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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. 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 Nonobtectomeran Superfamilies / 115 ABOUT THIS BOOK / xi and Moths Superfamily Schrecken- steinioidea / 115 Superfamily Epermenioidea / 116 Morphology / 3 Superfamily / 116 Biology / 13 Superfamily Pterophoroidea / 117 Significance in Natural Superfamily Choreutoidea / 121 and Human Communities / 17 Superfamily / 123 Fossil Record and Evolution / 19 Superfamily / 126 A History of Collectors Superfamily Tortricoidea / 128 in Western North America / 23 Superfamily / 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 / 167 / 201 Primitive Lineages / 33 Superfamily / 201 Zeugloptera Clade / 33 Superfamily / 203 Superfamily Micropterigoidea / 33 Superfamily Mimallonoidea / 232 , Homoneura Clade / 34 Superfamily / 232 Superfamily Eriocranioidea / 34 Superfamily / 236 Clade / 35 Superfamily / 247 Superfamily Clade / 36

Heteroneura, Clade / 37 SUGGESTIONS FOR COLLECTING Superfamily / 37 AND OBSERVING MOTHS / 321 Superfamily Incurvarioidea / 38 GLOSSARY / 329 / 44 Superfamily INDEX / 333 , Nonapoditrysian PLANT INDEX / 357 Superfamilies / 47 GENERAL INDEX / 367 Superfamily / 47 Superfamily / 53 Superfamily / 59 Color Plates / between pages 114 Superfamily Gelechioidea, and 115 Oecophorid Lineage / 60 Superfamily Gelechioidea, Elachistid Lineage / 64 Superfamily / 104

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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: , , 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: , . Figure 18. Male genitalia of a ditrysian moth (), 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- , , and Yponomeutoidea; males ven- val ditrysian moth, frontal aspect. tral or venterolateral aspect, females ventral aspect: , Figure 22. Chaetotaxy (setal map) of a larval ditrysian moth (Tor- Yponomeutidae, Argyresthiidae, Douglasiidae. tricidae). Figures 113–122. Genitalia of Yponomeutoidea and , 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): , , 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): , Alucitidae, , . 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): , neoidea and Gracillarioidea; , , Psychi- , , Megalopygidae. dae, , . Figures 165–174. Genitalia structure of exemplar species of Cram- Figure 48. Two-dimensional portrayal of phylogenetic relation- bidae, ventral aspect (except as noted): , 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: , Oecophori- bidae, ventral aspect (except as noted): Pyraustinae, Spi- dae, Depressariidae, Stenomatidae, Ethmiidae. lomelinae, Acentropiinae (Nymphulinae), 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- , ventral aspect, aedeagus to right (except as noted): chioidea. Pyralinae, Chrysauginae, Epipaschiinae, Phycitinae.

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Figures 193–200. Genitalia structure of exemplar species of Pyral- lonoidea, Lasiocampoidea, Bombycoidea, , Sphin- idae, Phycitinae, ventral aspect. gidae. Figure 201. Schematic representation of Geometridae FW with Figure 230. Schematic representation of FW with color color pattern elements used in diagnoses. pattern elements used in diagnoses. Figures 202–209. Genitalia structure of exemplar species of Figures 231–240. Genitalia structure of exemplar species of Noc- Geometridae, ventral aspect, aedeagus. tuoidea, ventral aspect: , Arctiinae, Lymantriidae. Figures 210–217. Genitalia structure of exemplar species of Figures 241–252. Genitalia structure of exemplar species of Noc- Geometridae, ventral aspect, aedeagus. tuidae, ventral aspect, except as noted. Figure 218. Tescalsia giulianiata Ferguson, adult female. Figure 253. Single specimen spreading blocks. Figures 219–229. Genitalia structure of exemplar species Figure 254. The Keifer cork board method of spreading small of bombycoid moths, ventral aspect, aedeagus: Mimal- Lepidoptera.

