SYSTEMATICS OF THE TRIBE (: : ) INCLUDING A REVIEW OF THE HARRIS

By

CLARE HILARY SCOTT

A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

2012

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© 2012 Clare H. Scott

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To my family whose support throughout this long journey has been invaluable

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ACKNOWLEDGMENTS

This research would not have been possible without the support of numerous

people and institutions. I would like to acknowledge the support and advice of my major

advisor and committee chair, Dr. Marc Branham, and the other members of my

graduate committee, Drs. Rebecca Simmons, Jackie Miller, David Reed, and Christine

Miller.

I would like to thank Drs. Susan Weller and Jennifer Zaspel for giving me the

opportunity to participate in their molecular phylogeny of Arctiinae. Furthermore, I wish

to acknowledge the National Science Foundation Award DEB#0919185 for the partial financial support of this research. Without the funding Drs. Weller and Zaspel received, the molecular phylogeny of Lithosiini would not have been completed. In addition, the work of Pablo Chialvo, Taylor Wardwell, and Elizabeth Phillippi on this study was invaluable. Pablo helped to amplify the COI and 28S gene fragments and willingly came into the lab on the weekend and holidays to help with this project. Taylor and Elizabeth helped to amplify the CytB and RpS5 gene fragments. In addition, Taylor dealt with submitting all fragments for sequencing.

I would like to acknowledge all of the collections listed in Chapters 2-5 for the material that they provided. Without the use of these specimens, the morphological studies would not have been possible, and the taxon sampling of the molecular study would not have been as broad. Further thanks are offered to the American Museum of

Natural History, the Carnegie Museum of Natural History, the Cornell University

Collection, the Florida Museum of Natural History, the Museum of Comparative Zoology

(Harvard), the United States Museum of Natural History, and the Yale Peabody

Museum. Each of these institutes was kind enough to allow me to visit and examine

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their holdings of several families of Lepidoptera, while looking for specimens of

Lycomorpha. The tribal phylogenetic analyses were conducted using the University of

Florida Phyloinformatics Cluster for High Performance Computing in the Life Sciences and the CIPRES Science Gateway (Miller et al. 2010). Without access to these computer clusters, the time needed to run the tribal phylogenetic analysis would have been prohibitively long.

This research was funded in part by the Museum of Comparative Zoology Ernst

Mayr Travel Grant in Systematics, the Systematics, Evolution, and Biodiversity

Section’s Student Travel Award of the Entomological Society of America, the University of Florida Alumni Fellowship, and the University of Florida SPICE fellowship.

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TABLE OF CONTENTS

page

ACKNOWLEDGMENTS ...... 4

LIST OF TABLES ...... 10

LIST OF FIGURES ...... 11

ABSTRACT ...... 14

CHAPTER

1 INTRODUCTION ...... 16

Literature Review ...... 16 Larval Feeding Behavior...... 17 Chemical Defense ...... 19 Courtship Behavior ...... 20 Research Objectives ...... 22 Chapter 2 ...... 22 Chapter 3 ...... 23 Chapter 4 ...... 24 Chapter 5 ...... 25 Chapter 6 ...... 26

2 A PRELIMINARY PHYLOGENY OF THE LICHEN MOTH TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE) BASED ON MORPHOLOGICAL CHARACTERS ...... 27

Background Information ...... 27 Materials and Methods...... 31 Taxon Sampling ...... 31 Morphology ...... 32 Characters Examined ...... 33 Characters and their Phylogenetic Usefulness ...... 33 Head ...... 33 Thorax ...... 34 Wings ...... 35 Abdomen...... 36 Male abdomen and genitalia ...... 37 Female abdomen and genitalia ...... 42 Phylogenetic Analysis ...... 44 Results ...... 45 Discussion ...... 49

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3 MOLECULAR PHYLOGENY OF THE TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE) ...... 83

Background Information ...... 83 Materials and Methods...... 87 Taxon Sampling ...... 87 DNA Extraction, Amplification, and Sequencing ...... 88 Phylogenetic Methods ...... 91 Results and Discussion...... 92

4 PHYLOGENY OF THE LICHEN MOTH GENUS LYCOMORPHA HARRIS (LEPIDOPTERA: EREBIDAE: ARCTIINAE) ...... 109

Background Information ...... 109 Materials and Methods...... 112 Taxon Sampling ...... 112 Collections Consulted ...... 113 Morphology ...... 114 Characters Examined ...... 115 Phylogenetic Analysis ...... 115 Results ...... 117 Phylogenetic Analysis and Taxonomic Implications ...... 117 Species Checklist of Lycomorpha ...... 122 Checklist of Species Described as Lycomorpha ...... 123 Lithosiini ...... 123 ...... 124 ...... 125 Incertae sedis ...... 126 Characters and their Phylogenetic Usefulness ...... 126 Head ...... 126 Thorax ...... 127 Wings ...... 127 Abdomen...... 128 Male abdomen and genitalia ...... 128 Female abdomen and genitalia ...... 134 Characters examined but excluded from the analysis ...... 137

5 A REVISION OF THE GENUS LYCOMORPHA HARRIS (LEPIDOPTERA: EREBIDAE: ARCTIINAE: LITHOSIINI) INCLUDING SPECIES FORMERLY PLACED IN THE GENUS PROPYRIA HAMPSON (LEPIDOPTERA: EREBIDAE: ARCTIINAE: LITHOSIINI) ...... 167

Background Information ...... 167 Taxonomic History of Lycomorpha and Propyria ...... 169 Synonymic Checklist of the Genus Lycomorpha ...... 172 Materials and Methods...... 173 Material Examined ...... 173

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Morphology ...... 174 Species Descriptions ...... 175 Systematic Entomology of the Genus Lycomorpha ...... 176 Lycomorpha atroxantha (Schaus) ...... 180 Lycomorpha concolor Scott New Species ...... 184 Lycomorpha fulgens (Henry Edwards) ...... 190 Lycomorpha grotei (Packard) ...... 197 Lycomorpha miniata Packard ...... 203 Lycomorpha morelosia (Schaus) ...... 209 Lycomorpha neomexicanus Scott New Species ...... 214 Lycomorpha normani (Schaus) ...... 218 (Drury) ...... 221 Lycomorpha ptychoglene (Hampson) ...... 227 Lycomorpha pulchra Dyar ...... 232 Lycomorpha regulus (Grinnell) ...... 238 Lycomorpha splendens Barnes and McDunnough ...... 244 Lycomorpha texanus Scott New Species ...... 248

6 DISCUSSION AND FUTURE DIRECTIONS ...... 268

Tribal Study ...... 268 Review of Lycomorpha ...... 270 Future Directions ...... 271 Tribal Systematics ...... 271 Chemical Defense ...... 272 Courtship Behavior of Lycomorpha ...... 273

APPENDIX

A MORPHOLOGICAL DATA MATRIX FOR TRIBAL PHYLOGENY ...... 274

B CHARACTER DIAGNOSTICS FOR STRICT CONSENSUS TREE ...... 279

C DATA MATRIX USED TO PRODUCE TREES BASED ON MOLECULAR DATA 281

D LIST OF CHARACTERS AND STATES IN PHYLOGENY OF LYCOMORPHA ... 335

Head ...... 335 Thorax ...... 335 Wings ...... 335 Abdomen ...... 335 Male Abdomen and Genitalia ...... 336 Male Abdomen ...... 336 Genital Capsule ...... 336 Phallus...... 338 Female Abdomen and Genitalia ...... 338

E ALL SPECIES DATA MATRIX FOR LYCOMORPHA PHYLOGENY ...... 341

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F COLLECTION LOCALITY DATA FROM SPECIMENS USED TO ILLUSTRATE THE ADULT HABITUS OF LYCOMORPHA ...... 342

LIST OF REFERENCES ...... 345

BIOGRAPHICAL SKETCH ...... 360

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LIST OF TABLES

Table page

2-1 Comparative table of generic groups of Lithosiini ...... 54

2-2 Ingroup taxon sampling ...... 55

2-3 Species examined in the phylogenetic analysis ...... 56

3-1 List of species used in the analysis ...... 99

3-2 PCR protocol data ...... 102

3-3 Optimal partitioning schemes ...... 103

3-4 Interpretation of the Bayes Factor ...... 104

4-1 Historical family treatment of Lycomorpha ...... 140

4-2 Species included in the phylogenetic analysis ...... 141

4-3 Character support for major ingroup clades...... 143

4-4 Results of Outgroup Jackknife Analysis ...... 145

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LIST OF FIGURES

Figure page

2-1 Sample of lithosiine species included in the study ...... 61

2-2 Lateral view, head ...... 62

2-3 Prothoracic leg ...... 63

2-4 Male hindwing ...... 64

2-5 A2 sternite ...... 65

2-6 Proximal margin of male eighth sternite, Part A...... 66

2-7 Proximal margin of male eighth sternite, Part B...... 67

2-8 Tegumen location ...... 68

2-9 Tegumen fusion ...... 69

2-10 Pleural sclerites ...... 70

2-11 Juxta ...... 71

2-12 External view, left valva ...... 72

2-13 Internal view, right valva ...... 73

2-14 External view, left valva (A,B) and Internal view, right valva (C,D) ...... 74

2-15 Lateral view, left side, phallus ...... 75

2-16 Abdominal pelt (A,C) and female genital capsule (B,D), Part A ...... 76

2-17 Abdominal pelt (A,C) and female genital capsule (B,D), Part B ...... 77

2-18 Abdominal pelt (A) and female genital capsule (B), Part C ...... 78

2-19 Ventral female genital capsule...... 79

2-20 Bursa copulatrix ...... 80

2-21 Strict consensus of the 4408 MP trees ...... 81

2-22 Majority rule consensus tree of the Bayesian analysis ...... 82

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3-1 Maximum likelihood tree from analysis of AIC, AICc optimal partitioning scheme ...... 105

3-2 Maximum likelihood tree from analysis of BIC optimal partitioning scheme ..... 106

3-3 Majority rule consensus tree of the Bayesian analysis using the AIC, AICc optimal partitioning scheme ...... 107

3-4 Majority rule consensus tree of the Bayesian analysis using the BIC optimal partitioning scheme ...... 108

4-1 Species mistakenly described as members of Lycomorpha ...... 146

4-2 Lycomorpha species with red forewings and primarily black hindwings ...... 147

4-3 Strict consensus of 3 trees (L=192, CI=0.68, RI=0.71) resulting from the MP analysis of the MF matrix ...... 148

4-4 Strict consensus of 3 trees (L=192, CI=0.68, RI=0.73) resulting from the MP analysis of the AS matrix ...... 149

4-5 Bayesian Inference consensus tree from the analysis of the AS matrix ...... 150

4-6 Strict consensus of 9 trees (L=191, CI=0.68, RI=0.69) resulting from the exclusion of outgroup taxon ...... 151

4-7 Strict consensus of 3 trees (L=184, CI=0.70, RI=0.72) resulting from the exclusion of outgroup taxon Ptychoglene ...... 152

4-8 Strict consensus of 6 trees (L=186, CI=0.68, RI=0.70) resulting from the exclusion of outgroup taxon ...... 153

4-9 Strict consensus of 3 trees (L=179, CI=0.70, RI=0.74) resulting from the exclusion of outgroup taxon Lycomorphodes ...... 154

4-10 Strict consensus of 6 trees (L=175, CI=0.70, RI=0.75) resulting from the exclusion of outgroup taxon Hypermaepha ...... 155

4-11 Strict consensus of 9 trees (L=190, CI=0.67, RI=0.70) resulting from the exclusion of outgroup taxon Dolichesia ...... 156

4-12 Strict consensus of 24 trees (L=198, CI=0.66, RI=0.71) resulting from constraining the monophyly of Lycomorpha ...... 157

4-13 Strict consensus of 30 trees (L=194, CI=0.67, RI=0.72) resulting from constraining the monophyly of Propyria ...... 158

4-14 Male antennal flagellomeres ...... 159

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4-15 Female antennal flagellomeres...... 160

4-16 State of gena ...... 161

4-17 Labial palps ...... 162

4-18 Forewing venation ...... 163

4-19 Hindwing venation ...... 164

4-20 Second abdominal sternite ...... 165

4-21 A7/A8 androconia ...... 166

5-1 Strict consensus of 3 most parsimonious trees (L=192, CI=0.68, RI=0.73) resulting from the maximum parsimony analysis of the all species (AS) dataset ...... 255

5-2 Images of the adult habitus of the genus Lycomorpha, Plate 1 ...... 256

5-3 Images of the adult habitus of the genus Lycomorpha, Plate 2 ...... 257

5-4 Images of the adult habitus of the genus Lycomorpha, Plate 3 ...... 258

5-5 Images of the adult habitus of the genus Lycomorpha, Plate 4 ...... 259

5-6 Images of the adult habitus of the genus Lycomorpha, Plate 5 ...... 260

5-7 Adult habitus of female L. pelopia ...... 260

5-8 Male flagellomere state ...... 261

5-9 Female flagellomere state ...... 262

5-10 Labial palp segment fusion ...... 263

5-11 Medial veins of the forewing ...... 264

5-12 Medial and cubital veins of hindwing ...... 265

5-13 Anterolateral process (ALP) form ...... 266

5-14 Androconia of the A7/A8 intersegmental membrane ...... 267

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Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy

SYSTEMATICS OF THE LICHEN MOTH TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE) INCLUDING A REVIEW OF THE GENUS LYCOMORPHA HARRIS

By

Clare Hilary Scott

December 2012

Chair: Marc A. Branham Major: Entomology and Nematology

The tribe Lithosiini is composed of an estimated 5,000 to 6,000 species, which are distributed worldwide. Lithosiini represents the most biodiverse lineage of lichen feeders within Lepidoptera. The tribe is well known for this larval feeding behavior, but no studies have been conducted to examine the evolutionary relationships among the genera of Lithosiini. Only one study has been completed that examined the systematics of a lithosiine genus. However, changes continue to be made to the of both the tribe and the genera that compose it. In this study, phylogenies of the tribe were constructed based on datasets composed of either morphological or molecular data.

These phylogenies were used to examine the evolutionary relationships among the genera of the tribe and to determine whether the subtribes are monophyletic. The datasets were analyzed using maximum parsimony, Bayesian inference, and maximum likelihood methods. The phylogenies obtained based on the adult morphology are almost entirely unresolved, but Lithosiini is recovered as monophyletic. The topologies recovered in the molecular phylogeny are more resolved. However, there is no nodal support for the deeper nodes, and Lithosiini is not found to be monophyletic. None of

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the phylogenies produced by either dataset recovers the subtribes of Lithosiini as monophyletic.

In addition to the higher-level studies of the tribe, a review of the genus

Lycomorpha was completed. A phylogeny was constructed based on adult morphological characters. This phylogeny demonstrates that Lycomorpha is not reciprocally monophyletic with respect to the lithosiine genus Propyria. Based on these findings the two genera are synonymized. The name Lycomorpha is given priority based on seniority. Following these changes in the classification of the genus, a revision of the genus Lycomorpha was completed. Descriptions are provided for both the genus and fourteen of the twenty species within Lycomorpha. Three new species are described as part of this study.

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CHAPTER 1 INTRODUCTION

Literature Review

The subfamily Arctiinae Leach is comprised of over 11,000 species, which are cosmopolitan in distribution. This subfamily has been hypothesized to contain at least five tribes, Lithosiini, Arctiini, Ctenuchini, Pericopini, and Thyretini, each of which has formerly been treated as a separate family. In order to assess the monophyly of these five tribes, Jacobson & Weller (2002) constructed a phylogeny of the subfamily using adult and larval morphological characters. The resulting phylogeny recovered Arctiinae

as a monophyletic lineage that was supported by two larval and three adult characters.

The tribe Lithosiini was found to be monophyletic, and eight synapomorphies defined

this lineage. Arctiini sensu latu was found to be paraphyletic with respect to Pericopini

and Ctenuchini. Jacobson & Weller (2002) redefined the tribe Arctiini to include these

lineages. In addition, the phylogeny demonstrated that Ctenuchini was neither

monophyletic nor natural. The Syntomiina of Ctenuchini was found to be sister to

Thyretini, while the remaining species of Ctenuchini were recovered in Arctiini. The

monophyletic Syntomiina+Thyretini clade, Syntomiini, was recovered sister to Lithosiini.

Previously, Bendib & Minet (1998) had hypothesized that Syntomiini was sister to

Arctiini. Although the phylogenetic analysis of the morphological characters found

Lithosiini and Syntomiini to be sister, studies conducted using molecular data (Zahiri et

al. 2011, 2012) have recovered Syntomiini sister to Arctiini. In addition, these molecular

phylogenies recovered Lithosiini as the basal most clade within Arctiinae. Despite

finding different relationships among the tribes, the monophyly of Lithosiini, Arctiini, and

Syntomiini has been supported by both morphological and molecular phylogenetic

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studies. However, the relationships among the genera that comprise these tribes remain in flux. This is especially true of the tribe Lithosiini, which is composed of 457 genera.

No empirical studies have been conducted that examine the systematics of this tribe.

However, authors (Bendib & Minet 1999; Birket-Smith 1965; Forbes 1960; Franclemont

1983) all suggested subtribal relationships among the genera of Lithosiini. This lack of resolved phylogenetic relationships among the genera of each tribe make it impossible to study the evolution of chemical defense or courtship behavior within these tribes.

