Table Of ContentTHE AMPHIBIAN TREE OF LIFE
DARREL R. FROST,1 TARAN GRANT,1,4 JULIA´N FAIVOVICH,1,4
RAOUL H. BAIN,1,2 ALEXANDER HAAS,5 CE´LIO F.B. HADDAD,6
RAFAEL O. DE SA´ ,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 VertebrateZoology (Herpetology),2Centerfor Biodiversityand Conservation, and
3Division of InvertebrateZoology, American Museum of Natural History
(DRF: [email protected]; TG: [email protected]; JF: [email protected];
RHB: [email protected]; CJR: [email protected]; WCW: [email protected]);
4Department of Ecology, Evolution, and EnvironmentalBiology,
Columbia University,New York, NY 10027;
5Biocenter Grindel and Zoological Museum Hamburg, Martin-Luther-King-Platz3,
D-20146 Hamburg, Germany([email protected]);
6Departamento de Zoologia, Instituto de Biocieˆncias, UniversidadeEstadual Paulista (UNESP),
Caixa Postal 199, 13506-900 Rio Claro, Sa˜o Paulo, Brazil ([email protected]);
7Department of Biology, Universityof Richmond, Richmond, VA 23173-0001 ([email protected]);
8Department of Biodiversityand Conservation Biology, Universityof the Western Cape,
Private Bag X17, Bellville 7535, South Africa ([email protected]);
9Department of Zoology, The Natural History Museum, CromwellRoad,
London SW7 5BD, UK ([email protected]);
10South Australia Museum, Evolutionary Biology Unit, North Terrace,
Adelaide 5000, South Australia ([email protected]);
11Department of Biology, Universityof Texas at Arlington, TX 76019-0001 ([email protected]);
12Division Herpetolog´ıa, Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’,
Angel Gallardo 470, 1405 Buenos Aires, Argentina ([email protected]);
13Wildlife ResearchLaboratory, Florida Fish and Wildlife ConservationCommission,
4005 South Main Street, Gainesville,FL 32601-9075 ([email protected]);
14Department of Herpetology, California Academyof Sciences, 875 Howard Street,
San Francisco, CA 94103-3009 ([email protected]);
15Museum of Zoology and Departmentof Ecology and Evolutionary Biology, Universityof Michigan,
1109 Geddes Avenue, Ann Arbor, MI 48109-1079 ([email protected]);
16Instituto de Ciencias Naturales, Universidad Nacional de Colombia,
Apartado 7495, Bogota´, 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 figures, 5 tables, 7 appendices
Issued March 15, 2006
CopyrightqAmericanMuseumofNaturalHistory2006 ISSN0003-0090
CONTENTS
Abstract ....................................................................... 8
Introduction ................................................................... 11
Materials and Methods ......................................................... 13
Conventions and Abbreviations ................................................ 13
General Analytical Approach: Theoretical Considerations ......................... 13
Taxon Sampling ............................................................. 14
Character Sampling .......................................................... 15
Laboratory Protocols ......................................................... 17
Sequence Preanalysis: Heuristic Error Checking ................................. 17
Molecular Sequence Formatting ............................................... 17
Analytical Strategy .......................................................... 18
Review of Current Taxonomy, the Questions, and Taxon Sampling .................. 22
Comparability of Systematic Studies ........................................... 22
Amphibia ................................................................... 23
Gymnophiona ............................................................... 24
Rhinatrematidae ........................................................... 25
Ichthyophiidae and Uraeotyphlidae .......................................... 25
Scolecomorphidae ......................................................... 25
Typhlonectidae ............................................................ 25
‘‘Caeciliidae’’ ............................................................. 25
Caudata .................................................................... 26
Sirenidae ................................................................. 26
Hynobiidae ............................................................... 28
Cryptobranchidae .......................................................... 28
Proteidae ................................................................. 28
Rhyacotritonidae .......................................................... 30
Amphiumidae ............................................................. 30
Plethodontidae ............................................................ 30
Salamandridae ............................................................ 36
Dicamptodontidae ......................................................... 37
Ambystomatidae .......................................................... 37
Anura ...................................................................... 38
‘‘Primitive’’ Frogs ........................................................... 41
Ascaphidae ............................................................... 41
Leiopelmatidae ............................................................ 41
Discoglossidae and Bombinatoridae .......................................... 44
‘‘Transitional’’ Frogs ......................................................... 45