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PREFACE

Few people appreciate the diversity of insect life, and most nat- sections give general features of the morphology of adults and uralists, including professionally trained biologists who work immature stages and summarize adult behavior, larval biology, with vertebrates or flowering plants, are surprised, even skep- and life cycles. In this book, inspired by I. F. B. Common’s Moths tical, when they hear estimated numbers of insect species. of Australia, we have taken an innovative approach as compared There are believed to be about 1.5 million named and described to previous treatments of Nearctic Lepidoptera. We attempt species of worldwide, which is more than five times the comparable treatment of all moth families, irrespective of the number of all other combined. Estimates of the num- size of the moths or their traditional popularity with collectors. ber in collections remaining to be distinguished and named We present a summary of morphology, larval biology, geo- vary widely—3 to 5 million or more—and of course most of graphical distribution, and diversity for superfamilies and fam- those, along with countless species never collected (estimates ilies and discuss about 25% of the species in each family range from 10 to 30 million), will have been exterminated as and larger known to occur in western North America. a result of human population growth within the next few Obviously, therefore, this is not an identification manual that decades. Of the total described, an estimated 11 to 12%, or illustrates all moth species. On the other hand, we think ento- 160,000 to 180,000 species, are Lepidoptera, most of which oc- mologists, ecologists, and recreational collectors will find it use- cur in tropical regions. About 13,000 live in North America ful in determining taxa to which species of interest belong, and north of Mexico, of which 94% are moths, the remainder but- we provide an entry into the literature—general references are terflies. We believe there are 7,500 to 8,000 named species of given at the end of the introduction, and major taxonomic and moths in the western states and provinces, along with at least larval biology references are cited at the end of each superfam- another 3,000 that have been collected but not yet named. ily. The text summarizes host-plant relationships and life history It has been more than a century since a volume treating the patterns, which we hope will be valuable to ecologists and other moths of North America was published, The Moth Book, by W. biologists. We believe our treatment can open doors to the world J. Holland. In its era, when there were about half as many of for amateur collectors and others interested named species as now, Holland did an admirable job of sum- in biological studies, a field that has lagged in attracting spe- marizing knowledge and illustrating the larger moths with fine cialists in North America. Moreover, we provide estimates of to- color photos, but his representation of small species, the so- tal species numbers in North America and the West and thereby called microlepidoptera, was sparse, only 6% of the described give insights into the challenge remaining in descriptive taxon- species. That volume, long unavailable, was reprinted in pa- omy in each family, subfamily, and larger genus. perback form by Dover Press in 1968 with no additions and Between us we have more than 100 years’ experience col- scant updating of the names. In 1926, E. O. Essig, in Insects of lecting moths and investigating their life history stages in the Western North America, gave accounts of more than 250 moth western United States, primarily in California, where many of species and their larval biologies, especially those of concern in the pioneers of moth study in the western Nearctic worked agriculture. This was a small selection of the western fauna, but (e.g., Henry Edwards, Lord Walsingham, Albert Koebele, W. S. Essig summarized life histories of species in many families, with Wright, F. X. Williams, J. A. Comstock, H. H. Keifer, C. M. greater emphasis on some of the microlepidoptera (Gracillari- Dammers, and J. W. Tilden). Our experience has dictated an ob- idae, Sesiidae, Tortricidae) and pyraloids than on many of the vious Californicentric bias in species discussed, but we have macromoths, so it served as a good general introduction. In also lived and studied for varying periods in Colorado, Virginia, 1984, Covell’s Field Guide to Moths of Eastern North America pro- Arizona, Mexico, Costa Rica, and Australia, all experiences that vided a much-needed illustrated manual to the macromoths, have contributed to the background of this book. which inspired many lepidopterists to expand their horizons A great deal of progress has been made during the past half beyond butterflies and begin developing local groups of moth century toward understanding North American Lepidoptera, enthusiasts. Again, however, the “micros,” which make up enabling development of this book. This era coincided with the about 45% of the described Lepidoptera in North America, were growth of the Lepidopterists’ Society, founded in 1947 by C. L. slighted, with only token representatives included, especially in Remington and H.K. Clench, and progress was accelerated by the leaf-mining families, which had only 2% of their species il- modern advances in travel and easy access to areas that were lustrated. remote, and by the use of black light and mercury vapor light In this text we summarize the diversity, , and lar- traps. It must be difficult for young students today to imagine val biology of western North American moths. The introductory the state of knowledge of western North American moths when