Larval Feeding Behavior

Within the subfamily Arctiinae, the larvae range from monophagous (e.g.

Utetheisa ornatrix (Linnaeus) feeds on only Crotalaria spp.) to highly polyphagous (e.g.

Estigmene acrea (Drury) feeds on more than sixty-five larval host plants (Hartmann et al. 2005)). Dyer et al. (2007) found that as latitude decreases, there is an increase in the number of monophagous species of Arctiinae. The larvae of the tribe Arctiini have been reported to feed on the tissue of both angiosperms and gymnosperms. Many of the species within this tribe are capable of consuming host plants that are defended with pyrrolizidine alkaloids (PA). In addition, some species can feed on plants containing cardiac glycosides (CG). The larvae that consume plants containing either PA or CG sequester these chemicals within their bodies for defense. Weller et al. (1999) hypothesized that the ability to sequester PA from the larval host plant was present in the ancestor of Arctiini. Although the ability to consume plants containing PA occurs throughout the tribe, not all species that sequester PA can successfully reach adulthood feeding only on plants containing these compounds (Bernays et al. 2002). Few reports exist for the larval feeding behavior of the small (~200 species) tribe Syntomiini. The larvae of this tribe have been found to feed on an assortment of plant families, dead

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plant material, and in some species, and (Weller et al. 2009). The tribe

Lithosiini, which is estimated to contain 5,000 to 6,000 species, is hypothesized to

represent the most biodiverse lineage of lichen feeders within Lepidoptera. Although

lichenivory has evolved several times within the ditrysian Lepidoptera (Rawlins 1984),

the majority of the lineages that practice this behavior only consist of of approximately

two-dozen species (Wagner et al. 2008). However, debate exists over whether the

larvae of Lithosiini are true lichen feeders or graze solely on the algal component of

lichens. In their feeding experiments with Eilema complana (Linnaeus), Hesbacher et al.

(1995) found that early instar larvae fed exclusively on the cortical and algal layers of

the lichen Cladonia pyxidata. By consuming these tissues of the lichen, the larvae

avoided feeding on the fungal component of the lichen. In other feeding experiments

conducted using Eilema species, the larvae were found to prefer those lichens that

possess fewer polyphenolic compounds (Pöykkö & Hyvärinen 2003; Pöykkö et al.

2005). The results of these feeding experiments suggest that lithosiine larvae are

algivores. However, Hesbacher et al. (1995) detected the presence of lichen phenolics

that are only produced by the fungal component of the lichen within the adults of eleven

lithosiine species representing five genera. Furthermore, Pöykkö & Hyvärinen (2003)

noted that while the larvae fed preferentially on a lichen species without polyphenolic

substances, they would forage on other lichens that contained polyphenolics. This

behavior suggests that larvae of Lithosiini selectively feed on the fungal components of

lichens to obtain defensive chemicals but require other nutritional factors to achieve

adulthood.

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Chemical Defense

The use of chemical defenses against predators and parasitoids is found throughout each of the three tribes of the subfamily Arctiinae. The chemicals used for defense can be sequestered by the larvae from their host, produced de novo by either the adults or larvae, or obtained by an adult visiting a plant that contains the desired defensive chemical (e.g. pharmacophagy, as defined by Boppré 1984). Within Arctiinae, the most commonly found defensive chemicals are PA, CG, iridoid glycosides (IG), and biogenic amines (BA) (Weller et al. 1999; Nishida 2002). Hristov & Conner (2005) found that species, which possessed CG, were the most unpalatable to the big brown bat,

Eptesicus fuscus, followed by species with PA. Species of Arctiinae that are chemically defended frequently advertise their distastefulness with bright color patterns. In addition, some of these species also use ultrasonic clicks to warn off auditory predators such as bats.

All of the previously mentioned chemicals can be found within different members of the tribe Arctiini. The ability to sequester PA is hypothesized to have been present in the ancestor of the tribe (Weller et al. 1999). PA can be obtained through the larval host plant or by pharmacophagy in the adults. Some species (e.g. E. acrea) are defended in

all life stages by PA (Hartmann et al. 2004). In addition, Hartmann et al. (2004) found

that males will transfer their PA load to the female while mating, and the females

transfer their PA load to the eggs. CG and IG are obtained from the larval host plant.

However, BA are produced de novo and provide the least protection (Weller et al. 1999;

Hristov & Conner 2005). Less is known about the chemical compounds found in the

tribe Syntomiini. The species phegea Linnaeus was found to possess BA, and

other species of Amata have been found to possess pyrazines (Weller et al. 1999). The

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only chemicals that have been found in the tribe Lithosiini are lichen phenolics.

Hesbacher et al. (1995) examined 103 adults of sixteen species representing eight lithosiine genera using HPLC analysis to test for the presence of lichen polyphenolics.

Individuals of eleven species representing five genera possessed known lichen phenolics and other lichen-derived compounds. An additional six species were found that possessed only lichen-derived compounds. However, this survey (Hesbacher et al.

1995) was not broad enough to determine how frequently the sequestration of lichen polyphenolics occurs within Lithosiini. Although the frequency of chemical defenses within Lithosiini has not been broadly documented, the adults of both brightly colored, mimetic lineages (e.g. Hypoprepia Hübner) and drably colored individuals (e.g. Agylla

Walker) have been found to be unpalatable to bats and birds (Acharya & Fenton 1992;

Collins & Watson 1983; Sargent 1995). Furthermore, Hristov & Conner (2005) found that Hübner was equally as unpalatable as Arctiini species that possess CG.

Courtship Behavior

Throughout the subfamily Arctiinae, the males of some species have been found to use scent (pheromones), sound, and/or a combination of both in their courtship rituals. When males use pheromones, these chemicals are released using eversible pheromone glands (coremata) or scent scales (androconia). Androconia occur in numerous forms throughout Arctiinae including hairbrushes on the male forelegs

(Schneider et al. 1999), patches of modified wing scales (Jacobson & Weller 2002;

Holloway 2002), and subabdominal pouches (Weller et al. 2000). The sounds produced in courtship occur at ultrasonic frequencies and are created by buckling modified cuticular plates present on the metathorax (tymbal organs). These are the same

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structures used to create the ultrasonic clicks that advertise the presence of chemical defenses to auditory predators.

Both types of courtship are found to occur in the tribe Arctiini. The pheromones used by almost all of the males of this tribe have been found to contain one of two dihydropyrrolizines: hydroxydanaidal or danaidal (Schulz 2009). Both of these compounds are derived from PA. Both monophagous and polyphagous species of

Arctiini that consume plants containing PA produce PA-derived pheromones. However, the larvae of polyphagous species do not always encounter plants containing PA. When the male larvae of these species (e.g. Creatonotus gangis Linnaeus, Creatonotus transiens (Walker), E. acrea) do not obtain PA, their coremata do not develop fully

(Boppré & Schneider 1985; Jordan et al. 2007; Schneider et al. 1982). Males that lack fully developed coremata do not engage in lekking behavior (Jordan et al. 2005; Willis &

Birch 1982; Wunderer et al. 1986). In addition, PA can be found in species whose larvae no longer feed on plants containing these compounds (e.g. Cisseps fulvicullis

Hübner). In the species C. fulvicullis, pharmacophagous adult males obtain the compounds, which are then transferred to the female as a nuptial gift (Doussourd 1986).

Although the use of pheromones derived from PA is the most common courtship strategy found within Arctiini, some species have been found to use pheromones that are not derived from PA (e.g. Cycnia in Conner 1987) or rely solely on ultrasound (e.g.

Syntomeida epilais Walker in Sanderford & Conner 1990, 1995).

Although the courtship behaviors of the tribe Arctiini have been well studied, very little is known of the courtship behaviors of the tribes Syntomiini and Lithosiini. Males of

Syntomiini possess androconial organs (Schneider et al. 1999). However, the chemicals

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present in their pheromones are unknown. Within the tribe Lithosiini, the males possess a wide range of scent disseminating structures that include coremata, modified wing scale patches, and androconial pouches on both the abdomen and genital capsule

(pers. obs.). As in Syntomiini, the pheromones contained in these structures are unknown. However, the Lithosiini have been found to be able to sequester lichen phenolic compounds for defense. It is possible that, like Arctiini, they have co-opted their defensive chemicals for courtship. In addition, adult Lithosiini frequently possess tymbal organs. Černý (1990) demonstrated that the ultrasound produced by species of

Setina Schrank was used in their courtship rituals. Until more data is obtained for the tribes Syntomiini and Lithosiini, the evolution of chemical defense and courtship in the subfamily Arctiinae cannot be studied. In addition, a resolved phylogeny of Lithosiini will be needed to study the evolution of these behaviors within this tribe.

Research Objectives

Chapter 2

The tribe Lithosiini is estimated to contain approximately 5,000 to 6,000 species.

These species are placed within 457 genera and are cosmopolitan in distribution.

Lithosiini is well known for the feeding behavior of its larvae: lichenivory. In addition, the

adults of some species have been found to sequester lichen-phenolics (Hesbacher et

al. 1995). The tribe Lithosiini is the only lichen-feeding lineage that sequesters these

compounds. It is hypothesized that the lichen phenolics are used for defense, and many

of the adults are aposematically colored. Although the tribe is well known for its

association with lichens, very little is known about the evolutionary relationships of the

tribe at any level. Phylogenetic studies completed using either morphological or

molecular data (Jacobson & Weller 2002; Zahiri et al. 2012) have recovered Lithosiini

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as one of three monophyletic tribes within the subfamily Arctiinae. However, the taxon sampling of these studies was not dense enough to make statements about the evolutionary relationships among the genera. Bendib & Minet (1999) proposed the existence of seven subtribes within Lithosiini based on larval, pupal, and adult morphological characters. However, less than a quarter of the genera of Lithosiini have been placed in a subtribe. The primary objective of this chapter was to construct a phylogeny of the tribe Lithosiini using adult morphological characters. The resulting phylogenies were used to examine the evolutionary relationships among genera belonging to each of the seven subtribes and genera that have not yet been placed in a subtribe. The specific questions addressed in this study were: 1) Is the tribe Lithosiini monophyletic? 2) Do the subtribes proposed by Bendib & Minet (1999) represent monophyletic clades? 3) Can the unplaced genera included in this analysis be associated with any of the seven subtribes? 4) What type of phylogenetic signal is present within adult morphological characters?

Chapter 3

To accurately study the evolution of behaviors and assess the monophyly of the subtribes, a resolved phylogeny of Lithosiini is needed. As noted in Chapter 2, no phylogenetic hypotheses have been proposed for the tribe. A phylogenetic study of

Lithosiini was completed using adult morphological characters (Chapter 2; Scott &

Branham In Press). However, the phylogenies recovered through the analysis of these characters were almost entirely unresolved. In other superfamilies, families, and subfamilies of Lepidoptera, phylogenetic analyses conducted using molecular datasets have been able to resolve the higher-level relationships that morphological studies could not (Regier et al. 2012; Ugelvig et al. 2011; Wahlberg et al. 2003, 2005; Zahiri et

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al. 2011, 2012). The primary objective of this chapter was to construct a phylogenetic hypothesis of Lithosiini using a molecular dataset composed of four gene fragments.

These gene fragments included two mitochondrial markers, cytochrome oxidase C subunit I (COI) and cytochrome oxidase B (CytB), and two nuclear markers, ribosomal protein S5 (RpS5) and the nuclear large subunit rRNA 28S D2 loop (28S). Each of these gene fragments has been shown to be phylogenetically informative at the higher- level within Lepidoptera (Wahlberg et al. 2009; Mutanen et al. 2010; Zahiri et al. 2011,

2012). The following questions were addressed in this research: 1) Is the phylogenetic signal present within the molecular dataset strong enough to resolve the higher-level relationships of Lithosiina? 2) Is the tribe Lithosiini monophyletic? 3) Are the subtribes

Cisthenina, Lithosiina, Nudariina, and Acsalina monophyletic? 4) What effect does the selection criterion used to partition the dataset and select models of evolution have on the phylogeny that is recovered?

Chapter 4

Phylogenetic hypotheses concerning the evolutionary relationships both among and within the genera of Lithosiini are lacking. Similar wing color patterns occur among some genera (e.g. Moore, Walker, and Adites Moore), which makes determining their generic limits difficult. In addition, other genera are known to represent non-monophyletic groups (e.g. Eilema Hübner). To date, only one phylogenetic study has been completed that assessed the systematics of a lithosiine genus (Jacobson

1995). However, authors (Durante 2008, 2009, 2012; Holloway 2002; Knowlton 1967) continue to publish revisionary works not based on phylogenetic analysis. The issues presented above can be seen in the lithosiine genus Lycomorpha Harris. The adults are aposematically colored and form mimicry complexes with other Lithosiini, Arctiini, and

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Zygaenidae as well as lycid beetles. In addition, the adult wing color patterns led to this genus being mistakenly placed in several families. Lycomorpha was transferred to

Lithosiini by Forbes (1960) who noted that the loss of an Sc vein was the only character that supported its placement in Euchromiidae. Jacobson & Weller (2002) confirmed this placement within the Lithosiini with their phylogenetic analysis of Arctiinae. Draudt

(1917) noted similarities between the color pattern and body shape of Lycomorpha and

the lithosiine genera Propyria Hampson and Ptychoglene Felder. In the morphological

phylogeny of the tribe, species of Lycomorpha and Propyria were found within a single

clade (Chapter 2; Scott & Branham In Press). No phylogenetic studies have been

completed that examine either the evolutionary relationships of the species of

Lycomorpha or the generic limits of this genus. The primary objective of this study was

to construct a phylogeny of the genus Lycomorpha using adult morphological

characters. The phylogeny that was obtained was used to address the following

questions: 1) Is Lycomorpha monophyletic? 2) What is the relationship between

Lycomorpha and Propyria? 3) What are the relationships among the species of

Lycomorpha?

Chapter 5

The phylogeny that was constructed for Lycomorpha using adult morphological

characters found the genus to be paraphyletic with respect to Propyria (Chapter 4).

Based on these findings, the two genera were synonymized. The older name,

Lycomorpha, was given priority. The primary objective of this chapter was to produce a

revision of the genus Lycomorpha. The goals of the revision were to produce a

description and diagnosis of the genus Lycomorpha, as well as fourteen of the twenty

species placed in it. As part of the revision, three new species were described. The

25

descriptions and diagnoses provide characters that can be used to place species within the genus and separate species with similar wing color patterns. Data on the distribution and biology of each species is provided in each species description.

Chapter 6

The objective of this chapter was to discuss the results that were obtained from the studies completed in Chapters 2-5. This chapter provided justification for the methods used in the phylogenetic analyses of the tribe. In addition, it offered directions that future research could pursue.

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CHAPTER 2 A PRELIMINARY PHYLOGENY OF THE LICHEN MOTH TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE) BASED ON MORPHOLOGICAL CHARACTERS

Background Information

Lepidopteran larvae have evolved a wide variety of feeding habits, which include consuming the fungal symbiont of lichens (lichenivory). This feeding behavior has arisen several times within the ditrysian Lepidoptera (Rawlins 1984). The majority of the licheniverous lineages are composed of less than two-dozen species, but the lichen moth tribe Lithosiini (Lepidoptera: Erebidae: Arctiinae) contains approximately 3,150 described species. Furthermore, the Lithosiini are the only lineage known to be capable of sequestering phenolics produced by the fungal symbiont of the lichen (Hesbacher et al. 1995; Wagner et al. 2008). Both the adults and larvae are thought to use the phenolics as defensive compounds (Weller et al. 1999). The adults of both brightly colored, mimetic lineages (e.g. Hypoprepia Hübner) and drably colored individuals (e.g.

Agylla Walker) have been found to be unpalatable to bats and birds (Acharya & Fenton

1992; Collins & Watson 1983; Sargent 1995). Although the tribe is well known for its larval feeding behavior, no work has been done to clarify the evolutionary relationships among the genera. Until a phylogeny is reconstructed, it will not be possible to study the evolution of lichenivory and other traits in the life history of these .

The classification of the tribe Lithosiini has undergone several changes since its original description. Stephens (1829) described the lichen moths as the family

Lithosiidae. Hampson (1900) reduced Lithosiidae to a subfamily of Arctiidae,

Lithosiinae. However, some authors continued to recognize the family Lithosiidae

(Hampson 1920; Kiriakoff 1951, 1963; Forbes 1960). Although Kiriakoff (1963) applied

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the name Lithosiidae, he noted that the tympanic structures of most lithosiine genera do not differ from the structures found in Arctiidae. Forbes (1960) stated that Lithosiidae was probably no more than a subfamily of Arctiidae, and his use of Lithosiidae was to maintain consistency with his earlier publications. More recently, during the course of remedying the paraphyly that had been identified within Noctuidae (Jacobson & Weller

2002; Mitchell et al. 1997, 2000; Weller et al. 1994), Lafontaine and Fibiger (2006) reduced the Arctiidae, Nolidae, Strepsimanidae, Lymantriidae, and Erebidae (sensu

Fibiger & Lafontaine 2005) to subfamilies of Noctuidae. As a result of this revision,

Lithosiinae was reduced to the tribe Lithosiini. In the most recent phylogenetic analyses of the superfamily (Zahiri et al. 2011, 2012), Erebidae was reassigned family status, and Arctiinae and Lithosiini are now treated as members of Erebidae.

