Early Flowers and Angiosperm Evolution

"The recent discovery of diverse fossil flowers and floral organs in Cretaceous strata has revealed astonishing details about the structural and systematic diversity of early angiosperms. Exploring the rich fossil record that has accumulated over the last three decades, this is a unique study of the evolutionary history of flowering plants from their earliest phases in obscurity to their dominance in modern vegetation. The discussion provides comprehensive biological and geological background information, before moving on to summarise the fossil record in detail. Including previously unpublished results based on research into Early and Late Cretaceous fossil floras from Europe and North America, the authors draw on direct palaeontological evidence of the pattern of angiosperm evolution through time. Synthesising palaeobotanical data with information from living plants, this unique book explores the latest research in the field, highlighting connections with phylogenetic systematics, structure and the biology of extant angiosperms"--Résumé de l'éditeur Title......Page 4 Copyright......Page 5 Contents......Page 6 Preface......Page 10 1.1.1 Anthophytes and alternative patterns......Page 12 1.1.2 Relationships among living angiosperms......Page 14 1.1.3 Origin of the angiosperm flower......Page 16 1.2.1 Vegetative features......Page 17 1.2.2 Reproductive features......Page 19 1.3.1 Pre-Cretaceous angiosperms?......Page 27 1.3.2 The fossil record......Page 28 1.4.1 Increasing angiosperm diversity and abundance......Page 30 1.4.2 Co-evolution with animals: herbivory, dispersal and pollination......Page 31 2.1 Understanding the plant fossil record......Page 34 2.1.1 Fossil wood......Page 37 2.1.2 Fossil leaves......Page 38 2.1.3 Fossil reproductive structures......Page 40 2.1.4 Fossil pollen......Page 46 2.2 The adequacy of the angiosperm fossil record......Page 49 3.1.1 Geologic and geographic changes during the Cretaceous......Page 50 3.1.2 Early and mid-Cretaceous......Page 51 3.1.3 Late Cretaceous......Page 52 3.2.1 Palaeontological indicators of climate......Page 56 3.2.2 Geological indicators of climate......Page 58 3.3 Climate change during the Cretaceous......Page 61 3.3.1 Early and mid-Cretaceous......Page 62 3.3.2 Late Cretaceous......Page 63 3.4 Implications for angiosperm diversification......Page 64 4.1 The stratigraphic framework......Page 66 4.2 Key areas for Cretaceous angiosperms......Page 67 4.3.1 Portugal: Early Cretaceous localities......Page 68 4.3.2 Portugal, Late Cretaceous localities......Page 74 4.3.3 Spain......Page 75 4.3.4 The Czech Republic......Page 76 4.3.5 Austria......Page 77 4.3.6 Germany......Page 78 4.3.7 Great Britain......Page 79 4.3.8 Sweden......Page 81 4.4 Eastern North America......Page 83 4.4.1 Virginia......Page 85 4.4.2 Maryland and Washington, D.C.......Page 87 4.4.5 North Carolina......Page 88 4.4.7. Alabama......Page 89 4.5.1 Kansas and Nebraska......Page 90 4.5.3 Colorado, Montana, North and South Dakota and New Mexico......Page 91 4.6 Alaska......Page 92 4.8 Israel, Jordan and Lebanon......Page 93 4.9.1 Egypt and Sudan......Page 95 4.10 West Africa and Brazil......Page 96 4.10.1 West Africa......Page 97 4.10.2 Brazil......Page 98 4.11 Asia......Page 99 4.11.1 Kazakhstan......Page 100 4.11.3 Siberia, Northeastern Russia and the Far East of Russia......Page 102 4.11.4 Mongolia......Page 103 4.11.5 China......Page 104 4.11.6 Japan......Page 106 4.12.3 India......Page 107 4.12.4 Australia......Page 108 4.12.5 Antarctica......Page 109 4.12.7 Southern South America......Page 110 5.1 Angiosperms among extant and