Evolutionary Biology VOLUME 31

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1 Evolutionary Biology VOLUME 31

2 A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon publication. Volumes are billed only upon actual shipment. For further information please contact the publisher.

3 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

4 The Library of Congress catalogued the first volume of this title as follows: Evolutionary biology, v New York, Appleton-Century-Crofts. v. illus., 24 cm annual. Editors: T. Dobzhansky and others. 1. Evolution Period. 2. Biology Period. I. Dobzhanksy, Theodosius Grigorievich, QH366.A1E9 575' ISBN ISBN (ebook) DOI / Springer Science+Business Media New York Originally published by Kluwer Academic/Plenum Publishers in 2000 Softcover reprint of the hardcover 1st edition 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

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

6 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 vii

7 Contents 1. Speciation Processes in the Adaptive Radiation of Hawaiian Plants and Animals Elysse M. Craddock Introduction The Geology of Hawaii and Biological Consequences... 3 Speciation in Hawaiian Plants The Silversword Radiation... 6 Radiation of the Genus Bidens Radiation of the Lobelioid Genus Cyanea... 9 Radiation of the Alsinoideae Speciation in Hawaiian Animals Speciation in Hawaiian Honeycreepers Speciation in Hawaiian Crickets Speciation in Hawaiian Planthoppers Speciation in Hawaiian Drosophila Speciation Processes in Hawaii The Role of Ecological Factors in Hawaiian Speciation The Role of Sexual Selection The Role of Founder Events Summary Acknowledgments References Codon Bias and the Context Dependency of Nucleotide Substitutions in the Evolution of Plastid DNA Brian R. Morton Introduction The Plastid Genome Codon Bias ix

8 x Contents Codon Bias in Unicellular Organisms Codon Usage Patterns in the Plastid Genome Selection on Codon Usage in Different Plastid Genomes Codon Bias of Euglena Chloroplast Genes Codon Bias and Expression Level Maintenance of Codon Bias Codon Usage of the Plant psba Gene Evolution of Selection on Codon Usage Mutation Dynamics Models for Nucleotide Substitution Neighboring Base Composition and Substitution Bias Context and Synonymous Substitution Rate Distribution of Synonymous Substitutions in rbel Context and Synonymous Substitution Rate of Chloroplast Genes Context Dependency in Lineages Other than the Angiosperms Conclusions Acknowledgments... _, References The Origin of the Mineral Skeleton in Chordates lerzy Dzik Introduction Architecture of Conodont Apparatuses Function of Conodont Apparatuses Growth and Function of Conodont Elements... '" Conodont Element Morphogenesis Relationships of Yunnanozoon Paraconodonts and the Evolutionary Origin of the Conodont Apparatus Dickinsonia and the Ancestry of Chordates The Problem of Protoconodonts Conclusions Acknowledgments References

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

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