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we started. One had to depend upon Forbes’ Lepidoptera of New on southern California noctuids), Ted Poling, Amanda Roe, Dan York; Holland’s Moth Book, if you could find a copy; and Essig’s Rubinoff, Alma Solis, Bruce Walsh, and Shen-Horn Yen. In ad- Insects of Western North America to even get started, and there dition to systematists listed in the preceding paragraph, many were no comprehensive museum collections of microlepi- other taxonomists have provided specimens, identifications, doptera and pyraloids in the western states. and collection records for our respective museums, aiding the development of our understanding of western moths, includ- ing Bill Bauer, Norris Bloomfield, Don Bowman, Terry Dickel, Acknowledgments Cliff Ferris, Todd Gilligan, Paul Johnson, Peter Jump, Ed Knud- Several persons served as our mentors, helping to show the way son, Ron Leuschner, John Nordin, Jack Powers, Ron Robertson, when the world of systematics and moths seemed an impon- Dan Rubinoff, J. B. Sullivan, Catherine Toschi (Tauber), Bruce derable labyrinth, by providing insights and encouragement, Walsh, Ron Wielgus, and Don Wright. including Richard Bohart, Charles Harbison, Paul Hurd, Jr., Hart- We are grateful for the efforts by generations of students and ford Keifer, Don MacNeill, Douglas Ferguson, and Edward Ross. others at the University of California, Berkeley, who assisted by We owe much of our knowledge to lepidopterists who paved collecting larvae and helping to maintain the Lepidoptera rear- the way to writing this book by their collections, publications, ing program and its database. This effort has been one of the and providing identifications during the past 50 years: Ron most comprehensive of its kind in the United States, including Hodges orchestrated the publication of The Moths of America collections and maintenance of nearly 15,000 rearing lots which North of Mexico series, including his treatments of the Sphingi- successfully produced adults and field host data for about 1,350 dae and parts of the Gelechioidea; and he spent several summers species during 1957–2006. Participants included John Brown, collecting in Arizona, Colorado, Utah, and western Texas. Don Don Burdick, John Chemsak, John De Benedictis, Robert Dietz Davis wrote monographs of primitive moths, Psychidae, and Car- IV, Deborah Green, Yu-Feng Hsu, John Lawrence, Paul Rude, posinidae. A publication in 1987 of Immature Insects, Volume 1, Brian Scaccia, Don Veirs, David Wagner, Jim Whitfield, and Jo orchestrated by Fred Stehr, was an important advance, as have Wolf. Much of this work was supported by research grants from been monographs by Annette Braun (Bucculatricidae, Tischeri- governmental agencies: The National Science Foundation, idae), Frank Hasbrouck (Acrolophidae), David Adamski (Blasto- “Comparative biology of Microlepidoptera “ in 1965–1970; “Sys- basidae), Don Duckworth (Stenomidae) and with Tom Eichlin tematics of Tortricine Moths” in 1985–1990; the U.S.D.A., For- (Sesiidae), Jean-Francois Landry (Coleophoridae and Scythridi- est Service, “Effects of Dylox on non-target organisms during a dae); Lauri Kaila (Elachistidae); Dalibor Povolny´, many contri- Modoc budworm outbreak” in 1974 and the CANUSA/West Re- butions in gnorimoschemine Gelechiidae; R. Gaedike search Agreement, “Taxonomic relationships and pheromone (Acrolepiidae, Epermeniidae, Douglasiidae); John Heppner isolation among western spruce budworms populations” in (); Yu-Feng Hsu (Heliodinidae); and C. Gielis 1978–1982; Bureau of Land Management, “Survey of Lepi- and Deborah Matthews (Pterophoridae). Margaret MacKay, T. N. doptera inhabiting three dune systems in the California