Acknowledgments x

1 Introduction 1

1.1 Changing paradigms in amphibian evolution 3

1.2 Paleobiology: data, methods, and time scales 5

1.3 Concepts and metaphors: how scientists “figure out” problems 7

1.4 Characters and phylogenies 8

1.5 What’s in a name? 8

References 11

2 The Amphibian World: Now and Then 13

2.1 Tetrapoda 14

2.1.1 The tetrapod skeleton 14

2.1.2 Tetrapod characters 23

2.1.3 Stem-tetrapods (Tetrapodomorpha) 25

2.1.4 Carboniferous tetrapods or tetrapodomorphs? 31

2.2 The amniote stem-group 32

2.2.1 Anthracosauria 33

2.2.2 Seymouriamorpha 37

2.2.3 Chroniosuchia 38

2.2.4 Lepospondyli 40

2.2.4.1 Lepospondyl characters 42

2.2.4.2 Microsauria 42

2.2.4.3 Lysorophia 44

2.2.4.4 Nectridea 44

2.2.4.5 Aïstopoda 45

2.2.4.6 Adelospondyli 46

2.2.4.7 Acherontiscidae 46

2.2.5 Gephyrostegida 46

2.2.6 Amniota 47

2.2.6.1 Stem-amniotes and early crown amniotes 48

2.3 The lissamphibian stem-group (Temnospondyli) 48

2.3.1 Edopoidea 51

2.3.2 Dendrerpeton and Balanerpeton 53

2.3.3 Dvinosauria 54

2.3.4 Dissorophoidea and Zatracheidae 54

2.3.5 Eryopoidea 56

2.3.6 Stereospondyli 57

2.4 Albanerpetontidae 58

2.5 Lissamphibia 59

2.5.1 Lissamphibian characters 61

2.5.2 Batrachia 62

2.5.2.1 Anura (frogs and toads) 62

2.5.2.2 Caudata (salamanders) 67

2.5.2.3 Gymnophiona (caecilians) 68

References 70

3 Amphibian Life Through Time 81

3.1 Aquatic predators prepare for land 83

3.2 Hot springs, scorpions, and little creepers 83

3.3 Life in the tropical coal forest 85

3.4 Neotenes explore unfavorable waters 89

3.5 Lowlands, uplands, and a cave 90

3.6 Hide and protect: extreme life in the hothouse 94

3.7 Predators in deltas, lakes, and brackish swamps 97

3.8 Stereospondyls in refugia, lissamphibians on the rise 97

3.9 Batrachians diversify, stereospondyls disappear 100

3.10 Lissamphibians expand into diverse habitats 101

References 102

4 The Amphibian Soft Body 106

4.1 How to infer soft tissues in extinct taxa 107

4.2 Fossil evidence: soft tissue preservation 109

4.3 Head and visceral skeleton 110

4.4 Respiratory organs 113

4.5 Lateral lines, electroreception, and ears 118

References 122

5 Evolution of Functional Systems 126

5.1 How paradigms and brackets give a functional scenario 127

5.2 Feeding and breathing under water 131

5.3 Decoupling breathing and feeding 134

5.4 Hearing: exapting the spiracle and hyomandibula 136

5.5 Respiration in early tetrapods 141

5.6 The evolution of terrestrial feeding 143

5.7 Transforming fins into limbs 144

5.8 Locomotion in paleozoic tetrapods 146

References 148

6 Development and Evolution 152

6.1 Ontogeny in modern amphibians 153

6.2 Fossil ontogenies 158

6.3 Ontogeny as a sequence: developmental trajectories 163

6.4 Histology: the skeleton as archive 167

6.5 Changing shape: allometry 171

6.6 Heterochrony: the evolution of development 174

6.7 Body plans: gene regulation and morphogenesis 179

References 184

7 Paleoecology 191

7.1 Lissamphibian ecology 192

7.2 Paleoecology: problems and perspectives 193

7.3 Paleozoic and Mesozoic amphibians 196

7.4 Amphibian evolution as a walk through trophic levels 203

References 205

8 Life History Evolution 208

8.1 Plasticity, reaction norm, and canalization 209

8.2 Reaction norms in extant amphibians 210

8.3 The biphasic life cycle in lissamphibians 211

8.4 Seymouriamorphs: biphasic life cycles without metamorphosis 213

8.5 Temnospondyls: flexible uni- and biphasic ontogenies 213

8.6 Lepospondyls: dwarfism and uniphasic life cycles 215

8.7 The evolution of metamorphosis 216

8.8 The evolution of neoteny 216

8.9 General features of life history evolution 217

References 219

9 Phylogeny 222

9.1 Phylogeny of amphibians 223

9.2 The big picture: tetrapod diversification 223

9.3 The origin of lissamphibians 224

References 231

10 Macroevolution 234

10.1 What is macroevolution? 235

10.2 Patterns of early tetrapod evolution 235

10.3 Major factors of amphibian evolution 240

10.4 Clades, space, and time 248

10.5 Diversity, disparity, and extinction 249

10.6 The evolution of terrestriality 252

References 254

Index 260

Plate section between pp. 124 and 125