biotic interactions in recent and fossil benthic communities978-1-4757-0740-3/1.pdf · volume 3...
TRANSCRIPT
TOPICS IN GEOBIOLOGY Series Editor: F. G. Stehli, University of Oklahoma
Volume 1 SKELETAL GROWTH OF AQUATIC ORGANISMS Biological Records of Environmental Change Edited by Donald C. Rhoads and Richard A. Lutz
Volume 2 ANIMAL-SEDIMENT RELATIONS The Biogenic Alteration of Sediments Edited by Peter L. McCall and Michael J. S. Tevesz
Volume 3 BIOTIC INTERACTIONS IN RECENT AND FOSSIL BENTHIC COMMUNITIES Edited by Michael J. S. Tevesz and Peter L. McCall
Biotic Interactions in Recent and Fossil Benthic Communities Edited by
Michael J. S. Tevesz Cleveland State University Cleveland, Ohio
and
Peter L. Me Call Case Western Reserve University Cleveland, Ohio
Springer Science+Business Media, LLC
Library of Congress Cataloging in Publication Data
Main entry under title:
Biotic interactions in recent and fossil benthic communities.
(Topics in geobiology; v. 3) 1. Paleoecology. 2. Biotic communities. 1. Tevesz, Michael J. S. II. McCall , P. L.
(Peter L.), 1948- . III. Series. QE720.B56 1983 560' .45 83-13953 ISBN 978-1-4757-0742-7 ISBN 978-1-4757-0740-3 (eBook) DOI 10.1007/978-1-4757-0740-3
© 1983 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1983
AII rights reserved
No part of this book may be reproduced. stored in a retrieval system. Of transmitted in any form or by any means. electronic, mechanical. photocopying. microfilming. recording, or otherwise , without written permission rrom the Publisher
Contributors
William I. Ausich Department of Geological Sciences, Wright State University, Dayton, Ohio 45435
Richard K. Bambach Department of Geological Sciences, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061
David J. Bottjer Department of Geological Sciences, University of South-ern California, Los Angeles, California 90089 ·
Richard Cowen Department of Geology, University of California, Davis, California 95616
David Jablonski Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85721
J. B. C. Jackson Department of Earth and Planetary Sciences, The Johns Hopkins University, Baltimore, Maryland 21218
Susan M. Kidwell Department of Geosciences, University of Arizona, Tucson, Arizona 85721
Jennifer A. Kitchell Department of Geology and Geophysics, University of Wisconsin, Madison, Wisconsin 53706
Andrew H. Knoll Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, Massachusetts 02138
David W. Larson Department of Geology, Franklin and Marshall College, Lancaster, Pennsylvania 17604. Present address: Department of Geology, Hamilton College, Clinton, New York 13323
Jere H. Lipps Department of Geology, University of California, Davis, California 95616
Peter L. McCall Department of Geological Sciences, Case Western Reserve University, Cleveland, Ohio 44106
David L. Meyer Department of Geology, University of Cincinnati, Cincinnati, Ohio 45221
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vi Contributors
Donald C. Rhoads Department of Geology and Geophysics, Yale University, New Haven, Connecticut 06520
J. John Sepkoski, Jr. Department of the Geophysical Sciences, University of Chicago, Chicago, Illinois 60637
Peter M. Sheehan Department of Geology, Milwaukee Public Museum, Milwaukee, Wisconsin 53233
Michael J. S. Tevesz Department of Geological Sciences, Cleveland State University, Cleveland, Ohio 44115
Charles W. Thayer Department of Geology, University of Pennsylvania and Academy of Natural Sciences, Philadelphia, Pennsylvania 19104
James W. Valentine Department of Geological Sciences, University of California, Santa Barbara, California 93106
Geerat J. Vermeij Department of Zoology, University of Maryland, College Park, Maryland 20742
Sarah Ann Woodin Department of Biology, University of South Carolina, Columbia, South Carolina 29208
Preface
When Frank Stehli approached us in 1978 to participate in the Topics in Geobiology series, the idea for this book came easily to mind, because the role of biotic interactions in controlling the distribution of fossil organisms is at once intriguing and problematical. After organizing our own thoughts on this diffuse subject, we contacted a number of people whom we knew or suspected were interested in the subject and would be willing to commit their thoughts to writing and scrutiny. Given the current paucity of knowledge in this area, we encouraged responsible speculation that might promote further investigation. To maintain some semblance of coherency, we have limited the contents of this volume to topics largely (but not exclusively) pertaining to benthic marine invertebrates, the organisms with the best fossil record. We have not tried to be encyclopedic, because this would expose an embarrassingly large number of lacunae in the field. In fact, to a large extent the book has been molded around the response of our colleagues to our requests for contributions.