desert” Freeman, N. S. Obraztsov, John Brown, Richard L. Brown, and in 1977-1978. The Agricultural Experiment Station, University Bill Miller contributed greatly to our knowledge of Tortricidae; of California, Berkeley, funded the research project “Taxonomy Marc Epstein to Zygaenoidea (especially Limacodidae); and Scott and biology of Microlepidoptera,” 1962–1994. Miller (Dalceridae). Eugene Munroe, Alma Solis, and H. H. Ne- Color images were provided by Evi Buckner-Opler, Rollin unzig have contributed extensively to Pyraloidea; W. C. McGuf- Coville, Lee Dyer, Scott Kinnee, Jean-Francois Landry, David fin, Douglas Ferguson, and Klaus Bolte to Geometridae; and Powell, Brian Scaccia, David Wagner, and Bruce Walsh. David Hardwick, Don Lafontaine, and Bob Poole to Noctuoidea. David Furth provided access and use of images made by the Jack Franclemont, who trained many of our contemporary lepi- late Douglas Ferguson, in the Smithsonian Institution, Depart- dopterists during his long career at Cornell University, published ment of Entomology illustration archives. We are indebted to the MONA treatment of bombycoid taxa and spent many sum- Steve Lew, Tina Mendez, Ainsley Seago, and Kip Will for assis- mers collecting in Arizona and Texas. Fred Rindge lived in tance with a Microptics Digital Imaging System, enabling pro- California, where he completed his graduate studies, and later duction of the specimen images of microlepidoptera and was curator of Lepidoptera at the American Museum of Natural pyraloids by Powell, to Jocelyn Gill for production of macro History, New York, he spent several summers collecting in the moth images in Ottawa, and to Don Lafontaine for providing Rocky Mountains and published numerous revisions of access and use of specimens in the CNC, Agriculture Canada. geometrid genera. Doug Ferguson lived in the Northeast but Jocelyn edited and composed the images on Plates 1–58. The made many collecting trips to the West and published revisions line drawings were made by Powell (Figs. 1–27) and by Ainsley of geometrids, saturniids, and arctiid genera. The singularly com- Seago (Figs. 28–252), redrawn from Powell drafts and literature prehensive volume on Lepidoptera in Handbook of Zoology, or- sources acknowledged and funded by the P. C. Powell Trust chestrated and edited by Niels Kristensen, published in 1999, Fund and the Essig Museum of Entomology. Figures 63, 64, and pooled knowledge of specialists worldwide and provided the sys- 86 are original drawings by Celeste Green and Figure 117 is by tematics framework upon which our text is based. With the in- Tina Jordan; we regret that neither lived to see their use here. creased literature access have come a legion of collectors, some Lastly, and most importantly, we sincerely acknowledge the of whom have posted lists and images on the internet. support provided by our wives, Liz Randal and Evi Buckner- We, along with Marc Epstein and James Adams, who wrote Opler, enabling completion of this project: To Liz for 30 years the Zygaenoidea, owe special thanks to the following people for of incomparable companionship with an admirable balance of providing helpful reviews of parts of the manuscript: David encouragement, tolerance, forbearance, and irreverence; and Adamski, Val Albu, Richard L. Brown, Charles Covell, Cliff Fer- Evi exhibited almost infinite patience over a three year-period ris, Chuck Harp, John Heppner, Ron Hodges, Boris Kondratieff, when most of the text was written, including taking an extra Jean-Francois Landry, Deborah Matthews, Noel McFarland, Don job to provide more quiet time for writing. There may be no Lafontaine, Tomas Mustelin (who also provided his manuscript way to adequately repay their kindness.