fossil seed plants......Page 112 5.2 Bennettitales-Erdtmanithecales-Gnetales (BEG) group......Page 115 5.3 Gnetales......Page 116 5.3.1 Temporal and spatial patterns of gnetalean radiation......Page 117 5.3.2 Elaterates......Page 119 5.3.3 Gnetalean mesofossils and macrofossils......Page 120 5.4 Erdtmanithecales......Page 125 5.5 Unassigned dispersed seeds of the BEG group......Page 130 5.6 Bennettitales (Cycadeoidales)......Page 135 5.6.1 Permineralised material......Page 139 5.6.2 Compression material......Page 140 5.7 Pentoxylales......Page 141 5.8.1 Corystospermales and potentially related plants......Page 142 5.8.2 Caytoniales......Page 145 5.8.3 Peltaspermales......Page 148 5.8.4 Glossopteridales......Page 149 5.8.5 Czekanowskiales......Page 150 6.1 Hypotheses of seed plant relationships......Page 152 6.1.1 Cladistic hypotheses based on morphology......Page 155 6.1.2 Cladistic hypotheses based on molecular data......Page 159 6.1.3 Current status of phylogenetic studies......Page 160 6.2.1 The angiosperm flower......Page 161 6.2.2 The angiosperm stamen......Page 162 6.2.3 The angiosperm ovule......Page 163 6.2.5 Vegetative structure of angiosperms......Page 165 6.3.1 Implications of hypotheses of relationships......Page 166 6.3.3 Angiosperm age based on palaeobotanical data......Page 167 6.4 Pre-Cretaceous angiosperm-like fossils......Page 169 7.2 Phylogenetic studies of angiosperms based on molecular data......Page 174 7.3.2 Eumagnoliids......Page 175 7.3.4 Eudicots......Page 177 7.4 Angiosperm phylogeny: future directions......Page 178 7.5 Assignment of fossils to extant groups......Page 179 8.1.1 Fossils of uncertain relationships at the ANITA grade......Page 180 8.3.1 Hydatellaceae......Page 182 8.3.2 Cabombaceae and Nymphaeaceae......Page 183 8.4 Austrobaileyales......Page 187 8.4.2 Schisandraceae (including Illiciaceae)......Page 188 8.4.3 Trimeniaceae......Page 190 8.5 Chloranthaceae......Page 191 8.5.1 Asteropollis......Page 192 8.5.2 Hedyosmum-like flowers......Page 193 8.5.3 Chloranthistemon......Page 194 8.6 Ceratophyllaceae......Page 196 9.1.1 Bevhalstia......Page 200 9.1.2 Lappacarpus, Ievlevia, Donlesia and Beipiaoa......Page 201 9.2 Fossil flowers attached to inflorescences and stems......Page 203 9.2.1 Archaefructus......Page 204 9.2.2 Xingxueina and Caspiocarpus......Page 206 9.2.3 The Koonwarra fossil......Page 207 9.2.4 Myricanthium, Zlatkocarpus and other inflorescences from the Peruc flora......Page 208 9.2.5 Inflorescences from the Dakota Formation......Page 209 9.2.6 Other inflorescences......Page 210 9.3.1 Lesqueria elocata and other isolated reproductive structures......Page 211 9.4 Permineralised flowers......Page 212 9.5 Isolated angiosperm mesofossils......Page 213 9.5.1 Isolated flowers......Page 214 9.5.2 Isolated fruits and seeds......Page 216 9.6.1 Afropollis and Schrankipollis......Page 219 9.6.2 Asteropollis and Clavatipollenites......Page 221 9.6.3 Retimonocolpites, Brenneripollis and Pennipollis......Page 222 9.6.5 Stellatopollis......Page 223 9.6.6 Transitoripollis and Tucanopollis......Page 225 9.7 Fossil leaves of uncertain relationships......Page 226 10.1 Classification of eumagnoliids......Page 230 10.1.1 Fossil eumagnoliids of uncertain relationships......Page 231 10.2.1 Fossil Magnoliales of uncertain relationships......Page 234 10.2.3 Annonaceae......Page 235 10.2.5 Magnoliaceae......Page 237 10.3 Laurales......Page 242 10.3.1 Fossil Laurales of uncertain relationships......Page 243 10.3.3 Calycanthaceae......Page 244 10.3.5 Hernandiaceae......Page 246 10.3.6 Lauraceae......Page 247 10.3.7 Monimiaceae......Page 254 10.4.1 Canellaceae......Page 