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MOTHS of WESTERN NORTH AMERICA UCPB004-3960G-FM Pi-Xiv.Qxd 30/1/09 7:24 Am Page Ii UCPB004-3960G-FM Pi-Xiv.Qxd UCPB004-3960G-FM_pi-xiv.qxd 30/1/09 7:24 am Page i UCPB004-3960G-FM_pi-xiv.qxd MOTHS OF WESTERN NORTH AMERICA UCPB004-3960G-FM_pi-xiv.qxd 30/1/09 7:24 am Page ii UCPB004-3960G-FM_pi-xiv.qxd This page intentionally left blank UCPB004-3960G-FM_pi-xiv.qxd 30/1/09 7:24 am Page iii UCPB004-3960G-FM_pi-xiv.qxd Moths of Western North America JERRY A. POWELL PAUL A. OPLER UNIVERSITY OF CALIFORNIA PRESS Berkeley Los Angeles London UCPB004-3960G-FM_pi-xiv.qxd 1/30/09 9:03 PM Page iv Aptara University of California Press, one of the most distinguished univer- Library of Congress Cataloging-in-Publication Data sity presses in the United States, enriches lives around the world by advancing scholarship in the humanities, social sciences, and natural Powell, Jerry A. sciences. Its activities are supported by the UC Press Foundation and Moths of western North America / Jerry A. Powell, Paul A. Opler. by philanthropic contributions from individuals and institutions. For p. cm. more information, visit www.ucpress.edu. Includes bibliographical references and index. ISBN 978-0-520-25197-7 (case : alk. paper) 1. Moths—West (U.S.) University of California Press 2. Moths—Northwest, Canadian. I. Opler, Paul A. II. Title. Berkeley and Los Angeles, California QL548.P69 2009 595.780970—dc22 University of California Press, Ltd. 2008048605 London, England Manufactured in China. © 2009 by The Regents of the University of California 17 16 15 14 13 12 11 10 09 10987654321 Photo/image credits: T. Arcand: 61.25–61.27, 61.31, 61.34, 62.10, 62.20, 62.23, 62.24, 63.12, 63.31, 63.36, 64.1, 64.24–64.26, 64.29. The paper used in this publication meets the minimum requirements E. Buckner-Opler: 63.3, 63.5, 63.21, 63.28–63.30. California Depart- of ANSI/NISO Z 39.48-1992 (R 1997)(Permanence of Paper). ment of Food & Agriculture: 61.6. Canadian National Collection (CNC): 61.28, 61.32, 62.13, 62.25, 62.32, 63.19, 63.20, 63.26, 63.27, 63.33, 63.35, 64.3–64.6, 64.9, 64.13, 64.20–64.23, 64.27, 64.28, Cover: Arctia caja (Linnaeus), California, Sierra Co., San Francisco 64.32–64.34. R. Coville: Last page color section, 59.35, 60.4, 60.5, State U., Sierra Nevada Field Campus, July 2007. Species is also shown 60.7, 60.17, 60.21–60.24, 60.27, 61.2. H. V. Daly: 60.1. L. Dyer: 61.7. in plate 47.8, which is from Elko Co., Nevada. The Sierra Nevada D. C. Ferguson: 61.11, 61.29, 61.30, 61.33, 61.36, 62.2, 62.3, population moths differ in color and pattern from the Rocky Moun- 62.5–62.8, 62.11, 62.14, 62.16, 62.26, 62.27, 62.32, 62.34, 64.7, 64.8, tain and Basin Ranges form. Photograph by Rollin Coville. 64.12, 64.14–64.16. C. Hansen: 61.13. P. G. Johnson II: First page color section, 59.2, 60.6, 60.9, 60.19, 60.30, 62.34, 63.22–63.25. N. McFarland: 61.9. L. Minor-Penland: 61.13. P. A. Opler: 61.23, 63.8, 63.9, 64.10, 64.11. S. Passoa: 64.19. D. J. Powell: 59.25, 59.33, 60.10, 60.15, 61.4, 61.5, 61.14, 61.15–61.22. J. A. Powell: 59.3, 59.4–59.22, 59.26–59.32, 59.34, 59.36, 60.2, 60.3, 60.8, 60.11–60.16, 60.18, 60.20, 60.25, 60.26, 60.28, 60.29, 60.31–60.36, 61.1, 61.3, 61.24, 62.33, 63.2, 64.17, 64.18, 64.35. E. S. Ross 59.23. B. Scaccia 61.35, 62.1, 62.4, 62.9, 62.12, 62.15, 62.17–62.19, 62.21, 62.22. D. L. Wagner 59.1, 61.8, 61.10. B. Walsh 62.28–62.31, 