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ABOUT THIS BOOK

Definition of Western North America Scientific and Common Names We define “the West” to include the states and Canadian names are governed by the International Code of Zoo- provinces and parts thereof west of a line drawn along the east- logical Nomenclature (ICZN), which is among the very few laws ern base of the western cordillera, from the British and Richard- agreed upon by all nations. The primary value of our scientific son mountains, Yukon Territory; and Mackenzie Mountains, name system, and the reason it has been used in the same for- Northwest Territory; along the Rocky Mountains from north- mat for 250 years, is its universality. The genus name must be eastern British Columbia, western Alberta, Montana, unique, different from every other animal’s generic name, and Wyoming, and Colorado; to New Mexico along the Sacra- the names are used in the same Latinized form worldwide. As mento Mountains; and western Texas along the Guadalupe, our knowledge and the number of described species increase, Davis, and Chisos mountains (Fig. 1). We mention transconti- species names change, especially because they are moved from nental and other extralimital ranges, but we do not treat one genus to another, or a species is found to be the same as species that are restricted to regions east of the defined part of an earlier described one, whose name takes priority. western North America. Animals are classified by a hierarchy of categories called taxa (singular: taxon), with each higher taxon containing one to many of the next subordinate category. Within the animal Selection of Species kingdom, there are six obligate ranks that define the position For each family, subfamily, and larger genus we aimed to dis- of every species: phylum, class, order, family, genus, and cuss and illustrate about 25% of the named moth species species. The scientific name of every animal is based on the two known to occur in the West. Criteria that we used to select the lowest-rank categories, genus (plural: genera) and species (plu- species included, not necessarily in order of priority, are that ral: species). Hence the name has two parts, the generic name, they be (1) widespread, well-known species, especially those the first letter of which is always capitalized, followed by the of economic importance in agriculture, forestry, garden and specific name, the first letter of which is not capitalized. Ani- turf management, and stored products; (2) species for which mal names usually are given in italics. Thus an animal’s name we have information on life cycle and biology, particularly the is more than a name because it is informative on relationships; larval foods; (3) distinctive or showy species and those other- the species is more closely related to other members of its wise liable to be easily recognized, especially in populated ar- genus than it is to those of other genera. Similarly, knowing eas; (4) species with special adaptations, behavioral or the name of the family to which a species belongs tells us morphological, even if they are obscure or local in occurrence; much about its relationships; genera in the same family are (5) a generic representation: ideally all genera might have been closer relatives than any of them is to genera in other families. treated, but including at least one species from every genus As an example, the largest moth in western North America, would have exceeded 25% of the total in some taxa, and many (Linnaeus), is classified as of those would be poorly known species. Therefore, we ex- Phylum Arthropoda, which includes all animals with a cluded many monotypic genera or those with only one or a hard exoskeleton and jointed appendages few species in the West for which scant information is avail- able. Some families with only one or a few species are pro- Class Insecta, with three body sections and portionately better represented (e.g., essentially all six pairs of legs , , , Adeli- Order Lepidoptera, the moths and butterflies; having dae, , Zygaenoidea, and Doidae). In sum- a covering of scales and sucking proboscis formed mary, we selected the better-known species for which we have from the maxillary laciniae biological or other interesting information.

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FIGURE 1. Map of North America with the western portion defined for faunal representation in ths book delineated by the bold, broken line.

Family Noctuidae, moths with thoracic tympana which contains one to many genera. Typically these interme- diate ranks are used primarily by specialists, enabling commu- Genus Ascalapha nication about additional levels of relationships. For example, Species odorata, a name proposed by Linnaeus there are 9,000 described species of the family Tortricidae in 1758, in the tenth edition of his Systema worldwide, and these are assigned to three subfamilies, which Naturae, which was later selected as the start- have 11, three, and eight tribes in which their genera are clas- ing base of our binomial nomenclature sified. Without the intermediate categories, comprehension of system. relationships among the more than a thousand genera would be exceedingly difficult, even for the most dedicated special- The author who originally proposed a species name usually is ist. Moreover, endings for the names of ranks between genus listed with the genus and species, to complete the name; the and order are spelled out in the ICZN. Thus, -oidea for super- author’s name is placed in parentheses if the species has been family, -idae for family, -inae for subfamily, and -ini for tribe assigned to a genus different from its original combination. are consistently used and provide immediate recognition of the In addition, several intermediate ranks are commonly em- relative position of the group within its lineage. ployed for insects because there are vast numbers of species. In Common or vernacular names for species vary from place Lepidoptera, more than 120 families are defined, which are to place for the same animal, particularly but not limited to grouped into about 44 superfamilies. Many families are divided species that occur in countries using different languages, and into subfamilies, and larger subfamilies into tribes, each of there are no rules governing their proposal or priority. Thus

