Evolution PDF
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Evolution EvoLUtion Carl T. Bergstrom University of Washington Lee Alan Dugatkin University of Louisville B W. W. NortoN & CompaNy NeW york • LoNdoN W. W. Norton & Company has been independent since its founding in 1923, when Wil- liam Warder Norton and Mary D. Herter Norton first published lectures delivered at the People’s Institute, the adult education division of New York City’s Cooper Union. The Nortons soon expanded their program beyond the Institute, publishing books by celebrated academ- ics from America and abroad. By mid-century, the two major pillars of Norton’s publishing program—trade books and college texts—were firmly established. In the 1950s, the Norton family transferred control of the company to its employees, and today—with a staff of four hundred and a comparable number of trade, college, and professional titles published each year—W. W. Norton & Company stands as the largest and oldest publishing house owned wholly by its employees. Copyright © 2012 by W. W. Norton & Company, Inc. All rights reserved Printed in the United States of America First Edition Editor: Betsy Twitchell Developmental editors: Sandy Lifland, Andrew Sobel Project editor: Carla L. Talmadge Editorial assistant: Cait Callahan Production manager: Chris Granville Managing editor, College: Marian Johnson Design director: Rubina Yeh Book designer: Lissi Sigillo Photo editor: Stephanie Romeo Photo researcher: Dena Digilio Betz Media editor: Patrick Shriner Associate editor, emedia: Callinda Taylor Media assistant: Carson Russell Marketing manager: John Kresse Composition: Preparé, Inc. Illustration studio: Precision Graphics Manufacturing: Quad Graphics Library of Congress Cataloging-in-Publication Data Bergstrom, Carl T. Evolution / Carl T. Bergstrom, Lee Alan Dugatkin.—1st ed. p. cm. Includes bibliographical references and index. ISBN 978-0-393-92592-0 (hardcover)—ISBN 978-0-393-91341-5 (pbk.) 1. Evolution (Biology) I. Dugatkin, Lee Alan, 1962– II. Title. QH366.2.B483 2012 576.8—dc23 2011036572 ISBN: 978-0-393-11941-1 (pdf ebook) W. W. Norton & Company, Inc., 500 Fifth Avenue, New York, NY 10110 www.wwnorton.com W. W. Norton & Company Ltd., Castle House, 75/76 Wells Street, London W1T 3QT 1 2 3 4 5 6 7 8 9 0 BriEF ContEntS part I Foundations of Evolutionary Biology 1 An overview of Evolutionary Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 Early Evolutionary ideas and Darwin’s insight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 3 natural Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 4 Phylogeny and Evolutionary History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 5 inferring Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 part II Evolutionary Genetics 6 transmission Genetics and the Sources of Genetic variation . . . . . . . . . . . . . . . . . . . . . . . 178 7 the Genetics of Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202 8 Evolution in Finite Populations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242 9 Evolution at Multiple loci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 290 10 Genome Evolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334 part III The History of Life 11 the origin and Evolution of Early life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378 12 Major transitions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 404 13 Evolution and Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432 14 Species and Speciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 454 15 Extinction and Evolutionary trends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 498 part IV Evolutionary Interactions 16 the Evolution of Sex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 538 17 Sexual Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 562 18 the Evolution of Sociality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 584 19 Coevolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 620 20 Evolution and Medicine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646 v ContEntS about the authors xix preface xxi part I Foundations of Evolutionary Biology CHapter 1 An Overview of Evolutionary Biology 2 1.1 a Brief Introduction to evolution and Natural Selection 5 Evolutionary Change and the Food We Eat 6 Evolutionary Change and Pharmaceuticals 8 Evolution and Conservation Biology 11 1.2 empirical and theoretical approaches to the Study of evolution 14 Empirical Approaches 14 theoretical Approaches 20 Box 1.1 A Mathematical Model of the Sex Ratio 23 theory and Experiment 24 Summary 25 key terms 25 review Questions 26 Suggested readings 27 CHapter 2 Early Evolutionary Ideas and Darwin’s Insight 28 2.1 the Nature of Science: Natural versus Supernatural explanations 31 Methodological naturalism 31 Hypothesis testing and logic 31 2.2 time and a Changing World 32 2.3 the origins of Life and Its diversity 35 2.4 organisms are Well-Suited to their environment 38 Paley’s natural theology 38 Jean-Baptiste lamarck and the inheritance of Acquired Characteristics 38 Patrick Matthew and natural Selection 39 2.5 darwin’s theory 41 Darwin’s two Fundamental insights 41 vii viii Contents Publication of On the Origin of Species 41 Means of Modification and Pigeon Breeding 43 Artificial Selection 44 Changing Species 45 2.6 darwin on Natural Selection 45 Darwin, variation, and Examples of natural Selection 46 the Power of natural Selection 47 Malthus and the Scope of Selection 47 transformational and variational Processes of Evolution 48 2.7 darwin on Common ancestry 50 the tree of life 50 Groups within Groups 52 Common Descent and Biogeography 53 2.8 problems with darwin’s theory 54 Problem 1: Accounting for Complex Structures with Multiple intricate Parts 54 Problem 2: Explaining traits and organs of Seemingly little importance 54 Problem 3: Why Doesn’t natural Selection run out of variation to Sort on? 55 2.9 the reaction to darwin and early History of the modern Synthesis 55 Summary 57 key terms 58 review Questions 58 Suggested readings 59 CHapter 3 Natural Selection 60 3.1 the Components of Natural Selection 63 natural Selection and Coat Color in the oldfield Mouse 66 3.2 adaptations and Fit to environment 72 Defining Adaptation 72 Adaptations and Fit to Environment 72 3.3 Natural Selection in the Field 74 3.4 Natural Selection in the Laboratory 78 lenski’s long-term Evolution Experiment 79 Box 3.1 Measuring Allele Frequencies and Fitnesses in E. coli 81 3.5 Constraints on What Natural Selection Can achieve 83 Physical Constraints 84 Evolutionary Arms races 88 natural Selection lacks Foresight 88 3.6 origin of Complex traits 89 intermediate Stages with Function Similar to Modern Function 90 novel Structures and Exaptations 92 novelty at the Molecular level 95 Summary 97 key terms 98 review Questions 98 Suggested readings 99
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