Michael J. Benton is Professor of Vertebrate Palaeontology at the University of Bristol. He is interested particularly in early reptiles, Triassic dinosaurs and macroevolution, and has published 50 books and over 400 scientific articles. He founded the Masters in Paleobiology degree at Bristol, which has now graduated over 400 students.

David A.T. Harper is a leading expert on fossil brachiopods, numerical methods in paleontology and Phanerozoic stratigraphy. He is Professor of Paleontology, and Principal of Van Mildert College in Durham University. He has published over 15 books and monographs, including a couple of influential textbooks, as well as over 300 scientific articles and, together with Øyvind Hammer, the widely-used software package PAST.

Preface xi

Acknowledgments xiii

1 Paleontology as a science 1

Introduction 2

Paleontology in the modern world 2

Paleontology as a science 3

Steps to understanding 12

Fossils and evolution 16

Paleontology today 17

Review questions 23

Further reading 23

References 24

2 Stratigraphy 25

Introduction 26

How stratigraphy works 26

New techniques, new tools 40

Geological time scale: a common language 46

Extraterrestrial stratigraphy 47

Review questions 48

Further reading 48

References 49

3 Paleogeography and paleoclimates 50

Paleobiogeography 50

Fossils in mountain belts 64

Paleoclimates 69

The Anthropocene 76

Review questions 77

Further reading 77

References 77

4 Paleoecology 80

Introduction 81

Taphonomic constraints: sifting through the debris 83

Populations: can groups of individuals make a difference? 85

Habitats and niches 86

Paleocommunities 95

Evolutionary paleoecology 100

Ecological ranking of mass extinctions 110

Full contents vii

Review questions 112

Further reading 112

References 112

5 Taphonomy and the quality of the fossil record 115

Introduction 116

Fossil preservation 116

Quality of the fossil record 130

Review questions 137

Further reading 138

References 138

6 Fossil form and function 140

Introduction 141

Growth and form 141

Evolution and development 147

Interpreting the function of fossils 152

Review questions 163

Further reading 164

References 164

7 Macroevolution and the tree of life 165

Introduction 166

Evolution by natural selection 167

Evolution and the fossil record 170

Trends and radiations 179

The tree of life 184

Review questions 191

Further reading 191

References 191

8 Biodiversity, extinction, and mass extinction 193

Introduction 194

The diversification of life 195

Mass extinctions 202

The "big five" mass extinction events 207

Extinction then and now 215

Review questions 220

Further reading 220

References 221

9 The origin of life 223

Introduction 224

The origin of life 224

Evidence for the origin of life 228

Life diversifies: eukaryotes 236

Review questions 245

Further reading 246

References 246

10 Protists 248

Introduction 249

Protozoa 252

Mystery protists of the proterozoic and paleozoic 260

Phytoplankton 266

Review questions 277

Further reading 277

References 277

11 Origin and expansion of the metazoans 279

Origins and classification 280

Invertebrate body and skeletal plans 284

Five key faunas 289

Review questions 304

Further reading 304

References 304

12 The basal metazoans: sponges and corals 306

Introduction 307

Porifera 307

Cnidaria 322

Review questions 341

Further reading 342

References 342

13 Lophophorates 1: brachiopods and bryozoans 344

Introduction 345

Brachiopoda 345

Bryozoa 360

Review questions 369

Further reading 370

References 370

14 Lophotrochozoans 2: mollusks and annelids 372

Mollusks 372

Introduction 373

Early mollusks 375

Class Bivalvia 380

Class Gastropoda 384

Class Cephalopoda 390

Class Scaphopoda 403

Class Rostroconcha 403

Evolutionary trends within the Mollusca 404

Annelids 406

Review questions 410

Further reading 410

References 410

15 Ecdysozoa: arthropods 412

Introduction 413

Early arthropod faunas 413

Subphylum Trilobitomorpha (artiopoda) 416

Subphylum Chelicerata 428

Subphylum Myriapoda 430

Subphylum Hexapoda 430

Subphylum Crustacea 433

Exceptional arthropod faunas through time 438

Review questions 442

Further reading 442

References 443

16 Deuterostomes: echinoderms and hemichordates 445

Introduction 446

Echinoderms 446

Hemichordates 468

Review questions 484

Further reading 484

References 484

17 Fishes and basal tetrapods 486

Introduction 487

Origin of the vertebrates 487

Jaws and fish evolution 492

Tetrapods 498

Reign of the reptiles 503

Review questions 510

Further reading 510

References 511

18 Dinosaurs and mammals 512

Introduction 513

Dinosaurs and their kin 513

Bird evolution 521

Rise of the mammals 522

The line to humans 530

Review questions 536

Further reading 536

References 537

19 Fossil plants and fungi 538

Introduction 539

Terrestrialization of plants 539

The great coal forests 547

Seed-bearing plants 552

Flowering plants 562

Review questions 568

Further reading 568

References 568

20 Trace fossils 570

Introduction 571

Understanding trace fossils 571

Trace fossils in sediments 578

Review questions 592

Further reading 592

References 592

Finale 594

Glossary 598

Appendix 1: Stratigraphic chart 617

Appendix 2: Paleogeographic maps 619

Index 620