255 10.4.2 Winteraceae......Page 256 10.5.2 Aristolochiaceae......Page 257 10.5.3 Lactoridaceae......Page 258 10.5.6 Hydnoraceae......Page 259 11.1 Classification of monocots......Page 260 11.3.1 Dispersed leaves......Page 261 11.3.3 Reproductive structures......Page 263 11.4 Acorales......Page 266 11.5.1 Putative fossil Alismatales......Page 267 11.5.2 Araceae......Page 268 11.5.3 Alismataceae......Page 273 11.5.4 Butomaceae and Hydrocharitaceae......Page 275 11.5.5 Potamogetonoids......Page 276 11.7.2 Cyclanthaceae......Page 277 11.9 Asparagales......Page 278 11.10.1 Arecales......Page 279 11.10.2 Poales......Page 281 11.10.4 Zingiberales......Page 283 12.1 Classification of eudicots......Page 286 12.2 Early-diverging eudicots......Page 287 12.4.1 Early tricolpate pollen in dispersed palynofloras......Page 288 12.4.2 Early tricolpate pollen in situ in reproductive structures......Page 291 12.4.3 Fossil reproductive structures of probable eudicot relationships......Page 294 12.5.1 Berberidaceae......Page 300 12.5.4 Lardizabalaceae......Page 301 12.5.6 Papaveraceae......Page 302 12.6.1 Nelumbonaceae......Page 303 12.6.2 Platanaceae......Page 304 12.7 Sabiaceae......Page 312 12.8 Buxales......Page 314 12.8.1 Buxaceae......Page 318 12.9.1 Trochodendraceae......Page 319 13.1 Classification of core eudicots......Page 322 13.3 Gunnerales......Page 323 13.5 Berberidopsidales......Page 324 13.6 Santalales......Page 325 13.7 Caryophyllales......Page 326 13.8 Saxifragales......Page 327 13.8.1 Unassigned Saxifragales......Page 328 13.8.2 Altingiaceae......Page 331 13.8.4 Hamamelidaceae......Page 332 13.8.5 Haloragaceae and Crassulaceae......Page 334 13.8.7 Grossulariaceae......Page 335 13.8.9 Paeoniaceae......Page 336 14.2 Fossil evidence of rosids......Page 338 14.5.1 Celastrales......Page 340 14.5.3 Malpighiales......Page 341 14.6 The nitrogen-fixing clade......Page 343 14.6.1 Fabales......Page 344 14.6.2 Rosales......Page 345 14.6.4 Fagales......Page 347 14.6.5 Fossil Fagales of uncertain relationships......Page 348 14.6.6 Normapolles complex......Page 349 14.6.7 Nothofagaceae and Fagaceae......Page 360 14.6.8 Ticodendraceae, Betulaceae and Casuarinaceae......Page 361 14.6.9 Myricaceae, Rhoipteleaceae and Juglandaceae......Page 363 14.7.2 Myrtales......Page 365 14.7.4 Sapindales......Page 368 14.7.6 Brassicales......Page 369 14.7.7 Malvales......Page 370 15.1.1 Asterid fossils of uncertain relationships......Page 372 15.2.1 Fossil Cornales of uncertain relationships......Page 374 15.2.2 Cornaceae......Page 375 15.3 Ericales......Page 376 15.3.1 Ericalean fossils of uncertain relationships......Page 377 15.3.2 Pentaphylacaceae and Theaceae......Page 383 15.3.3 Core Ericales: Clethraceae, Cyrillaceae and Ericaceae......Page 384 15.3.4 Core Ericales: Sarraceniaceae, Actinidiaceae and Roridulaceae......Page 385 15.3.5 Ebenaceae and primuloids......Page 386 15.3.6 Symplocaceae, Diapensiaceae and Styracaceae......Page 388 15.4 Lamiids (Euasterids I)......Page 389 15.5 Boraginaceae, Icacinaceae and Vahliaceae......Page 390 15.7 Gentianales......Page 392 15.9 Campanulids (Euasterids II)......Page 393 15.10 Aquifoliales, Escalloniales and Asterales......Page 394 15.11 Bruniales, Apiales, Paracryphiales and Dipsacales......Page 396 16 Patterns of structural diversification in angiosperm reproductive organs......Page 398 16.1 Inflorescence structure......Page 399 16.2.1 Sex of flowers......Page 402 16.2.2 Receptacle, floral phyllotaxis and merism......Page 403 16.2.3 Position of floral organs......Page 405 16.2.4 Perianth......Page 406 16.2.5 Androecium......Page 411 16.2.6 Pollen......Page 415 16.2.7 