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Molecular Biology
and Genetics
of the Lepidoptera
 CONTEMPORARY TOPICS in ENTOMOLOGY SERIES

THOMAS A. MILLER EDITOR

Insect Symbiosis
Edited by Kostas Bourtzis and Thomas A. Miller

Insect Sounds and Communication: Physiology, Behaviour, Ecology, and

Evolution
Edited by Sakis Drosopoulos and Michael F. Claridge

Insect Symbiosis, Volume 2

Edited by Kostas Bourtzis and Thomas A. Miller

Insect Symbiosis, Volume 3

Edited by Kostas Bourtzis and Thomas A. Miller

Food Exploitation by Social Insects: Ecological, Behavioral, and Theoretical

Approaches
Edited by Stefan Jarau and Michael Hrncir

Molecular Biology and Genetics of the Lepidoptera

Edited by Marian R. Goldsmith and František Marec
CONTEMPORARY TOPICS in ENTOMOLOGY SERIES

THOMAS A. MILLER EDITOR

Molecular Biology
and Genetics
of the Lepidoptera

Edited by
Marian R. Goldsmith
František Marec
CRC Press
Taylor & Francis Group
6000 Broken Sound Parkway NW, Suite 300
Boca Raton, FL 33487-2742
© 2010 by Taylor and Francis Group, LLC
CRC Press is an imprint of Taylor & Francis Group, an Informa business

No claim to original U.S. Government works

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International Standard Book Number: 978-1-4200-6014-0 (Hardback)

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Library of Congress Cataloging‑in‑Publication Data

Molecular biology and genetics of the Lepidoptera / editors, Marian R. Goldsmith and Frantisek
Marec.
p. cm. -- (Contemporary topics in entomology series)
Includes bibliographical references and index.
ISBN 978-1-4200-6014-0 (alk. paper)
1. Lepidoptera--Physiological genomics. 2. Lepidoptera--Molecular genetics. I. Goldsmith, Marian
R. II. Marec, Frantisek. III. Title. IV. Series.

QL562.2.M635 2010
595.78--dc22 2009012588

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Contents
Preface..............................................................................................................................................vii
About the Editors...............................................................................................................................xi
Contributors.................................................................................................................................... xiii

Chapter 1 Evolutionary Framework for Lepidoptera Model Systems...........................................1

Amanda D. Roe, Susan J. Weller, Joaquin Baixeras, John Brown,
Michael P. Cummings, Donald R. Davis, Akito Y. Kawahara,
Cynthia S. Parr, Jerome C. Regier, Daniel Rubinoff,
Thomas J. Simonsen, Niklas Wahlberg, and Andreas Zwick

Chapter 2 Recent Progress in Silkworm Genetics and Genomics...............................................25

Marian R. Goldsmith

Chapter 3 Rise and Fall of the W Chromosome in Lepidoptera................................................. 49

František Marec, Ken Sahara, and Walther Traut

Chapter 4 Sex Chromosomes and Sex Determination in Bombyx mori...................................... 65

Hiroaki Abe, Tsuguru Fujii, and Toru Shimada

Chapter 5 Evolutionary and Developmental Genetics of Butterfly Wing Patterns: Focus

on Bicyclus anynana Eyespots.................................................................................... 89
Patrícia Beldade and Suzanne V. Saenko

Chapter 6 Prospects for Locating Adaptive Genes in Lepidopteran Genomes: A Case

Study of Butterfly Color Patterns.............................................................................. 105
Simon W. Baxter, Owen McMillan, Nicola Chamberlain,
Richard H. ffrench-Constant, and Chris D. Jiggins

Chapter 7 Molecular and Physiological Innovations of Butterfly Eyes..................................... 121

Marilou P. Sison-Mangus and Adriana D. Briscoe

Chapter 8 Lepidopteran Circadian Clocks: From Molecules to Behavior................................. 137

Christine Merlin and Steven M. Reppert

Chapter 9 Lepidopteran Chemoreceptors.................................................................................. 153

Kevin W. Wanner and Hugh M. Robertson

v
vi Contents

Chapter 10 Sexual Communication in Lepidoptera: A Need for Wedding Genetics,

Biochemistry, and Molecular Biology....................................................................... 169
Fred Gould, Astrid T. Groot, Gissella M. Vásquez, and Coby Schal

