Evolution in Action (eBook)

Preface Radiations, or Evolution in Action 6
Introduction to the Priority Programme ``Radiations: Origins of Biological Diversity´´ 10
Contents 14
Contributors 18
Part I Approaches in Botany 28
Rapid Radiations and Neoendemism in the Madagascan Biodiversity Hotspot 29
1 Introduction 30
2 Research Approach 32
2.1 Taxonomy 32
2.2 Phylogeny 32
2.3 Nuclear Markers 32
2.4 Establishing Biogeographic Hypothesis 33
2.5 Divergence Time Estimates 33
2.6 Niche Modelling 34
3 Results 34
3.1 Taxonomic Revision 34
3.2 Scaly Tree Fern Phylogeny 34
3.3 Radiations of Scaly Tree Ferns in Madagascar 34
3.4 Comparing Current Distribution and Fundamental Distribution 37
3.5 Scaly Tree Ferns on the Mascarene Islands 37
4 Perspectives 39
References 40
Rapid Radiation in the Barley Genus Hordeum (Poaceae) During the Pleistocene in the Americas 42
1 Introduction 42
1.1 Framework of the Studies, Questions to Answer 45
2 Materials and Employed Methods 46
3 Results and Discussion 49
3.1 Phylogeny of Hordeum 49
3.2 Biogeography 50
3.3 Speciation Rates 51
3.4 Chloroplast Genealogy of Hordeum 52
3.5 The Eurasian Hordeum marinum Species Complex 53
3.6 North American H. californicum Group 54
3.7 Southern Patagonian Species 54
4 Conclusions 55
References 56
Studying Adaptive Radiation at the Molecular Level: A Case Study in the Macaronesian Crassulaceae-Sempervivoideae 59
1 Introduction 60
1.1 Speciation and Adaptive Radiation 60
1.2 Island Radiations and Macaronesian Crassulaceae-Sempervivoideae 60
1.3 Candidate Genes 63
2 Material and Methods 63
2.1 Selection of the Study Group 64
2.1.1 nrITS 64
2.1.2 Candidate Genes 64
2.2 Selection of Candidate Genes 64
2.2.1 Regulatory Genes 64
2.2.2 Structural Gene 64
2.3 Laboratory Work 65
2.4 Data Analysis 65
2.4.1 Phylogenetic Analyses 65
Ka/Ks Values 65
2.4.2 Dating 66
Molecular Dating 66
3 Results and Discussion 66
3.1 Are the MCS the Result of an Adaptive Radiation? 66
3.2 Gene Evolution 69
3.2.1 Orthologous/Paralogous Copies 69
General Pattern 72
3.2.2 Phylogenetic Significance 75
3.2.3 Evolution Rates of Candidate Genes 76
3.3 Evolution of MCS 77
4 Summary 79
References 79
Key Innovations Versus Key Opportunities: Identifying Causes of Rapid Radiations in Derived Ferns 84
1 Introduction 84
2 Identifying Evidence for Biological Radiations 85
3 Determining the Role of Key Innovations and/ or Key Opportunities 87
4 Case Studies in the Epiphytic Polypodiaceae 88
5 Key Innovations: Ants and Ferns 90
6 Key Innovations and Key Opportunities 93
7 Perspectives 93
References 95
Evolution of the Mating System in the Genus Capsella (Brassicaceae) 99
1 Introduction 99
1.1 The Mating System in Brassicaceae Genera 100
1.2 The Genus Capsella Is a Wild Relative of the Model Plant Arabidopsis 101
1.3 The Dynamic of the Mating System Provides Adaptive Potential 102
2 Materials and Methods 104
2.1 Plant Material 104
2.2 Identification of S-alleles in C. grandiflora and C. rubella 104
2.2.1 DNA Extraction PCR and Cloning
2.2.2 PCR-RFLP 105
2.2.3 Sequencing and Sequence Analyzes 105
2.3 Screening of BAC-Clones: Identification of S-locus in C. rubella 106
2.3.1 Analysis of the BAC Insert: Primer Walking 106
2.4 Crossing Experiments 106
2.4.1 Crossing Experiment I: Intraspecific Crossing 106
2.4.2 Crossing Experiment II: Interspecific Crossing 107
3 Results 107
3.1 Identification of S-Alleles 107
3.1.1 S-Allele Specific PCR-RFLP 109
3.2 Screening of BAC-Clones: Identification of S-Locus Genes in C. rubella 109
3.2.1 Identification of C. rubella 5-kb BAC fragment through comparison with A. thaliana 109
