, Evolutionary
Analysis FIFTH EDITION
Jon C. Herron
University of Washington
Scott Freeman
University of Washington
With contributions by
Jason Hodin
Hopkins Marine Station
of Stanford University
and University of Washington
Brooks Miner
Cornell University
Christian Sidor
University of Washington
Boston Columbus Indianapolis New York San Francisco Upper Saddle River
Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montreal Toronto
Delhi Mexico City Sao Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo
,Editor-in-Chief: Beth Wilbur Copyeditor: Chris Thillen
Senior Acquisitions Editor: Michael Gillespie Design Manager: Mark Ong
Executive Director of Development: Deborah Gale Cover and Interior Designer: Mark Ong
Project Editor: Laura Murray Art Developer, Illustrator: Robin Green
Assistant Editor: Eddie Lee Senior Photo Editor: Travis Amos
Manager, Text Permissions: Tim Nicholls Photo Research and Permissions Management:
Text Permissions Specialist: Kim Schmidt, S4Carlisle Bill Smith Group
Publishing Services Content Producer: Daniel Ross
Director of Production: Erin Gregg Media Project Manager: Shannon Kong
Managing Editor: Michael Early Director of Marketing: Christy Lesko
Production Project Manager: Lori Newman Executive Marketing Manager: Lauren Harp
Production Management Services: Progressive Manufacturing Buyer: Christy Hall
Publishing Alternatives Cover and Text Printer: Courier Kendalville
Cover Photo Credit: Auke Holwerda / Getty Images
Credits and acknowledgments for materials borrowed from other sources and reproduced, with
permission, in this textbook appear on the appropriate page within the text [or on p. 822].
Copyright ©2014, 2007, 2004, 2001, 1998 by Jon C. Herron and Scott Freeman. Published by Pearson
Education, Inc. All rights reserved. Manufactured in the United States of America. This publication is
protected by Copyright, and permission should be obtained from the publisher prior to any prohibited
reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic,
mechanical, photocopying, recording, or likewise. To obtain permission(s) to use material from this
work, please submit a written request to Pearson Education, Inc., Permissions Department, 1900
E. Lake Ave., Glenview, IL 60025. For information regarding permissions, call (847) 486-2635.
Readers may view, browse, and/or download material for temporary copying purposes only, provided
these uses are for noncommercial personal purposes. Except as provided by law, this material may not
be further reproduced, distributed, transmitted, modified, adapted, performed, displayed, published, or
sold in whole or in part, without prior written permission from the publisher.
Many of the designations used by manufacturers and sellers to distinguish their products are claimed as
trademarks. Where those designations appear in this book, and the publisher was aware of a trademark
claim, the designations have been printed in initial caps or all caps.
Library of Congress Cataloging-in-Publication Data is available upon request.
ISBN 10: 0-321-61667-7; ISBN 13: 978-0-321-61667-8 (Student Edition)
ISBN 10: 0-321-92799-0; ISBN 13: 0-321-92799-6 (Instructor’s Review Copy)
ISBN 10: 0-321-92816-4; ISBN 13: 978-0-321-92816-0 (Books a la Carte Edition)
1 2 3 4 5 6 7 8 9 10—CRK—17 16 15 14 13
Library of Congress Control Number: 2013943730
www.pearsonhighered.com
, Brief Contents
PART 1 PART 3
INTRODUCTION 1 ADAPTATION 369
CHAPTER 1 A Case for Evolutionary CHAPTER 10 Studying Adaptation:
Thinking: Understanding HIV 1 Evolutionary Analysis of
Form and Function 369
CHAPTER 2 The Pattern of Evolution 37
CHAPTER 11 Sexual Selection 407
CHAPTER 3 Evolution by Natural Selection 73
CHAPTER 12 The Evolution of Social Behavior 455
CHAPTER 4 Estimating Evolutionary Trees 109
CHAPTER 13 Aging and Other Life-History
Characters 491
CHAPTER 14 Evolution and Human Health 535
PART 2
MECHANISMS OF CHAPTER 15 Genome Evolution and the
EVOLUTIONARY CHANGE 147 Molecular Basis of Adaptation 581
CHAPTER 5 Variation Among Individuals 147
CHAPTER 6 Mendelian Genetics in PART 4
Populations I: Selection and THE HISTORY OF LIFE 609
Mutation 179
CHAPTER 7 Mendelian Genetics in CHAPTER 16 Mechanisms of Speciation 609
Populations II: Migration, Drift, CHAPTER 17 The Origins of Life
and Nonrandom Mating 233 and Precambrian Evolution 645
CHAPTER 8 Evolution at Multiple Loci: CHAPTER 18 Evolution and the Fossil Record 691
Linkage and Sex 291
CHAPTER 19 Development and Evolution 735
CHAPTER 9 Evolution at Multiple Loci:
Quantitative Genetics 329 CHAPTER 20 Human Evolution 769