Although the classification of the lichen moths has undergone several transitions, very little is known about the evolutionary relationships of the tribe at any taxonomic level. Three separate phylogenetic studies (Jacobson & Weller 2002; Wink & von

Nikisch-Rosenegk 1997; Zahiri et al. 2012) have recovered Lithosiini as a monophyletic

clade. Jacobson & Weller (2002) used morphological characters to assess the

monophyly of the five tribes of Arctiinae. Lithosiini was recovered as one of three

monophyletic tribes, and eight synapomorphies were identified. All of these

synapomorphies arise from the larval morphology. Wink and von Nikisch-Rosenegk

(1997) amplified the 16S rRNA gene for taxa from Arctiinae (including 5 species of

Lithosiini) and Nymphalidae to examine evolution of pyrrolizidine alkaloid and cardiac

glycoside sequestration. Zahiri et al. (2012) used 8 genes (1 mitochondrial and 7

nuclear) to examine the evolutionary relationships within the family Erebidae. All of

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these analyses found the Lithosiini to be monophyletic; however, the taxon sampling was insufficient (7, 5, and 5 genera, respectively) to infer relationships among the genera. Although Lithosiini has been recovered as one of three monophyletic tribes of

Arctiinae, there are two hypotheses about the evolutionary relationships among the tribes. Jacobson & Weller (2002) found that Lithosiini and Syntomiini were sister taxa.

This relationship was supported by six synapomorphies (3 adult and 3 larval

characters). The Lithosiini + Syntomiini clade was sister to Arctiini. However, Bendib &

Minet (1998) hypothesized that the Syntomiini were sister to Arctiini based on the

morphology of the female dorsal pheromone gland. Most recently, Zahiri et al. (2011,

2012) recovered Lithosiini as the sister taxa to a clade composed of Syntomiini and

Arctiini. The only other phylogenetic analysis conducted specifically on Lithosiini

examined the monophyly of the genus Agylla (Jacobson 1995). At the time of the study,

Agylla was composed of 138 species making it the fifth, largest genus within Lithosiini.

However, the cladistic analysis of morphological characters suggested that the genus

was paraphyletic.

Although no phylogenetic studies have been completed that examine the

evolutionary relationships among the genera of Lithosiini, several authors have

proposed the existence of relationships among groups of genera (Table 2-1). Forbes

(1939a) referred to generic groups but stated, “the classification of the Lithosiinae is too

uncertain to make discussion of value” (1939a p. 169). Based on his study of the

genitalia of lithosiine genera that occur in West Africa, Birket-Smith (1965) proposed the

existence of either a subtribe and two supergenera, or two subtribes, within Lithosiini.

However, he noted that given the amount of variation present in the genitalia these

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should be considered interim groupings until the entire tribe Lithosiini had been studied.

Franclemont (1983) also recognized Afridina as a subtribe of the Lithosiini. However,

Kitchings and Rawlins (1998) have since treated this subtribe as a subfamily of Nolidae.

Most recently, Bendib & Minet (1999) proposed the existence of seven subtribes

(Figs 2-1 A-V) within Lithosiini: Phryganopterygina, Acsalina, Lithosiina, Cisthenina,

Nudariina, Endrosina, and Eudesmiina. They provided definitions for all of the subtribes

except Lithosiina using larval, pupal, and adult morphological characters. Lithosiina was

left to be treated in a later work. They noted that larval characters, such as the presence

of a mandibular mola (a structure hypothesized to help the larvae grind the tougher

fungal tissue of lichens (Gardner 1943)), are the most reliable for accurately placing

taxa within Lithosiini. However, no analyses were conducted to determine whether the

morphological characters that were proposed to define the subtribes represent

synapomorphies, symplesiomorphies or are homoplasious. Holloway (2002) noted that

two of the adult characters defined as autapomorphies of the subtribe Nudariina have

evolved in parallel in other lithosiine subtribes. Also, Holloway (2002) was unable to

place all Bornean Lithosiini within subtribes using the descriptions provided by Bendib &

Minet (1999). Despite these complications, the classification proposed by Bendib &

Minet (1999) has been adopted in more recent publications (Ferguson & Opler 2006;

Holloway 2002; Powell & Opler 2009; Schmidt & Opler 2008). Based on the work of

Bendib & Minet (1999) and other authors who recognized generic groupings, 112

genera have been placed within subtribes. However, that leaves the majority of the

estimated 457 genera of Lithosiini (Jacobson & Weller 2002) unplaced (Figs 2-1W-Z).

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As noted by Bendib & Minet (1999) and demonstrated in Jacobson & Weller’s

(2002) phylogeny, larval morphological characters are the most consistent for placing species within the tribe Lithosiini. However, the larvae of many of the species that occur in the tropics have not been found (Holloway 2002). Adult characters, such as the wing venation and absence of ocelli, have also been used to place species into Lithosiini.

However, using the loss of ocelli to place species within Lithosiini led to aocellate phaegopterine genera (Pygoctenucha Grote and Lerina Walker) being mistakenly placed within the tribe. Furthermore, the use of wing venation caused the true lithosiine genus Lycomorpha Harris to be misplaced within Euchromiini (Forbes 1960).

In this paper, we report the first phylogenetic analysis of the tribe Lithosiini using

morphological characters coded from adult specimens. Species representing each of

the seven subtribes, as well as those not currently placed in a subtribe, are examined.

The contribution of the adult morphological characters to the resolution of the

evolutionary relationships within the tribe is assessed. In addition, the phylogeny is

examined to study the evolutionary relationships within Lithosiini, and determine

whether support existed for the subtribal relationships proposed by Bendib & Minet

(1999).

Materials and Methods

Taxon Sampling

Seventy-six species representing each subtribe of Lithosiini, as well as genera

that are not currently assigned to a subtribe were sampled (Table 2-2). When available, the type species of a genus was included in the analysis. Male-female pairs were available for all but five species. Outgroup taxa from the subfamily and the tribe Arctiini were included in the analysis. Aganainae has been recovered in the sister

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clade to the Arctiinae in several phylogenetic analyses (Kitching 1984; Zahiri et al. 2011,

2012). Furthermore, Zahiri et al. (2011, 2012) found the Lithosiini to be the most basal

lineage of Arctiinae. heliconia Linnaeus was included as the representative of this subfamily. Pagara simplex Walker, the other outgroup taxon, is considered an aocellate phaegopterine (Bendib & Minet 1999) that has previously been treated as a lithosiine

(Covell 1984; Forbes 1960; Franclemont; 1983). The larvae of P. simplex lack a

mandibular mola (Forbes 1960). The mandibular mola is a synapomorphy of the tribe,

and presently it is this character that is used to definitively place species within the tribe.

Ferguson and Opler (2006) transferred P. simplex to the tribe Arctiini. See Table 2-3 for

the full species list and the locality data for dissected specimens.

Morphology

Dissections of the genitalia were prepared using standard methods (Winter 2000)

after softening the abdomens in warm 10% potassium hydroxide aqueous solution for

30 to 90 minutes. Specimens were viewed in 20% ethanol, and the material was

subsequently stored in glycerol. External morphological characters were coded from

pinned, dried specimens. All characters were scored using a Nikon SMZ800 light

microscope. Pencil drawings were made to illustrate characters and states using a

camera lucida. The drawings were scanned and saved as PDFs. These files were

imported into Adobe Illustrator CS5 and inked using the pen tool. The terminology for

male and female genitalia follows Klots (1970), Kristensen (2003), and Forbes (1939b,

1954). Although Birket-Smith (1965) proposed a new nomenclature for the genitalia of

Lithosiini, homologous structures cannot be identified with confidence in subtribes other

than Lithosiina. The terminology for the wing venation follows Kristensen (2003).

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Characters Examined

Eighty-two morphological characters (68 binary and 14 multistate; 183 states) were scored for all of the species sampled (Appendix A). Morphological characters are as follows: head (7 characters, 16 states), thorax (5 characters, 11 states), wings (12 characters, 25 states), abdomen (3 characters, 7 states), male abdomen and genitalia

(38 characters, 85 states), and female abdomen and genitalia (17 characters, 39 states). Multistate characters were treated as unordered. When characters were linked, e.g. the presence of the radial vein and all of the radial sector veins (Character 13) and the branching pattern of R1 and Rs1-Rs4 (Character 14), resulting inapplicable

characters were coded as missing “?” (Strong & Lipscomb 1999). Linking the characters

allows the presence of the structure to contribute to the analysis and document variation

in the structure without overly simplifying characters (Pogue & Mickevich 1990).

Characters and their Phylogenetic Usefulness

Here we present the list of phylogenetic characters used in this study. We

discuss the phylogenetic usefulness of each character. The individual character indices

Consensus Index (CI) and Retention Index (RI) are included after each character.

Linked characters are indicated by a “*”.

Head

1. Male flagellomere shape. 0: Filiform, 1: Pectinate. (CI = 0.05, RI = 0.43).

2. * Male flagellomeres: sensilla chaetica. 0: Present, 1: Absent. (CI = 0.50, RI = 0.80).

3. * Location of sensilla chaetica on male flagellomeres. ?: Does not apply, 0: Proximally, before the medial ridge of the flagellomere, 1: Distally, at or beyond the medial ridge of the flagellomere, 2: Apices of the rami. (CI = 0.18, RI = 0.10).

4. Female flagellomere shape. 0: Filiform, 1: Pectinate. (CI = 0.06, RI = 0.21).

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5. * Female flagellomeres: sensilla chaetica. 0: Present, 1: Absent. (CI = 0.50, RI = 0.80).

6. * Location of sensilla chaetica on female flagellomeres. ?: Does not apply, 0: Proximally, before the medial ridge of the flagellomere, 1: Distally, at or beyond the medial ridge of the flagellomere, 2: Apices of the rami. (CI = 0.29, RI = 0).

7. Form of gena (when viewing the head in profile). 0: A continuous band around the base of the eye that joins the frons (Fig. 2-2A), 1: Occurring posteriorly to the eye as a band but not continuous around the base of the eye to the frons (Fig. 2- 2B). (CI = 0.11, RI = 0.11).

Summary of Head Characters: Although the indices for the head characters

were mostly low, some of the characters were useful for defining small clades, and one

character defined the tribe. The flagellomere shape varied between filiform and

pectinate in both the males and females. Sensilla chaetica were present in most species

of both sexes. Their loss defined the Euthyone+Bruceia clade and Eurylomia. Lithosiini

was defined by the placement of the sensilla chaetica distally in both males and

females. Within the tribe, the occurrence of the chaetica at the tips of the rami arises

independently several times. A gena that can be seen as a continuous band around the

eye defined the Lycomorpha+Propyria clade and the Ptychoglene clade. This state also

arose independently in several unrelated species.

Thorax

8. Tibial spur formula. 0: 0-2-2, 1: 0-2-3, 2: 0-2-4. (CI = 0.50, RI = 0.83).

9. Tarsal claw. 0: Simple, 1: Bifid. (CI = 0.04, RI = 0.40).

10. Epiphysis. 0: Long, two-thirds the length of the tibia or nearly the entire length of the tibia, 1: Short, approximately half the length of the tibia or less. (CI = 0.25, RI = 0.25).

11. Prothoracic tibia modified with elongate apical projection. 0: Present (Fig. 2-3A), 1: Absent (Fig. 2-3B). (CI = 0.50, RI = 0.50).

12. Tymbal organ. 0: Present, 1: Absent. (CI = 0.06, RI = 0.25).

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Summary of Thoracic Characters: Although thoracic characters have been found to be phylogenetically informative in higher-level studies of Notodontidae (Miller

1991) and Arctiinae (Jacobson & Weller 2002), they contributed little information about the evolutionary relationships of Lithosiini. The tibial spur formula was useful in defining small clades. Most species examined had a formula of 0-2-4. A formula of 0-2-3 defined the Lycomorpha+Propyria clade and Rhabdatomis. A 0-2-2 formula was unique to the

Euthyone+Bruceia clade. A long epiphysis defined the

Apistosia+Gnamptonychia+Inopsis+Eurylomia clade. It was lost once in Eurylomia. The long epiphysis also arose independently in cervicalis and the outgroup taxon

Pagara. The presence of an elongate apical projection on the protibia was unique to the

Apistosia+Gnamptonychia+Inopsis+Eurylomia clade. This state was reversed once in

Apistosia. The form of the tarsal claw and the presence of a tymbal organ were highly variable undergoing several losses and reversions.

Wings

13. Forewing: R1 free to costal margin and does not anastomose with Sc. 0: Present, 1: Absent. (CI = 0.06, RI = 0.29).

14. * Forewing: Radial vein and Radial Sector Veins. 0: R1, Rs1-Rs4 present, 1: Rs4 lost. (CI = 0.33, RI = 0).

15. * Forewing: radial vein branching pattern when R1, Rs1-Rs4 present. ?: Does not apply, 0: R1; Rs1; (Rs2 (Rs3, Rs4)), 1: R1; Rs1; ((Rs2, Rs3) Rs4). (CI = 0.07, RI = 0.55).

16. Forewing: M1. 0: Present, 1: Absent. (CI = 0.33, RI = 0).

17. Forewing: M2. 0: Present, 1: Absent. (CI = 0.50, RI = 0.80).

18. Forewing: M3. 0: Present, 1: Absent. (CI = 1.00)

19. Hindwing: Sc+R1. 0: Present, 1: Absent. (CI = 0.17, RI = 0.55).

20. Hindwing: Rs and M1. 0: Separate, 1: Stalked beyond distal margin of discal cell. (CI = 0.14, RI = 0.50).

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21. * Hindwing: M2. 0: Present, 1: Absent. (CI = 0.06, RI = 0.50).

22. * Hindwing: if present, M2 not fused with M3. ?: Does not apply, 0: Present, 1: Absent. (CI = 0.11, RI = 0.11).

23. Hindwing: M3 and CuA1. 0: Separate for entire distance to margin, 1: Stalked, 2: Fused. (CI = 0.09, RI = 0.36).

24. Male Hindwing: Anal angle enlarged and folded. 0: Present (Fig. 2-4A), 1: Absent (Fig. 2-4B). (CI = 0.20, RI = 0.20).

Summary of Wing Characters: Although variation was present within the

venation of the forewing, the characters coded from this region were only useful in

defining small groups or species. The loss of the M2 vein was unique to the largest

clade of Lithosiina taxa, which contains the genera , Eilema, , and

Pelosia. Within this clade, the state was reversed once in the species .

The loss of the M3 vein defined Crambidia.

Similarly, the variation present in the hindwing was only useful for defining

species or small clades. The Sc+R1 vein was lost six times within the tribe. These

losses corresponded to the species that compose the largest clade of Cisthenina

recovered by the Bayesian Inference analysis. However, this relationship was not

recovered in the Maximum Parsimony analysis. The M2 vein was lost 17 times in

Lithosiini. The occurrence of a CuA1 and M3 vein that were fused and stalk beyond the

distal margin of the discal cell arose 19 times.

Abdomen

25. * Anterolateral process (ALP) on A2 sternite. 0: Present (Figs 2-5 B-D), 1: Absent (Fig. 2-5A). (CI = 0.33, RI = 0).

26. * Location of the ALP. ?: Does not apply (Fig. 2-5A), 0: Firmly fused to the rest of the A2 (Figs 2-5C,D), 1: Emerging from the proximal end of apodeme (Fig. 2-5B). (CI = 0.11, RI = 0.47).

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27. * Form of ALP if it is fused to the rest of A2. ?: Does not apply (Figs 2-5A,B), 0: Flattened, sclerotized lobe (Fig. 2-5C), 1: Sclerotized bar (Fig. 2-5D), 2: Sclerotized knob that is shorter than the length of the apodeme. (CI = 0.17, RI = 0.38).

Summary of Abdominal Characters: Although variation was present in the

anterolateral process (ALP) on the second sternite, this variation was not

phylogenetically informative. The ALP was lost in three unrelated genera (Eilema,

Hypermaepha, and Hemipsilia). An ALP that was located at the proximal end of the apodeme arose independently eight times within Lithosiini. Most species examined possessed a flattened, sclerotized lobe ALP. The sclerotized bar form of the ALP arose independently ten times in the tribe.

Male abdomen and genitalia

28. Proximal margin of A8 sternite. 0: Extending dorsad to the edge of the A8 tergite (Fig. 2-7A), 1: Located entirely ventrad, with no portion extending onto the A8 pleurite or tergite. (Fig. 2-6A), 2: Extending onto the A8 pleurite but not beyond (Fig. 2-6B), 3: Fused with the proximal margin of the A8 tergite forming a continuous ring (Fig. 2-7B). (CI = 0.10, RI = 0.38).

29. Tegumen occurring entirely dorsad not extending ventrad of the meso-dorsal costal margins of the valvae. 0: Present (Fig. 2-8A), 1: Absent (Fig. 2-8B). (CI = 0.04, RI = 0.33).

30. * Fusion/connection between the two halves of the tegumen. 0: Absent (Fig. 2- 9A), 1: Present (Figs 2-9B,C). (CI = 0.25, RI = 0.25).