This book is divided into four parts. Part I documents the importance of biotic interactions in Recent benthic marine environments and the preservation of evidence of interactions. A major theme running through benthic ecological literature from the late 1800s to the present is the relative importance of physical environmental factors such as temperature, salinity, and subtratum type versus biological factors such as competition, predation, amensalism, disease, and parasitism. We think it is safe to say that until fairly recently, most workers, with few exceptions, subscribed to the notion that biologic interactions had little effect on distribution and abundance. The dominance of this view may be ascribed in large measures to the way bottom communities were studied: from the deck of a boat, with a grab sampler and large-mesh sieve that did not collect early life stages, and over an area of many square miles. In situ observations and small-scale observations were difficult to make. These older ideas still strongly influence many paleoecologists. The first several chapters show, however, that while physical factors related to phenomena such as seasonality affect benthic communities, biotic interactions profoundly affect species composition and abundance of both hard- and soft-bottom com~ munities.
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Part II documents that biotic interactions are widespread and influential not only in different environments (as demonstrated in Part I) but also among a number of highly distinct taxonomic groups: prokaryotes, siliceous phytoplankton (we have included them, contrary to our title, because of their excellent fossil record and preservation among the benthos), benthic foraminifera, and crinoids.
While we think it is apparent that biotic interactions are an important influence on the structure and dynamics of modern benthic communities, and are common among particular groups of organisms with excellent fossil records, the importance of these factors in helping shape ancient communities and their history is less clear. One goal of paleontology is to explain the causes behind faunal changes through time. For example, why did brachiopods decline? Why did trilobites disappear? What are the reasons for the explosive evolution in the Cambro-Ordovician? Some explanations invoke habitat removal or alteration by changing shelf area and plate motions. Other explanations, like those proposed for modern environments, center around changes in specific aspects of the physical environment such as temperature, salinity, or oxygen. These may all be important reasons, and there is often good evidence to support them. Other potentially important reasons for these changes, such as biotic interactions, are too little explored. Nevertheless, a few bold writers have claimed that local-level interactions have caused large-scale changes in the biosphere. For instance, the origin and initial adaptive radiations of metazoans and the rise of angiosperms are believed by some to have been caused by biotic interactions such as cropping and mutualism, respectively.
But hypotheses that give great importance to the role of biotic interactions in shaping ancient communities and their history may be attacked because the fossil record is too incomplete and imprecise to reconstruct many local-level interactions, and because large-scale changes in the biosphere occur over such a long time span that they may be more influenced by stochastic rather than deterministic factors. The authors of the final sections of the book attempt to show that biotic interactions of various sorts are pervasive throughout much of the history of life (Part III) and have been responsible for influencing a wide variety of phenomena. In Part IV, evidence is presented to show that biotic interactions have helped shape the structure, distribution, and evolution of marine benthic communities for much of the past 550 million years.
We hope that the several chapters provide an organized presentation and commentary on the present level of knowledge concerning biotic interactions and serve to stimulate more thought and research on these subjects, especially among paleontologists and paleoecologists.