Pipoidea .................................................................. 46
Rhinophrynidae ........................................................... 47
Pipidae ................................................................... 47
Pelobatoidea .............................................................. 48
Pelobatidae and Scaphiopodidae ............................................. 49
Pelodytidae ............................................................... 49
Megophryidae ............................................................. 49
‘‘Advanced’’ Frogs—Neobatrachia ............................................. 50
‘‘Hyloidea’’ ................................................................. 52
Heleophrynidae ........................................................... 52
Sooglossidae and Nasikabatrachidae ......................................... 53
Limnodynastidae, Myobatrachidae, and Rheobatrachidae ....................... 53
‘‘Leptodactylidae’’ ......................................................... 56
‘‘Ceratophryinae’’ ......................................................... 57
‘‘Cycloramphinae’’ ........................................................ 59
3
4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297
Eleutherodactylinae ........................................................ 60
Leptodactylinae ........................................................... 61
‘‘Telmatobiinae’’ .......................................................... 61
‘‘Hemiphractinae’’ ......................................................... 62
Brachycephalidae .......................................................... 63
Rhinodermatidae .......................................................... 63
Dendrobatidae ............................................................. 63
Allophrynidae ............................................................. 64
Centrolenidae ............................................................. 64
Hylidae .................................................................. 64
Hylinae .................................................................. 64
Pelodryadinae ............................................................. 65
Phyllomedusinae .......................................................... 65
Bufonidae ................................................................ 67
Ranoidea ................................................................... 70
Arthroleptidae, Astylosternidae, and Hyperoliidae ............................. 72
Arthroleptidae ............................................................. 72
Astylosternidae ............................................................ 72
Hyperoliidae .............................................................. 74
Hemisotidae .............................................................. 74
Microhylidae .............................................................. 75
Scaphiophyrninae .......................................................... 77
Asterophryinae and Genyophryninae ......................................... 77
Brevicipitinae ............................................................. 78
Cophylinae ............................................................... 78
Dyscophinae .............................................................. 79
Melanobatrachinae ......................................................... 79
Microhylinae .............................................................. 80
Phrynomerinae ............................................................ 80
‘‘Ranidae’’ ................................................................ 81
Ceratobatrachinae .......................................................... 81
Conrauinae ............................................................... 83
Dicroglossinae ............................................................ 83
Lankanectinae ............................................................. 89
Micrixalinae .............................................................. 90
Nyctibatrachinae ........................................................... 91
Petropedetinae, Phrynobatrachinae, and Pyxicephalinae ........................ 91
Ptychadeninae ............................................................. 93
‘‘Raninae’’ ................................................................ 93
Ranixalinae .............................................................. 106
Rhacophoridae and Mantellidae ............................................ 106
Results ...................................................................... 110
Sequence Length Variation and Notes on Analysis .............................. 110
Topological Results and Discussion ........................................... 111
Outgroup Relationships ...................................................... 112
Amphibia (Lissamphibia) and Batrachia ....................................... 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
Neobatrachia ............................................................. 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 Amphibians ............................................. 141
Taxonomic Accounts ........................................................ 147
