Paleobiology II

PALAEOBIOLOGY II The dinosaur Diplodocus as seen in the BBC’s acclaimed series ‘Walking with Dinosaurs’ — the world’s first natural history of dinosaurs © BBC Worldwide Ltd, 1999 PALAEOBIOLOGY II EDITED BY DEREK E.G BRIGGS Department of Geology & Geophysics Yale University New Haven Connecticut 06511 USA AND PETER R CROWTHER Keeper of Geology National Museums and Galleries of Northern Ireland Ulster Museum Botanic Gardens Belfast BT9 5AB OF THE PALAEONTOLOGICAL ASSOCIATION To the memory of J.J Sepkoski Jr © 2001, 2003 by Blackwell Science Ltd, a Blackwell Publishing company 350 Main Street, Malden, MA 02148-5018, USA 108 Cowley Road, Oxford OX4 1JF, UK 550 Swanston Street, Carlton South, Melbourne, Victoria 3053, Australia Kurfürstendamm 57, 10707 Berlin, Germany The right of the Authors to be identified as the Authors of the Editorial Material in this Work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988 All rights reserved No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the prior permission of the publisher First published 2001 First published in paperback 2003 Library of Congress Cataloging-in-Publication Data Palaeobiology II / edited by D.E.G Briggs, P.R Crowther; foreword by E.N.K Clarkson p cm Includes bibliographical references and index ISBN 0-632-05147-7 (hb : alk paper)—ISBN 0-632-05149-3 (pb : alk paper) Palaeobiology I Title: Palaeobiology two II Title: Palaeobiology III Briggs, D.E.G IV Crowther, Peter R QE719.8 P34 560 — dc21 2001 00-031211 A catalogue record for this title is available from the British Library Set by SNP Best-set Typesetter Ltd., Hong Kong Printed and bound in the United Kingdom by MPG Books Ltd, Bodmin, Cornwall For futher information on Blackwell Publishing, visit our website: http://www.blackwellpublishing.com Contents List of Contributors, ix Foreword, xv e.n.k clarkson Major Events in the History of Life 1.1 Early Life, 1.2 1.3 1.1.1 Origin of Life, a lazcano 1.1.2 Exploring for a Fossil Record of Extraterrestrial Life, j.d farmer 1.1.3 Life in the Archaean, 13 r buick 1.1.4 Late Proterozoic Biogeochemical Cycles, 22 g.a logan and r.e summons The Cambrian Radiation, 25 1.2.1 Metazoan Origins and Early Evolution, 25 d.h erwin 1.2.2 Significance of Early Shells, 31 s conway morris 1.2.3 Cambrian Food Webs, 40 n.j butterfield 1.2.4 The Origin of Vertebrates, 43 m.p smith and i.j sansom Palaeozoic Events, 49 1.3.1 Ordovician Radiation, 49 a.i miller 1.3.2 Rise of Fishes, 52 j.a long 1.3.3 Evolution of Reefs, 57 r.a wood 1.3.4 Early Land Plants, 63 d edwards 1.3.5 Afforestation — the First Forests, 67 s.e scheckler 1.3.6 Terrestrialization of Animals, 71 p.a selden 1.3.7 Origin of Tetrapods, 74 m.i coates 1.3.8 Carboniferous Coal-swamp Forests, 79 w.a dimichele 1.3.9 Rise and Diversification of Insects, 82 c.c labandeira 1.4 1.5 1.3.10 Origin of Mammals, 88 j.a hopson Mesozoic Events, 94 1.4.1 Mesozoic Marine Revolution, 94 p h k e l l e y and t.a hansen 1.4.2 Origin and Radiation of Angiosperms, 97 e.m friis, k.r pedersen and p.r crane 1.4.3 Rise of Birds, 102 l.m chiappe Cenozoic Events, 106 1.5.1 Evolution of Modern Grasslands and Grazers, 106 t.e cerling 1.5.2 Radiation of Tertiary Mammals, 109 c.m janis 1.5.3 Rise of Modern Land Plants and Vegetation, 112 m.e collinson 1.5.4 Early Primates, 115 k.d rose 1.5.5 Hominid Evolution, 121 b.a wood 1.5.6 Neandertals, 127 l.c aiello The Evolutionary Process and