Chapter 11 Genetics of Host Range in Lepidoptera.................................................................... 195

Sara J. Oppenheim and Keith R. Hopper

Chapter 12 Genetics and Molecular Biology of the Major Crop Pest Genus Helicoverpa......... 219
Karl Gordon, Wee Tek Tay, Derek Collinge,
Adam Williams, and Philip Batterham

Chapter 13 Molecular Genetics of Insecticide Resistance in Lepidoptera.................................. 239

David G. Heckel

Chapter 14 Innate Immune Responses of Manduca sexta.......................................................... 271

Michael R. Kanost and James B. Nardi

Chapter 15 Lepidopterans as Model Mini-Hosts for Human Pathogens and as a Resource

for Peptide Antibiotics............................................................................................... 293
Andreas Vilcinskas

Chapter 16 Intrahemocoelic Toxins for Lepidopteran Pest Management...................................307

Nina Richtman Schmidt and Bryony C. Bonning

Chapter 17 The Interactions between Polydnavirus-Carrying Parasitoids and Their

Lepidopteran Hosts................................................................................................... 321
Michael R. Strand

Chapter 18 Densovirus Resistance in Bombyx mori................................................................... 337

Keiko Kadono-Okuda

Index............................................................................................................................................... 349
Preface
Moths and butterflies, insects of the order Lepidoptera, are among the most diverse and species-rich
groups of organisms. (With more than 150,000 species, they comprise the second largest order of
animals, after beetles, Coleoptera.) Their primarily phytophagous caterpillars represent important
components of mainly terrestrial habitats all over the world. Many species have a significant impact
on human society, whether negative as major pests of agriculture and forestry or beneficial as pol-
linators and food sources for other animals. A special case is the domesticated silkworm, Bombyx
mori. For many years the rearing of silkworms for silk production, sericulture, was one of the most
important industries in more than 30 countries, especially in Asia (China, Japan, Korea, Thailand),
South Asia (India), Europe (France, Italy, Russia, Rumania, Bulgaria), and, in the twentieth century,
South America (Brazil).
The Lepidoptera are probably the most widely studied group of invertebrates. Butterflies particu-
larly, with their beautiful wing patterns, have attracted the attention of professional researchers and
amateur entomologists. However, other than the domesticated silkworm, only a few lepidopteran
species have attracted the deeper interest of geneticists. For example, the female heterogamety typi-
cal of all Lepidoptera had been first inferred at the beginning of the last century from sex-linked
inheritance in the magpie moth, Abraxas grossulariata, by L. Doncaster and G.H. Raynor before
the sex chromosomes were identified in several lepidopteran species by J. Seiler. The gypsy moth,
Lymantria dispar, had become well known for the development of intersexuality from studies by
R. Goldschmidt in the 1930s. Among early genetic models, we can certainly count the flour moth,
Ephestia kuehniella, with a collection of eye-pigment mutants that were used in numerous studies
on the ommochrome synthesis pathway by A. Kühn and colleagues and were used to develop the
one-gene-one-enzyme hypothesis by a member of Kühn’s group, E. Caspari. All these and other
genetic studies until the late 1960s on formal genetics including chromosome numbers for sev-
eral hundred species of moths and butterflies have been reviewed comprehensively in Lepidoptera
Genetics by R. Robinson, published by Pergamon Press in 1971. This book also presents in detail the
famous industrial melanism story, which began in 1848 with the discovery of the first carbonaria
morph of the peppered moth (Biston betularia) in Manchester, England. In this book, considerable
attention is paid to the evolution and genetics of Batesian mimicry in Hypolimnas and Papilio but-
terflies that mimic wing markings of poisonous milkweed butterflies of the genus Danaus. Today’s
textbook theory of Batesian mimicry is based on studies of a number of researchers (among others,
J.W.Z. Brower, C.A. Clarke, and P.M. Sheppard) and was further elaborated by D. Charlesworth and
B. Charlesworth in 1975.
In many ways this volume is a follow-up to Molecular Model Systems in the Lepidoptera
(Goldsmith and Wilkins 1995), a collection of articles published in 1995 that highlighted research
in selected current or emerging models with the aim of drawing the attention of the scientific com-
munity to their special qualities and experimental strengths. Adaptation of phytophagous lab-reared
animals to artificial diet and relatively large size were major conveniences at the time that placed
the silkworm in the forefront of basic research in countries where sericulture was deep-rooted and
established the tobacco hornworm, Manduca sexta, as the premier model in the United States for
many kinds of fundamental studies, though not genetics. Both insects remain central models in
the lepidopteran pantheon, with Bombyx remaining the most advanced in molecular genetics and
structural and functional genomics and Manduca still among the first in areas like innate immunity,
olfaction and neurobiology, endocrinology, and biochemistry. With advances in molecular tools and
genomic techniques, size and mass-rearing are no longer critical, so that the choice of lepidopteran
model systems, as seen in the current volume, increasingly reflects their advantages for illuminating