31. * Fusion extends more than half the length of the halves of the tegumen. ?: Does not apply (Fig. 2-9A), 0: Present (Fig. 2-9B), 1: Absent (Fig. 2-9C). (CI = 0.05, RI = 0.31).

32. * Sutures indicating fusion between the halves of tegumen. ?: Does not apply (Fig. 2-9A) 0: Present (Fig. 2-9C), 1: Absent (Fig. 2-9B). (CI = 0.10, RI = 0.10).

33. * Pleural Sclerites. 0: Present (Figs 2-10 B-D), 1: Absent (Fig. 2-10A). (CI = 0.17, RI = 0.44).

34. * Connection between the pleural sclerites and tegumen. ?: Does not apply (Fig. 2-10A), 0: Membranous tissue (Fig. 2-10B), 1: Thin, sclerotized bar (Fig. 2-10C), 2: Fused (Fig. 2-10D). (CI = 0.07, RI = 0.32).

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35. * Connection between the pleural sclerites and the vinculum. ?: Does not apply (Fig. 2-10A), 0: Membranous tissue (Fig. 2-10D), 1: Thin, sclerotized bar (Figs 2- 10B,C). (CI = 0.25, RI = 0.50).

36. * Scaphium. 0: Present, 1: Absent. (CI = 0.04, RI = 0.32).

37. * Scaphium form. ?: Does not apply, 0: Sclerotized bar running the length of the tuba analis, 1: Sclerotized triangular plate, 2: Two sclerotized bars forming a v- shape, 3: Rectangular sclerotized plate. (CI = 0.75, RI = 0).

38. Subscaphium. 0: Present, 1: Absent. (CI = 0.17, RI = 0.38).

39. Juxta occurring as a single, sclerotized plate undivided by membranous tissue. 0: Present (Fig. 2-11A), 1: Absent (Fig. 2-11B). (CI = 0.33, RI = 0.60).

40. Sclerotized portion of juxta extends dorsad toward the tegumen (not occurring as a flat plate). 0: Present, 1: Absent. (CI = 0.06, RI = 0.25).

41. Distal end of costal margin occurs as an identifiable structure. 0: Present (Figs 2- 12A,C), 1: Absent (Figs 2-12B,D and 2-14A). (CI = 0.07, RI = 0.41).

42. Processus basalis of costa. 0: Present (Fig. 2-12C), 1: Absent (Figs 2-12A,B,D and 2-14A,B). (CI = 0.11, RI = 0.20).

43. Mesad elongations of the costa (Tendon of Forbes 1939b). 0: Present (Figs 2- 12B,D and 2-14A), 1: Absent (Figs 2-12A and 2-14B). (CI = 0.05, RI = 0.10).

44. * Editum. 0: Present (Figs 2-12C, 2-13A,C, and 2-14C), 1: Absent (Figs 2- 13B,D). (CI = 0.04, RI = 0.15).

45. * Location of the editum based on the dorsal margin of the valva. ?: Does not apply (Figs 2-13B,D), 0: Proximally, before the midpoint of the valva (Figs 2-12C and 2-13A), 1: Distally, at or beyond the midpoint of the valva (Figs 2-13C and 2- 14C). (CI = 0.08, RI = 0.48).

46. Digitus emerging from the distal end of the costa. 0: Present (Figs 2-13 B-D), 1: Absent (Figs 2-13A and 2-14C,D). (CI = 0.08, RI = 0.37).

47. Valvulla occurring as fleshy tissue. 0: Present (Figs 2-12A,B,D, 2-13C, and 2- 14A,B), 1: Absent (Fig. 2-12C). (CI = 0.06, RI = 0.53).

48. * Form of cucullus. 0: Cucullus fused to both the costal and saccular margins of valva, no distinct ends (Fig. 2-12C), 1: Cucullus fused to costal margin but separated from saccular margin by valvulla (occurring as a separate fleshy lobe) (Fig. 2-14A), 2: Cucullus and valvulla fused into a contiguous structure (Fig 2- 12A,B,D, 2-13C, and 2-14B). (CI = 0.12, RI = 0.52).

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49. * Form of fusion between cucullus and valvulla when they form a single contiguous structure. ?: Does not apply (Figs 2-12C and 2-14A), 0: Cucullus forming slightly sclerotized dorsal margin of fleshy lobe of valvulla (Fig. 2-12D), 1: Sclerotized cucullus present on interior surface of valva while fleshy valvulla forms the external surface (Figs 2-12A and 2-13C), 2: Cucullus and valvulla form a fleshy lobe with no markings to distinguish either structure (Fig. 2-14B), 3: Cucullus heavily sclerotized dorsal portion of lobe and valvulla forms the fleshy ventral margin (Fig. 2-12B). (CI = 0.30, RI = 0.42).

50. * Corona of the cucullus. 0: Present (Figs 2-12A and 2-13C,D), 1: Absent (Figs 2- 13A,B and 2-14C,D). (CI = 0.05, RI = 0.21).

51. * Form of corona. ?: Does not apply (Figs 2-13A,B and 2-14C,D), 0: Spines (Figs 2-12A and 2-13C), 1: Setae (Fig. 2-13D). (CI = 0.33, RI = 0.50).

52. * Spacing of the spines/setae that form the corona. ?: Does not apply (Figs 2- 13A,B and 2-14C,D), 0: Dense, bases of separate spines/setae in contact (Fig. 2- 13C), 1: Not dense, bases of separate spines/setae not in contact (Fig. 2-13D). (CI = 0.13, RI = 0).

53. Inner face of the valvae: anellifera extending more than 2/3 the length of the valvae. 0: Present (Figs 2-13A and 2-14C), 1: Absent (Figs 2-13 B-D and 2-14D). (CI = 0.05, RI = 0.46).

54. * Inner face of valvae: clasper. 0: Present (Figs 2-13C and 2-14C), 1: Absent (Figs 2-13A,B,D and 2-14D). (CI = 0.13, RI = 0.53).

55. * Inner face of the valvae: form of clasper. ?: Does not apply (Figs 2-13A,B,D and 2-14D), 0: Well developed (Fig. 2-13C), 1: Reduced (Fig. 2-14C) (CI = 0.25, RI = 0.50).

56. Dorsal margin of the sacculus not extending onto the internal face of the valva. 0: Present (Figs 2-13A,B), 1: Absent (Figs 2-13C,D and 2-14C,D). (CI = 0.04, RI = 0.19).

57. * Process of the sacculus. 0: Present (Figs 2-12D, 2-13B,D, and 2-14A,B,D), 1: Absent (Figs 2-12A,C, 2-13A, and 2-14C). (CI = 0.06, RI = 0.50).

58. * Location where the saccular process arises on the ventral margin of the valva. ?: Does not apply (Figs 2-12A,C, 2-13A, and 2-14C), 0: Proximal half of the valva (Fig. 2-14D), 1: At or beyond the midpoint of the valva (Figs 2-12D, 2-13B,D and 2-14A). (CI = 0.20, RI = 0.33).

59. * Saccular process extending to the distal apex of the valva. ?: Does not apply (Figs 2-12A,C, 2-13A, and 2-14C), 0: Present (Figs 2-12D and 2-13B), 1: Absent (Figs 2-13D and 2-14 A,B,D). (CI = 0.13, RI = 0.36).

39

60. * Saccular process tapering to a point. ?: Does not apply (Figs 2-12A,C, 2-13A, and 2-14C), 0: Present (Figs 2-12D, 2-13B, 2-14A), 1: Absent (Figs 2-13D and 2- 14B,D). (CI = 0.08, RI = 0.08).

61. Basiphallus (caecum) form. 0: Well developed, ductus ejaculatorious simplex (DES) located entirely dorsad of the caecum (Fig. 2-15A), 1: Reduced, DES emerging from the anterior dorsal end of caecum (Fig. 2-15B), 2: Absent, DES emerges from the anterior end of the phallus (Fig. 2-15C). (CI = 0.10, RI = 0.40).

62. Ruggose patches on vesica. 0: Present, 1: Absent. (CI = 0.05, RI = 0.39).

63. Sclerotized bar/plate on vesica. 0: Present, 1: Absent. (CI = 0.08, RI = 0.29).

64. * Vesica ornamented with cornuti. 0: Present, 1: Absent. (CI = 0.05, RI = 0.18).

65. * Number of cornuti present. ?: Does not apply, 0: A single cornutus present, 1: More than one cornutus present. (CI = 0.10, RI = 0.44).

Summary of Male Characters: The male genitalia accounted for 38 of the 82

characters examined. Although the character indices within this suite of characters were

generally low, these characters illustrated the variation across genera. The only

character coded from the abdomen referenced the form of the proximal margin of the

eighth sternite. The tribe was defined by a proximal margin that extended onto the

pleurites but not beyond onto the tergite. All of the other states also occur within the

tribe. In addition, the character varied within some genera (e.g. Eilema). Although none

of the species examined possessed coremata (inflatable androconial structures

occurring between the seventh and eighth sternites Ferguson (1985)), pockets of

androconial scales were present on the abdominal segments of some males (e.g.

Clemensia). However, these structures were so varied that homologous states could not

be identified. Although these characters were not coded for this analysis, they could be

informative in revisions of genera.

The male genital capsule of the Lithosiini was highly variable. The valves and

tegumen were greatly modified across genera. In addition, the males of many genera

40

possess pleural sclerites that joined the tegumen and the vinculum. The pleural sclerite was thought to be lost in the subfamily Arctiinae (S. Weller pers. comm.). In some of the species examined, the pleural sclerite was found fused to the posterior margin of the tegumen (e.g. Lithosia quadra).

The two halves of the tegumen were fused for some or all of their length in the majority of the species examined. In the species where fusion was present, the tegumen could occur as a sclerotized plate that is entirely separate from the vinculum

(e.g. Lycomorpha). In addition, some species possessed sutures that could be used to identify the two halves of the tegumen. Although the presence of fusion was the most common state, the two halves of the tegumen were entirely separate in some species of

Lithosiini.

The valves of Lithosiini varied from having an external face that was completely sclerotized to one that had fleshy lobes present. The genera currently placed in

Acsalina, Cisthenina, and Eudesmiina possessed the former state. The latter state was found in the remaining subtribes. In the taxa with a sclerotized external face of the valve, it was not possible to identify the margins of the costa and the sacculus.

However, some species possessed a break in the sclerotization on the costal margin that could be used to identify the distal end of the costa. On the inner face of the valve, structures such as an editum, clasper, and corona of the cucullus were present in some species.

The phallus of Lithosiini was highly variable. Most species had a well-developed caecum on the basiphallus. The vesica was often ornamented with ruggose patches,

41

sclerotized bars/plates, or cornuti. However, ornamentation was completely absent in some species (e.g. Lycomorpha splendens Barnes & McDunnough).

Female abdomen and genitalia

66. * Segment A7 heavily sclerotized unlike preceding segments. 0: Present (Figs 2- 16 A-D and 2-17 A-D), 1: Absent (Fig. 2-18A,B). (CI = 0.04, RI = 0.12).

67. * Form of heavy sclerotization of A7. ?: Does not apply (Fig. 2-18A,B), 0: Continuous around entire segment (Figs 2-16A,B), 1: Membranous break occurring on sternite (Figs 2-16C,D), 2: Membranous break occurring on tergite (Figs 2-17A,B), 3: Membranous breaks occurring on pleurites (Figs 2-17C,D). (CI = 0.12, RI = 0.21).

68. A8 tergite does not extend beyond the distal margin of A7 tergite. 0: Present (2- 17B), 1: Absent (Figs 2-16B,D, 2-17D, and 2-18B). (CI = 0.07, RI = 0.13).

69. * Location of the ostium bursa. 0: A7 sternite (Fig. 2-19A), 1: Intersegmental membrane between A7 and A8 (Figs 2-19B,C), 2: A8 sternite (Fig. 2-19D). (CI = 0.20, RI = 0.11).

70. * Form of the distal margin of the A7 sternite when ostium bursa occurs is located in the A7 to A8 intersegmental membrane. ?: Does not apply (Figs 2-19A,D), 0: Nearly horizontal with no distinct indentations (Fig. 2-19B), 1: With a distinct, concave indentation (Fig. 2-19C). (CI = 0.05, RI = 0.26).

71. Anterior apophyses. 0: Short, less than or equal to length of A8 pleurite, 1: Long, greater than length of A8 pleurite. (CI = 0.05, RI = 0.32).

72. Posterior apophyses. 0: Short, less than or equal to the length of the A8 pleurite, 1: Long, greater than the length of the A8 pleurite. (CI = 0.04, RI = 0.13).

73. Dorsal pheromone glands within the A8-A9 intersegmental membrane. 0: Present, 1: Absent. (CI = 1.00, RI = 1.00).

74. * Sclerotization of ductus bursa. 0: Present (Figs 2-20A,B), 1: Absent (Fig. 2- 20C). (CI = 0.05, RI = 0.05).

75. * Sclerotization of ductus bursa located proximally to ostium bursa. ?: Does not apply (Fig. 2-20C), 0: Present (Fig. 2-20A), 1: Absent (Fig. 2-20B). (CI = 0.13, RI = 0.36).

76. Length of ductus bursa. 0: Short, less than or equal to the length of A7 tergite, 1: Long, greater than the length of A7 tergite. (CI = 0.06, RI = 0.29).

77. * Signa on the corpus bursa. 0: Present, 1: Absent. (CI = 0.17, RI = 0.17).

42

78. * Signa number. ?: Does not apply, 0: One, 1: Two, 2: More than two, 3: Many single signa covering large portions of the interior of the corpus bursa. (CI = 0.13, RI = 0.38).

79. * When more than one signum occur, all have the same form. ?: Does not apply, 0: Present, 1: Absent. (CI = 0.14, RI = 0.14).

80. * Signa extends from the corpus bursa into the ductus bursa. ?: Does not apply, 0: Present, 1: Absent. (CI = 0.13, RI = 0.42).

81. * Appendix bursa. 0: Present, 1: Absent. (CI = 0.05, RI = 0.30).

82. * Location of appendix bursa. ?: Does not apply, 0: Corpus bursa, 1: Ductus bursa. (CI = 0.10, RI = 0.40).

Summary of Female Abdomen and Genitalia Characters: Eighteen characters

were coded from the female genitalia. Similar to the male genitalia, these characters

helped to illustrate the variation among the genera.

The seventh abdominal segment of some species was found to be more heavily

sclerotized than the preceding segments. When the sclerotization was present, it could

extend around the segment unbroken or possess membranous regions on the sternite,

tergite, or pleurites.

The ostium bursa was located in the seventh sternite, the intersegmental

membrane between the seventh and eighth sternite, and the eighth sternite. When the

ostium bursa occurred in the intersegmental membrane of species with a heavily

sclerotized seventh segment, a deep indentation in the posterior margin of the seventh

sternite where the ostium bursa occurred was sometimes found (e.g. Lycomorpha grotei

(Packard)). Sclerotization of the ductus bursa was present in most species. The

presence of signa on the corpus bursa defined Lithosiini. Within the tribe, signa were

lost in five species. The number of signa present varied. It was possible to count the

separate signa in some species (e.g. two signa, L. grotei). However, some species

43

possessed many single signa that formed a large patch covered a broad area of the corpus bursa (e.g. Stigmatophora). This condition was coded as a separate state from the species that possessed three or more separate signa.

Phylogenetic Analysis

Phylogenetic trees were constructed using both parsimony and Bayesian

analyses. The Maximum Parsimony (MP) analysis was performed in Paup* 4.0b 10

(Swofford 2003). A heuristic search of 1000 random-taxa additions using the tree bisection reconnection (TBR) algorithm was performed to identify the most

parsimonious cladogram topology. Nodal support for the strict consensus of the MP

trees was evaluated using Bremer support indices (BS: Bremer 1988, 1994). Bremer

support indices were calculated in Paup* using a command file produced with

TreeRot.v3 (Sorenson & Franzosa 2007) that searched for trees inconsistent with the

constraint statement given for each node of the strict consensus tree. When discussing

our results, we define support values as giving weak (BS 1-2), moderate (BS 3-5), good

(BS 6-10), or strong (BS≥11) support (Wahlberg & Nylin 2003; Wahlberg et al. 2003,

2005). The Bayesian Inference (BI) analysis was conducted using MrBayes 3.1.2

(Ronquist & Huelsenbeck 2003). The Mk+Γ model (Lewis 2001) was used to analyze

the morphological dataset, which had been input as standard. The Γ parameter was

included in the model because Lewis (2001) considers it to be the most appropriate way

to allow for rate heterogeneity in morphological character evolution. The Bayesian

analyses were performed with four chains, one cold and three hot, using the default

temperature settings. Five simultaneous, independent runs of 20,000,000 generations

were conducted. The Markov chain Monte Carlo (MCMC) chains were started at

random trees. Samples were drawn from the cold chain every 1,000 generations. Five

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million generations were discarded as ‘burn-in’. The probabilities of the five runs were summarized, and the potential scale reduction factor (PSRF: Gelman & Rubin 1992) was calculated to confirm that the runs had converged. The PSRF value should approach 1 as the independent runs converge. The trees from the Bayesian analysis were summarized as a majority rule consensus tree. Posterior probability (PP) provided the clade credibility values for the tree. The consensus trees from both types of analysis were visualized using FigTree v1.3.1 (Rambaut 2010).