the Fossil Record 2.1 Species Evolution, 133 2.1.1 Speciation and Morphological Change, 133 d.b lazarus 2.1.2 Evolutionary Stasis vs Change, 137 a.h cheetham 2.1.3 Rapid Speciation in Species Flocks, 143 a.r m ccune Evolution of Form, 147 2.2.1 Developmental Genes and the Evolution of Morphology, 147 g.a wray 2.2.2 Constraints on the Evolution of Form, 152 p.j wagner 2.2.3 Occupation of Morphospace, 157 a.r.h swan 2.2 v vi 2.3 2.4 Contents Macroevolution, 162 2.3.1 Origin of Evolutionary Novelties, 162 d jablonski 2.3.2 Controls on Rates of Evolution, 166 s.m stanley 2.3.3 Competition in Evolution, 171 j.j sepkoski jr 2.3.4 Biotic Interchange, 176 d.r lindberg 2.3.5 Importance of Heterochrony, 180 k.j m cnamara 2.3.6 Hierarchies in Evolution, 188 t.a grantham 2.3.7 Phylogenetic Tree Shape, 192 p.n pearson 2.3.8 Contingency, 195 s.j gould 2.3.9 Selectivity during Extinctions, 198 m.l m c kinney 2.3.10 Biotic Recovery from Mass Extinctions, 202 d.j bottjer 2.3.11 Evolutionary Trends, 206 d w m cshea Patterns of Diversity, 211 2.4.1 Biodiversity through Time, 211 m.j benton 2.4.2 Late Ordovician Extinction, 220 p.j brenchley 2.4.3 Late Devonian Extinction, 223 g.r m cghee j r 2.4.4 End-Permian Extinction, 226 p.b wignall 2.4.5 Impact of K–T Boundary Events on Marine Life, 229 r.d norris 2.4.6 Impact of K–T Boundary Events on Terrestrial Life, 232 j.a wolfe and d.a russell 2.4.7 Pleistocene Extinctions, 234 k roy 3.2 3.3 3.4 Taphonomy 3.1 Fossilized Materials, 241 3.1.1 DNA, 241 h.n poinar and s pääbo 3.1.2 Proteins, 245 m.j collins and a.m gernaey 3.1.3 Lipids, 247 r.p evershed and m.j lockheart 3.1.4 Bacteria, 253 k liebig 3.1.5 Resistant Plant Tissues — Cuticles and Propagules, 256 p f van bergen 3.1.6 Animal Cuticles, 259 b.a stankiewicz and d.e.g briggs 3.1.7 Shells, 262 k.h meldahl 3.1.8 Bones, 264 c denys Fossilization Processes, 270 3.2.1 Decay, 270 p.a allison 3.2.2 Bioerosion, 273 e.n edinger 3.2.3 Preservation by Fire, 277 a.c scott 3.2.4 Role of Microbial Mats, 280 j.-c gall 3.2.5 Bioimmuration, 285 p.d taylor and j.a todd 3.2.6 Transport and Spatial Fidelity, 289 l.c anderson 3.2.7 Time-averaging, 292 k.w flessa Preservation in Different Ecological Settings, 297 3.3.1 Major Biases in the Fossil Record, 297 s.m kidwell 3.3.2 Benthic Marine Communities, 303 w.d allmon 3.3.3 Ancient Reefs, 307 j.m pandolfi 3.3.4 Marine Plankton, 309 r.e martin 3.3.5 Terrestrial Plants, 312 r.a gastaldo 3.3.6 Pollen and Spores, 315 j.m van mourik 3.3.7 Terrestrial Vertebrates, 318 a k b e h r e n sme y e r 3.3.8 Sphagnum-dominated Peat Bogs, 321 t.j painter 3.3.9 Archaeological Remains, 325 v straker Lagerstätten, 328 3.4.1 Exceptionally Preserved Fossils, 328 d.e.g briggs 3.4.2 Precambrian Lagerstätten, 332 a h k n ol l and shuhai xiao 3.4.3 Chengjiang, 337 j bergström 3.4.4 The Soom Shale, 340 r.j aldridge, s.e gabbott and j.n theron 3.4.5 The Rhynie Chert, 342 n.h trewin 3.4.6 Hunsrück Slate, 346 r raiswell, c bartels and d.e.g briggs Contents 3.4.7 La Voulte-sur-Rhône, 349 p.r wilby 3.4.8 The Santana Formation, 351 d.m martill 3.4.9 Las Hoyas, 356 j.l sanz, m.a fregenal-martínez, n melénd ez and f ortega 3.4.10 The Princeton Chert, 359 r.a stockey 3.4.11 Dominican Amber, 362 g.o poinar jr Palaeoecology 4.1 Fossils as Living Organisms, 367 4.1.1 Bringing Fossil Organisms to Life, 367 p.w skelton 4.1.2 Stromatolites, 376 m.r walter 4.1.3 Plant Growth Forms and Biomechanics, 379 t