839b; Bran- our sense (and that of Fischer von Waldheim, chiata Fitzinger, 1843; Ramibranchia A.H.A. Du- 1813) are Urodelia Ra®nesque, 1815; Gradientia meÂril, 1863; and Perennibranchiata Knauer, 1883. Merrem, 1820 (not Gradientia Laurenti, 1768); ANURA: For 60 years (Romer, 1945), Salientia Batrachoidei Mayer, 1849; Saurobatrachii Fatio, has been considered to contain the fossil taxon 1872; Mecodonta Wiedersheim, 1877; and Neo- Proanura and the extant crown group, Anura. This caudata Cannatella and Hillis, 1993 (at least under use has been followed by most workers (e.g., Ta- their cladographic de®nition as applied to our to- marunov, 1964a, 1964b; Trueb and Cloutier, pology). 1991; Cannatella and Hillis, 1993; Ford and Can- CRYPTOBRANCHOIDEI: Dunn (1922) ®rst recog- natella, 1993). We accept this usage, although for nized this monophyletic group as a superfamily, most of their history the names Salientia and An- Cryptobranchoidea, which was shortly thereafter ura were used interchangeably. Salientia, as ®rst (Noble, 1931) regarded as a suborder, albeit re- coined (Laurenti, 1768: 24), was the order con- taining the superfamily name ending. Tamarunov taining frogs but also sharing Proteus with Lau- (1964b) ®rst changed the name ending to avoid renti's Gradientia. Merrem (1820: 163), in his in- implying a regulated superfamily rank, an emen- ¯uential classi®cation, used Salientia for an order dation that we follow. of frogs alone and this usage was followed by DIADECTOSALAMANDROIDEI: The name Salaman- many subsequent workers (e.g., Wied-Neuwied, droidea (as an above-family-group name) has 1825; Hogg, 1839a; Gray, 1850a; GuÈnther, 1859 been applied by different authors to several dif- ``1858''). But, if the name Salientia is to be con- ferent concepts: (1) all salamanders, excluding strued as something other than the crown group, Amphiuma (Sarasin and Sarasin, 1890); (2) Sala- it must be given the authorship of Romer (1945), mandridae ϩ Amphiumidae ϩ Plethodontidae who provided the concept that is current today. (Noble, 1931; following Dunn, 1922, who used Cannatella and Hillis (1993) cladographically de- the name as a superfamily for Ambystomatidae ®ned Salientia to mean a taxon containing Anura [sensu lato], Salamandridae, and Plethodontidae); and all of its fossil relatives more closely related (3) restricted to the family Salamandridae (Regal, to it than to CaudataÐin other words, Salientia 1966; Laurent, 1986 ``1985''; Dubois, 2005); (4) Romer, 1945. Salamandridae, Amphiumidae, Plethodontidae, The concept Anura started as anoures (A.M.C. and Brachosauroididae (fossil taxon; Tamarunov, DumeÂril, 1806: 93), French vernacular for the 1964b, as Salamandroidei); and (5) all salaman- frog ``famille'' (and therefore unavailable for ders, excluding Sirenidae and Cryptobranchoidea above-family-group nomenclature, contra Dubois, (Duellman and Trueb, 1986). 2004b), which was subsequently Latinized and We could have rede®ned ``Salamandroidea'' for ranked as an order as Anuri by Fischer von Wald- a sixth time, this time to include sirenids. Rather heim (1813: 58), as Anuria by Ra®nesque (1815: than do this, and extend the confusion, we provide 78), as Anoura by Gray (1825: 213), and ulti- a new name to correspond to a new taxonomic mately as Anura by Hogg (1839a: 270). Syno- concept, Diadectosalamandroidei: all salamanders nyms of Anura (and of most uses of Salientia be- excluding Cryptobranchoidea. fore 1945) are Ecaudata Scopoli, 1777: 464; PERENNIBRANCHIA: Merrem (1820) provided the Ecaudati Fischer von Waldheim, 1813: 58; Acau- name Amphipneusta for Hypochthon (ϭ Proteus) data Knauer, 1883: 100; Batrachia Tschudi, 1838: and Siren. Unfortunately, he named this taxon as 25; Heteromorpha Fitzinger, 1835: 107; Miura a tribe, between order and genus, thereby imply- Van der Hoeven, 1833: 307; Pygomolgaei Ritgen, ing under the current International Code of Zoo- 1828: 278; and Raniformia Hogg, 1839a: 271. logical Nomenclature (ICZN, 1999) that it should LEIOPELMATIDAE: Had we retained two mono- be considered to be within the family group and typic familes for Ascaphus (Ascaphidae) and 358 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Leiopelma (Leiopelmatidae), the name Amphicoe- tent to Bombinatoridae and Alytidae in our us- la Noble, 1931, would have been available for this age), rendering Costati and Opisthocoela syno- taxon in unregulated nomenclature. Subsequent nyms in their original forms. We employ the older authors (e.g., Romer, 1945; Reig, 1958) have ex- name as emended by Stejneger (1907). tended the concept of this taxon to various fossil ALYTIDAE: Within the framework of regarding groups, and it remains an open question whether Discoglossidae and Bombinatoridae to be subfam- these taxa are internal or external to the original ilies of a larger Discoglossidae, Dubois (1987) ex- implied clade. plained the nomenclatural issues as well as the LALAGOBATRACHIA: The names available for history of his 1982 appeal to the International this taxon demonstrate the illusion of precision Commission of Zoological Nomenclature to give that can result from retaining cladographically op- Discoglossidae precedence over Bombinatoridae timized taxonomic names when the underlying to- (as well as over Alytae and Bombitatores). Dubois pology changes in ways that require a substantial (2005) used the older name, Alytidae, for this tax- reconceptualization of diagnosis and content. The on formerly known as Discoglossidae, noting the taxon whose cladographic position as de®ned by use of Alytini by SanchõÂz (1984), among others. Ford and Cannatella (1993) that corresponds to We follow this usage as consistent with the Inter- Lalagobatrachia in content and most closely ap- national Code of Zoological Nomenclature proximates it in diagnostic features is Discoglos- (ICZN, 1999). sanura (Discoglossidae and all frogs other than ACOSMANURA: Acosmanura is identical in con- Leiopelma and Ascaphus, the most recent com- tent with Ranoidei Dubois (1983), which is a ju- mon ancestor of this group and all of its descen- nior homonym of Ranoidei Sokol (1977) (ϭ dants), as understood on the preferred tree of Ford Acosmanura ϩ Xenoanura). Sokol applied the and Cannatella (1993). The name that their cla- name Ranoidei to the group containing all frogs dographic de®nition would require to be applied excluding his Discoglossoidei (Leiopelmatidae is Pipanura (Pipimorpha ϩ Neobatrachia), al- and Discoglossidae, both sensu lato). though the resulting content of this application is NEOBATRACHIA: Sarasin and Sarasin (1890: substantially different from that intended and the 245) coined a new taxon name, Neobatrachii, as diagnosis is entirely different. a subclass for all living amphibians. This name is XENOANURA: The name Pipoidea Ford and Can- homonymous with Neobatrachia Reig, which was natella (1993) had its placement de®ned clado- graphically as including Pipidae and Rhinophryn- coined by Reig as a taxon of frogs (Dubois, idae, their most recent common ancestor, and all 2004b, 2005). Because Neobatrachii Sarasin and of its descendants (which likely includes Paleo- Sarasin, 1890, is so unfamiliar, we see little batrachidae, a fossil taxon), while Xenoanura Sav- chance that this homonymy will cause any con- age (1973: 352) had as its original content Pipi- fusion (contra Dubois, 2004b, 2005). Regardless, dae, Rhinophrynidae, and Paleobatrachidae. should homonymy become an issue in unregulat- Therefore these two names are subjective syno- ed taxonomic names, this taxon will require a new nyms. We employ the older name. name. BATRACHOPHRYNIDAE:IfBatrachophrynus is SOKOLANURA: The cladographically de®ned names Bombinanura and Discoglossanura provid- not closely allied with Caudiverbera and Telma- ed by Ford and Cannatella (1993) optimize de®- tobufo, it is nomenclaturally unfortunate because nitionally on this branch, so in one sense they are Batrachophrynidae Cope, 1875, is the oldest name synonyms of our Sokolanura, except that the con- for the inclusive taxon as long as Batrachophry- tent and diagnoses of Bombinanura and Disco- nus is considered to be a member of the family glossanura under this optimization are signi®cant- group. We suspect that additional work will show ly different from that which was originally pro- Batrachophrynus to attach elsewhere in the gen- posed. Rather than engender considerable confu- eral cladogram, which will render the name of the sion we coin a new name. family containing only Telmatobufo and Caudiv- COSTATA: As originally coined (Nicholls, 1916: erbera as Reig, 1960. 86), Opisthocoela contained solely Discoglossidae BUFONIDAE: A number of the generic names (ϭ Bombinatoridae and Alytidae in our usage), used in our work require comment. rendering it a synonym of Costati Lataste, 1879. Chascax Ritgen, 1828 (type species: Bombi- Opisthocoela was subsequently used to contain nator strumosus Merrem, 1820), is a senior name Leiopelmatidae, Discoglossidae, and Pipidae for Peltophryne Fitzinger, 1843, but is a nomen (Ahl, 1930: 83) or Discoglossidae and Pipidae oblitum under Article 23.9 of the International (Noble, 1931: 486). The original use of Costati Code of Zoological Nomenclature (ICZN, 1999). (Lataste, 1879: 339) was as a taxon containing Epidalea Cope, 1864. Calamita Oken, 1816, is Discoglossidae and Alytidae (equivalent in con- a senior synonym of Epidalea Cope, 1864. How- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 359 ever, Calamita Oken, 1816, is unavailable accord- Gastrechmia (Hemisotidae), and included only ing to Opinion 417 (Anonymous, 1956). Phryniscidae, Dendrobatidae (excluding Colos- Rhinella Fitzinger, 1826. Other names that are tethidae), Engystomidae, and Brevicipitidae. If the available for Rhinella Fitzinger, 1826 (type spe- name Firmisternia were to be applied to this cies: Bufo (Oxyrhynchus) proboscideus Spix, branch in the cladogram, the intended content 1824, by monotypy); Otilophus Cuvier, 1829 would have to to come from Firmisternia Boulen- (type species: Rana margaritifera Laurenti, 1768, ger, 1882, not Firmisternia Cope, 1875. by original designation); Eurhina Fitzinger, 1843 Complicating the application of this name is (type species: Bufo (Oxyrhynchus) proboscideus Ranoidei Sokol, 1977. Sokol (1977) named Ran- Spix, 1824, by original designation); and Trachy- oidei as a suborder to include all taxa not in his cara Tschudi, 1845 (type species: Trachycara fus- Discoglossoidea (in his usage, composed of Dis- ca Tschudi, 1834, by original designation). Ox- coglossidae [sensu lato, including Bombinatori- yrhynchus Spix, 1824 (no type species designat- dae] and Leiopelmatidae [including Ascaphus]). In other words, Ranoidei Sokol, 1977, is com- ed) is not available for this taxon because it is a ϩ junior homonym of Oxyrhynchus Leach, 1818 (a posed of our Acosmanura Xenoanura. Subse- ®sh). quently, Dubois (1983) applied the name Ranoidei to all frogs, excluding his Pipoidei (ϭ Xenoanura Rhaebo Cope, 1862. A senior synonym of ϩ Anomocoela of our usage) and Discoglossoidei. Rhaebo Cope, 1862, is Phrynomorphus Fitzinger, This renders Ranoidei Dubois, 1983, a synonym 1843: 32 (type species: Bufo leschenaulti DumeÂril ϭ of Acosmanura. and Bibron, 1841 [ Bufo guttatus Schneider, Our inclination is to preserve as closely as pos- 1799]), which is a junior homonym of Phryno- sible the near-universal vernacular term for this morphus Curtis, 1831, an insect genus. group, ``ranoid''. We also would have liked to use RANOIDES: Two other names are available for the form ``Ranoidei'', but, unfortunately, this this taxon: Diplasiocoela Nicholls, 1916 (which would have engendered confusion with Ranoidei was coined as a synonym of Firmisternia sensu Sokol and Ranoidei Dubois. We therefore have Boulenger, 1882), and Firmisternia Boulenger, formed the name as Ranoides, a taxon name ex- 1882. Surprisingly, the ®rst use of the name Fir- plicitly above regulated nomenclature. To main- misternia (Cope, 1875: 8) explicitly excluded tain parallel spelling in its sister taxon, we also Raniformia (ranids, rhacophorids, petropedetids, form the new name for the old Hyloidea as Hy- hyperoliids, and Leptopelis in Cope's sense) and loides.