Gynoecium......Page 419 16.2.8 Ovules and seeds......Page 421 16.3.1 Flower size......Page 423 16.3.2 Protection in floral bud......Page 424 16.3.3 Synorganisation of floral parts......Page 425 17.1.1 Pollination in Ginkgo and cycads......Page 426 17.1.3 Pollination in Gnetales......Page 427 17.2.2 Abiotic pollination......Page 428 17.2.3 Biotic pollination......Page 429 17.3.1 Thrip pollination and the fossil history of Thysanoptera......Page 430 17.3.3 Bee/wasp pollination and the fossil history of Hymenoptera......Page 432 17.3.4 Fly pollination and the fossil history of Diptera......Page 434 17.3.5 Butterfly/moth pollination and the fossil history of Lepidoptera......Page 436 17.4.1 Bird pollination and the fossil history of birds......Page 437 17.4.2 Mammal pollination and the fossil history of mammals......Page 438 17.5.1 Pollination in pre-angiosperm vegetation......Page 439 17.5.2 Pollination in Early Cretaceous angiosperms......Page 443 17.5.3 Pollination in mid-Cretaceous angiosperms......Page 448 17.5.4 Pollination in Late Cretaceous angiosperms......Page 450 17.5.5 Pollination in Cenozoic angiosperms......Page 451 17.6 Large-scale trends in the history of angiosperm pollination......Page 452 18.1.1 Dispersal in cycads......Page 456 18.1.3 Dispersal in conifers......Page 457 18.2 Dispersal in extant angiosperms......Page 458 18.3 Animal dispersers......Page 459 18.3.2 Vertebrates......Page 460 18.4.1 Dispersal in pre-angiosperm vegetation......Page 461 18.4.2 Dispersal in Early Cretaceous angiosperms......Page 462 18.4.3 Dispersal in mid-Cretaceous angiosperms......Page 464 18.4.4 Dispersal in Late Cretaceous angiosperms......Page 466 18.5 Large-scale trends in the history of angiosperm dispersal......Page 467 19.1 Transition to angiosperm-dominated vegetation......Page 472 19.2.1 Free-sporing plants......Page 473 19.2.2 Bennettitales and Cycadales......Page 474 19.2.4 Cheirolepidiaceae......Page 476 19.3 Vegetation during the early diversification of angiosperms......Page 478 19.4 Early angiosperms: diversity in obscurity......Page 480 19.5 Mid-Cretaceous vegetation......Page 482 19.6 Late Cretaceous vegetation and floristic provinces......Page 483 20.1 Large-scale patterns in angiosperm diversification......Page 486 20.2 Patterns of angiosperm diversification: early lineages......Page 488 20.2.1 Nymphaeales......Page 489 20.2.2 Chloranthaceae......Page 491 20.3 Patterns of angiosperm diversification: eumagnoliids......Page 494 20.3.2 Laurales......Page 495 20.3.3 Magnoliales......Page 496 20.4 Patterns of angiosperm diversification: monocots......Page 497 20.5 Patterns of angiosperm diversification: eudicots......Page 499 20.5.1 Proteales......Page 500 20.5.2 Fagales......Page 502 20.6 Angiosperm evolution and global change through the Cenozoic......Page 506 20.6.2 Palaeogene vegetation......Page 507 20.6.3 Neogene vegetation and the origin of modern biomes......Page 508 20.7 Prospects......Page 509 References......Page 512 Index......Page 584 "The recent discovery of diverse fossil flowers and floral organs in Cretaceous strata has revealed astonishing details about the structural and systematic diversity of early angiosperms. Exploring the rich fossil record that has accumulated over the last three decades, this unique study follows the evolutionary history of flowering plants from their earliest phases in obscurity to their dominance in modern vegetation. The book provides comprehensive biological and geological background information, before moving on to summarise the fossil record in detail. Including previously unpublished results based on research into Early and Late