speck and n p ro w e 4.1.4 Sessile Invertebrates, 384 w.i ausich and d.j bottjer 4.1.5 Trilobites, 386 b.d.e chatterton 4.1.6 Trackways — Arthropod Locomotion, 389 s.j braddy 4.1.7 Durophagy in Marine Organisms, 393 r.b aronson 4.1.8 Buoyancy, Hydrodynamics, and Structure in Chambered Cephalopods, 397 d.k jacobs 4.1.9 Feeding in Conodonts and other Early Vertebrates, 401 m.a purnell 4.1.10 Locomotion in Mesozoic Marine Reptiles, 404 m.a taylor 4.1.11 Trackways — Dinosaur Locomotion, 408 m.g lockley 4.1.12 Dinosaur Ethology, 412 j.r horner 4.1.13 Predatory Behaviour in Maniraptoran Theropods, 414 a.d gishlick 4.1.14 Pterosaur Locomotion, 417 d.m unwin 4.1.15 Predation in Sabre-tooth Cats, 420 b van valkenburgh 4.1.16 Plant–Animal Interactions: Herbivory, 424 s ash 4.1.17 Plant–Animal Interactions: Insect Pollination, 426 w.l crepet 4.1.18 Plant–Animal Interactions: Dispersal, 429 j.j hooker and m.e collinson 4.2 4.3 vii Ancient Communities, 432 4.2.1 Ecological Changes through Geological Time, 432 m.l droser 4.2.2 Do Communities Evolve? 437 r.k bambach 4.2.3 Palaeobiogeography of Marine Communities, 440 g.r shi 4.2.4 Deep-sea Communities, 444 t oji 4.2.5 Ancient Hydrothermal Vent and Cold Seep Faunas, 447 c t s l i t t l e 4.2.6 Zooplankton, 451 s rigby and c.v milsom 4.2.7 Terrestrial Palaeobiogeography, 454 r.s hill 4.2.8 Epibionts, 460 h.l lescinsky 4.2.9 Fungi in Palaeoecosystems, 464 t.n taylor and e l tay l or Fossils as Environmental Indicators, 467 4.3.1 Taphonomic Evidence, 467 m.v.h wilson 4.3.2 Oxygen in the Ocean, 470 w oschmann 4.3.3 Carbon Isotopes in Plants, 473 d j b e e r l i n g 4.3.4 Bathymetric Indicators, 475 p.j orr 4.3.5 Atmospheric Carbon Dioxide — Stomata, 479 j c mc elwain 4.3.6 Climate — Wood and Leaves, 480 d.r greenwood 4.3.7 Climate — Modelling using Fossil Plants, 483 g.r upchurch jr 4.3.8 Climate — Quaternary Vegetation, 485 t we b b Systematics, Phylogeny, and Stratigraphy 5.1 Morphology and Taxonomy, 489 5.1.1 Quantifying Morphology, 489 r.e chapman and d rasskin-gutman 5.1.2 Morphometrics and Intraspecific Variation, 492 n.c hughes 5.1.3 Disparity vs Diversity, 495 m.a wills Calibrating Diversity, 500 5.2.1 Estimating Completeness of the Fossil Record, 500 m foote 5.2 viii 5.3 Contents 5.2.2 Analysis of Diversity, 504 a.b smith Reconstructing Phylogeny, 509 5.3.1 Phylogenetic Analysis, 509 m wilkinson 5.3.2 Fossils in the Reconstruction of Phylogeny, 515 p.l forey and r.a fortey 5.3.3 Stratigraphic Tests of Cladistic Hypotheses, 519 m.a norell 5.3.4 Molecular Phylogenetic Analysis, 522 j.p hu elsenbeck 5.3.5 Molecules and Morphology in Phylogeny — the Radiation of Rodents, 529 f.m catzeflis 5.3.6 Using Molecular Data to Estimate Divergence Times, 532 a cooper, n grassly and a rambaut 5.4 Fossils in Stratigraphy, 535 5.4.1 Stratigraphic Procedure, 535 p.f rawson 5.4.2 Calibration of the Fossil Record, 539 s.a bowring and m.w martin 5.4.3 Confidence Limits in Stratigraphy, 542 c.r marshall 5.4.4 High-resolution Biostratigraphy, 545 j backman 5.4.5 Sequence Stratigraphy and Fossils, 548 s.m holland Index, 555 List of Contributors L.C AIELLO Department of Anthropology, University College London, Gower Street, London WC1E 6BT, UK R.J ALDRIDGE Department of Geology, University of Leicester, University Road, Leicester LE1 7RH, UK P.A ALLISON T.H Huxley School for Environment, Earth Science & Engineering, Imperial College of Science, Technology & Medicine, Prince Consort Road, South Kensington, London SW7 5BP, UK W.D ALLMON Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, USA L.C ANDERSON Department of Geology & Geophysics, Louisiana State University, Baton Rouge, Louisiana 70803, USA R.B ARONSON Dauphin Island Sea Lab, 101 Bienville Boulevard, Dauphin Island, Alabama 36528, USA S ASH Department of Earth & Planetary Sciences, University of New Mexico, Albuquerque, New Mexico 87131, USA W.I AUSICH Department of Geological Sciences, Ohio State University, 155 South Oval Mall, Columbus, Ohio 43210-1397, USA J BACKMAN Department of Geology & Geochemistry, Stockholm University, S-106 91 Stockholm, Sweden R.K BAMBACH Department of Geological Sciences, Virginia Polytechnic Institute & State University, 4044 Derring Hall, Blacksburg, Virginia 24061-0420, USA C BARTELS Deutsches Bergbau-Museum, Am Bergbaumuseum 28, D-44791 Bochum, Germany D.J BEERLING Department of Animal & Plant Sciences, University of Sheffield, Alfred Denny Building, Western Bank, Sheffield S10 2TN, UK A.K BEHRENSMEYER Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0121, USA M.J BENTON Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queen's Road, Bristol BS8 1RJ, UK P.F van BERGEN Shell Global Solutions International BV, PO Box 38000, 1030 BN Amsterdam, The Netherlands J BERGSTRÖM Department of Palaeozoology, Swedish Museum of Natural History, PO Box 50007, S-104 05 Stockholm, Sweden D.J BOTTJER Department of Earth Sciences, University of Southern California, Los Angeles, California 90089, USA S.A BOWRING Department of Earth, Atmospheric & Planetary Sciences, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139, USA S.J BRADDY Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queen's Road, Bristol BS8 1RJ, UK P.J BRENCHLEY Department of Earth Sciences, University of Liverpool, PO Box 147, Liverpool L69 3GP, UK D.E.G BRIGGS Department of Geology & Geophysics, Yale University, New Haven, Connecticut 06511, USA R BUICK Department of Earth & Space Sciences and Astrobiology Program, University of Washington, Seattle, Washington 98195-1310, USA N.J BUTTERFIELD Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, UK F.M CATZEFLIS Institut des Sciences de l'Evolution, UMR 5554 CNRS & Université Montpellier 2, F-34095 Montpellier 05, France T.E CERLING Department of Geology & Geophysics, University of Utah, Salt Lake City, Utah 84103, USA R.E CHAPMAN Applied Morphometrics Laboratory (ADP) & Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA B.D.E CHATTERTON Department of Earth & Atmospheric Sciences, University of Alberta, Edmonton, Alberta, Canada T6G 2E1 A.H CHEETHAM Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA L.M CHIAPPE Department of Vertebrate Paleontology, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California, 90007, USA ix ... paper)—ISBN 0-632-05149-3 (pb : alk paper) Palaeobiology I Title: Palaeobiology two II Title: Palaeobiology III Briggs, D.E.G IV Crowther, Peter R QE719.8 P34 560 — dc21 2001 00-031211 A catalogue...PALAEOBIOLOGY II The dinosaur Diplodocus as seen in the BBC’s acclaimed series ‘Walking with Dinosaurs’ — the world’s first natural history of dinosaurs © BBC Worldwide Ltd, 1999 PALAEOBIOLOGY II EDITED... Calibrating Diversity, 500 5.2.1 Estimating Completeness of the Fossil Record, 500 m foote 5.2 viii 5.3 Contents 5.2.2 Analysis of Diversity, 504 a.b smith Reconstructing Phylogeny, 509 5.3.1