APPENDIX 7

NEW AND REVIVED COMBINATIONS AND CLARIFICATIONS REGARDING TAXONOMIC CONTENT Because many users of this work will not be va Lynch and Wake, 1989. Synonymy of Linea- familiar with gender agreement and other arcana triton Tanner, 1950, with Pseudoeurycea Taylor, of nomenclature, we present here the names of 1944, results in the three name changes: Pseu- species affected by generic changes made (and, in doeurycea lineola (Cope, 1865) new combination; some cases referenced) within this paper. In some P. orchileucos (Brodie, Mendelson, and Camp- places (noted) we provide the entire content of bell, 2002) new combination; and P. orchimelas certain taxa for clarity. (Brodie, Mendelson, and Campbell, 2002) new combination.

CAUDATA ANURA

PLETHODONTIDAE SOOGLOSSIDAE (1) Eurycea Ra®nesque, 1822. The synonymy (1) Sooglossus Boulenger, 1906. The placement of Haideotriton Carr, 1939, with Eurycea Ra®n- of Nesomantis Boulenger, 1909, into the synony- esque, 1822, results in Eurycea wallacei (Carr, my of Sooglossus Boulenger, 1906, results in one 1939) new combination. name change: Sooglossus thomasseti (Boulenger, (2) Pseudoeurycea Taylor, 1944. The synony- 1909) new combination. my of Ixalotriton Wake and Johnson, 1989, with LIMNODYNASTIDAE Pseudoeurycea Taylor, 1944, results in two name changes: Pseudoeurycea nigra (Wake and John- (1) Opisthodon Steindachner, 1867. Recogni- son, 1989) new combination; Pseudoeurycea par- tion of the Limnodynastes ornatus group as the 360 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 genus Opisthodon Steindachner, 1867, results in nation; C. guerreroensis (Lynch, 1967) new com- two resurrected combinations: Opisthodon orna- bination; C. gulosus (Cope, 1875 ``1876'') new tus (Gray, 1842); and O. spenceri (Parker, 1940). combination; C. hobartsmithi (Taylor, 1937) new combination; C. inachus (Campbell and Savage, 2000) new combination; C. jota (Lynch, 1980) BRACHYCEPHALIDAE new combination; C. laevissimus (Werner, 1896) (1) The partition of former Eleutherodactylus new combination; C. laticeps (DumeÂril, 1853) DumeÂril and Bibron, 1841, into the genera Crau- new combination; C. lauraster (Savage, Mc- gastor,``Eleutherodactylus'', ``Euhyas'', ``Pelor- Cranie, and Espinal, 1996) new combination; C. ius'', and Syrrhophus, results in the following new lineatus (Brocchi, 1879) new combination; C. loki or revived combinations. (Shannon and Werler, 1955) new combination; C. (a) Craugastor Cope, 1862 (all combinations longirostris (Boulenger, 1898) new combination; previously made by implication by Crawford and C. matudai (Taylor, 1941) new combination; C. Smith, 2005). Craugastor adamastus (Campbell, megacephalus (Cope, 1875 ``1876'') new combi- 1994) new combination; C. alfredi (Boulenger, nation; C. megalotympanum (Shannon and Werler, 1898) new combination; C. amniscola (Campbell 1955) new combination; C. melanostictus (Cope, and Savage, 2000) new combination; C. anciano 1875) new combination; C. merendonensis (Savage, McCranie, and Wilson, 1988) new com- (Schmidt, 1933) new combination; C. mexicanus bination; C. andi (Savage, 1974) new combina- (Brocchi, 1877) new combination; C. milesi tion; C. angelicus (Savage, 1975) new combina- (Schmidt, 1933) new combination; C. mimus tion; C. anomalus (Boulenger, 1898) new combi- (Taylor, 1955) new combination; C. monnichorum nation; C. aphanus (Campbell, 1994) new com- (Dunn, 1940) new combination; C. myllomyllon bination; C. augusti (DugeÁs In Brocchi, 1879) (Savage, 2000) new combination; C. necerus new combination; C. aurilegulus (Savage, Mc- (Lynch, 1975) new combination; C. noblei (Bar- Cranie, and Wilson, 1988) new combination; C. bour and Dunn, 1921) new combination; C. obe- azueroensis (Savage, 1975) new combination; C. sus (Barbour, 1928) new combination; C. occi- biporcatus (Peters, 1863) new combination; C. dentalis (Taylor, 1941) new combination; C. olan- bocourti (Brocchi, 1877) new combination; C. chano (McCranie and Wilson, 1999) new combi- bransfordii (Cope, 1886) new combination; C. nation; C. omiltemanus (GuÈnther, 1900) new brevirostris (Shreve, 1936) new combination; C. combination; C. omoaensis (McCranie and Wil- brocchi (Boulenger In Brocchi, 1882) new com- son, 1997) new combination; C. opimus (Savage bination; C. bufoniformis (Boulenger, 1896) new and Myers, 2002) new combination; C. palenque combination; C. catalinae (Campbell and Savage, (Campbell and Savage, 2000) new combination; 2000) new combination; C. cerasinus (Cope, 1875 C. pechorum (McCranie and Wilson, 1999) new ``1876'') new combination; C. chac (Savage, combination; C. pelorus (Campbell and Savage, 1987) new combination; C. charadra (Campbell 2000) new combination; C. persimilis (Barbour, and Savage, 2000) new combination; C. cheiro- 1926) new combination; C. phasma (Lips and plethus (Lynch, 1990) new combination; C. chry- Savage, 1996) new combination; C. podiciferus sozetetes (McCranie, Savage, and Wilson, 1989) (Cope, 1875) new combination; C. polymniae new combination; C. coffeus (McCranie and KoÈh- (Campbell, Lamar, and Hillis, 1989) new combi- ler, 1999) new combination; C. crassidigitus (Tay- nation; C. polyptychus (Cope, 1886) new combi- lor, 1952) new combination; C. cruzi (McCranie, nation; C. pozo (Johnson and Savage, 1995) new Savage, and Wilson, 1989) new combination; C. combination; C. psephosypharus (Campbell, Sav- cuaquero (Savage, 1980) new combination; C. age, and Meyer, 1994) new combination; C. punc- daryi (Ford and Savage, 1984) new combination; tariolus (Peters, 1863) new combination; C. pyg- C. decoratus (Taylor, 1942) new combination; C. maeus (Taylor, 1937) new combination; C. rani- emcelae (Lynch, 1985) new combination; C. em- formis (Boulenger, 1896) new combination; C. leni (Dunn and Emlen, 1932) new combination; ranoides (Cope, 1886) new combination; C. rayo C. epochthidius (McCranie and Wilson, 1997) (Savage and DeWeese, 1979) new combination; new combination; C. escoces (Savage, 1975) new C. rhodopis (Cope, 1867) new combination; C. combination; C. fecundus (McCranie and Wilson, rhyacobatrachus (Campbell and Savage, 2000) 1997) new combination; C. ®tzingeri (Schmidt, new combination; C. rivulus (Campbell and Sav- 1857) new combination; C. ¯eischmanni (Boett- age, 2000) new combination; C. rostralis (Werner, ger, 1892) new combination; C. galacticorhinus 1896) new combination; C. rugosus (Peters, 1873) (Canseco-MaÂrquez and Smith, 2004) new com- new combination; C. rugulosus (Cope, 1870) new bination; C. glaucus (Lynch, 1967) new combi- combination; C. rupinius (Campbell and Savage, nation; C. gollmeri (Peters, 1863) new combina- 2000) new combination; C. sabrinus (Campbell tion; C. greggi (Bumzahem, 1955) new combi- and Savage, 2000) new combination; C. saltuar- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 361 ius (McCranie and Wilson, 1997) new combina- combination; ``E.'' griphus (Crombie, 1986) new tion; C. sandersoni (Schmidt, 1941) new combi- combination; ``E.'' guanahacabibes Estrada and nation; C. sartori (Lynch, 1965) new combina- Rodriguez, 1985 new combination; ``E.'' gundla- tion; C. silvicola (Lynch, 1967) new combination; chi (Schmidt, 1920) new combination; ``E.'' ib- C. spatulatus (Smith, 1939) new combination; C. eria (Estrada and Hedges, 1996) new combina- stadelmani (Schmidt, 1936) new combination; C. tion; ``E.'' intermedia (Barbour and Shreve, 1937) stejnegerianus (Cope, 1893) new combination; C. new combination; ``E.'' jamaicensis (Barbour, stuarti (Lynch, 1967) new combination; C. taba- 1910) new combination; ``E.'' jaumei (Estrada sarae (Savage, Hollingsworth, Lips, and Jaslow, and Alonso, 1997) new combination; ``E.'' jugans 2004) new combination; C. talamancae (Dunn, (Cochran, 1937) new combination; ``E.'' junori 1931) new combination; C. tarahumaraensis (Dunn, 1926) new combination; ``E.'' karlschmid- (Taylor, 1940) new combination; C. taurus (Tay- ti (Grant, 1931) new combination; ``E.'' klini- lor, 1958) new combination; C. taylori (Lynch, kowskii (Schwartz, 1959) new combination; ``E.'' 1966) new combination; C. trachydermus (Camp- leoncei (Shreve and Williams, 