62.35, 62.36, 63.1, 63.4, 63.6, 63.7, 63.10, 63.11, 63.13–63.18, 64.2, 64.36. UCPB004-3960G-FM_pi-xiv.qxd 3/10/09 1:33 AM Page v Aptara CONTENTS LIST OF FIGURES / vii PART ONE Apoditrysia / 115 PREFACE / ix Introduction to Lepidoptera Nonobtectomeran Superfamilies / 115 ABOUT THIS BOOK / xi and Moths Superfamily Schrecken- steinioidea / 115 Superfamily Epermenioidea / 116 Morphology / 3 Superfamily Alucitoidea / 116 Biology / 13 Superfamily Pterophoroidea / 117 Significance in Natural Superfamily Choreutoidea / 121 and Human Communities / 17 Superfamily Sesioidea / 123 Fossil Record and Evolution / 19 Superfamily Cossoidea / 126 A History of Moth Collectors Superfamily Tortricoidea / 128 in Western North America / 23 Superfamily Zygaenoidea / 160 Marc E. Epstein and James K. Adams Unspecialized Obtectomera / 165 PART TWO Superfamily Copro- Classification and Natural History morphoidea / 166 of the Moths of Western North Superfamily Thyridoidea / 167 America Obtectomera / 167 Superfamily Pyraloidea / 167 Macrolepidoptera / 201 Primitive Lineages / 33 Superfamily Drepanoidea / 201 Zeugloptera Clade / 33 Superfamily Geometroidea / 203 Superfamily Micropterigoidea / 33 Superfamily Mimallonoidea / 232 Glossata, Homoneura Clade / 34 Superfamily Lasiocampoidea / 232 Superfamily Eriocranioidea / 34 Superfamily Bombycoidea / 236 Exoporia Clade / 35 Superfamily Noctuoidea / 247 Superfamily Hepialoidea Clade / 36 Heteroneura, Monotrysia Clade / 37 SUGGESTIONS FOR COLLECTING Superfamily Nepticuloidea / 37 AND OBSERVING MOTHS / 321 Superfamily Incurvarioidea / 38 GLOSSARY / 329 / 44 Superfamily Tischerioidea INSECT INDEX / 333 Ditrysia, Nonapoditrysian PLANT INDEX / 357 Superfamilies / 47 GENERAL INDEX / 367 Superfamily Tineoidea / 47 Superfamily Gracillarioidea / 53 Superfamily Gelechioidea / 59 Color Plates / between pages 114 Superfamily Gelechioidea, and 115 Oecophorid Lineage / 60 Superfamily Gelechioidea, Elachistid Lineage / 64 Superfamily Yponomeutoidea / 104 v UCPB004-3960G-FM_pi-xiv.qxd 30/1/09 7:24 am Page vi UCPB004-3960G-FM_pi-xiv.qxd This page intentionally left blank UCPB004-3960G-FM_pi-xiv.qxd 30/1/09 7:24 am Page vii UCPB004-3960G-FM_pi-xiv.qxd LIST OF FIGURES Figure 1. Map of North America Figure 64. Habitus drawing of flightless female Ethmia charybdis Figure 2. Schematic representation of the exoskeletal anatomy Powell (Ethmiidae), a winter moth. of a ditrysian moth, with prothoracic leg enlarged below. Figures 65–73. Genitalia structure of exemplar species of Gele- Figure 3. Descaled lepidopteran head, frontal aspect. chioidea, ventral aspect: Elachistidae, Coleophoridae, Mom- Figures 4–9. Wing venation: exemplar species of ancestral line- phidae. ages of Lepidoptera. Figures 74–86. Genitalia structure of exemplar species of Gele- Figures 10–17. Wing venation: exemplars of ditrysian moths. chioidea, aedeagus in situ: Scythrididae, Cosmopterigidae. Figure 18. Male genitalia of a ditrysian moth (Tortricidae), ven- Figure 87. Habitus drawing of female Areniscythris brachypteris tral aspect with valvae reflexed. Powell (Scythrididae). Figure 19. Female genitalia of a ditrysian moth (Tortricidae), ven- Figures 88–100. Genitalia structure of exemplar species of Gelechi- tral aspect; broken lines represent segments of abdominal pelt. idae, venterolateral or ventral or aspect, except as noted. Figure 20. Body forms of moth larvae. Figures 101–112. Genitalia structure