Amphibia Gray, 1825 ..................................................... 164
Gymnophiona Mu¨ller, 1832 ................................................ 165
Rhinatrematidae Nussbaum, 1977 .......................................... 165
Stegokrotaphia Cannatella and Hillis, 1993 ................................ 166
Ichthyophiidae Taylor, 1968 ............................................. 166
Caeciliidae Rafinesque, 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 Gu¨nther, 1859 ‘‘1858’’ ................................... 183
Sokolanura new taxon .................................................. 183
Costata Lataste, 1879 ................................................... 184
Alytidae 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
Hemiphractidae Peters, 1862 ............................................. 196
Meridianura new taxon ................................................. 196
Brachycephalidae Gu¨nther, 1858 ......................................... 197
Cladophrynia new taxon ................................................ 201
Cryptobatrachidae new family ........................................... 201
Tinctanura new taxon .................................................. 201
Amphignathodontidae Boulenger, 1882 .................................... 202
Athesphatanura new taxon .............................................. 202
Hylidae Rafinesque, 1815 ............................................... 202
Hylinae Rafinesque, 1815 ............................................... 203
Pelodryadinae Gu¨nther, 1858 ............................................ 204
Phyllomedusinae Gu¨nther, 1858 .......................................... 205
Leptodactyliformes new taxon ........................................... 205
Diphyabatrachia new taxon ............................................. 205
Centrolenidae Taylor, 1951 .............................................. 206
Leptodactylidae Werner, 1896 (1838) ..................................... 207
Chthonobatrachia new taxon ............................................ 208
Ceratophryidae Tschudi, 1838 ........................................... 208
Hesticobatrachia new taxon ............................................. 209
Cycloramphidae Bonaparte, 1850 ......................................... 210
Agastorophrynia new taxon ............................................. 210
Dendrobatoidea 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 Gu¨nther, 1858 (1843) ....................................... 224
Asterophryinae Gu¨nther, 1858 ........................................... 227
Cophylinae Cope, 1889 ................................................. 227
Dyscophinae Boulenger, 1882 ............................................ 228
Gastrophryninae Fitzinger, 1843 .......................................... 228
Melanobatrachinae Noble, 1931 .......................................... 229
Microhylinae Gu¨nther, 1858 (1843) ....................................... 229
Scaphiophryninae Laurent, 1946 ......................................... 230
Afrobatrachia new taxon ................................................ 231
Xenosyneunitanura new taxon ........................................... 231
Brevicipitidae Bonaparte, 1850 ........................................... 231
Hemisotidae Cope, 1867 ................................................ 232
Laurentobatrachia new taxon ............................................ 232
Hyperoliidae Laurent, 1943 .............................................. 232
Arthroleptidae Mivart, 1869 ............................................. 233
Natatanura new taxon .................................................. 234
Ptychadenidae Dubois, 1987 ‘‘1986’’ ..................................... 235
Victoranura new taxon ................................................. 235
Ceratobatrachidae 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
Petropedetidae Noble, 1931 .............................................. 238
Pyxicephalidae Bonaparte, 1850 .......................................... 240
Saukrobatrachia new taxon .............................................. 241
Dicroglossidae 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 Rafinesque, 1814 .............................................. 247
Nyctibatrachidae Blommers-Schlo¨sser, 1993 ............................... 247
Ranidae Rafinesque, 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 Clarifications Regarding
Taxonomic Content ......................................................... 359
ABSTRACT
The evidentiary basis of the currently accepted classification 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 deficiencies 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, ł 2,400 bp of mitochondrial sequences) and the nuclear genes
histone H3, rhodopsin, tyrosinase, and seven in absentia, and the large ribosomal subunit28S
(ł2,300bpofnuclearsequences;ca.1.8millionbasepairs;x¯ 53.7kb/terminal).Thedataset
includes 532 terminals sampled from 522 species 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
monophylyofall‘‘family-group’’taxa.Allcurrentlyrecognizednominalfamiliesandsubfam-
iliesweresampled,withtheexceptionofProtohynobiinae(Hynobiidae).Manyofthecurrently
recognized genera were also sampled. Although we discuss the monophyly of genera, and