1963) new combi- bell, 1994) new combination; C. uno (Savage, nation; ``E.'' limbata (Cope, 1862); ``E.'' lucioi 1984) new combination; C. vocalis (Taylor, 1940) (Schwartz, 1980) new combination; ``E.'' luteola new combination; C. vulcani (Shannon and Wer- (Gosse, 1851) new combination; ``E.'' maestren- ler, 1955) new combination; C. xucanebi (Stuart, sis (DõÂaz, CaÂdiz, and Navarro, 2005) new com- 1941) new combination; C. yucatanensis (Lynch, bination; ``E.'' minuta Noble, 1923 new combi- 1965) new combination; C. zygodactylus (Lynch nation; ``E.'' monensis (Meerwarth, 1901) new and Myers, 1983) new combination. combination; ``E.'' nubicola Dunn, 1926 new (b) ``Euhyas'' Fitzinger, 1843: ``Euhyas'' ac- combination; ``E.'' orcutti (Dunn, 1928); ``E.'' or- monis (Schwartz, 1960) new combination; ``E.'' ientalis (Barbour and Shreve, 1937) new combi- adela (Diaz, Cadiz, and Hedges, 2003) new com- nation; ``E.'' oxyrhynca (DumeÂril and Bibron, bination; ``E.'' albipes (Barbour and Shreve, 1841) new combination; ``E.'' pantoni (Dunn, 1937) new combination; ``E.'' alcoae (Schwartz, 1926); ``E.'' parabates (Schwartz, 1964) new 1971); ``E.'' alticola (Lynn, 1937) new combi- combination; ``E.'' paulsoni (Schwartz, 1964) nation; ``E.'' amadeus (Hedges, Thomas, and new combination; ``E.'' pentasyringos (Schwartz Franz, 1987) new combination; ``E.'' andrewsi and Fowler, 1973) new combination; ``E.'' pezo- (Lynn, 1937) new combination; ``E.'' apostates petra (Schwartz, 1960) new combination; ``E.'' (Schwartz, 1973) new combination; ``E.'' arms- pictissima (Cochran, 1935) new combination; trongi (Noble and Hassler, 1933) new combina- ``E.'' pinarensis (Dunn, 1926) new combination; tion; ``E.'' atkinsi (Dunn, 1925) new combination; ``E.'' planirostris (Cope, 1862) new combination; ``E.'' briceni (Boulenger, 1903) new combination; ``E.'' rhodesi (Schwartz, 1980) new combination; ``E.'' caribe (Hedges and Thomas, 1992) new ``E.'' richmondi (Stejneger, 1904) new combina- combination; ``E.'' casparii (Dunn, 1926) new tion; ``E.'' ricordii (DumeÂril and Bibron, 1841); combination; ``E.'' cavernicola (Lynn, 1954) new ``E.'' rivularis (Diaz, Estrada, and Hedges, 2001) combination; ``E.'' corona (Hedges and Thomas, new combination; ``E.'' schmidti (Noble, 1923) 1992) new combination; ``E.'' counouspea new combination; ``E.'' sciagraphus (Schwartz, (Schwartz, 1964) new combination; ``E.'' cubana 1973) new combination; ``E.'' semipalmata (Barbour, 1942) new combination; ``E.'' cuneata (Shreve, 1936) new combination; ``E.'' simulans (Cope, 1862) new combination; ``E.'' dimidiatus (Diaz and Fong, 2001); ``E.'' sisyphodemus (Cope, 1862) new combination; ``E.'' dolomedes (Crombie, 1977) new combination; ``E.'' syming- (Hedges and Thomas, 1992) new combination; toni (Schwartz, 1957) new combination; ``E.'' te- ``E.'' emiliae (Dunn, 1926) new combination; tajulia (Estrada and Hedges, 1996) new combi- ``E.'' etheridgei (Schwartz, 1958) new combina- nation; ``E.'' thomasi (Schwartz, 1959) new com- tion; ``E.'' furcyensis (Shreve and Williams, 1963) bination; ``E.'' thorectes (Hedges, 1988) new new combination; ``E.'' fusca (Lynn and Dent, combination; ``E.'' toa (Estrada and Hedges, 1943) new combination; ``E.'' glandulifer (Coch- 1991) new combination; ``E.'' tonyi (Estrada and ran, 1935) new combination; ``E.'' glandulifero- Hedges, 1997) new combination; ``E.'' turquinen- ides (Shreve, 1936) new combination; ``E.'' gla- sis (Barbour and Shreve, 1937) new combination; phycompus (Schwartz, 1973) new combination; ``E.'' varleyi (Dunn, 1925) new combination; ``E.'' glaucoreia (Schwartz and Fowler, 1973) ``E.'' ventrilineata (Shreve, 1936) new combina- new combination; ``E.'' goini (Schwartz, 1960) tion; ``E.'' warreni (Schwartz, 1976); ``E.'' wein- new combination; ``E.'' gossei (Dunn, 1926) new landi (Barbour, 1914) new combination; ``E.'' combination; ``E.'' grabhami (Dunn, 1926) new zeus (Schwartz, 1958) new combination; ``E.'' combination; ``E.'' grahami (Schwartz, 1979) zugi (Schwartz, 1958) new combination. new combination; ``E.'' greyi (Dunn, 1926) new (c) ``Pelorius'' Hedges, 1989. ``Pelorius'' chlo- 362 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297 rophenax (Schwartz, 1976) new combination; ler, 1963) new combination; L. granti (Boulenger, ``P.'' hypostenor (Schwartz, 1965) new combi- 1914) new combination; L. gularis (Parker, 1936) nation; ``P.'' inoptatus (Barbour, 1914) new com- new combination; L. humeralis (Boulenger, 1912) bination; ``P.'' nortoni (Schwartz, 1976) new new combination; L. kubori (Zweifel, 1958) new combination; ``P.'' parapelates (Hedges and combination; L. michaeltyleri new name;36 L. Thomas, 1987) new combination; ``P.'' ruthae montana (Peters and Doria, 1878) new combina- (Noble, 1923) new combination. tion; L. narinosa (Zweifel, 1958) new combina- (d) Syrrhophus Cope, 1878 (all resurrected tion; L. obsoleta (LoÈnnberg, 1900) new combi- combinations either previously formed or implied nation; L. oktediensis (Richards and Johnston, by prior authors). Syrrhophus angustidigitorum 1993) new combination; L. papua (Boulenger, (Taylor, 1940); S. cystignathoides (Cope, 1877); 1897) new combination; L. perimetri (Zweifel, S. dennisi Lynch, 1970; S. dilatus (Davis and Dix- 1958) new combination; L. persimilis (Zweifel, on, 1955); S. dixoni (Lynch, 1991); S. grandis 1958) new combination; L. pulchra (Wandolleck, (Dixon, 1957); S. guttilatus (Cope, 1879); S. in- 1911 ``1910'') new combination; L. rueppelli terorbitalis Langebartel and Shannon, 1956; S. le- (Boettger, 1895) new combination; L. semipal- prus Cope, 1879; S. longipes (Baird, 1859); S. mata (Parker, 1936) new combination; L. trachy- marnockii Cope, 1878; S. maurus (Hedges, 1989); dermis (Zweifel, 1983) new combination; L. zwei- S. modestus Taylor, 1942; S. nitidus (Peters, feli (Tyler, 1967) new combination. 1870); S. nivicolimae Dixon and Webb, 1966; S. pallidus Duellman, 1958; S. pipilans Taylor, 1940; LEPTODACTYLIDAE S. rubrimaculatus Taylor and Smith, 1945; S. ru- (1) Leptodactylus Fitzinger, 1826. The place- fescens (Duellman and Dixon, 1959); S. saxatilis ment of Adenomera Steindachner, 1867, as a syn- (Webb, 1962); S. syristes (Hoyt, 1965); S. tere- onym of Lithodytes Fitzinger, 1843, and Lithody- tistes Duellman, 1958; S. verrucipes Cope, 1885; tes as a subgenus of Leptodactylus, as well as the S. verruculatus (Peters, 1870). placement of Vanzolinius Heyer, 1974, as a syn- onym of Leptodactylus, results in the new or re- HYLIDAE:PELODRYADINAE vived combinations. (a) Adenomera Steindachner, 1867, and Lith- (1) Litoria Tschudi, 1838: The synonymy of odytes Fitzinger, 1843. Leptodactylus (Lithodytes) Nyctimystes Stejneger, 1916, and Cyclorana andreae MuÈller, 1923; L. (L.) araucarius (Kwet Steindachner, 1867 (with retention of Cyclorana and Angulo, 2002) new combination; L. (L.) bok- as a subgenus within Litoria), with Litoria Tschu- ermanni Heyer, 1973; L. (L.) diptyx Boettger, di, 1838, results in the following new or revived 1885; L. (L.) hylaedactylus (Cope, 1868); L. (L.) combinations. lineatus (Schneider, 1799); L. (L.) lutzi (Heyer, (a) Cyclorana Steindachner, 1867. Litoria (Cy- 1975) new combination; L. (L.) marmoratus clorana) alboguttata (GuÈnther, 1867); L. (C.) aus- (Steindachner, 1867); L. (L.) martinezi (Boker- tralis (Gray, 1842) new combination; L. (C.) brev- mann, 1956). ipes (Peters, 1871) new combination; L. (C.) cryp- (b) Vanzolinius Heyer, 1974. Leptodactylus dis- totis (Tyler and Martin, 1977) new combination; codactylus Boulenger, 1884 ``1883''. L. (C.) cultripes (Parker, 1940) new combination; L. (C.) longipes (Tyler and Martin, 1977) new DENDROBATIDAE combination; L. (C.) maculosa (Tyler and Martin, 1977) new combination; L. (C.) maini (Tyler and Ameerega Bauer, 1986. Although Ameerega Martin, 1977) new combination; L. (C.) manya Bauer, 1986, is an older name for the clade cur- (van Beurden and McDonald, 1980) new combi- rently referred to by other authors as Epipedoba- nation; L. (C.) novaehollandiae (Steindachner, tes Myers, 1987, an extensive revision of Dendro- 1867) new combination; L. (C.) platycephala batidae is under way, rendering a much different (GuÈnther, 1873) new combination; L. (C.) vagita generic taxonomy from that employed in this pa- (Tyler, Davies, and Martin, 1981) new combina- per (Grant et al., in press). For this reason we do tion; L. (C.) verrucosa (Tyler and Martin, 1977) not provide a list of new combinations for species new combination. of former Epipedobates. (b) Nyctimystes Stejneger, 1916. Litoria avo- 36 calis (Zweifel, 1958) new combination; L. chees- When placed in Litoria, Nyctimystes tyleri Zweifel, manae (Tyler, 1964) new combination; L. dayi 1983, becomes a secondary homonym of Litoria tyleri Martin, Watson, Gartside, Littlejohn, and Loftus-Hills, (GuÈnther, 1897) new combination; L. daymani 1979 ``1978''. Although we expect that ongoing work (Zweifel, 1958) new combination; L. disrupta will correct this nomenclatural anomaly, we propose Li- (Tyler, 1963) new combination; L. ¯uviatilis toria michaeltyleri nomen novum to replace Nyctimystes (Zweifel, 1958) new combination; L. foricula (Ty- tyleri Zweifel, 1983. 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 363