of exemplar species of Figure 21. Schematic representation of the head capsule of a lar- Gelechiidae, Douglasiidae, and Yponomeutoidea; males ven- val ditrysian moth, frontal aspect. tral or venterolateral aspect, females ventral aspect: Plutellidae, Figure 22. Chaetotaxy (setal map) of a larval ditrysian moth (Tor- Yponomeutidae, Argyresthiidae, Douglasiidae. tricidae). Figures 113–122. Genitalia of Yponomeutoidea and Sesiidae, Figure 23. Abdominal proleg crochet patterns. ventral aspect; aedeagus removed and shown to right Figure 24. Pupae of ditrysian moths, ventral aspect. (except as noted): Acrolepiidae, Heliodinidae, Bedelliidae, Figure 25. Two-dimensional portrayal of phylogenetic relation- Sesiidae. ships of extant lepidopteran superfamilies, based on parsi- Figure 123. Habitus drawing of Lithariapteryx elegans Powell, a di- mony analysis. urnal heliodinid that is endemic to the Monterey Bay and San Figure 26. Phylogenetic hypothesis of major lepidopteran line- Luis Obispo coastal dune systems in California. ages superimposed on the geologic time scale, with fossil oc- Figures 124–133. Genitalia structure of exemplar species of currences indicated. nonobtectomeran Apoditrysia, ventral aspect (except as Figure 27. Pictorial representation of a theoretical scenario for the noted): Epermeniidae, Alucitidae, Pterophoridae, Choreutidae. origins of angiosperm-feeding that led to the radiation of lep- Figures 134–141. Genitalia structure of exemplar species of idopteran during the Cretaceous. Tortricidae, Olethreutinae, ventral aspect. Figures 28–31, 33–38. Genitalia structures of ancestral lineages. Figures 142–154. Genitalia structure of exemplar species of Figure 32. Schematic representation of hepialid female terminal Tortricidae, Tortricinae, ventral aspect. segments, saggital section. Figures 155–164. Genitalia structure of exemplar species of Figures 39–47. Genitalia structure of exemplar species of Ti- Zygaenoidea, ventral aspect (except as noted): Zygaenidae, neoidea and Gracillarioidea; Tineidae, Acrolophidae, Psychi- Limacodidae, Dalceridae, Megalopygidae. dae, Gracillariidae, Bucculatricidae. Figures 165–174. Genitalia structure of exemplar species of Cram- Figure 48. Two-dimensional portrayal of phylogenetic relation- bidae, ventral aspect (except as noted): Scopariinae, Odonti- ships in Gelechioidea. inae, Glaphyriinae, Pyraustinae. Figures 49–62. Antennal, abdominal structures, and genitalia of Figures 175–184. Genitalia structure of exemplar species of Cram- exemplar species of Gelechioidea: Blastobasidae, Oecophori- bidae, ventral aspect (except as noted): Pyraustinae, Spi- dae, Depressariidae, Stenomatidae, Ethmiidae. lomelinae, Acentropiinae (ϭNymphulinae), Crambinae. Figure 63. Heads of ethmiid moths, showing the strongly up- Figures 185–192. Genitalia structure of exemplar species of curved labial palpus that is characteristic of most Gele- Pyralidae, ventral aspect, aedeagus to right (except as noted): chioidea. Pyralinae, Chrysauginae, Epipaschiinae, Phycitinae. vii UCPB004-3960G-FM_pi-xiv.qxd 30/1/09 7:24 am Page viii UCPB004-3960G-FM_pi-xiv.qxd Figures 193–200. Genitalia structure of exemplar species of Pyral- lonoidea, Lasiocampoidea, Bombycoidea, Saturniidae, Sphin- idae, Phycitinae, ventral aspect. gidae. Figure 201. Schematic representation of Geometridae FW with Figure 230.
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