provideremediesfornonmonophylywherepossible,wealsomakerecommendationsforfuture
research.
A parsimony analysis was performed under Direct Optimization, which simultaneouslyop-
timizes nucleotide homology (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), Salamandroidea
(paraphyletic with respect to Sirenidae), Leiopelmatanura (paraphyletic with respect to Asca-
phidae), Discoglossanura (paraphyletic with respect to Bombinatoridae), Mesobatrachia (par-
aphyleticwithrespecttoNeobatrachia),Pipanura(paraphyleticwithrespecttoBombinatoridae
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 1 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 1 Astylosternidae), Astylosternidae (paraphyletic
withrespecttoArthroleptinae),Ranidae(paraphyleticwithrespecttoRhacophoridaeandMan-
tellidae). In addition, many subsidiary taxa are demonstrated to be nonmonophyletic, such as
(1) Eleutherodactylus with respect to Brachycephalus; (2) Rana (sensu Dubois, 1992), which
is polyphyletic, with various elements falling far from each other on the tree; and (3) Bufo,
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,
caecilians/Gymnophiona, salamanders/Caudata, and frogs/Anura, form a monophyletic group,
towhichwerestrictthenameAmphibia;(2)GymnophionaformsthesistertaxonofBatrachia
(salamanders 1 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 assub-
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(CryptobranchidaeandHynobiidae)andDiadectosalamandroideinewtaxon(allother
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-
mandroideinewtaxon;(7)PerennibranchiaiscomposedofProteidaeandSirenidae;(8)Trep-
tobranchia new taxon is composed of two taxa, Ambystomatidae (including Dicamptodonti-
dae)andSalamandridae;(9)PlethosalamandroideinewtaxoniscomposedofRhyacotritonidae
andXenosalamandroideinewtaxon;(10)XenosalamandroideiiscomposedofPlethodontidae
and Amphiumidae; (11) Anura is monophyletic and composed of two clades, Leiopelmatidae
(includingAscaphidae)andLalagobatrachianewtaxon(allotherfrogs);(12)Lalagobatrachia
is composed of two clades, Xenoanura (Pipidae and Rhinophrynidae) and Sokolanura new
taxon(allotherlalagobatrachians);(13)BombinatoridaeandAlytidae(formerDiscoglossidae)
are each others’ closest relatives and in a clade called Costata, which, excluding Leiopelma-
tidae and Xenoanura, forms the sister taxon of all other frogs, Acosmanura;(14)Acosmanura
is composed of two clades, Anomocoela (5 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,togetherform-
ing Acosmanura; (16)Neobatrachia iscomposedoftwoclades,Heleophrynidae,andallother
neobatrachians,Phthanobatrachianewtaxon;(17)Phthanobatrachiaiscomposedoftwomajor
units,HyloidesandRanoides;(18)HyloidescomprisesSooglossidae(includingNasikabatrach-
idae) and Notogaeanura new taxon (the remaining hyloids); (19) Notogaeanura contains two
taxa, Australobatrachia new taxon and Nobleobatrachia new taxon; (20) Australobatrachiais
a clade composed of Batrachophrynidae and its sister taxon, Myobatrachoidea (Myobatrach-
idae and Limnodynastidae), which formsthe sister taxon ofallother hyloids,excludingsoog-
lossids; (21) Nobleobatrachia new taxon, is dominated at its base by frogs of a treefrog
morphotype, several with intercalary phalangeal cartilages—Hemiphractus (Hemiphractidae)
forms the sister taxon of the remaining members of this group, here termed Meridianura new
taxon; (22) Meridianura comprises Brachycephalidae (former Eleutherodactylinae 1 Brachy-
cephalus)andCladophrynianewtaxon;(23)Cladophryniaiscomposedoftwogroups,Cryp-
tobatrachidae (composed of Cryptobatrachus and Stefania, previously a fragmentof thepoly-
phyletic Hemiphractinae) and Tinctanura new taxon; (24) Tinctanura is composed of Am-
phignathodontidae (Gastrotheca and Flectonotus, 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)LeptodactyliformesiscomposedofDiphyabatrachianewtaxon(composedofCentrolen-
idae [including Allophryne] and Leptodactylidae, sensu stricto, including Leptodactylus and
relatives) and Chthonobatrachia new taxon; (27) Chthonobatrachia is composed of a refor-
mulated Ceratophryidae (which excludes such genera as Odontophrynus and Proceratophrys
and includes other taxa, such as Telmatobius) and Hesticobatrachia new taxon; (28) Hesti-
cobatrachia iscomposed ofa reformulatedCycloramphidae(whichincludesRhinoderma)and
Agastorophrynia new taxon; (29) Agastorophrynia is composed of Bufonidae (which is par-
tiallyrevised)andDendrobatoidea(DendrobatidaeandThoropidae);(30)Ranoidesnewtaxon
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
(ranidsandallies);(31)AllodapanuraiscomposedofMicrohylidae(whichispartiallyrevised)
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,whichincludesLeptopelinaeandformerAstylosternidae);(33)Natatanuranew
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 ofranoids,
Ametrobatrachia newtaxon;(36)AmetrobatrachiaiscomposedofAfricanuranewtaxonand
Saukrobatrachia new taxon; (37) Africanura is composed of two taxa: Phrynobatrachidae
(Phrynobatrachus, including Dimorphognathus and Phrynodon as synonyms) and Pyxice-
phaloidea; (38) Pyxicephaloidea is composed of Petropedetidae (Conraua, Indirana, Arthro-
leptides, and Petropedetes), and Pyxicephalidae (including a number of African genera, e.g.