3.3 Frequency and Dominance Relationships of S-Allele 111
3.3.1 Crossing Experiment I: Intraspecific Crossing 111
3.3.2 Crossing Experiment II: Interspecific Crossing 112
4 Discussion 114
4.1 The Chromosomal Location of the Capsella S-Locus is Similar to Arabidopsis 116
4.2 Natural Populations of Capsella Exhibit Flexible Mating Strategies 116
4.3 Evolutionary Consequences of Flexible Mating Strategies 118
5 Summary 118
References 119
Pollinator-Driven Speciation in Sexually Deceptive Orchids of the Genus Ophrys 123
1 Introduction 123
1.1 Sexual Deception 124
1.2 Specialised Pollination and Floral Isolation 125
1.3 Speciation 125
2 Case Study 1: Hybrid Speciation and Distinction of Species on Sardinia 128
3 Case Study 2: Pollinator-Driven Selection and Speciation in Ophrys lupercalis, Ophrys bilunulata and Ophrys fabrella on Majorca 130
4 Case Study 3: Speciation and Evolutionary Origin of Two Sympatrically Occurring Endemic Species, O. chestermanii and O. normanii, on Sardinia 133
5 Conclusions 136
References 137
Population Genetics of Speciation and Demographic Inference Under Population Subdivision: Insights from Studies on Wild Tomatoes (Solanum sect. Lycopersicon) 141
1 Introduction 142
2 Materials and Methods 143
2.1 Population Sampling and Sequenced Loci 143
2.2 The Isolation Speciation Model 146
2.3 Sampling Schemes, Population Subdivision, and Range Expansions 147
3 Results and Discussion 148
3.1 WH Model Parameter Estimation and the Impact of Population Subdivision 148
3.2 Assessing Postdivergence Gene Flow by Linkage Disequilibrium (LD) 151
3.3 Site Frequency Spectra in Samples from Nonequilibrium, Subdivided Populations 152
4 Summary 154
References 155
Genetic Diversity, Evolution and Domestication of Wheat and Barley in the Fertile Crescent 158
1 Introduction 159
2 Origins of Cultivated Plants and Agriculture: A Brief Historical Overview 159
3 Evolution and Domestication of Triticeae 161
3.1 Wheat Evolution and Domestication 165
3.1.1 Diploid Wheats 167
3.1.2 Tetraploid Wheats 173
3.1.3 Hexaploid Wheats-Bread Wheat 175
3.2 Barley Evolution and Domestication 176
4 Conclusions and Final Considerations 178
List of our publications resulting from the SPP 1127 179
Invited Lectures and Data Presented from the SPP 1127 180
Conferences Attended and Data Presented from the SPP 1127 180
Collaborations Resulted from the SPP 1127 181
References 181
Part II Host-Plant Interaction 188
Mechanisms of Speciation in Southeast Asian Ant-Plants of the Genus Macaranga (Euphorbiaceae) 189
1 Introduction 190
1.1 Co-Evolution in Mutualistic Associations: A Challenge for Evolutionary Biology 190
1.2 Radiation on Both Sides of a Myrmecophytic Interaction: The Macaranga-Crematogaster System 190
1.3 Aims of Our Study 193
2 What Does Phylogeny Tell Us About the Macaranga-Crematogaster Co-Evolution? 194
2.1 Monophyly of Macaranga Sections and Subsectional Groups with Myrmecophytes 194
2.2 Co-Adaptation and Host Shift are Major Determinants of the Macaranga-Crematogaster Co-Evolution 196
3 Which Roles Did Hybridization and Reticulate Evolution Play During the Evolution of Macaranga? 197
3.1 First Indications for Interspecific Gene Flow in Macaranga 197
3.2 Development of Markers to Analyze Species Boundaries and Gene Flow among Macaranga Species 197
3.3 Evidence for Introgression and Incomplete Lineage Sorting Among Macaranga Ant-Plants 198
3.4 Reticulate Evolution in Macaranga: Frequent Hybridization and Introgression but Little Evidence for Stable Hybrids 200
4 How Could the Ants Have Influenced Speciation Processes in Their Plant Partners? 202
