1 An introduction to amphibian cytogenetics 1
1.1 Amphibian systematics 1
1.2 The chromosomalapproach 2

2 Chromosomes as evolutionary markers 3
2.1 Chromosomal comparisons of higher taxa and the problem of establishing
the ancestral karyotype 6
2.2 Group comparisons and phylogenetic inference 7

3 Chromosome morphology and evolutionary relationships in the order
Anura 9

3.1 Suborder Archaeobatrachia: the ancient frogs 9
3 .1.1 Superfamily Discoglossoidea 9
3 .1.1 .1 Family Discoglossidae 9
3 .1.1.2 Family Leiopelmatidae 12
3.1.2 Superfamily Pelobatoidea 12
3.1.2.1 Family Pelobatidae 13
3.1.2.1.1 Subfamily Megophryinae 13
3.1.2.1.2 Subfamily Pelobatinae 14
3.1.2.2 Family Pelodytidae 14
3.1.3 Superfamily Pipoidea 14
3 .1.3.1 Family Pipidae 14
3.1.3.1.1 Subfamily Pipinae 14
3.1.3.1.2 Subfamily Xenopodinae 15
3.1.3.2 Family Rhinophrynidae 16

3.2 Suborder Neobatrachia: the modem frogs 16
3.2.1 Superfamily Bufonoidea 16
3.2.1.1 Family Bufonidae 17
3.2.1.2 Family Brachycephalidae 19
3.2.1.3 Family Centrolenidae 19
3.2.1.4 Family Dendrobatidae 20
3.2.1.5 Family Heleophrynidae 20
3.2.1.6 Family Hylidae 20
3.2.1.6.1 Subfamily Hemiphractinae 20
3.2.1.6.2 Subfamily Hylinae 21
3.2.1.6.3 Subfamily Pelodryadinae 23
3.2.1.6.4 Subfamily Phyllomedusinae 26
3.2.1.7 Family Leptodactylidae . 26
3.2.1.7.1 Subfamily Ceratophryinae 26
3.2.1.7.2 Subfamily Hylodinae 27
3.2.1.7.3 Subfamily Leptodactylinae 27
3.2.1.7.4 Subfamily Telmatobiinae 28
3.2.1.8 Family Myobatrachidae 30
3.2.1.8.1 Subfamily Limnodynastinae 30
3.2.1.8.2 Subfamily Myobatrachinae 32
3.2.1.9 Family Pseudidae 34
3.2.1.10 Family Rhinodermatidae 34

3.2.2 Superfamily Microhyloidea 34
3.2.2.1 Family Microbylidae 34
3.2.2.1.1 Subfamilies Asterophryinae and Genyophryninae 35
3.2.2.1.2 Subfamilies Brevicipitinae, Melanobatrachinae
and Phrynomerinae 36
3.2.2.1.3 Subfamily Cophylinae 37
3.2.2.1.4 Subfamily Dyscophinae 37
3.2.2.1.5 Subfamily Microhylinae 38

3.2.3 Superfamily Ranoidea 38
3.2.3.1 Family Arthroleptidae 38
3.2.3.1.1 Subfamily Arthroleptinae 39
3.2.3.1.2 Subfamily Astylosterninae 40
3.2.3.2 Family Hemisidae 40
3.2.3.3 Family Hyperoliidae 41
3.2.3.3.1 Subfamily Hyperoliinae 41
3.2.3.3.2 Subfamily Kassininae 41
3.2.3.3.3 Subfamily Leptopelinae 43
3.2.3.4 Family Ranidae 43
3.2.3.4.1 Subfamily Mantellinae 44
3.2.3.4.2 Subfamily Petropedetinae 44
3.2.3.4.3 Subfamily Raninae 45
3.2.3.5 Family Rhacophoridae 47
3.2.3.5.1 Subfamily Rhacophorinae 47
3.2.3.6 Family Sooglossidae 47

4 Chromosome morphology and evolutionary relationships in the order
Apode 48
4.1 The chromosome reduction hypothesis 50

5 Chromosome morphology and evolutionary relationships in the order
Caudata 52

5.1 Suborder Ambystomatoidea 52
5.1.1 Family Ambystomatidae 53
5.1.2 Family Dicamptodontidae 54
5.1.3 Family Plethodontidae 55
5.1.3.1 Subfamily Desmognathinae 55
5.1.3.2 Subfamily Plethodontinae 55
5.1.3.2.1 Tribe Bolitoglossini 55
5.1.3.2.2 Tribe Hemidactylini 56
5.1.3.2.3 Tribe Plethodontini 56

5.2 Suborder Cryptobranchoidea 57
5.2.1 Family Cryptobranchidae 57
5.2.2 Family Hynobiidae 57

5.3 Suborder Salamandroidea 57
5.3.1 Family Amphiumidae 58
5.3.2 Family Proteidae 60
5.3.3 Family Salamandridae 60
5.3.3.1 Hybridization studies in Triturus 61

5.4 Suborder Sirenoidea 62
5.4.1 Family Sirenidae 62

6 Chromosome banding and substructure studies patterns of change 63

6.1 C-banding 63

6.2 Fluorescence banding 72

6.3 G- and it-banding 74

6.4 Lampbrush chromosomes 80

7 Molecular evolution of the amphibian genome 84

7.1 The Plethodontidae 86

7.2 The genus Triturus 88

7.3 The genus Rana 91

7.4 The genus Xenopus 92

8 Nucleolus organiser evolution in Amphibians 92

8.1 Structural aspects 92
8.1.1 In-situ hybridisation with 18S + 28S ribosomal probes 92
8.1.2 Silver staining analysis 96
8.1.3 C-banding analysis 98
8.1.4 Fluorescence analysis 100

8.2 Secondary constrictions as an evolutionary marker 101
8.2.1 The genus Litoria 104
8.2.2 Subfamily Myobatrachinae 106
8.2.3 Subfamily Limnodynastinae 106
8.2.4 The possible convergence of NOR form in Cyclorana 107

8.3 The chromosomal location of 5S RNA cistrons 109

9 Evolution of genome size in Amphibians 111
9.1 The nature of genome size variation 111
9.2 The relationship between changes in genome size and other parameters 116

10 Sex chromosome evolution in the Amphibia 117
10.1 Heteromorphic sex chromosomes 117
10.2 Sex reversal experimentation and the nature of sex determination in
Amphibians 127

11 Polyploidy as an evolutionary mechanism in the Amphibia 129
11.1 The genus Xenopus 132
11.2 The Hyla versicolor complex 135
11.3 The leptodactyloid polyploids 136
11.4 The genus Neobatrachus 138
11.5 The Rana esculenta complex 140
11.6 The genus Ambystoma 142

12 Chromosomal evolution in the Amphibia: an overview 144
12.1 Modes of chromosomal repatterning in Amphibians 145
12.2 Chromosomal change and speciation 147
12.3 A phylogenetic perspective on the Amphibians 150
Acknowledgements 157
References 158
Appendices 190
Species index 235