BUFONIDAE (3) Anaxyrus Tschudi, 1845. Recognition of this major clade of Nearctic ``Bufo'' as a genus The extensive generic rearrangements result in requires a number of new name combinations: many name changes: Anaxyrus americanus (Holbrook, 1836) new com- (1) Altiphrynoides Dubois, 1987 ``1986''. The bination; A. baxteri (Porter, 1968) new combina- synonymy of Altiphrynoides Dubois, 1987 tion; A. boreas (Baird and Girard, 1852) new ``1986'', and Spinophrynoides Dubois, 1987 combination; A. californicus (Camp, 1915) new ``1986'', results in a single name change: Alti- combination; A. canorus (Camp, 1916) new com- phrynoides osgoodi (Loveridge, 1932) new com- bination; A. cognatus (Say in James, 1823) new bination. combination; A. compactilis (Wiegmann, 1833) (2) Amietophrynus new genus. The components new combination; A. debilis (Girard, 1854) new of Amietophrynus come from several former spe- combination; A. exsul (Myers, 1942) new com- cies groups of ``Bufo'', all exhibiting the 20-chro- bination; A. fowleri (Hinckley, 1882) new com- mosome condition, with the exception of the A. bination; A. hemiophrys (Cope, 1886) new com- pardalis group, which has reversed to the 22- bination; A. houstonensis (Sanders, 1953) new chromosome condition (Cunningham and Cherry, combination; A. kelloggi (Taylor, 1938) new com- 2004): A. asmarae (Tandy, Bogart, Largen, and bination; A. mexicanus (Brocchi, 1879) new com- Feener, 1982) new combination; A. blanfordii bination; A. microscaphus (Cope, 1867) ``1866'' (Boulenger, 1882) new combination; A. brauni new combination; A. nelsoni (Stejneger, 1893) (Nieden, 1911) new combination; A. buchneri new combination; A. punctatus (Baird and Girard, (Peters, 1882) new combination; A. camerunensis 1852) new combination; A. quercicus (Holbrook, (Parker, 1936) new combination; A. chudeaui 1840) new combination; A. retiformis (Sanders (Chabanaud, 1919); A. cristiglans (Inger and and Smith, 1951) new combination; A. speciosus Menzies, 1961) new combination; A. danielae (Girard, 1854) new combination; A. terrestris (Perret, 1977) new combination; A. djohongensis (Bonnaterre, 1789) new combination; A. wood- (Hulselmans, 1977) new combination; A. fuligin- housii (Girard, 1854) new combination. atus (de Witte, 1932) new combination; A. funer- (4) Bufo Laurent, 1768. This taxon is Bufo (sen- eus (Bocage, 1866) new combination; A. garmani su stricto). All other species in Bufo (sensu lato) (Meek, 1897) new combination; A. gracilipes should have the generic name Bufo placed in quo- (Boulenger, 1899) new combination; A. gutturalis tation marks (i.e., ``Bufo'') inasmuch as they are (Power, 1927) new combination; A. kassasii (Baha not part of the clade formally called Bufo, sensu El Din, 1993) new combination; A. kerinyagae stricto. Members of Bufo sensu stricto are Bufo (Keith, 1968) new combination; A. kisoloensis andrewsi Schmidt, 1925; B. aspinius (Yang, Liu, (Loveridge, 1932) new combination; A. langan- and Rao, 1996); B. bankorensis Barbour, 1908; B. oensis (Largen, Tandy, and Tandy, 1978) new bufo (Linnaeus, 1758); B. gargarizans Cantor, combination; A. latifrons (Boulenger, 1900) new 1842; B. japonicus Temminck and Schlegel, 1838; combination; A. lemairii (Boulenger, 1901) new B. kabischi Herrmann and KuÈhnel, 1997; B. min- combination; A. maculatus (Hallowell, 1854) new shanicus Stejneger, 1926; B. spinosus Daudin, combination; A. pantherinus (Smith, 1828) new 1803; B. tibetanus Zarevskij, 1926; B. torrenticola combination; A. pardalis (Hewitt, 1935) new Matsui, 1976; B. tuberculatus Zarevskij, 1926; B. combination; A. perreti (Schiùtz, 1963) new com- verrucosissimus (Pallas, 1814); B. wolongensis bination; A. poweri (Hewitt, 1935) new combi- Herrmann and KuÈhnel, 1997. nation; A. rangeri (Hewitt, 1935) new combina- (5) Former ``Bufo'' species not allocated to ge- tion; A. reesi (Poynton, 1977) new combination; nus (see discussion under ``Taxonomy'' and above A. regularis (Reuss, 1833) new combination; A. under Bufo) are: steindachneri (Pfeffer, 1893) new combination; A. (a) Nomina dubia. ``Bufo'' brevirostris Rao, superciliaris (Boulenger, 1888) new combination; 1937; ``B.'' simus Schmidt, 1857. A. taiensis (RoÈdel and Ernst, 2000) new combi- (b) Unassigned to group. ``B.'' koynayensis So- nation (including Bufo amieti Tandy and Perret, man, 1963; ``B.'' ocellatus GuÈnther, 1858. 2000, according to S. Stuart, personal commun.); (c) ``Bufo'' arabicus group, ``Bufo'' arabicus A. togoensis (Ahl, 1924) new combination; A. tub- Heyden, 1827; ``B.'' dhufarensis Parker, 1931; erosus (GuÈnther, 1858) new combination; A. tur- ``B.'' dodsoni Boulenger, 1895; ``B.'' scorteccii kanae (Tandy and Feener, 1985) new combina- Balletto and Cherchi, 1970. tion; A. urunguensis (Loveridge, 1932) new com- (d) ``Bufo'' mauritanicus group: ``Bufo'' maur- bination; A. villiersi (Angel, 1940) new combi- itanicus Schlegel, 1841). nation; A. vittatus (Boulenger, 1906) new (e) ``Bufo'' pentoni group, ``Bufo'' pentoni An- combination; A. xeros (Tandy, Tandy, Keith, and derson, 1893; ``B.'' tihamicus Balletto and Cher- Duff-MacKay, 1976) new combination. chi, 1973. 364 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(f) ``Bufo'' scaber group: ``Bufo'' atukoralei (Duellman and Toft, 1979) new combination; C. Bogert and Senanayake, 1966; ``B.'' kotagamai ornatus (Spix, 1824) new combination; C. poep- Fernando, Dayawansa, and Siriwardhane, 1994; pigii (Tschudi, 1845) new combination; C. pom- ``B.'' parietalis Boulenger, 1882; ``B.'' scaber bali (Baldissera, Caramaschi, and Haddad, 2004) Schneider, 1799; ``B.'' silentvalleyensis Pillai, new combination; C. pygmaeus (Myers and Car- 1981. valho, 1952) new combination; C. quechua (Gal- (g) ``Bufo'' stejnegeri group. ``Bufo'' ailaoanus lardo, 1961) new combination; C. rubescens Kou, 1984; ``B.'' cryptotympanicus Liu and Hu, (Lutz, 1925) new combination; C. rubropunctatus 1962; ``B.'' pageoti Bourret, 1937; ``B.'' stejne- (Guichenot, 1848) new combination; C. rumbolli geri Schmidt, 1931. (Carrizo, 1992) new combination; C. schneideri (h) ``Bufo'' stomaticus group. ``Bufo'' beddomii (Werner, 1894) new combination; C. spinulosus GuÈnther, 1876; ``B.'' hololius GuÈnther, 1876; (Wiegmann, 1834) new combination; C. vellardi ``B.'' olivaceus Blanford, 1874; ``B.'' stomaticus (Leviton and Duellman, 1978) new combination; LuÈtken, 1864; ``B.'' stuarti Smith, 1929; ``B.'' su- C. veraguensis (Schmidt, 1857) new combination. matranus Peters, 1871 (provisional allocation); (7) Cranopsis Cope, 1875. Recognition of the ``B.'' valhallae Meade-Waldo, 1909 (provisional- Middle American clade of ``Bufo'' as Cranopsis ly allocated). Cope, 1875, renders the following new or revived (6) Chaunus Wagler, 1828: Recognition of this combinations: Cranopsis alvaria (Girard in Baird, major clade of predominantly Neotropical ``Bufo'' 1849) new combination; C. aucoinae (O'Neill and (excluding Rhinella) as a genus requires a number Mendelson, 2004) new combination; C. bocourti of new