4.1 The ``Allopatric Speciation Hypothesis´´: Limited Effective Seed Dispersal Could Enhance Genetic Differentiation in Myrmec. 202
4.2 Population Structure of Myrmecophytic Versus Non-Myrmecophytic Macaranga Species: No Support for the “Allopatric Speciation Hypothesis” 203
4.3 Putative Role of Vicariance 206
4.4 The Relative Role of Seed and Pollen Dispersal 207
5 Conclusions 207
References 208
Speciation in Obligately Plant-Associated Crematogaster Ants: Host Distribution Rather than Adaption Towards Specific Hosts Drives the Process 212
1 Introduction 213
2 Species Delimitation of Crematogaster (Decacrema) Ants Associated with Macaranga 216
3 Co-evolution Between Crematogaster (Decacrema) and Macaranga Hosts: Adaptations of the Ants Towards Their Hosts 221
3.1 Choosing and Finding a Host 221
3.2 Wax Running 222
3.3 Entering the Host: Queen Size Matters 222
3.4 Available Food and Nesting Space 223
3.5 Are All Ant Partners Equal? - The Plant Perspective 223
4 Radiation in Crematogaster (Decacrema) Ants: Is It Driven By Adaptation Towards Different Host Species? 224
4.1 Population Genetic Studies on A Local Scale 224
4.2 Vicariant Evolution - The Broader Geographic Scale 227
5 Conclusion and Outlook 228
References 230
Radiation, Biological Diversity and Host-Parasite Interactions in Wild Roses, Rust Fungi and Insects 233
1 Introduction: Radiation, Biodiversity and Host-Parasite Interaction in the Rosa-System 234
2 Dog Roses are Allopolylpoids: Genetic Constitution of Section Caninae 236
3 Character Inheritance in the Heterogamous System of Dog Roses 237
4 Glandular Trichomes Matter: Rust Fungi on Rosa 239
5 Evolution and Diversity of Plant-Pathogen-Insect Foodwebs on Dog Roses 239
5.1 Are Invertebrate Communities Affected by Leaf Trichome Traits of Hosts? 241
5.2 Do Rhagoletis alternata and Diplolepis rosae Differ in Density Between the Three Rose Species? 243
5.3 Does Rhagoletis alternata Form Host Races on the Three Dog Rose Species? 248
6 How are the Differences Between the Three Closely Related Dog Rose Species Translated Into Higher Trophic Levels? 249
7 Conclusion 251
References 252
Speciation via Differential Host-Plant Use in the Tephritid Fly Tephritis conura 257
1 Introduction 257
2 Natural History of the T. conura - Cirsium System 259
2.1 T. conura 259
2.2 The Host Plants 261
3 Geographic Speciation 262
4 Gene Flow and Signs of Selection 265
5 Host-Race Adaptations 267
5.1 Plant Recognition and Willingness to Mate 268
5.2 Survival on C. oleraceum 269
5.3 Adaptation of Ovipositor Length 270
6 Does Plant History Matter in Host-Race Evolution? 271
7 Conclusions 273
References 274
Part III Approaches in Zoology 279
Solar Powered Seaslugs (Opisthobranchia, Gastropoda, Mollusca): Incorporation of Photosynthetic Units: A Key Character Enhancing Radiation? 280
1 Introduction 281
2 Materials and Methods 285
3 Results 286
3.1 Phyllodesmium 286
4 Sacoglossa 289
5 Discussion 292
6 Can We Consider the Incorporation of Photosynthetic Units (PUs) an Adaptive Radiation? 295
7 Conclusion 296
8 Summary 296
References 297
Are Cuticular Hydrocarbons Involved in Speciation of Fungus-Growing Termites (Isoptera: Macrotermitinae)? 300
1 Introduction 300
2 Cuticular Hydrocarbons and Nestmate Recognition 303
3 Variation of Cuticular Hydrocarbons Within and Between Termite Species 306
4 Heritability of Cuticular Hydrocarbons and Environmental Variation 309
5 Cuticular Hydrocarbons and Agonistic Behavior 312
6 Cuticular Hydrocarbons and Genetic Differentiation Between Phenotypes 315
7 Conclusions 316
References 318
