N J a u m “The publication of James and Nunnallee, or ‘the Daves’ as we know them, is n e n a matter for unreserved celebration, not only for lepidopterists and nature s a l lovers of all stripes, but for anyone who cares about our butterflies’ lives, a l n futures, conservation management, and the plants with which they have e d e co-evolved.” ROBERT MICHAEL PYLE , author of Mariposa Road: The First Butterfly Big Year L i f e H i s t Life o r i e s Histories Life Histories of Cascadia Butterflies presents, for the first time in North o America, the life stages of the entire butterfly fauna of a geographic region. f In exceptional and riveting detail, this authoritative field guide describes C and illustrates the life histories of the butterflies of the Pacific Northwest. a s of Virtually all of the 158 butterfly species occurring in southern British c Columbia, Washington, northern Idaho, and northern Oregon are included a in the book. Color photographs of each stage of life—egg, every larval instar, d pupa, adult—accompany information on the biology, ecology, and rearing of i each species. a Cascadia B Life Histories of Cascadia Butterflies is an unparalleled resource on the nat - u ural history of immature stages of butterflies in the Pacific Northwest—and t beyond, as many of Cascadia’s butterflies occur in other parts of North t Butterflies America as well as Europe and Asia. It provides interested citizens with a e basic understanding of the region’s rich butterfly life and serves as an r f invaluable reference for scientists and lepidopterists. l i e s Oregon State University Press ISBN 978-0-87071-626-3 Cover photographs by David G. James and David Nunnallee Cover images depict life stages of Euphydryas colon 9 7 8 0 8 7 0 7 1 6 2 6 3 OSU Cover design by David Drummond PRESS David G. James and David Nunnallee Life Histories of Cascadia Butterfl ies David G. James and David Nunnallee For more information or to purchase the book, visit http://osupress.oregonstate.edu/book/life-histories-of-cascadia-butterflies Oregon State University Press Corvallis Cascadia Butterflies.indb 3 8/12/11 4:14 PM The paper in this book meets the guidelines for permanence and durability of the Committee on Production Guidelines for Book Longevity of the Council on Library Resources and the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48-1984. Library of Congress Cataloging-in-Publication Data James, David G. Life histories of Cascadia butterflies / David G. James, David Nunnallee ; foreword by Robert Michael Pyle. p. cm. Includes bibliographical references and index. ISBN 978-0-87071-626-3 (alk. paper) -- ISBN 978-0-87071-648-5 (ebook) 1. Butterflies--Washington (State) 2. Butterflies--Northwest, Pacific. 3. Butterflies--Washington (State)--Pictorial works. 4. Butterflies--Northwest, Pacific--Pictorial works. I. Nunnallee, D. II. Pyle, Robert Michael. III. Title. QL551.W2J36 2011 595.78’909795--dc23 2011020404 © 2011 David G. James and David Nunnallee All rights reserved. First published in 2011 by Oregon State University Press Printed in China Oregon State University Press 121 The Valley Library Corvallis, OR 97331-4501 541-737-3166 • fax 541-737-3170 http://oregonstate.edu/dept/press Cascadia Butterflies.indb 4 8/12/11 4:14 PM Dedication To my father, Alan James, and mother, Doreen James, for supporting and encouraging my interest in butterflies as an 8-year-old, which led to a lifelong passion for insects and their biology. Also to my entomologist wife, Tanya, and our gorgeous daughters, Jasmine Vanessa and Rhiannon Vanessa, for making this endeavor a truly enjoyable and family affair! —David G. James To my wife, Jo, for her unwavering support, companionship, and assistance, both in the field and at home. —Dave Nunnallee Cascadia Butterflies.indb 5 8/12/11 4:14 PM Table of Contents Foreword by Robert Michael Pyle 8 Introduction 11 The Life History of a Butterfly 13 Life Strategies 18 Overwintering, Oversummering, and Diapause 23 Natural Enemies 28 Defense 34 Variation 41 Butterfly Habitats in Cascadia 44 Fieldwork 49 Rearing Techniques 53 Photographic Techniques 58 How to Use This Book 59 Quick Photo Guides to Eggs, Larvae, and Pupae 62 Cascadia Butterflies.indb 6 8/12/11 4:14 PM Species Accounts 68 Parnassians and Swallowtails 68 Whites and Sulphurs 86 Coppers 124 Hairstreaks 140 Blues and Metalmark 182 Fritillaries 228 Checkerspots and Crescents 266 Other Brushfoots, Admirals, and Monarch 288 Satyrs 324 Spreadwing Skippers 342 Monocot (Grass) Skippers 366 Glossary 404 Bibliography 409 Photo Credits and Data 419 Acknowledgments 430 Index 432 Cascadia Butterflies.indb 7 8/12/11 4:14 PM 11 Introduction Th is book is the fulfi llment of the authors’ long-time dream to document all the life stages of all the butterfl ies of a signifi cant geographic region, that of Cascadia. For David James the dream of comprehensively documenting