name combinations. Although we do not (Brocchi, 1877) new combination; C. campbelli reject the use of species groups (see Blair, 1972a; (Mendelson, 1994) new combination; C. canali- Duellman and Schulte, 1992), we think that these fera (Cope, 1877) new combination; C. cavifrons require considerable reevaluation regarding their (Firschein, 1950) new combination; C. coccifer monophyly and utility. Recommended changes (Cope, 1866) new combination; C. conifera are Chaunus abei (Baldissera, Caramaschi, and (Cope, 1862) new combination; C. cristata (Wieg- Haddad, 2004) new combination; C. achalensis mann, 1833) new combination; C. cycladen (Cei, 1972) new combination; C. achavali (Ma- (Lynch and Smith, 1966) new combination; C. neyro, Arrieta, and de SaÂ, 2004) new combina- fastidiosa (Cope, 1875) new combination; C. tion; C. amabilis (Pramuk and Kadivar, 2003) new gemmifer (Taylor, 1940) new combination; C. combination; C. amboroensis (Harvey and Smith, holdridgei (Taylor, 1952) new combination; C. 1993) new combination; C. arborescandens ibarrai (Stuart, 1954) new combination; C. leu- (Duellman and Schulte, 1992) new combination; comyos (McCranie and Wilson, 2000) new com- C. arenarum (Hensel, 1867) new combination; C. bination; C. luetkenii (Boulenger, 1891) new com- arequipensis (Vellard, 1959) new combination; C. bination; C. macrocristata (Firschein and Smith, arunco (Molina, 1782) new combination; C. ata- 1957) new combination; C. marmorea (Wieg- camensis (Cei, 1962) new combination; C. beebei mann, 1833) new combination; C. mazatlanensis (Gallardo, 1965) new combination; C. bergi (CeÂs- (Taylor, 1940) new combination; C. melanochlora pedez, 2000) new combination; C. chavin (Lehr, (Cope, 1877) new combination; C. nebulifer (Gi- KoÈhler, Aguilar, and Ponce, 2001) new combina- rard, 1854) new combination; C. occidentalis (Ca- tion; C. cophotis (Boulenger, 1900) new combi- merano, 1879) new combination; C. periglenes nation; C. corynetes (Duellman and Ochoa-M., (Savage, 1967); C. peripatetes (Savage, 1972) 1991) new combination; C. crucifer (Wied-Neu- new combination; C. perplexa (Taylor, 1943) new wied, 1821) new combination; C. diptychus combination; C. pisinna (Mendelson, Williams, (Cope, 1862) new combination; C. dorbignyi (Du- Sheil, and Mulcahy, 2005) new combination; C. meÂril and Bibron, 1841) new combination; C. fer- porteri (Mendelson, Williams, Sheil, and Mulca- nandezae (Gallardo, 1957) new combination; C. hy, 2005) new combination; C. signifera (Men- ®ssipes (Boulenger, 1903) new combination; C. delson, Williams, Sheil, and Mulcahy, 2005) new gallardoi (Carrizo, 1992) new combination; C. combination; C. spiculata (Mendelson, 1997) new gnustae (Gallardo, 1967) new combination; C. combination; C. tacanensis (Smith, 1852) new granulosus (Spix, 1824) new combination; C. combination; C. tutelaria (Mendelson, 1997) new henseli (Lutz, 1934) new combination; C. icteri- combination; C. valliceps (Wiegmann, 1833) new cus (Spix, 1824) new combination; C. inca (Stej- combination. neger, 1913) new combination; C. jimi (Stevaux, (8) Duttaphrynus new genus. Recognition of 2002) new combination; C. justinianoi (Harvey the former ``Bufo'' melanostictus group as a genus and Smith, 1994) new combination; C. limensis requires the following name changes: Duttaphry- (Werner, 1901) new combination; C. marinus nus crocus (Wogan, Win, Thin, Lwin, Shein, Kyi, (Linnaeus, 1758) new combination; C. nesiotes and Tun, 2003) new combination; D. cyphosus 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 365

(Ye, 1977) new combination; D. himalayanus 1941); P. gundlachi (Ruibal, 1959); P. lemur (GuÈnther, 1864) new combination; D. melanostic- Cope, 1869 ``1868''; P. longinasus (Stejneger, tus (Schneider, 1799) new combination; D. micro- 1905); P. peltocephala (Tschudi, 1838); P. taladai tympanum (Boulenger, 1882) new combination; (Schwartz, 1960). D. noellerti (Manamendra-Arachchi and Pethiya- (14) Phrynoidis Fitzinger, 1843. Recognition of goda, 1998) new combination. the former ``Bufo'' asper group as Phrynoidis re- (9) Epidalea Cope, 1864. Recognition as a ge- sults in two new combinations: Phrynoidis aspera nus of the former ``Bufo'' calamita group renders (Gravenhorst, 1829); P. juxtaspera (Inger, 1964) one revived name: Epidalea calamita (Laurenti, new combination. 1768). (15) Poyntonophrynus new genus. Recognition (10) Ingerophrynus new genus. Recognition of of the former ``Bufo'' vertebralis group as a genus the former ``Bufo'' biporcatus group ϩ allies re- results in the following new combinations: Poyn- sults in the following new combinations: Ingero- tonophrynus beiranus (Loveridge, 1932) new phrynus biporcatus (Gravenhorst, 1829) new combination; P. damaranus (Mertens, 1954) new combination; I. celebensis (GuÈnther, 1859 combination; P. dombensis (Bocage, 1895) new ``1858'') new combination; I. claviger (Peters, combination; P. fenoulheti (Hewitt and Methuen, 1863) new combination; I. divergens (Peters, 1912) new combination; P. grandisonae (Poynton 1871) new combination; I. galeatus (GuÈnther, and Haacke, 1993) new combination; P. hoeschi 1864) new combination; I. kumquat (Das and (Ahl, 1934) new combination; P. kavangensis Lim, 2001) new combination; I. macrotis (Bou- (Poynton and Broadley, 1988) new combination; lenger, 1887) new combination; I. parvus (Bou- P. lughensis (Loveridge, 1932) new combination; lenger, 1887) new combination; I. philippinicus P. parkeri (Loveridge, 1932) new combination; P. (Boulenger, 1887) new combination; I. quadri- vertebralis (Smith, 1848) new combination. porcatus (Boulenger, 1887) new combination. (16) Pseudepidalea new genus. Recognition of (11) Mertensophryne Tihen, 1960. Placing Ste- the former ``Bufo'' viridis group as a genus results phopaedes Channing, 1979 ``1978'' (as a subge- in the following new combinations: Pseudepida- nus) and the ``Bufo'' taitanus group within Mer- lea brongersmai (Hoogmoed, 1972) new combi- tensophryne provides the following new combi- nation; P. latastii (Boulenger, 1882) new combi- nations. nation; P. luristanica (Schmidt, 1952) new com- (a) Mertensophryne, unassigned to subgenus bination; P. oblonga (Nikolskii, 1896) new com- (the former ``Bufo'' taitanus group): Mertenso- bination; P. pewzowi (Bedriaga, 1898) new phryne lindneri (Mertens, 1955) new combina- combination; P. pseudoraddei (Mertens, 1971) tion; M. lonnbergi (Andersson, 1911) new com- new combination; P. raddei (Strauch, 1876) new bination; M. melanopleura (Schmidt and Inger, combination; P. surda (Boulenger, 1891) new 1959) new combination; M. micranotis (Loverid- combination; P. taxkorensis (Fei, 1999) new com- ge, 1925) new combination; M. mocquardi (An- bination; P. viridis (Laurenti, 1768) new combi- gel, 1924) new combination; M. nyikae (Lover- nation. idge, 1953) new combination; M. schmidti (Gran- (17) Rhaebo Cope, 1862. Recognition of the dison, 1972); M. taitana Peters, 1878 new com- former ``Bufo'' guttatus group results in the fol- bination; M. uzunguensis (Loveridge, 1932) new lowing name changes: Rhaebo anderssoni (Melin, combination. 1941) new combination; R. blombergi (Myers and (b) Mertensophryne, subgenus Stephopaedes: Funkhouser, 1951) new combination; R. caeru- Mertensophryne (Stephopaedes) anotis (Boulen- leostictus (GuÈnther, 1859) new combination ger, 1907) new combination; M. (S.) howelli [placed here on the basis of comments by Hoog- (Poynton and Clarke, 1999) new combination; M. moed, 1989a]; R. glaberrimus (GuÈnther, 1869) (S.) loveridgei (Poynton, 1991) new combination; new combination; R. guttatus (Schneider, 1799) Mertensophryne (Stephopaedes) usambarae new combination; R. haematiticus (Cope, 1862); (Poynton and Clarke, 1999). R. hypomelas Boulenger, 1913 (placed here pro- (12) Nannophryne GuÈnther, 1870: With the res- visionally). urrection of Nannophryne, the single species, (18) Rhinella Fitzinger, 1826. Recognition of Nannophryne variegata GuÈnther, 1870, takes its the former ``Bufo'' margaritifer group as the ge- original form. nus Rhinella results in the following name chang- (13) Peltophryne Fitzinger, 1843. Resurrection es: Rhinella acutirostris (Spix, 1824) new com- of Peltophryne results in the following names be- bination; R. alata (Thominot, 1884) new combi- ing revived: Peltophryne cataulaciceps (Schwartz, nation; R. castaneotica (Caldwell, 1991) new 1959); P. empusa Cope, 1862; P. ¯uviatica combination; R. ceratophrys (Boulenger, 1882) (Schwartz, 1972); P. fracta (Schwartz, 1972); P. new combination; R. cristinae (VeÂlez-Rodriguez fustiger (Schwartz, 1960); P. guentheri (Cochran, and Ruiz-Carranza, 2002) new combination; R. 366 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