Amietia[includingAfrana],Arthroleptella,Pyxicephalus,Strongylopus,andTomopterna);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
taxonomythatilluminatestheproblemsinourunderstandingofphylogeny,sothatfuturework
willbemadeeasier.Theserevisionsare:(1)placementofIxalotritonandLineatriton(Caudata:
Plethodontidae: Bolitoglossinae) into the synonymy of Pseudoeurycea, to render a monophy-
letic Pseudoeurycea; (2) placement of Haideotriton (Caudata: Plethodontidae: Spelerpinae)
intothesynonymyofEurycea,torenderamonophyleticEurycea;(3)placementofNesomantis
(Anura:Sooglossidae)intothesynonymyofSooglossus,toassureamonophyleticSooglossus;
(4)placementofCycloranaandNyctimystes(Anura:Hylidae:Pelodryadinae)intoLitoria,but
retaining Cyclorana as a subgenus, to provide a monophyletic Litoria; (5) partition of ‘‘Lim-
nodynastes’’ (Anura: Limnodynastidae) into Limnodynastes and Opisthodon to render mono-
phyletic genera; (6) placement of Adenomera, Lithodytes, and Vanzolinius (Anura: Leptodac-
tylidae) into Leptodactylus, to render a monophyletic Leptodactylus; (7) partition of ‘‘Eleuth-
erodactylus’’ (Anura: Brachycephalidae) into Craugastor, ‘‘Eleutherodactylus’’, ‘‘Euhyas’’,
‘‘Pelorius’’, and Syrrhophus to outline the taxonomic issues relevant to the paraphyly 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 newgenus,Anaxyrus,Chaunus,Cran-
opsis, Duttaphrynus new genus, Epidalea, Ingerophrynus new genus, Nannophryne, Pelto-
phryne, Phrynoidis, Poyntonophrynus new genus; Pseudepidalea new genus, Rhaebo, Rhi-
nella,Vandijkophrynusnewgenus);(9)placementofthemonotypicSpinophrynoides(Anura:
Bufonidae) into the synonymy of (formerly monotypic) Altiphrynoides 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)intothesynonymyofXenorhinatorender
a monophyletic Xenorhina; (12) transfer of a number of species from Plethodontohyla to
Rhombophryne (Microhylidae: Cophylinae) to render a monophyletic Plethodontohyla; (13)
placement of Schoutedenella (Anura: Arthroleptidae) into the synonymy of Arthroleptis; (14)
transfer of Dimorphognathus and Phrynodon (Anura: Phrynobatrachidae) into the synonymy
ofPhrynobatrachustorenderamonophyleticPhrynobatrachus;(15)placementofAfranainto
thesynonymyofAmietia(Anura:Pyxicephalidae)torenderamonophyletictaxon;(16)place-
mentofChaparanaandPaaintothesynonymyofNanorana(Anura:Dicroglossidae)torender
a monophyletic genus; (17) recognition as genera of Ombrana and Annandia(Anura:Dicrog-
lossidae: Dicroglossinae) pending placement of them phylogenetically; (18) return of Phry-
noglossusintothesynonymyofOccidozygatoresolvetheparaphylyofPhrynoglossus(Anura:
Dicroglossidae: Occidozyginae); (19) recognition of Feihyla new genus for Philautus palpe-
bralis to resolve the polyphyly of ‘‘Chirixalus’’; (20) synonymy of ‘‘Chirixalus’’ with Chi-
romantis to resolve the paraphyly of ‘‘Chirixalus’’; (21) recognition of the genus Babina,
composed of the former subgenera of Rana, Babina and Nidirana (Anura: Ranidae); (22)
recognition of the genera Clinotarsus, Humerana, Nasirana, Pelophylax, Pterorana, Pul-
chrana, and Sanguirana, formerly considered subgenera of Rana (Anura: Ranidae), with no
specialrelationshiptoRana(sensustricto);(23)considerationofGlandirana(Anura:Ranidae),
formerly a subgenus of Rana, as a genus, with Rugosa as a synonym; (24) recognition of
Hydrophylax (Anura: Ranidae) as a genus, with Amnirana and most species of former Chal-
corana included in this taxon as synonyms; (25) recognition of Hylarana (Anura: Ranidae)
as a genus and its content redefined; (26) redelimitation of Huia to include as synonyms
Eburana and Odorrana (both formersubgenera of Rana);(27) recognitionofLithobates(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 formersub-
genera; (29) redelimitation of the genus Sylvirana (Anura: Ranidae), formerly a subgenus of
Rana, with Papurana and Tylerana included as synonyms.
Description:(Hylinae, Pelodryadinae, and Phyllomedusinae, and excluding former Hemiphractinae,