vii
viii Preface

experimental questions peculiar to butterflies and moths. We have not attempted to be comprehen-
sive, but again to provide a showcase for projects representing fundamental biological problems
and practical ones arising from a pressing need to find new ways to control lepidopteran pests. We
apologize for omitting important model systems and key areas of investigation, and trust that the
research presented here will inspire readers to look further into the literature for additional fascinat-
ing and inspiring stories.
This volume begins with an overview of the current status of lepidopteran phylogenetics
(Chapter 1), which is essential for placing what we know in an evolutionary context, with a focus
on clades representing model systems. Subsequent chapters review the richness and diversity of
genomic and post-genomic resources now available or under development for individual species
or genera, including the silkworm (Chapter 2), the remarkable Müllerian mimics, the Heliconius
butterflies (Chapter 6), and the major crop pest genus, Helicoverpa (Chapter 12). An important
new theme that has emerged since the earlier volume is the role of sexual dimorphism on many
aspects of lepidopteran biology, including chromosome structure, sex chromosome systems,
and sex determination (Chapters 3 and 4), and mating behavior associated with butterfly vision
(Chapter 7), circadian clocks (Chapter 8), and pheromone production and perception (Chapter 10).
Similarly, in the past decade significant progress has been made in uncovering genes controlling
the development of butterfly wings and wing patterns, which is gradually yielding information in
an evolutionary context (Chapters 5 and 6). Knowledge of the diversity in structure and function
of lepidopteran chemoreceptors in comparison with those of other major insect orders (Chapter
9) represents another recent breakthrough that has depended on the extensive genome sequencing
project in Bombyx.
Turning to another dominant theme, many topics covered in this volume are connected directly
or indirectly with the critical area of insect control. These include using Lepidoptera to explore
the genetics and neurobiology of host range specificity (Chapter 11), to define mechanisms of
insecticide resistance (Chapter 13), to augment already well-established fundamental knowledge
of the innate insect immune response (Chapter 14), and to explore the interactions between polyd-
navirus-carrying parasitoids and their lepidopteran hosts (Chapter 17). Examples of practical
applications in pest control include testing lepidopteran-derived antimicrobial peptides and viru-
lence factors as potential therapeutic agents against human pathogens or to produce disease-resis-
tant plants (Chapter 15), and delivering a variety of intrahemocoelic toxins by different means
(Chapter 16). Encompassing the final theme of virus delivery systems and function, we end with
the first reported case of map-based or positional cloning in a lepidopteran, the successful isola-
tion of nsd-2, a gene conferring resistance to the densovirus BmDNV-2, a serious pathogen in
sericulture (Chapter 18). This, like so many of the projects described in this volume, represents a
high point of molecular genetic research in Lepidoptera, using strategies, methods, and sequence
information that were barely conceived in the mid-1990s. With the recent publication of “build
2” of the Bombyx genome (see Special Issue on the Silkworm Genome, Insect Biochemistry
and Molecular Biology, 38(12), December 2008) and the prospects of many more lepidopteran
genome sequencing projects to follow, we look forward to being able to report on even more rapid
progress in the near future.
In closing we would like to express heartfelt thanks to Tom Miller, the editor for the Contemporary
Topics in Entomology Series of CRC Press, who conceived of the new Lepidoptera book project and
recruited us as editors; without his cheerful encouragement, constant cajoling, gentle arm twisting,
creative ways of helping us keep up momentum, and faith in us it is doubtful we could have got-
ten the job done. We would also like to thank John Sulzycki, senior editor at CRC Press, for his
infinite patience, and Pat Roberson, our Taylor & Francis Group production coordinator, for hers;
her helpful and timely responses to our questions moved the project from conception to reality in
ways she will probably never know. Our thanks are also due to Gail Renard for preparation of the
proofs, and to Kelly Pennoyer, who performed endless tedious tasks to get the manuscript ready for
the first stage of publication. And finally, grateful thanks to our many authors for their hard work,
Preface ix

timely submissions, gracious responses to all of our comments and suggestions, and enthusiastic
support of the project, despite busy schedules and occasional communication gaps on our end. Your
contributions are invaluable.