M. J. S. Te\·esz P. L. McCall
Contents
I. Recent Interactions and Their Preservation
Chapter 1 • Biotic Interactions in Recent Marine Sedimentary Environments
Sarah Ann Woodin
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 1.1. Common Units of Measure . . . . . . . . . . . . . . . . . . . 5 1.2. Organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 1.3. Categories of Infauna. . . . . . . . . . . . . . . . . . . . . . . . 7
2. Competition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 2.1. Competition in the Recent. . . . . . . . . . . . . . . . . . . . 10 2.2. Competition in the Paleozoic . . . . . . . . . . . . . . . . . 15
3. Predation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 3.1. Types of Predators in the Recent............... 19 3.2. Predation in the Paleozoic. . . . . . . . . . . . . . . . . . . . 25
4. Pattern Differentiation: Competition or Predation? . . . . . . 27 5. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Chapter z • Biological Determinants of Present and Past Sessile Animal Distributions
J. B. C. Jackson
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 2. Organisms and Their Environments . . . . . . . . . . . . . . . . . 41 3. Causes of Distributions . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
3 .1. The Larval Pool. . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 7 3.2. Larval Habitat Selection and Interactions with
Previously Settled Organisms . . . . . . . . . . . . . . . . . 48 3.3. Competition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 3.4. Predation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 3.5. Mutualism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
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3.6. Life Histories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 4. Fossil Evidence for Causes of Distributions. . . . . . . . . . . . 81
4.1. Some Criteria for Recognition of Past Habitat Selection and Mortality Processes in Fossils. . . . . . 81
4.2. Examples of Ancient Interactions . . . . . . . . . . . . . . 89 5. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Chapter 3 • Seasonality: Effects in Marine Benthic Communities
James W. Valentine
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 1.1. Solar Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 3 1.2. Circulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
2. Seasonal Parameters with Primary Density-Independent Effects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 7 2.1. Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 2.2. Salinity and Other Physical Variables . . . . . . . . . . 131 2.3. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
3. Seasonal Parameters with Primarily Density-Dependent Effects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 5 3.1. Light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135 3.2. Nutrient Elements . . . . . . . . . . . . . . . . . . . . . . . . . 135 3.3. Primary Production . . . . . . . . . . . . . . . . . . . . . . . . 136 3.4. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
4. Processes That Mediate Density-Dependent Effects. . . . . . 142 4.1. General Theory . . . . . . . . . . . . . . . . . . . . . . . . . . . 142 4.2. Reproduction and Development: Seasonal
Strategies in z . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 4.3. Niche Expansion: Seasonal Strategies in X . . . . . . . 145 4.4. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
5. Consequences for Biotic Patterns. . . . . . . . . . . . . . . . . . . . 147 5.1. Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 5.2. Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148 5.3. Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Chapter 4 • Soft-Bottom Succession and the Fossil Record
1. 2.
Peter L. McCall and Michael J. S. Tevesz
Introduction What Is Succession? ............................ .
157 158
Contents xi
3. What Use Is It? 160 4. Nearshore Benthic Succession . . . . . . . . . . . . . . . . . . . . . 161
4.1. Biotic Interactions . . . . . . . . . . . . . . . . . . . . . . . . . 161 4.2. Study Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
5. Succession in Other Environments . . . . . . . . . . . . . . . . . . 169 6. Preservation of Soft-Bottom Succession. . . . . . . . . . . . . . . 171
6.1. Taphonomic Losses and Mixing . . . . . . . . . . . . . . . 171 6.2. Diagenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175 6.3. Comparison Of Life and Death Assemblages. . . . . . 176