dapsilis (Myers and Carvalho, 1945) new com- ARTHROLEPTIDAE bination; R. intermedia (GuÈnther, 1858) new com- (1) Arthroleptis Smith, 1849. The synonymy of bination; R. iserni JimeÂnez de la Espada, 1875; R. Schoutedenella de Witte, 1921, with Arthroleptis margaritifer (Laurenti, 1768) new combination; Smith, 1849, results in the following revived com- R. nasica (Werner, 1903) new combination; R. binations: Arthroleptis crusculum Angel, 1950; A. proboscidea (Spix, 1824); R. roqueana (Melin, discodactyla (Laurent, 1954); A. hematogaster 1941) new combination; R. scitula (Caramaschi (Laurent, 1954); A. lameerei de Witte, 1921; A. and Niemeyer, 2003) new combination; R. scler- loveridgei de Witte, 1933; A. milletihorsini Angel, ocephala (Mijares-Urrutia and Arends-R., 2001) 1922; A. mossoensis (Laurent, 1954); A. nimbaen- new combination; R. stanlaii (LoÈtters and KoÈhler, sis Angel, 1950; A. phrynoides (Laurent, 1976); 2000) new combination; R. sternosignata (GuÈn- A. pyrrhoscelis Laurent, 1952; A. schubotzi Nie- ther, 1858). den, 1911 ``1910''; A. spinalis Boulenger, 1919; (19) Vandijkophrynus new genus. Recognition A. sylvatica (Laurent, 1954); A. troglodytes Poyn- of the former ``Bufo'' angusticeps group as Van- ton, 1963; A. vercammeni (Laurent, 1954); A. xen- dijkophrynus results in the following name chang- ochirus Boulenger, 1905; A. xenodactyla Boulen- es: Vandijkophrynus amatolicus (Hewitt, 1925) ger, 1909; A. xenodactyloides Hewitt, 1933; A. new combination; V. angusticeps (Smith, 1848) zimmeri Ahl, 1925 ``1923''. new combination; V. gariepensis (Smith, 1848) new combination; V. inyangae (Poynton, 1963) new combination; V. robinsoni (Branch and HYPEROLIIDAE Braacke, 1996) new combination. (1) Hyperolius Rapp, 1842. The synonymy of Nesionixalus Perret, 1976, with Hyperolius Rapp, MICROHYLIDAE:ASTEROPHRYINAE 1842 (and recognition of Nesionixalus as a sub- genus), results in the following revived combi- (1) Xenorhina Peters, 1863. The synonymy of nations: Hyperolius (Nesionixalus) molleri (Be- Xenobatrachus Peters and Doria, 1878, with Xe- driaga, 1892); Hyperolius (Nesionixalus) thomen- norhina Peters, 1863, results in the following sis Bocage, 1886. changes: Xenorhina anorbis (Blum and Menzies, 1989 ``1988'') new combination; X. arfakiana (Blum and Menzies, 1989 ``1988'') new combi- CERATOBATRACHIDAE nation; X. bidens van Kampen, 1909; X. fuscigula (1) Ingerana Dubois, 1987 ``1986''. Treatment (Blum and Menzies, 1989 ``1988'') new combi- of Liurana Dubois, 1987 ``1986'', as a synonym nation; X. gigantea van Kampen, 1915; X. huon of Ingerana results in three name changes: Inger- (Blum and Menzies, 1989 ``1988'') new combi- ana alpina (Huang and Ye, 1997) new combina- nation; X. macrops van Kampen, 1913; X. mehelyi tion; I. medogensis (Fei, Ye, and Huang, 1997) (Boulenger, 1898) new combination; X. multisica new combination; I. reticulata (Zhao and Li, (Blum and Menzies, 1989 ``1988'') new combi- 1984) new combination. nation; X. obesa (Zweifel, 1960) new combina- tion; X. ocellata van Kampen, 1913; X. ophiodon PHRYNOBATRACHIDAE (Peters and Doria, 1878) new combination; X. rostrata (MeÂhely, 1898); X. scheepstrai (Blum (1) Phrynobatrachus GuÈnther, 1862. Placing and Menzies, 1989 ``1988'') new combination; X. Dimorphognathus Boulenger, 1906, into the syn- schiefenhoeveli (Blum and Menzies, 1989 onymy of Phrynobatrachus GuÈnther, 1862, ren- ``1988'') new combination; X. subcrocea (Men- ders Phrynobatrachus africanus (Hallowell, 1858 zies and Tyler, 1977) new combination; X. tumula ``1857'') new combination. Placing Phrynodon (Blum and Menzies, 1989 ``1988'') new combi- Parker, 1935, into the synonymy of Phrynobatra- nation; X. zweifeli (Kraus and Allison, 2002) new chus GuÈnther, 1862, renders Phrynobatrachus combination. sandersoni (Parker, 1935) new combination.

MICROHYLIDAE:COPHYLINAE PYXICEPHALIDAE (1) Rhombophryne Boettger, 1880. Transfer of (1) Amietia Dubois, 1987 ``1986''. Placing Af- several species of ``Plethodontohyla'' Boulenger, rana Dubois, 1992, into the synonymy of Amietia 1882, into Rhombophryne Boettger, 1880, renders Dubois, 1987 ``1986'', results in the following the following name changes: Rhombophryne al- name changes: Amietia amieti (Laurent, 1976) luaudi (Mocquard, 1901) new combination; new combination; A. angolensis (Bocage, 1866) Rhombophryne coudreaui (Angel, 1938) new new combination; A. desaegeri (Laurent, 1972) combination; Rhombophryne laevipes (Mocquard, new combination; A. dracomontana (Channing, 1895) new combination. 1978) new combination; A. fuscigula (DumeÂril 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 367 and Bibron, 1841) new combination; A. inyangae the synonymy of Occidozyga Kuhl and Van Has- (Poynton, 1966) new combination; A. johnstoni selt, 1822, presents the following revived combi- (GuÈnther, 1894 ``1893'') new combination; A. ru- nations: Occidozyga baluensis (Boulenger, 1896); wenzorica (Laurent, 1972) new combination; A. O. borealis (Annandale, 1912); O. celebensis vandijki (Visser and Channing, 1997) new com- Smith, 1927; O. diminutivus (Taylor, 1922); O. bination. ¯oresianus Mertens, 1927; O. laevis (GuÈnther, 1858); O. magnapustulosus (Taylor and Elbel, DICROGLOSSIDAE:DICROGLOSSINAE 1958); O. martensii (Peters, 1867); O. semipal- matus Smith, 1927; O. sumatrana (Peters, 1877); (1) Annandia Dubois, 1992. Treatment as a ge- O. vittatus (Andersson, 1942). nus produces a single new combination: Annandia delacouri (Angel, 1928) new combination. (This RHACOPHORIDAE combination was implied by Dubois, 2005.) (2) Eripaa Dubois, 1992. Treatment as a genus (1) Chiromantis Peters, 1854. The synonymy of produces a single new combination: Eripaa fas- Chirixalus Boulenger, 1893, with Chiromantis Pe- ciculispina (Inger, 1970) new combination. ters, 1854, presents the following new combina- (3) Nanorana GuÈnther, 1896. Placement of tions: Chiromantis cherrapunjiae (Roonwal and Chaparana Bourret, 1939, and Paa Dubois, 1975, Kripalani, 1966 ``1961'') new combination; C. into the synonymy of Nanorana GuÈnther, 1896, doriae (Boulenger, 1893) new combination; C. provides the following name changes: Nanorana dudhwaensis (Ray, 1992) new combination; C. aenea (Smith, 1922) new combination; N. annan- hansenae (Cochran, 1927) new combination; C. dalii (Boulenger, 1920) new combination; N. ar- laevis (Smith, 1924); C. nongkhorensis (Cochran, noldi (Dubois, 1975) new combination; N. bar- 1927) new combination; C. punctatus (Wilkinson, moachensis (Khan and Tasnim, 1989) new com- Win, Thin, Lwin, Shein, and Tun, 2003) new bination; N. blanfordii (Boulenger, 1882) new combination; C. shyamrupus (Chanda and Ghosh, combination; N. bourreti (Dubois, 1987 ``1986'') 1989) new combination; C. simus (Annandale, new combination; N. chayuensis (Ye, 1977) new 1915) new combination; C. vittatus (Boulenger, combination; N. conaensis (Fei and Huang, 1981) 1887) new combination. new combination; N. ercepeae (Dubois, 1974 (2) Feihyla new genus. We place only Philau- ``1973'') new combination; N. fansipani (Bourret, tus palpebralis Smith, 1924, into Feihyla,asFeih- 1939) new combination; N. feae (Boulenger, yla palpebralis (Smith, 1924), although we expect 1887) new combination; N. hazarensis (Dubois other species to be placed in this genus as data and Khan, 1979) new combination; N. liebigii emerge. (GuÈnther, 1860) new combination; N. liui (Du- (3) Kurixalus Ye, Fei, and Dubois, 1999. Be- bois, 1987 ``1986'') new combination; N. macu- cause the content of Kurixalus is controversial losa (Liu, Hu, and Yang, 1960) new combination; (see Delorme et al., 2005), we list the species we N. medogensis (Fei and Ye, 1999) new combina- regard as being in this monophyletic group: Ku- tion; N. minica (Dubois, 1975) new combination; rixalus eif®ngeri (Boettger, 1895); K. idiootocus N. mokokchungensis (Das and Chanda, 2000) new (Kuramoto and Wang, 1987) new combination; combination; N. polunini (Smith, 1951) new com- and provisionally Kurixalus verrucosus (Boulen- bination; N. quadranus (Liu, Hu, and Yang, 1960) ger, 1893) new combination (based on the tree, new combination; N. rarica (Dubois, Matsui, and data undisclosed, presented by Delorme et al., Ohler, 2001) new combination; N. robertingeri 2005). (Wu and Zhao, 1995) new combination; N. ros- tandi (Dubois, 1974 ``1973'') new combination; RANIDAE N. sternosignata (Murray, 1885) new combina- tion; N. taihangnica (Chen and Jiang, 2002) new Because of the extensive changes in ranid tax- combination; N. unculuanus (Liu, Hu, and Yang, onomy, we provide a listing of all recognized gen- 1960) new combination; N. vicina (Stoliczka, era and species, noting new combinations. 