butterfl y life histories extends back to the 1960s in England, when he fi rst began rearing butterfl ies—at the age of 8! We defi ne Cascadia for the purposes of this book as Washington State and the adjacent areas of northern Oregon, southern British Columbia, and the Idaho panhandle. Our coverage is virtually complete for Washington and nearly complete for these adjacent areas. Th is work is the result of approximately twenty years of combined eff ort by the two authors to produce the information and photographs for this book. Initially both authors began this project independently, working separately and unbeknownst to each other for several years. Th e authors met early in 2005 and, recognizing that they had very similar goals, pooled their resources Area of coverage. and eff orts and have been working together ever since. David James is an associate professor in the Department of Entomology at Washington State University, Prosser, located in the Yakima Valley of eastern Washington. David Nunnallee is an engineer by occupation and a naturalist by avocation who resides in western Washington State near Seattle. Life Histories of Cascadia Butterfl ies is a book about the immature stages—the eggs, larvae, and pupae—of the butterfl ies of the Pacifi c Northwest. Th e vast majority of the photographs and information were obtained by diligent and, usually, multiple rearings of individual species. Th is book is unique in the Americas in describing and illustrating with high-quality photographs all the immature stages of virtually all the butterfl y species within a substantial geographic area. While there are other American publications that deal with butterfl y larvae, there is no other publication in the New World, or to our knowledge in the Western world, with such a complete scope. Other publications either have complete life histories for a few butterfl y species, or cover only the fi nal instar (stage) of caterpillars for a wider selection of species; however, because many butterfl y larvae change radically during their development, an illustration of only the fi nal stage is oft en inadequate for identifi cation or comparative studies. Several converging factors underscore the need for a book of this kind. First, there has been a great surge of public interest in butterfl ies over the past decade or so. As members of the birding community have expanded their interests in recent years to butterfl ies, native plants, and dragonfl ies, they continue to search for interesting new pursuits, not to replace their natural history interests but to expand them. Th e study of butterfl y larvae is a “natural” for this audience. Now that many people are becoming comfortable with identifi cation of adult butterfl ies, they are ready to move on to Cascadia Butterflies.indb 11 8/12/11 4:14 PM 12 LIFE HISTORIES OF CASCADIA BUTTERFLIES immature stages. They want to be able to identify caterpillars they find, to locate larvae to rear with their children, or simply to expand their knowledge of the natural world. In addition, growing numbers of gardeners and horticulturists want not only to attract butterflies but also to provide a complete life history habitat for them and to be able to identify the larvae they find in their gardens. There is also a great deal of interest in the immature stages of butterflies within the scientific community as a taxonomic Youngsters are our future. Rhiannon James tool for clarifying species relationships, for enjoys her “butterfly nose.“ comparison of geographically separated populations, and as indicators of habitat degradation and climate change. Larvae are currently recognized, perhaps even more now than in earlier decades, for their value in differentiating butterfly species. In this book we hope that the considerable new information we present will help solve longtime problems in butterfly taxonomy and species identification. Professional and amateur entomologists are increasingly interested in the biology and ecology of butterflies because they provide excellent subjects for laboratory and field research. Butterflies are dependent on their environment, most particularly on plants, and as human encroachment compromises their environments, many butterfly species are struggling to survive. As a result, in Cascadia several butterfly species have been listed as endangered within the past few years, and still others have been placed on candidate species lists. Such listings typically require recovery plans that in turn may include captive rearing programs. State agencies, zoos, universities, and conservation organizations are currently cooperating to rear some of the listed species for reintroduction to the wild. We cannot protect what we do not understand. We hope this book will increase our understanding of butterfly life histories and that this will lead to more effective preservation programs.