Results

Analyses were based on 82 adult morphological characters sampled for 78 species. The Maximum Parsimony (MP) analysis resulted in 4408 MP trees with a length (L) of 736, consistency index (CI) of 0.10, and a retention index (RI) of 0.34. The

Bayesian Inference (BI) analysis produced a phylogram with a mean tree length (TL) of

16.36 with a variance of 1.08. Both analyses were rooted on the subfamily Aganainae

(Asota heliconia). Although all of the lithosiine species were found in a single lineage in the optimal cladogram from the parsimony analysis, there was no nodal support for this clade. Although the MP analysis was not able to recover the tribe as monophyleteic, the

Bayesian analysis did (PP ≥ 0.66). Neither of the analyses was able to entirely resolve the evolutionary relationships among the genera of the tribe. The strict consensus of the

MP trees produced a cladogram with no resolution of the deeper nodes (Fig. 2-21). The majority of the clades that were found on this tree received only moderate Bremer support. The low level of resolution in the MP cladogram, as well as the low nodal support may be due to high levels of homoplasy present in the morphological characters

(Appendix B). Although 81 of the characters were parsimony informative, most had very low CI and RI values. The BI phylogram (Fig. 2-22) had a somewhat more resolved

45

topology than the MP cladogram, and several of the nodes received strong posterior probability support values. Despite the low level of resolution present in each cladogram, both analyses recovered identical relationships among certain genera and species. Although the two cladograms contain similarities, novel evolutionary relationships were also recovered by both phylogenetic methods.

Our analyses found genera representing the subtribe Cisthenina present in seven separate clades. Cisthenina is currently composed of 45 genera that are found in both the Old and New World. However, there were several of the species representing the subtribe Cisthenina whose evolutionary relationships are entirely unresolved. The largest of the Cisthenina clades was comprised of the brightly colored, New World genera Lycomorpha and Propyria Hampson with strong support (PP ≥ 0.98, BS = 3).

Both genera are apparent mimics of other arctiines, zygaenids, and lycid beetles.

Furthermore, a strongly supported polytomy (PP ≥ 0.98) was found between this clade, the genus Dolichesia Schaus, and the weakly supported clade (PP ≥ 0.53) composed of

Hypermaepha Hampson, Lycomorphodes Hampson, and Talara Walker. Strong support

(PP = 1, BS = 3) was found for a clade containing the Neotropic genus Euthyone

Watson and the genera Bruceia Neumoegen and Haematomis Schaus. Bruceia was found to be paraphyletic as defined, and there was strong support (PP ≥ 0.96, BS = 3) for a sister relationship between Bruceis pulverina Neumoegen and Haematomis mexicana (Druce). Metalobosia Hampson and Odozana Walker were also recovered as sister (PP ≥ 0.79, BS = 2). The remaining Cisthenina clades provided support for monophyly of genera with the exception of Ptychoglene Felder. A sister relationship was found between Pt. erythrophora Felder and Pt. sanguineola (Boisduval) (PP = 1, BS =

46

4). However, the placement of the remaining species Pt. coccinea (Edwards) was unresolved. In addition, Packard was recovered as a monophyletic lineage with strong support (PP ≥ 0.99, BS = 7). There was also weak support (BS = 1) for a sister relationship between Clemensia and a clade formed of the only Old World

Cisthenina present in the analysis, Aemene Walker, and the unplaced Old World genus

Heliosia Hampson (BS = 2). A sister relationship between the Cisthenina genus

Rhabdatomis Dyar, which was strongly supported (PP = 1, BS = 3) as monophyletic, and the unplaced genus Epeiromulona Field was also found (PP ≥ 0.54).

The genera representing Lithosiina, a subtribe composed of 38 genera that are both Old and New World in distribution, were found in three clades. The genera Lithosia

Fabricius, Eilema Hübner, Crambidia Packard, and Hübner formed a clade with strong support (PP ≥ 0.97, BS = 3). Within this clade, however, the relationships among the genera were entirely unresolved in both analyses. Weak support (PP ≥ 0.52) was found for a sister relationship between this clade and the genus Cybosia Hübner. The remaining two Lithosiina clades also contained genera that have not received subtribal placement. A strongly supported (PP ≥ 0.99, BS = 4) relationship was present between the Lithosiina genera Apistosia Hübner, Gnamptonychia Hampson, and Inopsis Felder and the unplaced genus Eurylomia Felder. The brightly colored, large adults of each of these genera are mostly Neotropical in distribution. Within this clade, Gnamptonychia and Inopsis were found to be sister (PP ≥ 0.9) and were weakly associated (PP ≥ 0.65) with Apistosia. Finally, the unplaced species Ardonea moria (Walker) was moderately supported (PP ≥ 0.75, BS = 3) as sister to Gardinia Kirby.

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There was little resolution of the evolutionary relationships among the genera representing the two subtribes, Endrosina and Nudariina, composed entirely of Old

World genera. The relationships that were recovered for the genera representing each of these subtribes were almost entirely unique between the two analyses. Endrosina is composed of two genera that are Palearctic and Oriental in distribution. Both analyses recovered the sister relationship between the two species of Setina Schrank (PP ≥ 0.61,

BS = 3). However, no relationship was found between this clade and the other endrosine genus Stigmatophora Staudinger. The relationships recovered between the genera representing Nudariina (21 genera) were entirely different between our two analyses. The MP cladogram recovered a weakly supported clade (BS = 1) containing

Miltochrista Hübner, Hemipsilia Hampson and Asura Walker. Within the clade,

Miltochrista was found to be paraphyletic, and Asura was polyphyletic. The BI phylogram recovered only a weakly supported (PP ≥ 0.61) sister relationship between

Paidia Hübner and Cyana Walker.

The entirely New World subtribe Eudesmiina is presently composed of four genera. With the exception of Euryptidia ira (Druce), the species representing

Eudesmiina were recovered within a single, moderately supported clade (PP ≥ 0.86, BS

= 1). Within this clade, the genus Hübner was found to be paraphyletic as presently defined. The species of Josioides Felder and Eudesmia were recovered as a weakly supported monophyletic clade (PP ≥ 0.67).

The two monogeneric subtribes, Acsalina and Phryganopterygina, were represented in our analyses by a single species each. Their position in both the MP cladogram and BI phylogram was entirely unresolved. The evolutionary relationships of

48

the species representing the genera that have not been placed within a subtribe were mostly unresolved. However, some of these genera (e.g. Eurylomia and Heliosia) and the species Ardonea moria were found in clades composed of genera representing one of the subtribes.

Discussion

This is the first phylogenetic study conducted on the tribe Lithosiini using morphological data and broad taxon sampling. Previously, the use of adult morphological characters to place species within the tribe has proved problematic. The use of these characters has included true genera of Arctiini but resulted in the exclusion of Lithosiini genera. Our analyses indicated that the adult morphological characters did not provide a strong enough phylogenetic signal to resolve the older, higher-level

relationships within the tribe. In addition, these characters were only able to recover

Lithosiini as monophyletic in the BI analysis. It is possible that the inclusion of

characters coded from the endoskeleton sensu Snodgrass (1993) of the adults may

help to resolve the deeper nodes. These endoskeleton characters have been found to

be informative at the tribal level within Arctiinae (Jacobson & Weller 2002; DaCosta &

Weller 2005).

With the low level of deeper node resolution present in our results, it was not

possible to assess the monophyly of any of the subtribes proposed by Bendib & Minet

(1999). The genera representing each of the subtribes were either found in small clades

or their placement was unresolved. However, the clades that included individuals from

multiple genera contained only genera from a single subtribe or those genera that have

not yet been placed in a subtribe.

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The results found genera of the largest subtribe Cisthenina in several clades.

Four of these clades contained representatives of multiple genera. The largest clade was a strongly supported (PP ≥ 0.98) polytomy among the genus Dolichesia, the

Lycomorpha and Propyria clade, and a Lycomorphodes, Talara, and Hypermaepha

clade. DNA barcoding has also recovered a close relationship among Lycomorpha,

Lycomorphodes, and Talara (C. Schmidt pers. comm.). Within the Lycomorpha and

Propyria clade, our results from the Bayesian analysis suggested that the two genera were synonymous. The adults of these two genera share similar color patterns, and

Propyria schausi (Dyar) was originally described as a species of Lycomorpha. This was not the only example of paraphyly that the results found within a Cisthenina genus.

Bruceia was found to be paraphyletic with respect to Haematomis mexicana. The ranges of Bruceia and H. mexicana overlap within the southwestern United States.

However, H. mexicana is not the type species of the Haematomis. Until the type species of Haematomis can be included in an analysis of these genera, it will be unclear whether they are synonyms or H. mexicana is misplaced. The only Old World

Cisthenina included in the analysis, Aemene altaica (Lederer), was found to be sister to the unplaced Old World Heliosia. Furthermore, this clade was recovered as sister to the

New World Clemensia. Forbes (1939a) considered Clemensia to be related to the Old

World Nudaria generic group. Forbes later (1960) hypothesized that the genus was

closely related to genera, such as Siccia Walker, found within East Asia. Siccia is

presently placed within Cisthenina, and Bendib & Minet (1999) considered Aemene to

be a synonym of Siccia. However, Holloway (2002) has resurrected Aemene as a

separate genus.

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Genera representing the second largest subtribe, Lithosiina, were found in three clades. The type genus of the subtribe, Lithosia, was found in a strongly supported (PP

≥ 0.99, BS = 3) clade containing the genera Eilema, Crambidia, and Pelosia. Forbes

(1960) noted that the New World genus Crambidia and the New World Eilema bicolor

(Grote) were closely related to Lithosia. The relationships within this clade were not

resolved among the genera or among the species representing each genus. However,

as noted by Birket-Smith (1965) and Holloway (2002) the genus Eilema, as presently

defined, does not comprise a natural group.

The results recovered entirely different relationships among the genera of

Nudariina, the third largest subtribe, depending on the type of analysis that was

conducted. The results from the MP analysis recovered a weakly supported (BS = 1)

clade containing the genera Miltochrista, Hemipsilia, and Asura. Within this clade,

Miltochrista was recovered as paraphyletic, and Asura was found to be polyphyletic.

Both of these genera, as currently defined, are very large and possess similar color

patterns, which are also found in other Old World lithosiine genera (e.g. Lyclene Moore,

Barsine Walker, and Adites Moore). Bendib & Minet (1999) specifically stated that the

genus Asura was not monophyletic. Holloway (2002) noted that the present definition of

this genus overlaps with a previous definition of Miltochrista (Hampson). The Bayesian

analysis recovered only a weakly supported (PP ≥ 0.61) sister relationship between

Cyana and . Bendib & Minet (1999) noted a close relationship between these two

genera based on two wing characters (CuA1 and M3 stalked or coincident and a

projecting lobe near the base of R2 on the male forewing) and one larval character

(fusion of D and SD2 into a single verruca).

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Our results recovered very little information on the relationships within the smaller and monogeneric subtribes with the exception of Eudesmiina. All of the species representing Eudesmiina except Euryptidia ira were found within a single clade. Within the clade, the genus Eudesmia was paraphyletic. Furthermore, the relationships between Eudesmia and Josioides suggested that the two genera may be synonymous.

This was not unexpected. The adults of the genera that are placed within Eudesmiina are brightly colored with similar patterns. Bendib & Minet (1999) considered an

autapomorphy of the subtribe to be the contrasting black/brown and orange/yellow

areas present on the wings of Eudesmiina species. However as noted by Powell &

Opler (2009), these color patterns create difficulty in separating species. Within the

subtribe Endrosina, the genus Setina was found to be monophyletic. However, no

relationship recovered between it and the other endrosine genus Stigmatophora. No

relationships were observed for either monogeneric subtribe, Acsalina or

Phryganopterygina.

We found support for relationships between some of the genera that have not

been placed in subtribes and clades of genera from Cisthenina and Lithosiina. In each

instance, the unplaced genus was found to be related to a genus or genera that are

distributed in the same area of the world. However, the evolutionary relationships of

most of the unplaced genera were not recovered. Holloway (2002) noted that he was

not able to place all of the genera of Lithosiini that are found in Borneo into the subtribes

proposed by Bendib & Minet (1999).

Our study found that the adult external morphology does not provide a

phylogenetic signal of sufficient strength to resolve the higher-level relationships within

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Lithosiini. This result was not unexpected given that all the synapomorphies previously identified for the tribe arise within the larval morphology, and the use of adult morphological characters to assign species to Lithosiini has previously resulted in several misplacements. The use of other external morphological characters and internal characters that can only be examined through whole body dissections may help to clarify the evolutionary relationships. The inclusion of larval characters could also provide informative phylogenetic signals. Unfortunately, the larvae of most tropical lithosiines have not been found (Holloway 2002). It may also be necessary to use

molecular data to resolve the relationships. However, although it was not possible to

assess the monophyly of the subtribes proposed by Bendib & Minet (1999), the

relationships that were recovered by the analyses do provide support for the subtribes

and the evolutionary hypotheses of other authors.

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Table 2-1. Comparative table of generic groups of Lithosiini, including the location of the taxa the author was studying. Börner Forbes Kiriakoff 1920, 1932 1939a 1951 Germany Panama World ARCTIIDAE ARCTIIDAE LITHOSIIDAE Lithosiinae Lithosiinae Endrosiinae Lithosiini Lithosia group Nudariini Hypoprepia group LITHOSIIDAE Lithosiinae Nudariinae Endrosiinae

Birket-Smith Franclemont Bendib and Minet 1965 1983 1999 West Africa North America World ARCTIIDAE ARCTIIDAE ARCTIIDAE Lithosiinae Lithosiinae Lithosiinae Lithosiini Lithosiini Lithosiini (spelt Lithosini) Acsalini Acsalini Lithosiina Afridini Cisthenini (spelt Lithosinia) Nudariini Eilemina Eudesmiini (spelt Eileminia) Endrosini Phryganopterygini

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Table 2-2. Ingroup taxon sampling including the total number of genera placed in each subtribe Genera Species Subtribe Total # # Examined # Examined Acsalina 1 1 1 Cisthenina 45 21 37 Endrosina 2 2 3 Eudesmiina 4 3 7 Lithosiina 38 9 12 Nudariina 21 5 7 Phryganopterygina 1 1 1 Unplaced 345 7 8 457 49 76

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Table 2-3. Species examined in the phylogenetic analysis including subtribe, locality, and collection that provided the material Taxa (Genus species Author) Sex Collection Locality EREBIDAE Arctiinae Lithosiini Acsalina Acsala anomala Benjamin Male CUIC Yukon Territory Female CNC Yukon Territory Cisthenina Aemene altaica (Lederer) Male CUIC Tateshima, Japan Female CUIC Tateshima, Japan Barsinella mirabilis Butler Male CUIC Moengo, Suriname Bruceia hubbardi Dyar Male FLMNH Cochise Co., AZ, USA Female FLMNH Cochise Co., AZ, USA Br. pulverina Neumoegen Male CUIC Coconino Co., AZ, USA Female CUIC Coconino Co., AZ, USA plumbea Stretch Male FLMNH Liberty Co., FL, USA Female FLMNH Alachua Co., FL, USA Ci. subjecta Walker Male FLMNH Alachua Co., FL, USA Female FLMNH Ocean City, FL, USA Packard Male FLMNH Oldham Co., KY, USA Female FLMNH Oldham Co., KY, USA Cl. leopardina Schaus Male CUIC Chapare, Bolivia Female CUIC Chapare, Bolivia Dolichesia falsimonia Schaus Male MCZ Barro Colorado, Panama Female MCZ Barro Colorado, Panama Euthyone simplex (Walker) Male CUIC Barro Colorado, Panama Female CUIC Barro Colorado, Panama Eu. grisescens (Schaus) Male MCZ Barro Colorado, Panama Female MCZ Barro Colorado, Panama Gaudeator paidicus Dyar Male CUIC Barro Colorado, Panama Female CUIC Barro Colorado, Panama Haematomis mexicana (Druce) Male CUIC Santa Cruz Co., AZ, USA Female CUIC Cochise Co., AZ, USA Hypermaepha maroniensis Male CUIC Moengo, Suriname Schaus Female CUIC Moengo, Suriname Hypoprepia cadaverosa Streck Male FLMNH Torrance Co., NM, USA Female FLMNH Jeff Davis Co., TX, USA Hy. fucosa Hübner Male UofW Collier Co., FL, USA Female FLMNH Putnum Co., FL, USA Hy. inculta Edwards Male CUIC Cochise Co., AZ, USA Female CUIC Santa Cruz Co., AZ, USA Lycomorpha fulgens (Edwards) Male AMNH Socorro Co., NM, USA Female AMNH Pima Co., AZ, USA