1872) new combination; N. yunnanensis (Ander- (1) Amolops Cope, 1865. Amolops bellulus Liu, son, 1879 ``1878'') new combination. Yang, Ferraris, and Matsui, 2000; A. chakrataen- (4) Ombrana Dubois, 1992. Treatment as a ge- sis Ray, 1992; A. chunganensis (Pope, 1929); A. nus produces a single new combination: Ombrana cremnobatus Inger and Kottelat, 1998; A. daiyu- sikimensis (Jerdon, 1870) new combination. nensis (Liu and Hu, 1975); A. formosus (GuÈnther, 1876 ``1875''); A. gerbillus (Annandale, 1912); A. granulosus (Liu and Hu, 1961); A. hainanensis DICROGLOSSIDAE:OCCIDOZYGINAE (Boulenger, 1900 ``1899''); A. himalayanus (Bou- (1) Occidozyga Kuhl and Van Hasselt, 1822. lenger, 1888); A. hongkongensis (Pope and Rom- Replacement of Phrynoglossus Peters, 1867, into er, 1951); A. jaunsari Ray, 1992; A. jinjiangensis 368 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Su, Yang, and Li, 1986; A. kangtingensis (Liu, Fei, and Ye, 2001) new combination; H. khalam 1950); A. kaulbacki (Smith, 1940); A. larutensis (Stuart, Orlov, and Chan-ard, 2005) new combi- (Boulenger, 1899); A. liangshanensis (Wu and nation; H. kuangwuensis (Liu and Hu, 1966) new Zhao, 1984); A. lifanensis (Liu, 1945); A. loloen- combination; H. leporipes (Werner, 1930) new sis (Liu, 1950); A. longimanus (Andersson, 1939 combination; H. livida (Blyth, 1856 ``1855'') new ``1938''); A. mantzorum (David, 1872 ``1871''); combination; H. lungshengensis (Liu and Hu, A. marmoratus (Blyth, 1855); A. mengyangensis 1962) new combination; H. margaretae (Liu, Wu and Tian, 1995; A. monticola (Anderson, 1950) new combination; H. masonii (Boulenger, 1871); A. nepalicus Yang, 1991; A. ricketti (Bou- 1884); H. megatympanum (Bain, Lathrop, Mur- lenger, 1899); A. spinapectoralis Inger, Orlov, and phy, Orlov, and Ho, 2003) new combination; H. Darevsky, 1999; A. tormotus (Wu, 1977); A. tor- melasma Stuart and Chan-ard, 2005; H. modigli- rentis (Smith, 1923); A. tuberodepressus Liu and anii (Doria, Salvidio, and Tavano, 1999); H. mor- Yang, 2000; A. viridimaculatus (Jiang, 1983); A. afkai (Bain, Lathrop, Murphy, Orlov, and Ho, wuyiensis (Liu and Hu, 1975). 2003) new combination; H. narina (Stejneger, (2) Babina Thompson, 1912. Babina adeno- 1901) new combination; H. nasica (Boulenger, pleura (Boulenger, 1909) new combination; B. 1903); H. nasuta Li, Ye, and Fei, 2001 new com- caldwelli (Schmidt, 1925) new combination; B. bination; H. schmackeri (Boettger, 1892) new chapaensis (Bourret, 1937) new combination; B. combination; H. sinica (Ahl, 1927 ``1925'') new daunchina (Chang, 1933) new combination; B. combination; H. sumatrana Yang, 1991; H. su- holsti (Boulenger, 1892) new combination; B. lini pranarina (Matsui, 1994) new combination; H. (Chou, 1999) new combination; B. pleuraden swinhoana (Boulenger, 1903) new combination; (Boulenger, 1904) new combination; B. psaltes H. tabaca (Bain and Nguyen, 2004) new combi- (Kuramoto, 1985) new combination; B. subaspera nation; H. tiannanensis (Yang and Li, 1980) new (Barbour, 1908). combination; H. trankieni (Orlov, Ngat, and Ho, (3) Clinotarsus Mivart, 1869. Clinotarsus cur- 2003) new combination; H. utsunomiyaorum tipes (Jerdon, 1854 ``1853''). (Matsui, 1994) new combination; H. versabilis (4) Glandirana Fei, Ye, and Huang, 1991 (Liu and Hu, 1962) new combination; H. wuch- ``1990''. Glandirana emeljanovi (Nikolskii, 1913) uanensis (Xu, 1983) new combination. new combination; G. minima (Ting and T'sai, (6) Humerana Dubois, 1992. Humerana hu- 1979); G. rugosa (Temminck and Schlegel, 1838) meralis (Boulenger, 1887) new combination; Hu- new combination; R. tientaiensis (Chang, 1933 merana miopus (Boulenger, 1918) new combina- ``1933±1934'') new combination. tion; Humerana oatesii (Boulenger, 1892) new (5) Huia Yang, 1991. Huia absita Stuart and combination. Chan-ard, 2005; H. amamiensis (Matsui, 1994) (7) Hydrophylax Fitzinger, 1843. Hydrophylax new combination; H. andersonii (Boulenger, albolabris (Hallowell, 1856); H. albotuberculatus 1882) new combination; H. anlungensis (Liu and (Inger, 1954) new combination; H. amnicola (Per- Hu, 1973) new combination; H. archotaphus (In- ret, 1977) new combination; H. asperrima (Perret, ger and Chan-ard, 1997) new combination; H. 1977) new combination; H. chalconotus (Schle- bacboensis (Bain, Lathrop, Murphy, Orlov, and gel, 1837) new combination; H. crassiovis (Bou- Ho, 2003) new combination; H. banaorum (Bain, lenger, 1920) new combination; H. darlingi (Bou- Lathrop, Murphy, Orlov, and Ho, 2003) new com- lenger, 1902) new combination; H. everetti (Bou- bination; H. cavitympanum (Boulenger, 1893); lenger, 1882) new combination; H. galamensis Huia chapaensis (Bourret, 1937) new combina- (DumeÂril and Bibron, 1841); H. igorotus (Taylor, tion; H. chloronota (GuÈnther, 1876 ``1875'') new 1922) new combination; H. kampeni (Boulenger, combination; H. daorum (Bain, Lathrop, Murphy, 1920) new combination; H. lemairei (de Witte, Orlov, and Ho, 2003) new combination; H. exiliv- 1921) new combination; H. lepus (Andersson, ersabilis (Li, Ye, and Fei, 2001) new combination; 1903) new combination; H. longipes (Perret, H. grahami (Boulenger, 1917) new combination; 1960) new combination; H. luzonensis (Boulen- H. graminea (Boulenger, 1900 ``1899'') new com- ger, 1896) new combination; H. macrops (Bou- bination; H. hainanensis (Fei, Ye, and Li, 2001) lenger, 1897) new combination; H. malabaricus new combination; H. hejiangensis (Deng and Yu, (Tschudi, 1838); H. occidentalis (Perret, 1960) 1992) new combination; H. hmongorum (Bain, new combination; H. parkerianus (Mertens, Lathrop, Murphy, Orlov, and Ho, 2003) new com- 1938); H. raniceps (Peters, 1871) new combina- bination; H. hosii (Boulenger, 1891) new combi- tion; H. tipanan (Brown, McGuire, and Diesmos, nation; H. iriodes (Bain and Nguyen, 2004) new 2000) new combination. combination; H. ishikawae (Stejneger, 1901) new (8) Hylarana Tschudi, 1838. Hylarana ery- combination; H. jingdongensis (Fei, Ye, and Li, thraea (Schlegel, 1837); H. guentheri (Boulenger, 2001) new combination; H. junlianensis (Huang, 1882); H. macrodactyla GuÈnther, 1858; H. taipe- 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 369 hensis (Van Denburgh, 1909) new combination; ther, 1872); M. kinabaluensis (Inger, 1966); M. Hylarana tytleri Theobald, 1868. macrophthalmus (Matsui, 1986); M. orphnocnem- (9) Lithobates Fitzinger, 1843. L. areolatus is (Matsui, 1986); M. phaeomerus (Inger and Gri- (Baird and Girard, 1852) new combination; Lith- tis, 1983); M. poecilus (Inger and Gritis, 1983); obates berlandieri (Baird, 1859) new combina- M. whiteheadi (Boulenger, 1887). tion; Lithobates blairi (Mecham, Littlejohn, Old- (11) Nasirana Dubois, 1992. Nasirana alticola ham, Brown, and Brown, 1973) new combination; (Boulenger, 1882) new combination. (Sanders, 1973); L. bwana (12) Pelophylax Fitzinger, 1843.37 Pelophylax (Hillis and de SaÂ, 1988) new combination; L. cap- bedriagae (Camerano, 1882 ``1881'') new com- ito (LeConte, 1855) new combination; L. cates- bination; P. bergeri (GuÈnther, 1986) new combi- beianus (Shaw, 1802) new combination; L. chi- nation; P. cerigensis (Beerli, Hotz, Tunner, Hep- chicuahutla (Cuellar, MeÂndez-De La Cruz, and pich, and Uzzell, 1994) new combination; P. cho- VillagraÂn-Santa Cruz, 1996) new combination; L. senicus (Okada, 1931) new combination; P. cre- chiricahuensis (Platz and Mecham, 1979) new tensis (Beerli, Hotz, Tunner, Heppich, and Uzzell, combination; L. clamitans (Latreille, 1801) new 1994) new combination; P. demarchii (Scortecci, combination; L. dunni (Zweifel, 1957) new com- 1929) new combination; P. epeiroticus (Schnei- bination; L. ®sheri (Stejneger, 1893) new combi- der, So®anidou, and Kyriakopoulou-Sklavounou, nation; L. forreri (Boulenger, 1883) new combi- 1984) new combination; P. fukienensis (Pope, nation; L. grylio (Stejneger, 1901) new combina- 1929) new combination; P. hubeiensis (Fei and tion; L. heckscheri (Wright, 1924) new combina- Ye, 1982); P. kurtmuelleri (Gayda, 1940 tion; L. johni (Blair, 1965) new combination; L. ``1939''); P. lateralis (Boulenger, 1887) new com- juliani (Hillis and de SaÂ, 1988) new combination; bination; P. lessonae (Camerano, 1882 ``1881'') L. lemosespinali (Smith and Chiszar, 2003) new new combination; P. nigrolineatus (Liu and Hu, combination; L. macroglossa (Brocchi, 1877) new 1960 ``1959''); P. nigromaculatus (Hallowell, combination; L. maculatus (Brocchi, 1877) new 1861 ``1860''); P. perezi (Seoane, 1885); P. plan- combination; L. magnaocularis (Frost and Bag- cyi (Lataste, 1880) new combination; P. porosus nara, 1974) new combination; L. megapoda (Tay- (Cope, 1868) new combination; P. ridibundus lor, 1942) new combination; L. miadis (Barbour (Pallas, 1771); P. saharicus (Boulenger, 1913) and Loveridge, 1929) new combination; L. mon- new combination; P. shqipericus (Hotz, Uzzell, tezumae (Baird, 1854) new combination; L. neo- GuÈnther, Tunner, and Heppich, 1987) new com- volcanicus (Hillis and Frost, 1985) new combi- bination; P. shuchinae (Liu, 1950); P. tengger- nation; L. okaloosae (Moler, 1985) new combi- ensis (Zhao, Macey, and Papenfuss, 1988) new nation; L. omiltemanus (GuÈnther, 1900) new com- combination; P. terentievi (Mezhzherin, 1992) bination; L. onca (Cope, 1875) new combination; new combination. L. palmipes (Spix, 1824); L. palustris (LeConte, (13) Pterorana Kiyasetuo and Khare, 1986. 1825) new combination; L. pipiens (Schreber, Pterorana khare Kiyasetuo and Khare, 1986. 1782) new combination; L. psilonota (Webb, (14) Pulchrana Dubois, 1992. P. banjarana 2001) new combination; L. pueblae (Zweifel, (Leong and Lim, 2003) new combination; P. bar- 1955) new combination; L. pustulosus (Boulenger, amica (Boettger, 1900) new combination; P. de- 1883) new combination; L. septentrionalis (Baird, bussyi (van Kampen, 1910) new combination; P. 1854) new combination; L. sevosus (Goin and glandulosa (Boulenger, 1882) new combination; Netting, 1940) new combination; L. sierramad- P. grandocula (Taylor, 1920) new combination; P. rensis (Taylor, 1939 ``1938'') new combination; laterimaculata (Barbour and Noble, 1916) new L. spectabilis (Hillis and Frost, 1985) new com- combination; P. luctuosa (Peters, 1871) new com- bination; L. sphenocephalus (Cope, 1886) new bination; P. mangyanum (Brown and Guttman, combination; L. sylvaticus (LeConte, 1825) new 2002) new combination; P. melanomenta (Taylor, combination; L. tarahumarae (Boulenger, 1917) 1920) new combination; P. moellendorf® (Boett- new combination; L. taylori (Smith, 1959) new ger, 1893) new combination; P. picturata (Bou- combination; L. tlaloci (Hillis and Frost, 1985) lenger, 1920) new combination; P. siberu (Dring, new combination; L. vaillanti (Brocchi, 1877) new combination; L. vibicarius (Cope, 1894) new 37 We concur with Bogart (2003) that named hybri- combination; L. virgatipes (Cope, 1891) new dogens/kleptons are composed of hybrids, not covered combination; L. warszewitschii (Schmidt, 1857) by regulated Linnaean nomenclature. This does not new combination; L. yavapaiensis (Platz and mean that we reject the utilitarian naming conventions suggested by Dubois (1982: e.g., Pelophylax kl. escu- Frost, 1984) new combination; L. zweifeli (Hillis, lentus, Pelophylax kl. gra®), for denoting kinds of frog, Frost, and Webb, 1984) new combination. only that these names do not represent taxa in any evo- (10) Meristogenys Yang, 1991: Meristogenys lutionary/phylogenetic sense, but instead are ``kinds'' of amoropalamus (Matsui, 1986); M. jerboa (GuÈn- frogs. 370 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

McCarthy, and Whitten, 1990) new combination; (18) Sylvirana Dubois, 1992. Sylvirana arfaki P. signata (GuÈnther, 1872) new combination; P. (Meyer, 1875 ``1874'') new combination; S. attig- similis (GuÈnther, 1873) new combination. ua (Inger, Orlov, and Darevsky, 1999) new com- (15) Rana Linnaeus, 1758. bination; S. aurantiaca (Boulenger, 1904) new Boulenger, 1886; R. arvalis Nilsson, 1842; R. combination; S. aurata (GuÈnther, 2003) new com- asiatica Bedriaga, 1898; R. aurora Baird and Gi- bination; S. bannanica (Rao and Yang, 1997) new rard, 1852; R. boylii Baird, 1854; R. camerani combination; S. celebensis (Peters, 1872) new Boulenger, 1886; R. cascadae Slater, 1939; R. combination; S. chitwanensis (Das, 1998) new chaochiaoensis Liu, 1946; R. chensinensis David, combination; S. cubitalis (Smith, 1917) new com- 1875; R. chevronta Hu and Ye, 1978; R. dalma- bination; S. daemeli (Steindachner, 1868) new tina Fitzinger In Bonaparte, 1839; R. draytonii combination; S. danieli (Pillai and Chanda, 1977) Baird and Girard, 1852; R. dybowskii GuÈnther, new combination; S. elberti (Roux, 1911) new 1876; R. graeca Boulenger, 1891; R. holtzi Wer- combination; S. faber (Ohler, Swan, and Daltry, ner, 1898; R. huanrenensis Fei, Ye, and Huang, 2002) new combination; S. ¯orensis (Boulenger, 1991 ``1990''; R. iberica Boulenger, 1879; R. it- 1897) new combination; S. garoensis (Boulenger, alica Dubois, 1987 ``1985''; R. japonica Boulen- 1920) new combination; S. garritor (Menzies, ger, 1879; R. johnsi Smith, 1921; R. kukunoris 1987) new combination; S. gracilis (Gravenhorst, Nikolskii, 1918; R. kunyuensis Lu and Li, 2002; 1829) new combination; S. grisea (van Kampen, R. latastei Boulenger, 1879; R. longicrus Stejne- 1913) new combination; S. jimiensis (Tyler, 1963) ger, 1898; R. luteiventris Thompson, 1913; R. ma- new combination; S. kreffti (Boulenger, 1882) crocnemis Boulenger, 1885; R. multidenticulata new combination; S. latouchii (Boulenger, 1899) Chou and Lin, 1997; R. muscosa Camp, 1917; R. new combination; S. leptoglossa (Cope, 1868) okinavana Boettger, 1895; R. omeimontis Ye and new combination; S. maosonensis (Bourret, 1937) Fei, 1993; R. ornativentris Werner, 1903; R. pirica new combination; S. margariana (Anderson, 1879 Matsui, 1991; R. pretiosa Baird and Girard, 1853; ``1878'') new combination; S. milleti (Smith, R. pyrenaica Serra-Cobo, 1993; Rana sakuraii 1921) new combination; S. moluccana (Boettger, Matsui and Matsui, 1990; R. sangzhiensis Shen, 1986; R. sauteri Boulenger, 1909; R. tagoi Okada, 1895) new combination; S. montivaga (Smith, 1928; R. temporaria Linnaeus, 1758; R. tsushi- 1921) new combination; S. mortenseni (Boulen- mensis Stejneger, 1907; R. weiningensis Liu, Hu, ger, 1903) new combination; S. nigrotympanica and Yang, 1962; R. zhengi Zhao, 1999; R. zhen- (Dubois, 1992) new combination; S. nigrovittata haiensis Ye, Fei, and Matsui, 1995. (Blyth, 1856 ``1855'') new combination; S. no- (16) Sanguirana Dubois, 1992. Sanguirana vaeguineae (van Kampen, 1909) new combina- sanguinea (Boettger, 1893). new combination; S. tion; S. papua (Lesson, 1831) new combination; varians (Boulenger, 1894) new combination. S. persimilis (van Kampen, 1923) new combina- (17) Staurois Cope, 1865. Staurois latopalma- tion; S. spinulosa (Smith, 1923) new combination; tus (Boulenger, 1887); S. natator (GuÈnther, 1858); S. supragrisea (Menzies, 1987) new combination; S. nubilis (Mocquard, 1890); S. tuberilinguis Bou- S. temporalis (GuÈnther, 1864) new combination; lenger, 1918. S. volkerjane (GuÈnther, 2003) new combination.