7. Relation to Geologic Examples . . . . . . . . . . . . . . . . . . . . . 183 8. Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
Chapter 5 • Taphonomic Feedback: Ecological Consequences of Shell Accumulation
Susan M. Kidwell and David Jablonski
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195 2. Recent and Fossil Examples of Taphonomic Feedback. . . 196
2.1. Taphonomic Facilitation . . . . . . . . . . . . . . . . . . . . 197 2.2. Taphonomic Inhibition . . . . . . . . . . . . . . . . . . . . . 206
3. Expected Patterns in the Stratigraphic Record. . . . . . . . . . 208 3.1. Sediment Aggradation . . . . . . . . . . . . . . . . . . . . . . 212 3.2. Sediment Starvation . . . . . . . . . . . . . . . . . . . . . . . . 213 3.3. Sediment Bypassing . . . . . . . . . . . . . . . . . . . . . . . . 215 3.4. Erosional Truncation . . . . . . . . . . . . . . . . . . . . . . . 217 3.5. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
4. Case Study: Neogene Chesapeake Group, Atlantic Coastal Plain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220 4.1. Evidence of Taphonomic Feedback. . . . . . . . . . . . . 223 4.2. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
5. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233 References 235
II. Interactions among Selected Taxa
Chapter 6 • Biological Interactions and Precambrian Eukaryotes
Andrew H. Knoll
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251 2. Hypotheses of Eukaryotic Origins . . . . . . . . . . . . . . . . . . . 253
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3. A Scenario for Early Eukaryotic Evolution . . . . . . . . . . . . 258 4. The Fossil Record of Early Eukaryotes . . . . . . . . . . . . . . . 264
4.1. "Spot" Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265 4.2. Tetrahedral Tetrads . . . . . . . . . . . . . . . . . . . . . . . . 268 4.3. Filamentous Microfossils . . . . . . . . . . . . . . . . . . . . 268 4.4. Size Distribution Data . . . . . . . . . . . . . . . . . . . . . . 269 4.5. Acritarchs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 70 4.6. Vase-Shaped Microfossils . . . . . . . . . . . . . . . . . . . . 273 4.7. Precambrian Macrofossils . . . . . . . . . . . . . . . . . . . . 273
5. Ecological Consequences of Early Eukaryote Evolution. . . 274 6. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
Chapter 7 • Biotic Interactions and Siliceous Marine Phytoplankton: An Ecological and Evolutionary Perspective
Jennifer A. Kitchell
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285 2. Competition and Coexistence . . . . . . . . . . . . . . . . . . . . . . 287
2.1. Nutrient Uptake Kinetics . . . . . . . . . . . . . . . . . . . . 287 2.2. Coexistence of Competing Species . . . . . . . . . . . . . 291 2.3. The Paradox of Enrichment . . . . . . . . . . . . . . . . . . 294 2.4. Nonequilibrium Theories of Competitive
Coexistence. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296 3. Predator-Prey Interactions . . . . . . . . . . . . . . . . . . . . . . . . 297
3.1. Predation as a Selective Force. . . . . . . . . . . . . . . . . 297 3.2. Predation and the Maintenance of Diversity . . . . . . 302 3.3. Predation and the Sediment Record . . . . . . . . . . . . 303
4. Life History Strategies . . . . . . . . . . . . . . . . . . . . . . . . . . . 304 5. Evolutionary Mode of Phytoplankton . . . . . . . . . . . . . . . . 309 6. Paleontological Applications . . . . . . . . . . . . . . . . . . . . . . 312
6.1. Competitive Displacement in Evolutionary Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313
6.2. Character Divergence and Convergence. . . . . . . . . . 313 6.3. A Case Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 320
Chapters • Biotic Interactions in Benthic Foraminifera
Jere H. Lipps
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331
Contents xiii
2. Trophic Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 332 2.1. Food of Foraminifera . . . . . . . . . . . . . . . . . . . . . . . 332 2.2. Trophic Mechanisms in Foraminifera . . . . . . . . . . . 334 2.3. Consumption of Foraminifera by Other
Organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 52 3. Substrate Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359
3.1. Foraminifera as Epibionts . . . . . . . . . . . . . . . . . . . . 359 3.2. Foraminfera as Substrata . . . . . . . . . . . . . . . . . . . . 363
4. Competition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 363 5. Bioturbation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 364 6. Taphonomic Aspects of Foraminiferal Biotic
Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366 7. Future Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 70
Chapter g • Biotic Interactions among Recent and among Fossil Crinoids
David L. Meyer and William I. Ausich
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 2. Predation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378
2.1. Sources of Predation on Living Crinoids. . . . . . . . . 378 2.2. Possible Antipredator Adaptations of Living
Crinoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380 2.3. Predation on Ancient Crinoids . . . . . . . . . . . . . . . . 381 2.4. Possible Antipredator Morphology in Ancient
Crinoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 2.5. Regeneration and Nonlethal Predation . . . . . . . . . . 385
3. Competition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 3.1. Possible Mechanisms of Competition . . . . . . . . . . . 387 3.2. Niche Differentiation among Living and among
Ancient Crinoids. . . . . . . . . . . . . . . . . . . . . . . . . . . 388 4. Associations of Living Crinoids with Other
Organisms. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392 4.1. Polychaetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395 4.2. Molluscs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396 4.3. Crustaceans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396 4.4. Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397
5. Associations of Ancient Crinoids with Other Organisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 398 5.1. Nature of Associations . . . . . . . . . . . . . . . . . . . . . . 398 5.2. Commensalism . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399 5.3. Stereomic Malformations . . . . . . . . . . . . . . . . . . . . 406
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5.4. Crinoids as Epizoans . . . . . . . . . . . . . . . . . . . . . . . 411 5.5. Parasitism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 412
6. Other Interactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 414 7. Habitat Modification by Crinoids . . . . . . . . . . . . . . . . . . . 415
7.1. Contribution to Sediment . . . . . . . . . . . . . . . . . . . . 415 7.2. Effects on Substrata . . . . . . . . . . . . . . . . . . . . . . . . 416 7.3. Consequences for Community Succession........ 418
8. Role of Biotic Interactions in Crinoid Evolution . . . . . . . . 418 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 420
III. Biotic Interactions through Time
Chapter 10 • Algal Symbiosis and Its Recognition in the Fossil Record
Richard Cowen
1. Symbiosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431 2. Algal Symbiosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432
2.1. The Algal Symbionts . . . . . . . . . . . . . . . . . . . . . . . 433 2.2. Phyletic Distribution of Hosts of Algal
Symbionts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435 2.3. Geographical and Ecological Distribution . . . . . . . . 435
3. The Origin of Algal Symbiosis . . . . . . . . . . . . . . . . . . . . . 436 4. Characters Associated with Symbiosis. . . . . . . . . . . . . . . . 439
4.1. Characters That Promote Symbiosis . . . . . . . . . . . . 440 4.2. Characters That Result from Symbiosis . . . . . . . . . . 446
5. Recognition of Algal Symbiosis in the Fossil Record. . . . . 449 5.1. Usable Criteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . 449 5.2. Fossil Corals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 453 5.3. Fossil Foraminifera . . . . . . . . . . . . . . . . . . . . . . . . 454 5.4. Fossil Cardiacean Bivalves . . . . . . . . . . . . . . . . . . . 456 5.5. Rudist Bivalves . . . . . . . . . . . . . . . . . . . . . . . . . . . 456 5.6. Other Large Bivalves . . . . . . . . . . . . . . . . . . . . . . . 460 5.7. Recalcitrant Groups: The Planktonic Syndrome . . . 462
6. Case Study of Symbiosis in Permian Brachiopods. . . . . . . 462 6.1. The Feeding Mechanism of the
Richthofeniacea. . . . . . . . . . . . . . . . . . . . . . . . . . . . 463 6.2. Symbiosis in the Richthofeniacea . . . . . . . . . . . . . . 465 6.3. Symbiosis in the Teguliferinidae . . . . . . . . . . . . . . 471 6.4. Symbiosis in the Lyttoniacea . . . . . . . . . . . . . . . . . 472 6.5. The Origins of Symbiosis in Brachiopods . . . . . . . . 473 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 7 4
Contents XV
Chapter 11 • Sediment-Mediated Biological Disturbance and the Evolution of Marine Benthos
Charles W. Thayer
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480 2. Disturbance of Recent Sediments: Villains, Victims and