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Table 2-3. Continued Taxa (Genus species Author) Sex Collection Locality Ly. fulgens nr. Male AMNH Catron Co., NM, USA Female NHMLAC Cochise Co., AZ, USA Ly. grotei (Packard) Male NHMLAC Garfield Co., CO, USA Female NHMLAC Los Angeles Co., CA, USA Ly. pholus (Drury) Male USNM Ottawa Co., Quebec, Female CMNH Canada Pendleton Co., WV, USA Ly. splendens Barnes & Male USNM Washington Co., UT, McDunnough Female NHMLAC USA Yavapai Co., AZ, USA Lycomorphodes correbioides Male CMNH Zamora, Ecuador Schaus Female CMNH Zamora, Ecuador Metalobosia elis (Druce) Male CUIC Nova Teutonia, Brazil Female CUIC Nova Teutonia, Brazil Nodozana jucunda Jones Male AMNH Arima Valley, Trinidad Female AMNH Arima Valley, Trinidad Odozana floccosa Druce Male CUIC Moengo, Suriname Female CUIC Moengo, Suriname Prepiella aurea (Butler) Male AMNH Rodônia, Brazil Female AMNH Rodônia, Brazil Pre. radicans Hampson Male CUIC Moengo, Suriname Female CUIC Moengo, Suriname Propyria morelosia Schaus Male USNM Zacualpan, Mexico Female AMNH Barranca, Mexico Pro. ptychoglene Hampson Male USNM Jalapa, Mexico Female USNM Coatepec, Mexico Ptychoglene coccinea (Edwards) Male USNM Cochise Co., AZ, USA Female USNM Cochise Co., AZ, USA Pt. erythrophora Felder Male USNM Morelos, Mexico Female USNM Cordoba, Mexico Pt. sanguineola (Boisduval) Male USNM Jalapa, Mexico Female USNM Veracruz, Mexico Rhabdatomis coroides (Schaus) Male MCZ Barro Colorado, Panama Female MCZ Barro Colorado, Panama Rh. pueblae (Draudt) Male AMNH Sacatepequez, Female AMNH Guatemala Solola, Guatemala Talara coccinea Butler Male CMNH Hyatanahan, Brazil Female CMNH Oyapok River, French Guiana Ta. megaspila Walker Male AMNH Amazonas, Venezuela

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Table 2-3. Continued Taxa (Genus species Author) Sex Collection Locality Endrosina Setina aurita (Esper) Male AMNH Saint Véran, France Se. irrorella (Linnaeus) Male AMNH TL Viella, Spain Female YPM Co. Clare, Ireland Stigmatophora flava (Bremer and Male AMNH Guizhou Prov., China Grey) Female AMNH Guizhou Prov., China

Eudesmiina Eudesmia arida (Skinner) Male FLMNH Jeff Davis Co., TX, USA Female FLMNH Maricopa Co., AZ, USA Eu. praxis (Druce) Male AMNH Durango, Mexico Female AMNH Durango, Mexico Eu. ruficollis (Donovan) Male MCZ São Paulo, Brazil Female MCZ Aguas Blancas, Argentina Euryptidia ira (Druce) Male USNM Tiacolula, Mexico Female USNM Cuernavaca, Mexico Eu. univitta Hampson Male AMNH Pelotas, Brazil Female AMNH Pelotas, Brazil Josioides celena (Walker) Male AMNH Kamkusa, Guyana Female AMNH Shudihar River, Guyana Jo. myrrha (Cramer) Male CUIC Suapure, Venezuela Female CUIC Moengo, Suriname Lithosiina Apistosia judas Hübner Male CUIC Viçosa, Brazil Female CUIC Barro Colorado, Panama Packard Male FLMNH Alachua Co., FL, USA Female FLMNH Alachua Co., FL, USA Cybosia mesomella (Linnaeus) Male YPM Sussex, England Female YPM Cheshire, England Eilema bicolor (Grote) Male FLMNH Grand Co., CO, USA Ei. bipuncta (Hübner) Male CUIC Harbel, Liberia Female CUIC Harbel, Liberia Ei. heimi de Toulgoët Male AMNH Antsiranana, Madagascar Female AMNH Antsiranana, Madagascar Gardinia anoploa Hering Male AMNH Cochise Co., AZ, USA Female AMNH Pima Co., AZ, USA Gnamptonychia flavicollis (Druce) Male AMNH Les Puentes, Mexico Female AMNH Zacualpan, Mexico Inopsis modulata (Edwards) Male AMNH Jalapa, Mexico Female AMNH Jalapa, Mexico Lithosia quadra (Linnaeus) Male MCZ Female MCZ France

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Table 2-3. Continued Taxa (Genus species Author) Sex Collection Locality Li. sororcula (Hüfnagel) Male AMNH Tyrol Female AMNH Medí (Hüfnagel) Male MCZ Germany Female MCZ France Nudariina Asura cervicalis Walker Male AMNH NSW, Australia Female AMNH Victoria, Australia As. strigipennis (Herrich-schäffer) Male CUIC Kuraru, Taiwan Female CUIC Kuraru, Taiwan Hemipsilia coa-vestis (Hampson) Male CUIC Bukai, Taiwan Female CUIC Arisan, Taiwan Cyana interrogationis (Poujade) Male CUIC Ha Giang Prov., Vietnam Female CUIC South China Miltochrista gratiosa (Guerin) Male CUIC Kuraru, Taiwan Female CUIC Kuraru, Taiwan Mi. flexuosa Leech Male AMNH Les, Spain Female AMNH Les, Spain Paidia murina (Hübner) Male AMNH Seva, Spain

Phryganopterygina Phryganopteryx convergens de Male AMNH Antsiranana, Madagascar Toulgoët Female AMNH Antsiranana, Madagascar Unplaced Ardonea morio Walker Male AMNH Jalapa, Mexico Female AMNH Vera Cruz, Mexico Ar. peculiaris (Druce) Male CUIC Chapare, Bolivia Female CUIC Moengo, Suriname Balbura intervenata Schaus Male AMNH Cocle, Panama Female AMNH Barro Colorado, Panama Epeiromulona icterinus Field Male AMNH Barro Colorado, Panama Female AMNH San Blas, Panama Eurylomia cordula (Boisduval) Male AMNH Jalapa, Mexico Female AMNH Jalapa, Mexico Heliosia jucunda (Walker) Male CUIC N. Qld, Australia Female CUIC N. Qld, Australia Oeonistis entella (Cramer) Male CUIC Papua New Guinea Female MCZ Australia Paramulona nephalistis Male AMNH Santiago, Cuba (Hampson) Female AMNH Santiago, Cuba Arctiini Pagara simplex Walker Male CUIC Manatee Co., FL, USA Female CUIC Manatee Co., FL, USA

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Table 2-3. Continued Taxa (Genus species Author) Sex Collection Locality Aganainae Asota heliconia Linnaeus Male UMD Qld, Australia Female UMD Qld, Australia Collections: AMNH (American Museum of Natural History), CMNH (Carnegie Museum of Natural History), CNC (Canadian National Collection), CUIC (Cornell University Insect Collection), FLMNH (Florida Museum of Natural History), MCZ (Museum of Comparative Zoology, Harvard), NHMLAC (Natural History Museum of LA County), USNM (United States Museum of Natural History), UofW (University of Wisconsin – Oshkosh), UMD (University of , LepATOL Collection), YPM (Yale Peabody Museum)

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Figure 2-1. Sample of lithosiine species included in the study. Subtribe Acsalina: A) Acsala anomala ♂ and B) Acsala anomala ♀. Subtribe Cisthenina: C) , D) Propyria ptychoglene, E) Clemensia albata, F) Ptychoglene erythrophora, G) Bruceia pulverina, H) Gaudeator paidicus, and I) Barsinella mirabellis. Subtribe Endrosina: J) Setina aurita and K) Setina irrorella. Subtribe Eudesmiina: L) Euryptidia univitta, M) Eudesmia ruficollis, and N) Josiodes myrrha. Subtribe Lithosiina: O) Eilema heimi, P) Lithosia quadra ♂, Q) Gardinia anoploa, and R) Apistosia judas. Subtribe Nudariina: S) Paidia murina, T) Miltochrista gratiosa, and U) Asura cervicalis. Subtribe Phryganopterygina: V) Phryganopteryx convergens. Unplaced Genera: W) Paramulona nephalistis, X) Ardonea peculiaris, Y) Balbura intervenata, and Z) Eurylomia cordula.

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Figure 2-2. Lateral view, head. A) Lycomorpha pholus; G-gena present as a continuous band that joins F-frons. B) Bruceia hubbardi; G-gena not present as a continuous band that joins F-frons. The scale bar is equivalent to 1mm.

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Figure 2-3. Prothoracic leg. A) Eurylomia cordula; AP-apical projection. B) Hypoprepia fucosa. The scale bar is equivalent to 1mm.

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Figure 2-4. Male hindwing. A) Balbura intervenata; AA-anal angle enlarged and folded. B) Setina irrorella; AA-anal angle unmodified. The scale bar is equivalent to 1mm.

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Figure 2-5. A2 sternite. A) Eilema bipuncta; A-apodeme, anterior lateral process (ALP) absent. B) Setina irrorella; AAA-ALP attached to apodeme, A-apodeme. C) Ptychoglene erythrophora; ALP fused to A2, FSL-flattened sclerotized lobe form of ALP, A-apodeme. D) Lycomorpha pholus; ALP fused to A2, SB- sclerotized bar form of ALP, A-apodeme. The scale bar is equivalent to 1mm.

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Figure 2-6. Proximal margin of male eighth sternite, Part A: A) Acsala anomala; VO-8S ventrad only, P-pleurite, 6S-sixth sternite, 6T-sixth tergite, 7S-seventh sternite, 7T-seventh tergite, 8S-eighth sternite, 8T-eighth tergite; B) Eudesmia ruficollis; EP-8S extends onto pleurites, P-pleurite, 6S-sixth sternite, 6T-sixth tergite, 7S-seventh sternite, 7T-seventh tergite, 8S-eighth sternite, 8T-eighth tergite. The scale bar is equivalent to 1mm.

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Figure 2-7. Proximal margin of male eighth sternite, Part B: A) Lycomorpha pholus; TE- 8S up to 8T edge, P-pleurite, 6S-sixth sternite, 6T-sixth tergite, 7S-seventh sternite, 7T-seventh tergite, 8S-eighth sternite, 8T-eighth tergite; B) Gardinia anoploa; FT-8S fused to 8T, P-pleurite, 6S-sixth sternite, 6T-sixth tergite, 7S- seventh sternite, 7T-seventh tergite, 8S-eighth sternite, 8T-eighth tergite. The scale bar is equivalent to 1mm.

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Figure 2-8. Tegumen location. A) Lycomorpha pholus; TD-tegumen dorsad, C-costa, PS-pleural sclerite, U-uncus. B) Acsala anomala; TEV-tegumen extending ventrad, C-costa, U-uncus. The scale bar is equivalent to 1mm.

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Figure 2-9. Tegumen fusion. A) Apistosia judas; NF-no fusion of tegumen halves. B) Clemensia albata; TF- tegumen halves fused, FL-fusion long, NS-no sutures from fusion. C) Gardinia anoploa; TF-tegumen halves fused, FS-fusion short, S-sutures indicating location of fusion. The scale bar is equivalent to 1mm.

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Figure 2-10. Pleural sclerites. A) Acsala anomala; NS-no sclerites. B) Lycomorpha pholus; PS- pleural sclerites, MT-membranous tissue, SB-sclerotized bar. C) Lithosia quadra; PS-pleural sclerites, FT-fused to tegumen, MT-membranous tissue. D) Propyria morelosia; PS-pleural sclerites, SB-sclerotized bar. The scale bar is equivalent to 1mm.

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Figure 2-11. Juxta. A) Lithosia quadra; juxta SP-solid plate. B) Josiodes celena; juxta DM-divided by membrane. The scale bar is equivalent to 1mm.

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Figure 2-12. External view, left valva. A) Inopsis modulata; distal margin of C-costa occurring as an identifiable break, S-sacculus, CV-cucullus and valvulla fused into a contiguous structure, FV-fleshy valvulla found on external surface of valva, SC-spiny corona. B) Clemensia albata; C-costa and cucullus are a continuous structure, T-tendon, S-sacculus, CV-cucullus and valvulla fused into a contiguous lobe, cucullus forming heavily sclerotized dorsal portion and valvulla forms fleshy ventral margin. C) Lycomorpha fulgens nr.; distal margin of C-costa occurring as an identifiable break, PB-processus basalis, PE- proximal editum, S-sacculus, CU-cucullus fused to both C-costa and S- sacculus with no distinct ends. D) Lithosia quadra; C-costa and cucullus are a continuous structure, T-tendon, S-sacculus, DSP-distal saccular process extends to the distal apex of the valva and tapers to a point, CV-cucullus and valvulla fused into a contiguous structure with cucullus forming slightly sclerotized dorsal margin of the fleshy lobe of the valvulla. The scale bar is equivalent to 1mm.

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Figure 2-13. Internal view, right valva. A) Lycomorpha fulgens nr.; LA-long anellifer extends more than two-thirds the length of valva, PE-proximal editum, S- sacculus does not extend onto internal face of valva, T-tendon. B) Lithosia quadra; D-digitus emerges from distal end of costa, SA-short anellifer extends two-thirds the length of valva or less, S-sacculus, DSP-distal saccular process extends to distal apex of the valva and tapers to a point, T-tendon. C) Inopsis modulata; well developed CL-clasper, sclerotized CU-cucullus present on internal face of valva, D-digitus emerges from distal end of costa, DE-distal editum, DM-dorsal margin of sacculus extends onto internal face of valva, SA- short anellifer, dense SC-spined corona. D) Eilema bipuncta; not dense CS- corona of seta, D-digitus emerges from distal end of costa, DM-dorsal margin of sacculus extends onto internal face of valva, SA-short anellifer. The scale bar is equivalent to 1mm.

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Figure 2-14. External view, left valva (A,B) and Internal view, right valva (C,D). A) Miltochrista flexuosa; CU-cucullus is fused to the C-costa, T-tendon, V- valvulla a fleshy lobe that separates CU-cucullus from S-sacculus, DSP-distal saccular process tapers to a point but does not extend to the distal apex of valva. B) Eilema bipuncta; CV-cucullus and valvulla form a fleshy lobe, C- costa, DSP-distal saccular process does not extend to the distal apex of valva or taper to a point, S-sacculus. C) Josiodes celena; CL-clasper reduced, DE- distal editum, DM-dorsal margin of sacculus extends onto internal face of valva, LA-long anellifer, T-tendon. D) Clemensia albata; CV-cucullus and valvulla, DM-dorsal margin of sacculus extends onto internal face of valva, PSP-proximal saccular process does not extend to the distal apex of valva or taper to a point, SA-short anellifer, T-tendon. The scale bar is equivalent to 1mm.

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Figure 2-15. Lateral view, left side, phallus. A) Eilema bipuncta; DC-developed caecum with DES-ductus ejaculatorious simplex located dorsad of caecum, V-vesica. B) Miltochrista flexuosa; RC-reduced caecum with DES-ductus ejaculatorious simplex emerging from the anterior dorsal end of caecum, C-cornuti, V- vesica. C) Lycomorpha fulgens nr.; CA- caecum absent, DES-ductus ejaculatorious simplex emerges from the anterior end of phallus, C-cornuti, V- vesica. The scale bar is equivalent to 1mm.

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Figure 2-16. Abdominal pelt (A,C) and female genital capsule (B,D), Part A. A) Lycomorpha fulgens nr.; T-tergum, S-sternum; (B) Lycomorpha fulgens nr.; Seventh abdominal segment HS-heavily sclerotized without breaks in the sclerotization, eighth tergite (8T) extends beyond the distal margin of seventh tergite. C) Josiodes celena; T-tergum, S-sternum; D) Josiodes celena; Seventh abdominal segment heavily sclerotized with MS-membranous sternite, eighth tergite (8T) extends beyond the distal margin of seventh tergite. Stippling indicates differences in degree of sclerotization. The scale bar is equivalent to 1mm.

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Figure 2-17. Abdominal pelt (A,C) and female genital capsule (B,D), Part B. A) Lithosia sororcla; T-tergum, S-sternum; B) Lithosia sororcula; Seventh abdominal segment heavily sclerotized with MT-membranous tergite, eighth tergite (8T) does not extend beyond the distal margin of seventh tergite. C) Lycomorpha pholus; T-tergum, S-sternum; D) Lycomorpha pholus; Seventh abdominal segment heavily sclerotized with MP-membranous pleurites, eighth tergite (8T) extends beyond distal margin of seventh tergite. Stippling indicates differences in degree of sclerotization. The scale bar is equivalent to 1mm.

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Figure 2-18. Abdominal pelt (A) and female genital capsule (B), Part C. A) Hypoprepia cadaverosa; T-tergum, S-sternum; B) Hypoprepia cadaverosa; Sclerotization of seventh abdominal segment equivalent to preceding segments, eighth tergite (8T) extends beyond the distal margin of seventh tergite. Stippling indicates differences in degree of sclerotization. The scale bar is equivalent to 1mm.

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Figure 2-19. Ventral female genital capsule. A) Gardinia anoploa; OB-ostium bursa located in the seventh sternite (7S). B) Lycomorpha fulgens; OB-ostium bursa located in the membrane between the seventh and eighth sternite, distal margin of the seventh sternite (7S) NH-nearly horizontal. C) Lycomorpha grotei; OB-ostium bursa located in the membrane between the seventh and eighth sternite, distal margin of the seventh sternite (7S) DI-deeply indented. D) Lycomorpha pholus; OB-ostium located in the eighth sternite (8S), seventh sternite (7S). The scale bar is equivalent to 1mm.