iii
Analysis FIFTH EDITION
Jon C. Herron
University of Washington
Scott Freeman
University of Washington
With contributions by
Jason Hodin
Hopkins Marine Station
of Stanford University
and University of Washington
Brooks Miner
Cornell University
Christian Sidor
University of Washington
Boston Columbus Indianapolis New York San Francisco Upper Saddle River
Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montreal Toronto
Delhi Mexico City Sao Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo
,Editor-in-Chief: Beth Wilbur Copyeditor: Chris Thillen
Senior Acquisitions Editor: Michael Gillespie Design Manager: Mark Ong
Executive Director of Development: Deborah Gale Cover and Interior Designer: Mark Ong
Project Editor: Laura Murray Art Developer, Illustrator: Robin Green
Assistant Editor: Eddie Lee Senior Photo Editor: Travis Amos
Manager, Text Permissions: Tim Nicholls Photo Research and Permissions Management:
Text Permissions Specialist: Kim Schmidt, S4Carlisle Bill Smith Group
Publishing Services Content Producer: Daniel Ross
Director of Production: Erin Gregg Media Project Manager: Shannon Kong
Managing Editor: Michael Early Director of Marketing: Christy Lesko
Production Project Manager: Lori Newman Executive Marketing Manager: Lauren Harp
Production Management Services: Progressive Manufacturing Buyer: Christy Hall
Publishing Alternatives Cover and Text Printer: Courier Kendalville
Cover Photo Credit: Auke Holwerda / Getty Images
Credits and acknowledgments for materials borrowed from other sources and reproduced, with
permission, in this textbook appear on the appropriate page within the text [or on p. 822].
Copyright ©2014, 2007, 2004, 2001, 1998 by Jon C. Herron and Scott Freeman. Published by Pearson
Education, Inc. All rights reserved. Manufactured in the United States of America. This publication is
protected by Copyright, and permission should be obtained from the publisher prior to any prohibited
reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic,
mechanical, photocopying, recording, or likewise. To obtain permission(s) to use material from this
work, please submit a written request to Pearson Education, Inc., Permissions Department, 1900
E. Lake Ave., Glenview, IL 60025. For information regarding permissions, call (847) 486-2635.
Readers may view, browse, and/or download material for temporary copying purposes only, provided
these uses are for noncommercial personal purposes. Except as provided by law, this material may not
be further reproduced, distributed, transmitted, modified, adapted, performed, displayed, published, or
sold in whole or in part, without prior written permission from the publisher.
Many of the designations used by manufacturers and sellers to distinguish their products are claimed as
trademarks. Where those designations appear in this book, and the publisher was aware of a trademark
claim, the designations have been printed in initial caps or all caps.
Library of Congress Cataloging-in-Publication Data is available upon request.
ISBN 10: 0-321-61667-7; ISBN 13: 978-0-321-61667-8 (Student Edition)
ISBN 10: 0-321-92799-0; ISBN 13: 0-321-92799-6 (Instructor’s Review Copy)
ISBN 10: 0-321-92816-4; ISBN 13: 978-0-321-92816-0 (Books a la Carte Edition)
1 2 3 4 5 6 7 8 9 10—CRK—17 16 15 14 13
Library of Congress Control Number: 2013943730
www.pearsonhighered.com
, Brief Contents
PART 1 PART 3
INTRODUCTION 1 ADAPTATION 369
CHAPTER 1 A Case for Evolutionary CHAPTER 10 Studying Adaptation:
Thinking: Understanding HIV 1 Evolutionary Analysis of
Form and Function 369
CHAPTER 2 The Pattern of Evolution 37
CHAPTER 11 Sexual Selection 407
CHAPTER 3 Evolution by Natural Selection 73
CHAPTER 12 The Evolution of Social Behavior 455
CHAPTER 4 Estimating Evolutionary Trees 109
CHAPTER 13 Aging and Other Life-History
Characters 491
CHAPTER 14 Evolution and Human Health 535
PART 2
MECHANISMS OF CHAPTER 15 Genome Evolution and the
EVOLUTIONARY CHANGE 147 Molecular Basis of Adaptation 581
CHAPTER 5 Variation Among Individuals 147
CHAPTER 6 Mendelian Genetics in PART 4
Populations I: Selection and THE HISTORY OF LIFE 609
Mutation 179
CHAPTER 7 Mendelian Genetics in CHAPTER 16 Mechanisms of Speciation 609
Populations II: Migration, Drift, CHAPTER 17 The Origins of Life
and Nonrandom Mating 233 and Precambrian Evolution 645
CHAPTER 8 Evolution at Multiple Loci: CHAPTER 18 Evolution and the Fossil Record 691
Linkage and Sex 291
CHAPTER 19 Development and Evolution 735
CHAPTER 9 Evolution at Multiple Loci:
Quantitative Genetics 329 CHAPTER 20 Human Evolution 769
iii