Marian R. Goldsmith and František Marec

Kingston and České Budějovice
About the Editors
Marian R. Goldsmith, Ph.D., is professor and chair of the Department of Biological Sciences
at the University of Rhode Island. Together with Fotis Kafatos, in 1988 she cofounded the
International Workshop on the Molecular Biology and Genetics of the Lepidoptera, which is held
every two to three years in Kolympari, Crete, bringing together lepidopteran researchers from
around the world and serving as the only forum of its kind. Working primarily as a geneticist to
bring molecular tools and techniques to bear on the domesticated silkworm as a model system, she
has established strong collaborations with silkworm groups in Japan, France, and China, starting
in her postdoctoral years when she first brought Bombyx mori to the laboratory of Fotis Kafatos at
Harvard University in 1972 to study regulation and expression of chorion genes after a two-month
stay in the laboratories of Toshio Ito at the Sericultural Experiment Station (now National Institute
of Agrobiological Sciences [NIAS]) and Yataro Tazima at the National Institute of Genetics in
Misima, sponsored by a National Science Foundation fellowship. This was followed by many sab-
batical leaves and shorter stays during which she established collaborations with Bungo Sakaguchi
and Hiroshi Doira at Kyushu University, Hideaki Maekawa at the National Institute of Health in
Tokyo, Toshiki Tamura and Wajirou Hara at the National Institute of Sericultural and Entomological
Science in Tsukuba, and Masahiko Kobayashi and Toru Shimada at Tokyo University, where she
was a visiting professor in 1997–1998. At that time she began a long-standing collaboration with
Kazuei Mita at NIAS helping to bring the silkworm genome project to fruition. More recently she
served as a guest professor at the Shanghai Institute of Plant Physiology and Ecology in the labora-
tory of Yongping Huang and Xuexia Miao, developing tools for mapping and positional cloning of
complex traits. She is a fellow of the American Association for the Advancement of Science, and
in 2002 received the Sanshi-gaku-sho Annual Prize of the Japan Society of Sericultural Science
for “Scientific Achievement Related to the Molecular Genetic Study on Chorion Gene Groups in
Bombyx mori.” She coedited Molecular Model Systems in the Lepidoptera (Cambridge University
Press) with Adam Wilkins in 1995.

František Marec, CSc., is senior researcher at the Institute of Entomology, Biology Centre of
the Academy of Sciences of the Czech Republic, and professor of molecular and cell biology and
genetics at the Faculty of Science, University of South Bohemia, both in České Budějovice, Czech
Republic. Since 1990 he has been a research fellow of the Alexander von Humboldt Foundation,
Bonn, Germany. He has had a long-standing collaboration with Walther Traut at the Institute of
Biology, Medical University of Lübeck, Germany, where he worked as a Humboldtian in 1991–1992
and 1998. At the end of the nineties he also established a strong collaboration with Ken Sahara at
the Laboratory of Applied Molecular Entomology at the Hokkaido University, Sapporo, Japan. Like
many geneticists, he started his career in Drosophila, being interested in mutation genetics, but
soon become fascinated by Lepidoptera genetics. His early research interests included chemical and
radiation mutagenesis targeted to the development of genetic control strategies against lepidopteran
pests, chromosomal mechanisms of resistance of lepidopteran species to ionizing radiation, and
meiotic chromosome pairing including formation of the synaptonemal complex, a unique nuclear
structure mediating intimate association of two homologous chromosomes. Most of his early work
was done in the flour moth Ephestia kuehniella, historically the second model of Lepidoptera genet-
ics after Bombyx mori. More recent projects have focused on molecular composition of insect telom-
eres and phylogeny of insect telomeric DNA repeats, and on the study of molecular differentiation
and evolution of sex chromosomes in Lepidoptera using advanced methods of molecular cytogenet-
ics. In addition, his co-operation with the Insect Pest Control Section, Joint FAO/IAEA Division