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Figure 2-20. Bursa copulatrix. A) Propyria morelosia; DB-ductus bursa sclerotized, SP- sclerotization proximal to the OB-ostium bursa, CB-corpus bursa. B) Lithosia sororcula; DB-ductus bursa sclerotized, sclerotization is not proximal to the OB-ostium bursa, CB-corpus bursa. C) Lycomorpha fulgens; DB-ductsu bursa not sclerotized, OB-ostium bursa, CB-corpus bursa. Stippling indicates the presence of sclerotization. The scale bar is equivalent to 1mm.

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Figure 2-21. Strict consensus of the 4408 MP trees (L=736) L=1053, CI=0.10, RI=0.34 for the parsimony analysis rooted with Asota heliconia. BS values given above branches.

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Figure 2-22. Majority rule consensus tree of the Bayesian analysis rooted with Asota heliconia. Posterior probability support values given above the branches.

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CHAPTER 3 MOLECULAR PHYLOGENY OF THE TRIBE LITHOSIINI (LEPIDOPTERA: EREBIDAE: ARCTIINAE)

Background Information

The tribe Lithosiini (Lepidoptera: Erebidae: Arctiinae) presently contains approximately 3,150 species. However, it is estimated that the total species number could reach 5,000 to 6,000. These species are placed in 457 genera that are cosmopolitan in distribution. Although members of Lithosiini are found worldwide, the biodiversity of the tribe is highest within the tropics (Scoble 1992; Holloway 2002). In addition, the highest number of endemic genera occurs within Australia (Common

1990). Within the tribe, Bendib & Minet (1999) have proposed the existence of seven subtribes. However, no phylogenetic analysis has been conducted to determine whether the subtribes constitute natural groups. Furthermore, only 112 genera of Lithosiini have been placed in a subtribe. Adult Lithosiini are commonly known as lichen moths. This name is derived from the hypothesized feeding behavior of the larvae: lichenivory.

However, debate exists as to whether the larvae are lichen feeders or algal grazers

(Hesbacher et al. 1995; Wagner et al. 2008). Although the true larval feeding behavior is uncertain, Pöykkö and Hyvärinen (2003) observed later instar larvae of Eilema species

consuming the fungal thalli. In addition, the Lithosiini are also capable of sequestering

phenolics produced by the fungal symbiont of the lichen (Hesbacher et al. 1995;

Wagner et al. 2008). Although the tribe is known for its association with lichens, it will

not be possible to study the evolution of this behavior until the evolutionary relationships

among the genera are understood.

The tribe Lithosiini has been recovered as monophyletic in three separate studies

(Jacobson & Weller 2002; Wink & von Nikisch-Rosenegk 1997; Zahiri et al. 2012).

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However, the taxon sampling (7, 5, and 5 genera respectively) was not large enough to make inferences about the evolutionary relationships among the genera. Scott &

Branham (In Press) conducted the first phylogenetic analysis of the tribe using adult morphological characters and dense taxon sampling. Although they recovered some nodal support for relationships among small species groups, the higher-level relationships within the tribe were unresolved. This result was unsurprising since Bendib

& Minet (1999), Jacobson & Weller (2002), and Holloway (2002) all noted that the larval morphology was the most reliable for accurately placing taxa within Lithosiini. Although a study of the tribe conducted using both larval and adult morphological characters might provide more resolution to the higher-level relationships, the larvae of most lithosiine species that occur in the tropics are unknown (Holloway 2002). Although the

adult morphology of the tribe was not able to resolve the higher-level relationships

within Lithosiini, no studies have been conducted that attempt to resolve these

relationships using molecular data.

Recently molecular data has been found to be useful in resolving the higher-

level relationships within the order Lepidoptera (Regier et al. 2009; Mutanen et al. 2010;

Zahiri et al. 2011) and the families that comprise this order (Wahlberg et al. 2003, 2009,

2010; Rota 2011; Zahiri et al. 2012). These studies frequently combine multiple,

independent gene markers selected from both the mitochondrial DNA (mtDNA) and

nuclear DNA (nDNA). These two sources of DNA experience different rates of change.

The mtDNA is only transmitted through the maternal line, is non-recombinant, and

evolves at a higher rate than the nDNA (Simon et al. 1994). Each of these

characteristics of mtDNA allow it to aid in the identification of cryptic species and

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resolution of relationships among closely related species and genera (Lafontaine &

Schmidt 2010). However, the rapid rate of evolution in the mtDNA sometimes lead to overestimates of species numbers (Schmidt & Sperling 2008). In addition, the use of the full mitochondrial genome of several insect orders was unable to recover robust support for the deeper nodes of Arthropoda (Cameron et al. 2004). Although the mtDNA evolves at a rapid rate, nDNA has a slower mutation rate and undergoes recombination during meiosis. The slower rate of evolution within nDNA has made it useful in recovering the higher-level relationships within Lepidoptera (Regier et al. 2009; Mutanen et al. 2010).

When a dataset is composed of multiple genes, the difference in the rates of

evolution between mtDNA and nDNA provides biological justification for partitioning the

dataset (Rota 2011). However, as the number of genes included in an analysis

increases, the number of possible partitioning strategies exponentially increases (Lafear

et al. 2012). In addition, when a model-based method of phylogenetic analysis is used

to analyze these datasets, proper partitioning has been shown to increase the accuracy

of the phylogeny (Brown & Lemmon 2007). A commonly used way to select among

different partitioning strategies is the use of Bayes factors (Brandley et al. 2005; Brown

& Lemmon 2007). The Bayes factor is interpreted as the posterior probability of partition

scheme 1 over partition scheme 2 in a Bayesian analysis where each scheme has an

equal possibility of being true (Kass & Raftery 1995). After a partitioning scheme has

been selected, appropriate models of evolution must be selected for each partition of

the data. Selecting more complex models of evolution for each partition increases the

computational time of an analysis. However, the use of under-parameterized models has been found to lead to bias in Bayesian analyses (Huelsenbeck & Rannala 2004;

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Lemmon & Moriarty 2004). Several different statistical criteria have been developed to aid researchers in selecting appropriate models (Sullivan & Joyce 2005). Two criteria

that are frequently used to select the model of evolution are the Akaike information

criterion (AIC; Akaike 1974) and the Bayesian information criterion (BIC; Schwarz

1978). In addition, the AIC criterion has been modified to address small sample sizes:

the corrected Akaike information criterion (AICc: Sugiura 1978; Hurvich & Tsai 1989).

Each of these criteria is able to compare nested and non-nested models

simultaneously. However, the BIC penalizes overparameterization more strongly than

then AIC or AICc, and the BIC is less likely to increase the number of parameters as the

number of sites in the dataset increases. Although some researchers have shown the

BIC performs better in data simulations where the true model is known (Luo et al. 2010),

this disregards the fact that the AIC and AICc do not assume that any of the candidate

models are true (Sullivan & Joyce 2005). Therefore, the models selected by the three

criteria should be compared using Bayes factors since the model of evolution can only

truly be known in simulation studies.

The aim of this study is to use four gene fragments to analyze the evolutionary

relationships within Lithosiini. The gene fragments examined include two mitochondrial

markers, cytochrome oxidase C subunit I (COI) and cytochrome oxidase B (CytB), and

two nuclear markers, ribosomal protein S5 (RpS5) and the nuclear large subunit rRNA

28S D2 loop (28S). Each of the fragments is from a protein-coding gene, except for

28S. These genes have been shown to be phylogenetically informative at the higher-

level in Lepidoptera (Wahlberg et al. 2009; Mutanen et al. 2010; Zahiri et al. 2011,

2012). These genes are sequenced for forty-six species of Lithosiini representing four

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subtribes, as well as species not yet placed in a subtribe. Model-based phylogenetic methods, maximum likelihood and Bayesian inference, are used to examine the dataset. The phylogenies obtained are examined to study the deeper relationships within Lithosiini and assess the monophyly of the four subtribes included in the analysis.

In addition, the effect of different selection criterion (AIC, AICc, and BIC) for obtaining the optimal model of evolution is studied using Bayes factors (Kass & Raftery 1995).

Materials and Methods

Taxon Sampling

Forty-six species of Lithosiini representing twenty-six genera were included in the analysis (Table 3-1). Both Old World and New World species were sampled. Australia, an area with a high number of endemic genera of Lithosiini (Common 1990; Edwards

1996), was represented by 15 species that are placed in eight genera. However, the

Neotropics, Eastern Europe and Asia were represented by only eight species from seven genera. The species that were sampled are presently placed in four of the lithosiine subtribes proposed by Bendib & Minet (1999). The subtribes that were represented are Acsalina (1 species, 1 genus), Cisthenina (13 species, 7 genera),

Lithosiina (17 species, 9 genera), and Nudariina (10 species, 5 genera). In addition, species that have not yet been placed in a subtribe were also included in the analysis (5 species, 5 genera). Of the three subtribes that were not sampled, one subtribe,

Phryganopterygina, is monogeneric and restricted to Madagascar. The genera that comprise the other two, unrepresented subtribes are found in the regions for which few specimens suitable for DNA extraction (thirty years old or less) were available:

Neotropics (Eudesmiina) and the Palearctic and Oriental region (Endrosina).

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To test the monophyly of Lithosiini, outgroup taxa were selected from the subfamily Aganainae and the remaining two tribes of Arctiinae, Arctiini and Syntomiini.

Aganainae was represented by three species, which are placed in two genera. Kitching

(1984) and Zahiri et al. (2011, 2012) found Aganainae to be sister to Arctiinae. Pagara simplex Walker was chosen to represent the tribe Arctiini. Previously, P. simplex has

been treated as a member of Lithosiini (Covell 1984; Forbes 1960; Franclemont 1983).

However, Bendib & Minet (1999) considered this species to be an aocellate

phaegopterine, and Ferguson and Opler (2006) transferred it to Arctiini. In a

phylogenetic analysis of Lithosiini based on morphological data (Scott & Branham In

Press), P. simplex was recovered outside of the tribe. The tribe Syntomiini was

represented by Amata aperta (Meyrick). In previous phylogenetic analyses, the position

of Syntomiini within Arctiinae has varied based on the data type used in the analysis.

Using morphological data, Jacobson & Weller (2002) recovered Syntomiini sister to

Lithosiini. Zahiri et al. (2011, 2012) recovered Syntomiini sister to Arctiini using

molecular data. In this study, all analyses were rooted on the aganaine species Asota

heliconia (Linnaeus).

DNA Extraction, Amplification, and Sequencing

A voucher number was assigned to each specimen used for DNA extraction. This

voucher was placed in a vial or on a pin with the remaining moth body. This number

was also used to identify the DNA extract from the same specimen. DNA was extracted

from three legs taken from the right side of the thorax of specimens that were frozen or

stored in 96% ethanol. If a specimen had been allowed to dry, DNA was extracted from

the abdomen of the specimen using the method proposed by Knölke et al. (2005) and

Hundsdoerfer and Kitching (2010). The DNA extraction was performed using the

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DNeasy blood and tissue kit (Qiagen, Valencia, CA) with modifications noted below.

Unless stated otherwise, incubations were completed at 56˚C. When legs were used as a source of DNA, they were placed in a 1.5ml microcentrifuge tube containing 180µl

Buffer ATL and 20µl proteinase K, and a plastic tissue grinder (Axygen*, Union City,

CA) was used to homogenize the tissue in the tube. After homogenization, the tube was incubated for 12 to 14 hours. When an abdomen was used as a source of DNA, the abdomen was removed from the specimen and placed in a 1.5ml microcentrifuge tube in 140μl of Buffer ATL. The tube was incubated for 30 minutes. After the initial incubation period, 7µl of proteinase K was pipetted into the basal opening where the abdomen had been attached to the thorax. The tube was incubated for an additional 90 minutes. After the second incubation period, another 7μl of proteinase K was added to the basal opening of the abdomen. On female specimens, an incision was made down the pleurites on one side of the abdomen to prevent rupturing before the second aliquot of proteinase K was added. The tube was further incubated for 12 to 14 hours.

Following this incubation, the abdomen was placed onto a piece of parafilm in 40μl of

ATL buffer. Remaining soft tissue was removed from the abdomen using forceps that had been sterilized using a Spectrolinker XL-1000 UV crosslinker (Spectronics

Corporation). The abdominal pelt and genital capsule were stored in a vial that contained 20% ethanol to be dissected later. The soft tissue and ATL buffer were returned to the microcentrifuge tube and 6μl of proteinase K was added. The tube was returned to the water bath and incubated until all of the soft tissue was digested (30 minutes to several hours). Following the digestion of the tissue, the same extraction procedure was used for both the leg and abdominal tissue. The DNA was bound to a

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spin column membrane following the DNeasy blood and tissue kit protocol. After the

DNA had been bound to the spin column, it was washed twice with 500µl of AW1 buffer.

The sample was also washed twice with 500µl of AW2 buffer. After the washes were completed, the spin column was transferred to a new 1.5ml tube. To elute the DNA from the spin column, 100µl of Buffer AE was added to the center of the spin column membrane. After incubating for one minute at room temperature, the tube was centrifuged for one minute at ≥ 6,000 x g. This was step was repeated to obtain 200µl of

DNA extract. The DNA extracts were quantified using a Nanodrop ND-1000 spectrophotometer (ThermoScientific).

Polymerase chain reactions (PCR) were used to amplify two mitochondrial gene fragments, cytochrome c oxidase subunit I (COI, 690bp) and cytochrome oxidase b

(CytB, 660bp), and two nuclear gene fragments, ribosomal protein S5 (RpS5, 660bp) and the nuclear large subunit rRNA 28S D2 loop (28S, 680bp). PCR was conducted using a high fidelity mastermix, Accuzyme (Bioline, Randolph, MA), to amplify the fragments from 1.0µl of template DNA. The final volume of the reaction was 25.0µl. The primers, reagents, and thermocycler profiles used for each gene can be found in Table

3-2. The products obtained from PCR were separated using agarose gel electrophoresis (1% TBE gels). The DNA was stained using GelRed (Phenix, Candler,

NC) and visualized with ultraviolet light. The gene fragments were purified using a

QIAquick PCR Purification Kit (Qiagen, Valencia, CA). The purified products were sequenced in both the forward and reverse directions at the Biomedical Genomics

Center at the University of Minnesota Core Facility. The sequences were edited, aligned

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using MUSCLE (Edgar 2004), and concatenated in Geneious v5.6.5 (Drummond et al.

2011). The final concatenated file was 2878bp long.

Phylogenetic Methods

The concatenated dataset was analyzed using PartitionFinder v1.0.1 (Lanfear et al. 2012) to determine the optimal data partitioning strategy and model of evolution for each partition. The optimal partitioning scheme was determined using the Akaike information criterion (AIC; Akaike 1974), the corrected Akaike information criterion

(AICc: Sugiura 1978; Hurvich & Tsai 1989), and the Bayesian information criterion (BIC;

Schwarz 1978) (Table 3-3).

Phylogenetic trees were constructed using maximum likelihood (ML) and

Bayesian inference (BI) methods. In order to employ both optimal partitioning schemes identified with PartitionFinder, the ML and BI analyses were conducted twice. The ML analysis was conducted using RAxML – HPC2 v7.3.2 (Stamatikis 2006; Stamatikis et al.

2008) with the default settings. The GTR + G model of evolution was applied for each data partition in the analysis. Nodal support was calculated using 1000 bootstrap (BS) pseudoreplicates (Felsenstein 1985). These values were mapped onto the most likely topology identified by RAxML. The Bayesian Inference (BI) analysis was conducted using MrBayes v3.1.2 (Ronquist & Huelsenbeck 2003). The models of evolution used for each data partition can be found in Table 3-3. The Bayesian analyses were performed with four chains, one cold and three hot, using the default temperature settings. Two simultaneous, independent runs of 20,000,000 generations were conducted. The Markov chain Monte Carlo (MCMC) chains were started at random trees. Samples were drawn from the cold chain every 1,000 generations. Five million generations were discarded as ‘burn-in’. The probabilities of the two runs were

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summarized, and the potential scale reduction factor (PSRF: Gelman & Rubin 1992) was calculated to confirm that the runs had converged. The PSRF value should approach 1.000 as the independent runs converge. The trees from the Bayesian analysis were summarized as a majority rule consensus tree. Posterior probability (PP) provided the clade credibility values for the tree. Both the ML and BI analysis were conducted using the CIPRES Science Gateway (Miller et al. 2010). The trees from both

types of analysis were visualized using FigTree v1.3.1 (Rambaut 2010).

Bayes factors (BF; Kass & Raftery 1995) were used to compare the optimal

partitioning schemes identified using PartitionFinder. In order to calculate the BF, the

harmonic mean (HM) of the likelihood score was obtained from each BI analysis by

summing the probabilities of the two runs in the analysis after the burn-in had been

discarded. In order to calculate BF, the formula 2ln(BF1) = 2[ln(HM1) – ln(HM2)] was

used (Brandley et al. 2005; Brown & Lemon 2007; Rota 2011). In this formula, HM1 was

the harmonic mean of the first partitioning strategy, and HM2 was the harmonic mean of

the second partitioning strategy. The significance of the BF that were calculated using

this formula was determined by the values presented by Kass and Raftery (1995) were

a BF > 150 provides strong support against the second partitioning strategy (Table 3-4).