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Ascalapha odorata, mentioned above, is called the “black witch” ate-sized moths (. 6 to 20 mm). Measurements in centime- in the United States but has a different common name, and ters accurate to 0.2 cm were made for larger moths, using probably more than one, in Mexico and other Spanish-speak- vernier calipers. ing countries. A species often has more than one common name, depending upon where it is applied or in what circum- stance. For example the noctuid moth Heliocoverpa zea is called Abbreviations the “corn earworm”, “tomato fruitworm,” and “cotton boll- Generic names of both plants and insects often are abbreviated worm” in different places, depending on which crop it is in- by their first letter if already spelled out in the same paragraph. festing. In this book, we used the common names approved by Body parts with abbreviations are illustrated in Figs. 2, 3, and the Entomological Society of America and those established in 18 to 24. Abdominal segments are numbered 1 to 10 (or A1 to various books for some other species, which, however, some- A10), basal to caudal. In addition to morphological terms cited times contradict one another. We did not propose new com- in the figures, the following abbreviations are used: mon names for any of the vast majority of moth species that lack them. FW fore wing(s) HW hind wing(s) Sc, R, M, Cu, A wing vein systems: subcostal, radial, Literature medial, cubital, anal (Figs. 4–17) We refer to authors of major works, such as the chapters in cm centimeter (1 inch 2.54 cm) Kristensen’s 1999 volume in Handbook of Zoology, or a sys- mm millimeter (10 mm 1 cm) tematic revision of a particular group, as in the Moths of Amer- m meter (100 cm 1 m 3.28 feet) ica North of Mexico, but do not cite references in the text. km kilometer (1,000 m 1 km 0.61 miles) General references that treat many families (e.g., Prentice and mi mile others in Forest Lepidoptera of Canada) appear in a bibliogra- Co. County phy at the end of this introduction. Taxonomically restricted MONA Moths of America North of Mexico publications on individual families or larger genera are cited Mt., Mtn., Mtns. mount, mountain, mountains at the end of each chapter. These contain the sources of much MyBP million years before the present of our text and will provide an entry into the literature for feet (elevation) most taxa. Of course, we have referred to hundreds of other publications in scientific journals that treat individual species Institutional collection acronyms: or specific topics, citation of which is beyond the practical scope of this book. AMNH American Museum of Natural History, New York CNC Canadian National Collection, Ottawa Figures and Color Plates CSAC California State Collection, Most of the line drawing figures depict moth genitalia, detailed Department of Food and Agriculture, comparison of which often is essential for identifying genera Sacramento and species. We provide exemplar illustrations representing CSUC C. P. Gillette Museum of Arthropod Diversity, both sexes of one to several genera for most families. In some Colorado State University, Ft. Collins cases, these depict species for which there are no accounts in EME Essig Museum of Entomology, University of our text because better illustrations were available, from which California, Berkeley our drawings were made, than was true for discussed species. LACM Los Angeles County Museum of Natural See color section for an explanation of how to use the History, Los Angeles plates. PMY Peabody Museum, Yale University, New Haven, Conn. SBNHM Santa Barbara Natural History Museum, Santa Measurements Barbara, Calif. SDMNH San Diego Museum of Natural History, Size of adults is estimated by fore-wing length, measured from San Diego base to apex including the fringe. Using a micrometer disk, we UCD Bohart Museum of Entomology, University of measured five or more of the largest and smallest specimens California, Davis available to obtain a range of fore-wing lengths, to 0.1 mm ac- USNM U.S. National Museum of Natural History, curacy for tiny moths, and to 0.5 mm accuracy for intermedi- Washington, D.C.

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