Modi Operandi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481 2.1. Definition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487 2.2. Modes of Sediment-Mediated Interaction . . . . . . . . 487 2.3. Ranking of Modes of Disturbance . . . . . . . . . . . . . . 490 2.4. Effects of Bulldozing on IMOUS . . . . . . . . . . . . . . . 500 2.5. Determinants of Bioturbation Rates . . . . . . . . . . . . 501 2.6. Significance of Trophic Group Amensalism . . . . . . 505
3. Rise of the Bulldozers: Paleontological Perspective and Neontological Insight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507 3.1. Diversification and Deductions from Recent
Reworking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507 3.2. Evidence from Morphology, Traces, and Behavior
517 4. More Evidence from Trace Fossils: Tracking Villains
through Geologic Time . . . . . . . . . . . . . . . . . . . . . . . . . . . 520 5. Physical Disturbance and Biotic Stabilization: How
Shifting Were the Sands of Time?. . . . . . . . . . . . . . . . . . . 522 6. Phanerozoic Patterns: Restructuring the Benthos. . . . . . . . 524
6.1. Brachiopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 531 6.2. Bivalves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 535 6.3. Other Taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 536 6.4. Statistical Summary of Diversity Data. . . . . . . . . . . 537 6.5. Abundant Taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539 6.6. Abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539 6.7. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544
7. Extinctions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544 8. Archaic IMOUS in Recent Refugia: Avoiding the
Bulldozers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 547 8.1. Hard Substrata . . . . . . . . . . . . . . . . . . . . . . . . . . . . 547 8.2. Mobility . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 548 8.3. Deep Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 549 8.4. Littoral Zone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 552 8.5. Refuge in Size . . . . . . . . . . . . . . . . . . . . . . . . . . . . 552 8.6. Corais: Refugia in Excelcis . . . . . . . . . . . . . . . . . . . 553
9. Geological and Paleontological Consequences . . . . . . . . . 553 9.1. Sedimentology and Stratigraphy . . . . . . . . . . . . . . . 553 9.2. Biogeochemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . 554
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9.3. Preservation of Fossils . . . . . . . . . . . . . . . . . . . . . . 555 10. Speculation on Causes and Consequences. . . . . . . . . . . . . 557
10.1. Land Plants and Their Ramifications. . . . . . . . . . . . 557 10.2. Other Causes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 558 10.3. Infaunal vs. Epifaunal Suspension-Feeders. . . . . . . 559 10.4. Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 560 10.5. Brachiopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 560 10.6. Substrate Specificity . . . . . . . . . . . . . . . . . . . . . . . 561 10.7. Hardgrounds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 561
11. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 562 11.1. Potential Bias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 562 11.2. Alternatives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 563 11.3. Implications for Paleoecologic Methods: Generic
Duration, Diversity, and Abundance. . . . . . . . . . . . 564 11.4. Random Patterns? . . . . . . . . . . . . . . . . . . . . . . . . . . 565 11.5. Future Work . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 566 Appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 568 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 595
Chapter 12 • The Evolution of Infaunal Communities and Sedimentary Fabrics
David W. Larson and Donald C. Rhoads
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 627 2. Infaunal Life and Sediment Reworking. . . . . . . . . . . . . . . 628 3. Comparison of Sedimentary Fabrics . . . . . . . . . . . . . . . . . 629
3.1. Sedimentary Facies . . . . . . . . . . . . . . . . . . . . . . . . 631 3.2. Effect of Bioturbation . . . . . . . . . . . . . . . . . . . . . . . 633
4. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642 4.1. Additional Evidence . . . . . . . . . . . . . . . . . . . . . . . 642 4.2. Evolution of Infaunal Communities . . . . . . . . . . . . 642
5. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646
Chapter 13 • Shell-Breaking Predation through Time
Geerat J. Vermeij
