Evolutionary Biology VOLUME 31

Evolutionary Biology VOLUME 31

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Evolutionary Biology VOLUME 31 Edited by MAX K. HECHT Queens College of the City University of New York Flushing, New York ROSS J. MACINTYRE Cornell University Ithaca, New York and MICHAEL T. CLEGG University of California, Riverside Riverside, California SPRINGER SCIENCE+BUSINESS MEDIA, LLC

The Library of Congress catalogued the first volume of this title as follows: Evolutionary biology, v. 1-1967- New York, Appleton-Century-Crofts. v. illus., 24 cm annual. Editors: 1967- T. Dobzhansky and others. 1. Evolution Period. 2. Biology Period. I. Dobzhanksy, Theodosius Grigorievich, 1900- QH366.A1E9 575'.005 67-11961 ISBN 978-1-4613-6877-9 ISBN 978-1-4615-4185-1 (ebook) DOI 10.1007/978-1-4615-4185-1 2000 Springer Science+Business Media New York Originally published by Kluwer Academic/Plenum Publishers in 2000 Softcover reprint of the hardcover 1st edition 2000 10 9 8 7 6 5 4 3 2 A CLP. record for this book is available from the Library of Congress All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher

Contributors Elysse M. Craddock Division of Natural Sciences, Purchase College, State University of New York, Purchase, New York 10577-1400 Jerzy Dzik Institute of Paleobiology PAN, 00-818 Warsaw, Poland Brian R. Morton Department of Biological Sciences, Barnard College, Columbia University, New York, New York 10027 Kurt Schwenk Department of Ecology and Evolutionary Biology, UlJ,iversity of Connecticut, Storrs, Connecticut 06269-3043 GOOter P. Wagner. Department of Ecology and Evolutionary Biology and Center for Computational Ecology, Yale University, New Haven, Connecticut 06520-8106 v

Preface This volume is the thirty-second in this series, which includes thirty-one numbered volumes and one unnumbered supplement. The editors continue to focus on critical reviews, commentaries, original papers, and controversies in evolutionary biology. The topics of the review range from developmental biology to paleobiology. Recent volumes have included a broad spectrum of chapters on such subjects as molecular phylogenetics, homology and embryonic development, and paleobiological and developmental aspects of limb development. The editors continue to solicit manuscripts in all areas of evolutionary biology. Manuscripts should be sent to anyone of the following: Max K. Hecht, Department of Biology, Queens College of CUNY, Flushing, New York 11367; Ross 1. MacIntyre, Department of Genetics and Development, Cornell University, Ithaca, New York 14853; or Michael T. Clegg, Department of Botany and Plant Sciences, University of California, Riverside, California 92521. vii

Contents 1. Speciation Processes in the Adaptive Radiation of Hawaiian Plants and Animals Elysse M. Craddock Introduction............................................. 1 The Geology of Hawaii and Biological Consequences... 3 Speciation in Hawaiian Plants.............................. 6 The Silversword Radiation... 6 Radiation of the Genus Bidens........................... 8 Radiation of the Lobelioid Genus Cyanea... 9 Radiation of the Alsinoideae... 11 Speciation in Hawaiian Animals............................ 13 Speciation in Hawaiian Honeycreepers... 14 Speciation in Hawaiian Crickets.......................... 17 Speciation in Hawaiian Planthoppers... 20 Speciation in Hawaiian Drosophila... 24 Speciation Processes in Hawaii............................. 36 The Role of Ecological Factors in Hawaiian Speciation...... 36 The Role of Sexual Selection... 39 The Role of Founder Events... 42 Summary... 44 Acknowledgments... 44 References... 44 2. Codon Bias and the Context Dependency of Nucleotide Substitutions in the Evolution of Plastid DNA Brian R. Morton Introduction.............................................. 55 The Plastid Genome...................................... 56 Codon Bias... 59 ix

x Contents Codon Bias in Unicellular Organisms..................... 59 Codon Usage Patterns in the Plastid Genome.............. 61 Selection on Codon Usage in Different Plastid Genomes... 68 Codon Bias of Euglena Chloroplast Genes................. 72 Codon Bias and Expression Level........................ 73 Maintenance of Codon Bias... 75 Codon Usage of the Plant psba Gene.................... 79 Evolution of Selection on Codon Usage................... 82 Mutation Dynamics....................................... 84 Models for Nucleotide Substitution....................... 84 Neighboring Base Composition and Substitution Bias....... 85 Context and Synonymous Substitution Rate... 90 Distribution of Synonymous Substitutions in rbel... 92 Context and Synonymous Substitution Rate of Chloroplast Genes... 95 Context Dependency in Lineages Other than the Angiosperms... 96 Conclusions... 98 Acknowledgments... _,........... 98 References... 99 3. The Origin of the Mineral Skeleton in Chordates lerzy Dzik Introduction............................................. 105 Architecture of Conodont Apparatuses... 107 Function of Conodont Apparatuses......................... 112 Growth and Function of Conodont Elements... '".... 113 Conodont Element Morphogenesis... 122 Relationships of Yunnanozoon... 128 Paraconodonts and the Evolutionary Origin of the Conodont Apparatus........................................... 133 Dickinsonia and the Ancestry of Chordates.................. 137 The Problem of Protoconodonts... 141 Conclusions... 144 Acknowledgments... 146 References... 146

Contents xi 4. Evolutionarily Stable Configurations: Functional Integration and the Evolution of Phenotypic Stability Gunter P. Wagner and Kurt Schwenk Introduction............................................. 155 The Evolutionarily Stable Configuration: Overview............ 157 Case-Study: Lingual Feeding in Iguanian Lizards... 161 The ESC: Characterization and Postulates... 175 Postulate 1: Mutational Connectedness.................... 177 Postulate 2: Functional Uniqueness...... 177 Postulate 3: Functional Integration........................ 178 Postulate 4: Trade-offs in Performing Other Functions....... 178 Postulate 5: Functional Closure........................... 178 Origin of an ESC......................................... 179 Escaping from an ESC.................................... 180 Overriding Selection Pressures and Pleiotropic Effects... 181 Redundancy in Performing the Proper Function............ 183 Environmental Change and Modification of Internal Optimality Rules................................... 185 Character Analysis, Phylogenetic Patterns, and Comparative Methods... 188 Phylogenetic Signature of an ESC........................ 188 Patterns of Character Evolution within an ESC... 193 Phylogenetic Conclusions................................ 194 A Brief Review of Relevant Concepts: Functional Integration and Internal Selection... 196 Developmental and Functional Constraint................... 203 Problems................................................ 205 Some General Implications of the ESC Concept.............. 206 Acknowledgments... 207 Appendix 1: Millikan's (1984) "Proper Function"............. 207 Appendix 2: Character Evolution within an ESC... 208 No Phylogenetic Signal... 208 Phylogenetic Signal... 209 References... 211 Index... 219