Moths of Western North America

517 Pages·2009·27.36 MB·English

by  Jerry A. Powell, Paul A. Opler

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M OTHS OF WESTE R N NORTH AM E R ICA
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Moths of Western
North America
JERRY A. POWELL
PAUL A. OPLER
UNI V E R S I T Y O F C A L I F O R N I A P R E S S
Berkeley Los Angeles London
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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
10 9 8 7 6 5 4 3 2 1
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.
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CONTENTS
LIST OF FIGURES / vii PART ON E 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
Superfamily Tischerioidea / 44
INSECT INDEX / 333
Ditrysia, Nonapoditrysian
PLANT INDEX / 357
Superfamilies / 47
GENERAL INDEX / 367
Superfamily Tineoidea / 47
Superfamily Gracillarioidea / 53
Color Plates / between pages 114
Superfamily Gelechioidea / 59
Superfamily Gelechioidea, and 115
Oecophorid Lineage / 60
Superfamily Gelechioidea,
Elachistid Lineage / 64
Superfamily Yponomeutoidea / 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: 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 (⫽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- Pyralidae, 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, Saturniidae, Sphin-
idae, Phycitinae, ventral aspect. gidae.
Figure 201. Schematic representation of Geometridae FW with Figure 230. Schematic representation of Noctuidae 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: Notodontidae, 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 insects worldwide, which is more than five times the comparable treatment of all moth families, irrespective of the
number of all other animals 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 genus 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 microlepidoptera 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, taxonomy, 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
ix