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 649 2. Breakage as Agent of Mortality and Selection . . . . . . . . . . 650 3. Adaptations against Breakage . . . . . . . . . . . . . . . . . . . . . . 652 4. Gastropod Shell Form through Geological Time . . . . . . . . 655
Contents xvii
5. Armor in Other Groups. . . . . . . . . . . . . . . . . . . . . . . . . . . 660 6. The Geological Record of Shell-Breakers. . . . . . . . . . . . . . 661 7. Alternative and Additional Interpretations . . . . . . . . . . . . 663
Refurences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 664
IV. Effects of Interactions on Community Evolution
Chapter 14 • Diversification, Faunal Change, and Community Replacement during the Ordovician Radiations
J. John Sepkoski, Jr., and Peter M. Sheehan
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 673 2. Patterns of Diversification and Faunal Change during the
Ordovician . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 7 4 2.1. Global Diversity and the Three "Great
Evolutionary Faunas" . . . . . . . . . . . . . . . . . . . . . . . 675 2.2. Modeling Paleozoic Diversity Patterns . . . . . . . . . . 680
3. Distributional Ecology of the Ordovician Radiations. . . . . 684 3.1. Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 684 3.2. Analytic Methodology . . . . . . . . . . . . . . . . . . . . . . 687 3.3. Cluster Analysis of Cambro-Ordovician
Communities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 689 3.4. Factor Analysis of Cambro-Ordovician
Communities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695 4. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697
4.1. Generality of Results . . . . . . . . . . . . . . . . . . . . . . . 699 4.2. Mechanisms of Onshore-Offshore Change . . . . . . . 701
5. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 705 Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 710
Chapter 15 • Ecospace Utilization and Guilds in Marine Communities through the Phanerozoic
R. K. Bambach
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 719 1.1. Diversity Change through the Phanerozoic . . . . . . . 720 1.2. The Question of an Ecologic Role in Controlling
Diversity Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . 721 2. General Pattern of Ecospace Utilization. . . . . . . . . . . . . . . 722
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2.1. Turnover of Class-Level Taxa through Time . . . . . . 722 2.2. Change in General Ecospace Utilization . . . . . . . . . 725
3. The Guild Concept and Its Application to Paleocommunities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 728 3.1. Extension of the Guild Concept. . . . . . . . . . . . . . . . 728 3.2. Defining Guilds in Paleocommunities. . . . . . . . . . . 730
4. Guilds in Paleocommunities . . . . . . . . . . . . . . . . . . . . . . . 733 4.1. The Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 733 4.2. Differences in Guild Structures of Paleozoic and
Neogene Communities. . . . . . . . . . . . . . . . . . . . . . . 733 4.3. Similarities in Species Distribution within
Guilds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 736 4.4. "Superguilds" . . . . . . . . . . . . . . . . . . . . . . . . . . . . 740
5. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 742 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 44
Chapter 16 • Soft-Bottom Epifaunal Suspension-Feeding Assemblages in the Late Cretaceous: Implications for the Evolution of Benthic Paleocommunities
David Jablonski and David J. Bottjer
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 747 2. Late Cretaceous Offshore Benthic Assemblages. . . . . . . . . 749
2.1. Gulf and Atlantic Coastal Plain. . . . . . . . . . . . . . . . 749 2.2. Other Chalk Faunas . . . . . . . . . . . . . . . . . . . . . . . . 762
3. Late Cretaceous Nearshore Benthic Assemblages. . . . . . . . 771 3.1. Gulf and Atlantic Coastal Plain................ 771 3.2. Other Nearshore Faunas . . . . . . . . . . . . . . . . . . . . . 775
4. Structure of Late Cretaceous Assemblages. . . . . . . . . . . . . 776 4.1. The Ecologic Pattern . . . . . . . . . . . . . . . . . . . . . . . . 776 4.2. The Taphonomic Overprint. . . . . . . . . . . . . . . . . . . 778
5. Evolutionary History and Mechanisms . . . . . . . . . . . . . . . 781 5.1. Evolutionary History . . . . . . . . . . . . . . . . . . . . . . . 781 5.2. Evolutionary Mechanisms . . . . . . . . . . . . . . . . . . . 787
6. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 793 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 795
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 797