Cretaceous fossil floras from Europe and North America, the authors draw together direct palaeontological evidence of the pattern of angiosperm evolution through time. Synthesising palaeobotanical data with information from living plants, this book explores the latest research in the field and highlights connections with phylogenetic systematics as well as the structure and the biology of extant angiosperms"-- Provided by publisher. "The recent discovery of diverse fossil flowers and floral organs in Cretaceous strata has revealed astonishing details about the structural and systematic diversity of early angiosperms. Exploring the rich fossil record that has accumulated over the last three decades, this unique study follows the evolutionary history of flowering plants from their earliest phases in obscurity to their dominance in modern vegetation. The book provides comprehensive biological and geological background information, before moving on to summarise the fossil record in detail. Including previously unpublished results based on research into Early and Late Cretaceous fossil floras from Europe and North America, the authors draw together direct palaeontological evidence of the pattern of angiosperm evolution through time. Synthesising palaeobotanical data with information from living plants, this book explores the latest research in the field and highlights connections with phylogenetic systematics as well as the structure and the biology of extant angiosperms"-- Résumé de l'éditeur Machine generated contents note: Preface; 1. Introduction to angiosperms; 2. The nature of the angiosperm fossil record; 3. The environmental context of early angiosperm evolution; 4. Stratigraphic framework and key areas for Cretaceous angiosperms; 5. Angiosperms in context: extant and fossil seed plants; 6. Origin and age of angiosperms; 7. Phylogenetic framework and the assignment of fossils to extant groups; 8. Fossils near the base of the angiosperm tree; 9. Early fossil angiosperms of uncertain relationships; 10. Early fossils of eumagnoliids; 11. Fossils of monocots; 12. Fossils of eudicots: early diverging groups; 13. Fossils of core eudicots: basal lineages; 14. Fossils of core eudicots: rosids; 15. Early fossils of eudicots: asterids; 16. Patterns of structural diversification in angiosperm reproductive organs; 17. History and evolution of pollination in angiosperms; 18. History and evolution of dispersal in angiosperms; 19. Vegetational context of early angiosperm diversification; 20. The accumulation of angiosperm diversity; References; Index. "The recent discovery of diverse fossil flowers and floral organs in Cretaceous strata has revealed astonishing details about the structural and systematic diversity of early angiosperms. Exploring the rich fossil record that has accumulated over the last three decades, this is a unique study of the evolutionary history of flowering plants from their earliest phases in obscurity to their dominance in modern vegetation. The discussion provides comprehensive biological and geological background information, before moving on to summarise the fossil record in detail. Including previously unpublished results based on research into Early and Late Cretaceous fossil floras from Europe and North America, the authors draw on direct palaeontological evidence of the pattern of angiosperm evolution through time. Synthesising palaeobotanical data with information from living plants, this unique book explores the latest research in the field, highlighting connections with phylogenetic systematics, structure and the biology of extant angiosperms"--Résumé de l'éditeur