xi
xii About the Editors

of Nuclear Techniques in Food and Agriculture, International Atomic Energy Agency, Vienna,
Austria, resulted in a new approach for the construction of genetic sexing strains in lepidopteran
pests. Based on this approach, he works on the development of transgenic genetic sexing strains in
the codling moth, Cydia pomonella, aimed at generating male-only progeny for the control of this
pest using sterile insect technique or inherited sterility strategies. This work is done in collaboration
with Lisa G. Neven at the USDA—Agricultural Research Service, Yakima Agricultural Research
Laboratory, Wapato, Washington.
Contributors
Hiroaki Abe Nicola Chamberlain
Department of Biological Production School of Biological Sciences
Faculty of Agriculture University of Exeter in Cornwall
Tokyo University of Agriculture and Technology Penryn, United Kingdom
Tokyo, Japan
Derek Collinge
Joaquin Baixeras CSIRO Entomology
Cavanilles Institute of Biodiversity and
Canberra, Australia
Evolutionary Biology
and
University of Valencia
School of Biochemistry and Molecular Biology
Valencia, Spain
Australia National University
Philip Batterham Canberra, Australia
Department of Genetics
Bio21 Institute Michael P. Cummings
University of Melbourne Center for Bioinformatics and Computational
Parkville, Victoria, Australia Biology
University of Maryland
Simon W. Baxter College Park, Maryland
Department of Zoology
School of Biology Donald R. Davis
University of Cambridge Department of Entomology
Cambridge, United Kingdom
National Museum of Natural History
Patrícia Beldade Smithsonian Institution
Evolutionary Biology Group Washington, D.C.
Institute of Biology
Leiden University Richard H. ffrench-Constant
Leiden, The Netherlands School of Biological Sciences
University of Exeter in Cornwall
Bryony C. Bonning Penryn, United Kingdom
Department of Entomology
Iowa State University Tsuguru Fujii
Ames, Iowa Graduate School of Agriculture and Life
Science
Adriana D. Briscoe
The University of Tokyo
Department of Ecology and Evolutionary
Tokyo, Japan
Biology
University of California, Irvine
Marian R. Goldsmith
Irvine, California
Biological Sciences Department
John Brown University of Rhode Island
Systematic Entomology Laboratory Kingston, Rhode Island
USDA
National Museum of Natural History Karl Gordon
Smithsonian Institution CSIRO Entomology
Washington, D.C. Canberra, Australia

xiii
xiv Contributors

Fred Gould Owen McMillan

Department of Entomology and the Keck Department of Genetics
Center for Behavioral Biology North Carolina State University
North Carolina State University Raleigh, North Carolina
Raleigh, North Carolina
Christine Merlin
Astrid T. Groot Department of Neurobiology
Max Planck Institute for Chemical Ecology University of Massachusetts Medical School
Department of Entomology Worcester, Massachusetts
Jena, Germany
James B. Nardi
Department of Entomology
David G. Heckel
University of Illinois
Department of Entomology
Urbana, Illinois
Max Planck Institute for Chemical Ecology
Jena, Germany Sara J. Oppenheim
Department of Entomology
Keith R. Hopper North Carolina State University
United States Department of Agriculture Raleigh, North Carolina
Agricultural Research Service
Newark, Delaware Cynthia S. Parr
Human-Computer Interaction Lab
Chris D. Jiggins University of Maryland
Department of Zoology College Park, Maryland
School of Biology
University of Cambridge Jerome C. Regier
Cambridge, United Kingdom University of Maryland Biotechnology Institute
University of Maryland
Keiko Kadono-Okuda College Park, Maryland
National Institute of Agrobiological Sciences
Tsukuba, Japan Steven M. Reppert
Department of Neurobiology
Michael R. Kanost University of Massachusetts Medical School
Department of Biochemistry Worcester, Massachusetts
Kansas State University
Hugh M. Robertson
Manhattan, Kansas
Department of Entomology
University of Illinois at Urbana-Champaign
Akito Y. Kawahara
Urbana, Illinois
Department of Entomology
University of Maryland Amanda D. Roe
College Park, Maryland Department of Entomology
University of Minnesota
František Marec St. Paul, Minnesota
Biology Centre ASCR
Institute of Entomology Daniel Rubinoff
and Department of Plant and Environmental
Faculty of Science Protection Sciences
University of South Bohemia University of Hawaii
České Budějovice, Czech Republic Honolulu, Hawaii
Contributors xv

