Phylogeny and Evolution of Angiosperms

Soltis FM 5/13/05 12:01 PM Page iii

DOUGLAS E. SOLTIS University of Florida, Gainesville

PAMELA S. SOLTIS University of Florida, Gainesville

PETER K. ENDRESS University of Zurich

MARK W. CHASE Royal Botanic Gardens, Kew

SINAUER ASSOCIATES, INC. PUBLISHERS Sunderland, Massachusetts

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. Soltis FM 5/13/05 12:01 PM Page vi

Contents

Chapter 1 Molecular Analyses 45 Relationships of the Angiosperms Combined DNA and Morphology 47 Combined DNA Datasets 48 to Other Seed Plants 1 Amborellaceae, Nymphaeaceae, and Introduction 1 Austrobaileyales 50 Phylogenetic Studies: Extant Taxa 2 Other Basal Angiosperms: Monocots, Phylogenetic Studies: Integrating Fossils 10 Ceratophyllaceae, and Chloranthaceae 54 Future Studies 19 Other Basal Angiosperms: The Magnoliids 57 Chapter 2 Character Evolution in Basal Angiosperms 59 Phylogeny of Angiosperms: An Genetics of Floral Development: Overview 21 Implications for the Early Diversification Introduction 21 of the Flower 82 History of Efforts to Reconstruct The Appearance of Angiosperms in the Angiosperm Phylogeny 22 Fossil Record 85 Angiosperm Topology: Overview and Reconstructing the Earliest Angiosperms 86 Comparison with Modern Classifi- Future Research 89 cations 27 Sequence Variation: General Patterns and Rapid Radiations 33 Chapter 4 Timing of Diversification Events 35 Monocots 91 Future Studies 39 Introduction 91 Phylogenetic Analyses Based on Morphology 93 Chapter 3 Molecular Analyses of Monocot Basal Angiosperms 43 Relationships 94 Introduction 43 Current Understanding of Relationships Morphological Cladistic Analyses 44 Within Monocots 95

Fossil History of Monocots 116 Euasterids 213 Future Research 116 Euasterid I (Lamiids) 214 Euasterid II (Campanulids) 221 Chapter 5 Unplaced Euasterid II 228 Unplaced Asterids 229 Early-Diverging Eudicots 119 Patterns of Character Evolution in Introduction 119 Asterids 229 Ranunculales 124 Future Research 236 Proteales 125 Sabiaceae 125 Buxaceae 126 Chapter 10 Trochodendraceae 126 Angiosperm Classification 237 Character Evolution in Early- Introduction 237 Diverging Eudicots 126 The Angiosperm Phylogeny Group Floral Developmental Genetics 136 (APG) Classification 238 Future Research 137 Ranked versus Rank-Free Classification 239 Chapter 6 Classification of Angiosperms (Modified from APG II 2003) 243 Core Eudicots: Introduction and Smaller Lineages 139 Chapter 11 Introduction 139 Gunnerales 141 Parallel and Convergent “Berberidopsidales” 142 Evolution 249 Saxifragales 142 Introduction 249 Santalales 153 Parasitism 250 Future Research 156 Carnivory 256

C4 Photosynthesis 263 Chapter 7 Future Directions 266 Caryophyllales 157 Introduction 157 Chapter 12 Phylogeny of Caryophyllales 159 Floral Diversification 269 Character Evolution in Caryophyllales 163 Introduction 269 Future Research 167 Pathways of Evolutionary Innovations 270 Evolutionary Radiations 271 Chapter 8 Floral Phyllotaxis 272 Floral Merism 273 Rosids 169 Floral Symmetry 273 Introduction 169 Perianth 274 Vitaceae 171 Androecium 277 Eurosid I (Fabids) 171 Gynoecium 280 Eurosid II (Malvids) 189 Ovules 282 Other Rosids 200 Morphological Elaborations in Flowers Parasitism in the Rosids 204 Pollinated by Animals 283 Future Research 205 Morphological Reduction in Flowers that are not Animal-Pollinated 284 Chapter 9 Evolutionary Diversification in Flowers of the Major Angiosperm Groups 285 Asterids 207 Future Directions 286 Introduction 207

Chapter 13 Appendix 303 Evolution of Genome Size and Base Chromosome Number 287 References 311 Introduction 287 Genome Size 288 Taxonomic Index 353 Base Chromosome Number 298 Future Studies 302 Subject Index 365