Electric Organ Discharge Divergence Promotes Ecological Speciation in Sympatrically Occurring African Weakly Electric Fish (Campylomormyrus) 324
1 Introduction 325
2 Taxonomy of Campylomormyrus 326
3 Indication for Ecological Speciation 330
4 Electric Organ Discharge and Mate Choice 333
5 Electric Organ Discharge: A Magic Trait for Speciation? 334
References 336
Ongoing Phenotypic and Genotypic Diversification in Adaptively Radiated Freshwater Crabs from Jamaica 339
1 Introduction 340
2 Phenotypic Modification of Body Form in Response to Cave-Living 342
2.1 Materials and Methods 343
2.1.1 Morphometric Data 343
2.1.2 Genetic Data 344
2.2 Results 345
2.2.1 Morphometrics 345
2.2.2 Genetics 346
3 Genotypic Diversification Within Three Species of Freshwater Crabs 348
3.1 Material and Methods 350
3.2 Results 352
3.2.1 Sesarma dolphinum 352
3.2.2 Sesarma windsor and Sesarma meridies 355
3.3 Discussion 357
4 Additional Findings and Outlook 358
References 362
The Herring Gull Complex (Larus argentatus - fuscus - cachinnans) as a Model Group for Recent Holarctic Vertebrate Radiations 366
1 Introduction 366
2 State of the Art 369
3 Recent Progress 374
3.1 Phylogenetic Framework for the Herring Gull Complex 374
3.2 Population History of ``polyphyletic´´ Taxa in the Mitochondrial Gene Tree 375
3.2.1 Evolution of Herring Gulls in Europe 379
3.2.2 Phylogeographic History of Circumpolar Breeding Glaucous Gulls 379
3.2.3 Colonization Pattern of Greater Black-Backed Gulls 381
3.2.4 Summary 381
4 Future Perspectives 382
References 383
Genetic Divergence and Evolution of Reproductive Isolation in Eastern Mediterranean Water Frogs 387
1 Introduction 389
2 Materials and Methods 390
2.1 Taxon Sampling 390
2.2 Estimation of Genetic Divergence 390
2.3 Estimation of Rates and Times of Divergence Using a Non-constant Molecular Clock 393
2.4 Crossing Experiments 393
2.5 Bioacoustic Investigations 394
2.6 Female Choice Experiments 394
3 Results and Discussion 394
3.1 Genetic Diversity of Eastern Mediterranean Water Frogs 394
3.2 Genetic Divergence Between Population Pairs 397
3.3 Estimation of Confidence Limits for Divergence Times of Basic Lineages Using a Nonconstant Molecular Clock 401
3.4 Evolution of Anatolian Water Frog Populations 402
3.5 Genetic Incompatibilities 403
3.6 Antihybridization Mechanisms 406
3.7 Implications for Water Frog Systematics 409
3.8 Conclusions and Prospects 411
References 412
Inferring Multiple Corsican Limax (Pulmonata: Limacidae) Radiations: A Combined Approach Using Morphology and Molecules 418
1 Introduction 419
2 Materials and Methods 420
2.1 Collection and Treatment of Specimens 420
2.2 DNA Sequence Analysis 421
3 Results 422
3.1 Morphological and Copulation Studies 422
3.2 Sequence Analysis 426
3.3 Distribution 431
4 Discussion 431
4.1 Biogeographical Scenarios 431
4.2 Species Boundaries 437
4.3 Evolutionary Considerations 437
5 Conclusions 438
References 438
Supplement: List of Material 441
Appendix: Two New Species and One New Name of Peri-Tyrrhenian Limax 445
Limax giustii n. sp. 445
Limax ilvensis n. sp. 446
Limax vizzavonensis n. nom. 447
Additional References 447
Palaeogeography or Sexual Selection: Which Factors Promoted Cretan Land Snail Radiations? 449
1 Introduction 449
2 Systematics of the Xerocrassa Radiation on Crete 450
3 Ecological Differentiation of the Cretan Xerocrassa Species 453
4 Geographic Mode of Speciation 454
5 Evolution of Genitalia by Genetic Drift Versus Selection 456
6 Lock-and-Key Hypothesis Versus Sexual Selection 456
7 Influence of Evolution of Genitalia on Speciation in Xerocrassa 457
8 Sexual Selection and Non-adaptive Radiation 458