Suzanne V. Saenko Walther Traut

Evolutionary Biology Group Universität Lübeck
Institute of Biology Zentrum für Medizinische Strukturbiologie
Leiden University Institut für Biologie
Leiden, The Netherlands Lübeck, Germany

Ken Sahara Gissella M. Vásquez

Laboratory of Applied Molecular Entomology Department of Entomology and the Keck
Research Institute of Agriculture Center for Behavioral Biology
Hokkaido University North Carolina State University
Sapporo, Japan Raleigh, North Carolina

Coby Schal Andreas Vilcinskas

Department of Entomology and the Keck Institute of Phytopathology and Applied
Center for Behavioral Biology Zoology
North Carolina State University Interdisciplinary Research Center
Raleigh, North Carolina Justus-Liebig-University of Giessen
Giessen, Germany
Nina Richtman Schmidt
Department of Entomology Niklas Wahlberg
Iowa State University Department of Biology
Ames, Iowa University of Turku
Turku, Finland
Toru Shimada
Graduate School of Agriculture and Life Kevin W. Wanner
Science Department of Plant Sciences and Plant
The University of Tokyo Pathology
Tokyo, Japan Montana State University
Bozeman, Montana
Thomas J. Simonsen
Department of Biological Sciences Susan J. Weller
University of Alberta Department of Entomology
Edmonton, Alberta, Canada University of Minnesota
St. Paul, Minnesota
Marilou P. Sison-Mangus
Department of Ecology and Evolutionary Adam Williams
Biology Department of Genetics
University of California, Irvine Bio21 Institute
Irvine, California University of Melbourne
Parkville, Victoria, Australia
Michael R. Strand
Department of Entomology Andreas Zwick
University of Georgia Department of Entomology
Athens, Georgia University of Maryland
College Park, Maryland
Wee Tek Tay
CSIRO Entomology
Canberra, Australia
 1 Evolutionary Framework for
Lepidoptera Model Systems
Amanda D. Roe, Susan J. Weller, Joaquin Baixeras,
John Brown, Michael P. Cummings, Donald R. Davis,
Akito Y. Kawahara, Cynthia S. Parr, Jerome C. Regier,
Daniel Rubinoff, Thomas J. Simonsen, Niklas Wahlberg,
and Andreas Zwick

Contents
Introduction.........................................................................................................................................1
Phylogenetics and Model Systems.................................................................................................3
Overview of Lepidopteran Phylogeny...........................................................................................4
Overview of Selected Ditrysian Superfamilies..............................................................................4
Tortricoidea...............................................................................................................................5
Pyraloidea..................................................................................................................................7
Papilionoidea and Hesperioidea................................................................................................8
Bombycoidea.............................................................................................................................9
Noctuoidea.............................................................................................................................. 13
Current Research and Future Directions........................................................................................... 14
Molecular Ditrysian Phylogenies and the Role of Fossils........................................................... 14
Virtual Community Building in Lepidopteran Systematics......................................................... 17
Summary and Future Model Systems.......................................................................................... 18
Acknowledgments............................................................................................................................. 18
References......................................................................................................................................... 19

INTRODUCTION
Lepidoptera are among the most diverse and easily recognized organisms on the planet, with at
least 150,000 described species (Kristensen and Skalski 1998). They are one of the four mega-
diverse orders of holometabolous insects, together with Diptera (flies), Coleoptera (beetles), and
Hymenoptera (wasps, bees, and ants). Butterflies alone are more numerous than birds, Class Aves,
with approximately 18,000 species (Kristensen and Skalski 1998). Generally, Lepidoptera are char-
acterized by the presence of scaled wings, elongate sucking mouthparts (proboscis), and complete
(holometabolous) development where the larval stages are commonly referred to as “caterpillars.”
Historically, species of Lepidoptera have proven invaluable model systems in the fields of develop-
ment, genetics, molecular biology, physiology, evolution, and ecology (e.g., Bates 1861; Müller 1879;
Ford 1964; Ehrlich and Raven 1967; Kettlewell 1973). Interest in Lepidoptera species as model
systems stems from a number of biological characteristics that render this group amenable for study
(Bolker 1995). Lepidopterans are charismatic, due mainly to their striking variety of wing color
patterns and larval morphologies, and they are avidly collected by professionals and amateurs alike

1
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