References 459
Non-Ecological Radiations in Acoustically Communicating Grasshoppers? 463
1 Introduction 464
2 Acoustic Communication and Speciation 465
2.1 Species Diversity and Acoustic Communication 465
2.2 Molecular Phylogeny 467
2.3 Evolution of Leg Movement Patterns 469
2.4 Song Evolution in Closely Related Species 471
3 Mechanisms of Speciation 472
3.1 Divergence in Allopatry 472
3.2 Evolution of New Songs by Hybridization 473
4 Conclusions 474
References 474
Beyond Sympatric Speciation: Radiation of Sailfin Silverside Fishes in the Malili Lakes (Sulawesi) 477
1 Introduction 478
2 Speciation Research in Adaptive Radiations 479
3 Sailfin Silversides in the Malili Lakes 479
4 Patterns of Hybridization 481
5 A Key Role of River Petea? 483
6 Sympatric Speciation in Lake Matano 484
7 On the Mechanisms Driving Speciation Processes 487
8 Sexual Selection and the Evolution of Colour Polymorphisms 488
9 Perspectives 489
References 491
The Species Flocks of the Viviparous Freshwater Gastropod Tylomelania (Mollusca: Cerithioidea: Pachychilidae) in the Ancient Lakes of Sulawesi, Indonesia: The Role of Geography, Trophic Morphology and Color as Driving Forces in Adaptive Radiation 496
1 Introduction 497
1.1 Ancient lakes as ``Natural Laboratories´´ 497
1.2 The Gastropods in the ``Ancient Lakes´´ on Sulawesi 497
1.3 Scope and Aims 499
2 The Species Flocks of Tylomelania in Lakes on Sulawesi: Species Diversity and Endemism 500
3 A Molecular Phylogeny of Tylomelania: Lake Colonization, Species, and Introgression 505
4 Coevolution with Crabs 508
5 Adaptive Radiation Through Trophic Specialization 509
6 Speciation Patterns 512
7 The Distribution and Role of Body Coloration 513
8 Patterns of Radiation: Looking Beyond Tylomelania and the Sulawesi Lakes 515
9 Conservation 515
10 Conclusions and Outlook 516
Appendix1. 519
References 520
Speciation and Radiation in a River: Assessing the Morphological and Genetic Differentiation in a Species Flock of Viviparous Gastropods (Cerithioidea: Pachychilidae) 524
1 Introduction 525
2 The Systematic Framework: Phylogeny of the SE Asian Pachychilidae 526
3 The Kaek River: Geographical and Environmental Settings 530
4 River Capture: Paleogeography and Palaeohydrology 532
5 Sampling Design and Collection Sites 533
6 Patterns of Shell Variation Among and Within the Kaek River Species 536
7 Radular Morphology and Substrate Usage 542
8 Phylogenetic Relationships Inferred by Analyses of Mitochondrial Genes 546
9 Towards an Evolutionary Explanation: Conclusions from Incongruence 550
10 Dispersal or Vicariance: Genetic Exchange Between River Faunas and the Relevance of River Captures Within the Mekong Drainage System 554
11 Speciation and Radiation of Brotia in the Kaek River 555
References 557
The Neglected Side of the Coin: Non-adaptive Radiations in Spring Snails (Bythinella spp.) 562
1 Introduction 563
2 The Spring Snail Genus Bythinella 566
3 Identifying Radiations in the Genus Bythinella 569
4 Bythinella spp. and the Criteria for Non-adaptive Radiations 573
4.1 Ecological (Niche) Variation within Bythinella Radiations 573
4.2 Phenotypical Variation among and within Bythinella Radiations 574
4.3 Sympatry versus Allopatry of Bythinella spp. 577
5 Non-adaptive Radiations in Bythinella spp. 578
5.1 What Drives Non-adaptive Radiations? 578
5.2 Potential Mechanisms of Non-adaptive Radiations in Bythinella spp. 580
5.3 Adaptive and Non-adaptive Radiations: Discrete Processes or Not? 581
5.4 Perspectives in Studying Non-adaptive Radiations 583
6 Take Home Message 583
References 585
Index 590