Results and Discussion

Analyses were based on 2878bp of sequence obtained from four gene

fragments, two mitochondrial and two nuclear, sampled for 46 species of Lithosiini.

Each of the analyses was conducted twice in order to compare the optimal partitioning

schemes and models of evolution using the AIC, AICc, and BIC selection criteria. The

Bayes factors (BF) were calculated comparing the harmonic mean from the Bayesian

analysis using the BIC scheme (HM1 = -19918.22) to the harmonic mean from the

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analysis using the AIC, AICc scheme (HM2 = -19870.87). The BF obtained from this

comparison was 1.00238. Based on the values present by Kass and Raftery (1995;

Table 3-4) the difference in the probability of the two schemes was not significant.

Although the probability of the partitioning schemes identified by the selection criteria

were not found to be significantly different, the optimal scheme identified using the BIC

contained 24 fewer parameters than the one identified using the AIC and AICc. This is

congruent with the AIC and AICc not penalizing additional parameters as strongly as

BIC.

Both the maximum likelihood (ML) analysis (Figs 3-1, 3-2) and the Bayesian inference (BI) analysis (Figs 3-3, 3-4) recovered Arctiinae as a moderately to strongly supported monophyletic clade (BSAIC ≥ 79, BSBIC ≥ 78, PPAIC ≥ 0.99, PPBIC = 1).

However, none of the analyses recovered Lithosiini as a monophyletic group. Instead,

Lithosiini was found to be paraphyletic with respect to Syntomiini and Arctiini. In each

analysis, Pagara simplex (Arctiini) and Amata aperta (Syntomiini) were recovered in a

weakly supported (PPAIC ≥ 0.53, PPBIC ≥ 0.57) clade, which also contained Asura

polyspila Turner and Eilema sp.2. The relationships among these species were

unresolved in the BI analysis. However, the relationships within this clade were resolved

in the ML analysis, but there was no nodal support. In addition, the clade containing

Arctiini and Syntomiini was recovered in Clade D sister to the Miltochrista species

included in the analyses. Confusion has existed about the placement of P. simplex.

Although Covell (1984), Forbes (1960), and Franclemont (1983) treated it as a member

of Lithosiini, Bendib & Minet (1999) considered this species to be an aocellate

phaegopterine. However, S. Weller (pers. comm.) notes that the arrangement of the

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male valvae of P. simplex is inverted from the typical phaegopterine valvae. The males of P. simplex possess an entirely membranous costa and a sclerotized sacculus.

Although these analyses found Lithosiini to be paraphyletic to Arctiini and Syntomiini, both of those subtribes were represented by a single species. An analysis that includes several representatives of Arctiini and Syntimiini will help to clarify the relationships among the three tribes.

Four of the subtribes proposed by Bendib & Minet (1999) were represented in this analysis. In addition, five genera that have not yet been placed in a subtribe were also included. Three of the subtribes (Cisthenina, Lithosiina, and Nudariina) were represented by multiple species and genera. Both the ML and BI analyses recovered each of these subtribes to be polyphyletic. The majority of the species and genera representing these subtribes were recovered in one of four clades (Clades A – D).

These clades contained the same species in both partitioning schemes and

phylogenetic analyses. In addition, with the exception of Clade B and Clade D these

clades contained species from only one subtribe and unplaced species. The subtribe

Acsalina was represented by the single species that defines the subtribe, Acsala

anomala Benjamin. A. anomala was recovered sister to Ptychoglene coccinea

(Edwards) within the Clade B, which is composed primarily of species placed in

Cisthenina. This relationship received moderate support in both ML analyses but was

strongly supported in both of the BI analyses (BSAIC ≥ 75, BSBIC ≥ 75, PPAIC ≥ 0.99,

PPBIC ≥ 0.99). This result suggests that, as defined, Acsalina does not represent a

subtribe.

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All but two species of Lithosiinia were recovered in Clade A. This clade was weakly supported in all of the analyses (BSAIC ≥ 52, BSBIC ≥ 50, PPAIC ≥ 0.67, PPBIC ≥

0.64). Within Clade A, Eilema Hübner was found to be polyphyletic. Birket-Smith (1965) and Holloway (2002) noted that the genus Eilema, as presently defined, does not comprise a natural group. A moderately to strongly supported relationship was recovered between the type species of the type genus of Lithosiina, Lithosia quadra

(Linnaeus), and Eilema sp.1 (BSAIC ≥ 70, BSBIC ≥ 69, PPAIC ≥ 0.97, PPBIC ≥ 0.93). In

addition, this clade was found sister to the unplaced species Calamidia hirta (Walker)

(BSAIC ≥ 63, BSBIC ≥ 59, PPAIC ≥ 0.8, PPBIC ≥ 0.76). Another unplaced species,

gyge Druce, was also recovered within Clade A. However, the placement of H. gyge

varied based on both on the type of analysis and the partitioning scheme used.

Most of the species representing Cisthenina were found in Clade B and Clade C.

Clade B received moderate support in both of the BI analyses (PPAIC ≥ 0.86, PPBIC ≥

0.86). However, this clade was only weakly supported in the ML analysis using the AIC

partition strategy (BSAIC ≥ 51). All representatives of Cisthene Walker, except Cisthene

subjecta Walker, were found in this clade. Within Clade B, these species were divided

between two clades. The species of Cisthene range from North America through the

Neotropics. All of the species of Cisthene included in this analysis occur in North

America. Although Knowlton (1967) completed a revision of the species that occur north

of Mexico, no phylogenetic analysis was done to assess the generic limits of Cisthene.

The findings of this study suggest that, as currently defined, Cisthene is polyphyletic.

The unplaced species Meterhythosia sangala (Druce) was recovered in Clade B in all

analyses. M. sangala was recovered sister to the well supported Bruceia hubbardi Dyar

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+ Prepiella aurea nr. clade (BSAIC ≥ 98, BSBIC ≥ 98, PPAIC = 1, PPBIC = 1). Although this

relationship was recovered in all of the analyses, it received only moderate support in

the BI analyses (PPAIC ≥ 0.72, PPBIC ≥ 0.72). Clade C, which contains the remainder of

the species of Cisthenina, has no nodal support in either ML analysis but receives

moderate to strong support in the BI analyses (PPAIC ≥ 0.94, PPBIC ≥ 0.67). All of the

species of Hypoprepia Hübner were found in this clade as well as Bruceia pulverina

Neumoegen. A strongly supported (BSAIC ≥ 89, BSBIC ≥ 90, PPAIC = 1, PPBIC ≥ 0.69)

sister relationship was recovered between B. pulverina and Hypoprepia cadaverosa

Streck.

Clade D contained most of the species and three of the five genera representing

Nudariina. This clade received moderate support in the BI analyses (PPAIC ≥ 0.67, PPBIC

≥ 0.65). Of the three nudariine genera found in Clade D, only Miltochrista Hübner was

recovered as monophyletic (BSAIC = 100, BSBIC = 100, PPAIC = 1, PPBIC = 1). The

Miltochrista clade was recovered sister to the clade containing the representatives of

Syntomiini and Arctiini in each analysis (PPAIC ≥ 0.6, PPBIC ≥ 0.65). The genus Asura

Walker was found to be polyphyletic in all of the analyses. Bendib & Minet (1999) stated

that Asura was not monophyletic as defined. Lyclene Moore was found to be

paraphyletic with respect to Asura cervicalis Walker and the unplaced species Prinasura

quadrilineata (Pagenstecher). This relationship was strongly supported (BSAIC ≥ 0.95,

BSBIC ≥ 0.95, PPAIC = 1, PPBIC = 1). This finding was not unexpected since Lyclene and

Asura possess similar color patterns.

The evolutionary relationships of five of the species of Lithosiini (Cisthene

subjecta, Cyana meyricki (Rothschild & Jordan), Gardinia anoploa Hering, Heliosia

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jucunda (Walker), Schistophleps albida Walker) were unresolved among the analyses.

The placement of C. subjecta, the type species of the type genus of Cisthenina, and S. albida was dependent upon the type of analysis. In both ML analyses, C. subjecta was found in the basal most clade of the tree sister to the nudariine species S. albida.

However, there was no nodal support for this relationship. In addition, this relationship

was not recovered in either of the BI analyses. The nudariine species C. meyricki was

recovered sister to Clade D in the ML analyses, but its position was unresolved in the BI

analyses. G. anoploa and H. jucunda were both found to be related to Clade C in the

ML analysis. Again, these relationships were not recovered in the BI analysis.

This study found that molecular data were better able to resolve the evolutionary

relationships among the genera of Lithosiini. However, this dataset was still unable to

provide nodal support for the deeper nodes within the tribe. The majority of the species

of Lithosiini included in this analysis were missing data from at least one gene. In eleven

of the species, only the 28S gene marker was successfully sequenced. These gaps in

the data may be contributing to the lack of support for the higher-level relationships. In

addition, increasing the number of nDNA markers used in the analysis could aid in the

resolution of these relationships. Half of the gene markers used in this analysis were

obtained from the mtDNA, and the studies that have recovered the higher-level

relationships of families and tribes with robust nodal support (Wahlberg et al. 2003,

2009, 2010; Zahiri et al. 2011, 2012) included more nDNA fragments than mtDNA

fragments. Also, the study did not recover the tribe Lithosiini as monophyletic with

respect to the other two tribes of Arctiinae. This finding may be due to the gaps in the

data as well as the used of a single representative for the tribes Arctiini and Syntomiini.

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As with the study of the tribe based on morphological data, the subtribes proposed by

Bendib & Minet (1999) were not recovered as monophyletic groups. Although three of the subtribes were found to be polyphyletic, four clades composed almost entirely of species from single subtribes were recovered. This result suggests that with modification, the subtribes of Bendib & Minet could define natural groups. This study also identified several genera that are polyphyletic as presently defined. This finding was not surprising since wing color patterns have been used to place species in both

Cisthene and Asura. The use of this type of character to place species within a genus resulted in 19 species being mistakenly described as members of Lycomorpha Harris.

Although the use of molecular data provided more resolution of the relationships among the genera of Lithosiini, further analyses will be needed before the evolution of lichen feeding behaviors within the tribe can be studied.

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Table 3-1. List of species used in the analysis including the subtribe, locality, assigned voucher number, collection that provided the material, and the gene fragments that were successfully amplified. Taxa (Genus species) Locality Voucher #, Collection Gene Fragments 28S COI CytB RpS5 EREBIDAE Arctiinae Lithosiini Acsalina Acsala anomala Benjamin Yukon Territory JZCS133, CNC 28S - - RpS5 Cisthenina Bruceia hubbardi Dyar Arizona, USA PC015, CHS 28S COI - - B. pulverina Neumoegen Wyoming, USA JZCS132, CNC 28S COI - - Cisthene juanita Barnes & Benjamin Arizona, USA PC017, CHS 28S COI - - C. plumbea Stretch , USA JZCS139, CNC 28S COI CytB - C. subjecta Walker , USA PC011, CHS 28S COI - - C. tenuifascia Harvey Arizona, USA PC014, CHS 28S - - - Hypoprepia cadaverosa Streck Colorado, USA JZCS131, CNC 28S COI CytB - H. fucosa Hübner Tennessee, USA PC019, CHS 28S - - - H. fucosa tricolor (Fitch) Minnesota, USA JZCS062, UWO 28S COI CytB RpS5 Lycomorpha pholus (Drury) Connecticut, USA PC022, YPM 28S - - - Lycomorphodes sordida (Butler) Guatemala JZCS140, CNC 28S - - - Prepiella aurea nr. Guatemala JZCS141, CNC 28S - - - Ptychoglene coccinea (Edwards) Arizona, USA PC084, USNM 28S - - - Lithosiina Agylla septentrionalis Barnes & McDunnough Arizona, USA JZCS125, CNC - COI - - rubricollis (Linnaeus) Denmark JZCS135, CNC 28S - CytB RpS5 Crambidia lithosioides Dyar Tennessee, USA PC018, CHS 28S - - - Cybosia mesomella (Linnaeus) Denmark JZCS137, CNC 28S - - - Eilema bicolor (Grote) Minnesota, USA JZCS060, UWO 28S COI CytB RpS5 E. complana (Linnaeus) Denmark JZCS136, CNC 28S COI - RpS5 E. dorsalis (Walker) Australia JZCS040, UMD 28S COI CytB - E. griseola (Hübner) France JZCS134, CNC 28S COI CytB RpS5

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Table 3-1. Continued Taxa (Genus species) Locality Voucher #, Collection Gene Fragments E. plana (Boisduval) Australia JZCS028, UMD 28S COI - - Eilema sp.1 Australia JZCS082, UMD 28S COI CytB RpS5 Eilema sp.2 Australia JZCS087, UMD 28S COI - RpS5 Eilema sp.3 Russia/Nepal JZCS107, 28S COI CytB - UWO Eilema sp.4 Russia/Nepal JZCS108, 28S COI - - UWO Gardinia anoploa Hering Arizona, USA JZCS126, CNC 28S COI - RpS5 Gnamptonychia flavicollis (Druce) Arizona, USA JZCS127, CNC 28S COI CytB RpS5 Inopsis modulata (Edwards) Arizona, USA JZCS128, CNC 28S COI - - Lithosia quadra (Linnaeus) Russia JZCS105, 28S COI - - UWO Nudariina Asura cervicalis Walker Australia JZCS047, UMD 28S COI - - A. polyspila Turner Australia JZCS033, UMD 28S COI CytB RpS5 Cyana meyricki (Rothschild & Jordan) Australia JZCS001, UMD 28S COI CytB RpS5 Lyclene pyraula (Meyrick) Australia JZCS020, UMD 28S COI CytB - L. reticulata (Felder) Australia JZCS041, UMD - COI CytB RpS5 Lyclene sp.1 Australia JZCS038, UMD 28S COI - - Lyclene sp.1 nr. Australia JZCS048, UMD 28S COI - RpS5 (Forster) Denmark JZCS138, CNC - COI CytB RpS5 Miltochrista sp.1 Russia/Nepal JZCS109, 28S COI - - UWO Schistophleps albida (Walker) Australia JZCS056, UMD 28S COI - - Unplaced Calamidia hirta (Walker) Australia JZCS009, UMD 28S COI - RpS5 Heliosia jucunda (Walker) Australia JZCS071, UMD 28S - - - Hiera gyge Druce Costa Rica JZCS130, CNC 28S - - - Meterhythosia sangala (Druce) Guatemala JZCS129, CNC 28S - - - Prinasura quadrilineata Australia JZCS019, UMD 28S COI - - (Pagenstecher)

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Table 3-1. Continued Taxa (Genus species) Locality Voucher #, Collection Gene Fragments Syntomiini Amata aperta (Meyrick) Australia JZCS081, UMD 28S COI - RpS5 Arctiini Pagara simplex Walker Tennessee, USA PC004, CHS 28S - - - Aganainae Asota heliconia Australia JZCS029, UMD 28S COI CytB - (Linnaeus) As. orbona (Vollenhoven) Australia JZCS078, UMD 28S COI CytB - Neochera dominia Australia JZCS008, UMD 28S COI CytB - (Cramer) Collections: CHS (Collection of C. Scott), USNM (United States Museum of Natural History), UWO (University of Wisconsin – Oshkosh), UMD (University of Maryland), YPM (Yale Peabody Museum)

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Table 3-2. PCR protocol data including thermocycler profiles, primer sequences for each gene used, and the reagents used. Thermocycler Profiles COI: 95˚C/3 min; 33 cycles: 95˚C/1 min, 48˚C/1 min, 72˚C/1 min; A final extension of 95˚C/3 min

CytB: 94˚C/5 min; 29 cycles: 94˚C/1 min, 45˚C/1 min, 72˚C/1 min; One final extension cycle: 94˚C/1 min, 45˚C/1 min, 72˚C/10 min.

28S D2 Loop: 94˚C/2 min 30 sec; 40 cycles: 94˚C/30 sec, 65˚C/30 sec, 72.0˚C/30 sec; A final extension of 72.0˚C/2 min.

RpS5: 95˚C/5 min; 40 cycles: 94˚C/30 sec, 55˚C/30 sec, 72˚C/1 min 30 sec; A final extension of 72˚C/10 min.

Primer Name Sequence from 5’ to 3’ COI DNA Barcode Segment LepF1 5’ – ATTCAACCAATCATAAAGATATTGG – 3’ LepR1 5’ – TAAACTTCTGGATGTCCAAAAAATCA – 3’

CytB REVCB2H 5’ – TGAGGACAAATATCATTTTGAGGW – 3’ REVCBJ 5’ – ACTGGTCGAGCTCCAATTCATGT – 3’

28S D2 Loop LEE 28S_D2-F1 5’ – GAGTACGTGAAACCGTTCAG – 3’ 28S_D2-R1 5’ – CTGACCAGGCATAGTTCAC – 3’

RpS5 HybrpS5degF 5’ – ATGGCNGARGARAAYTGGAAYGA – 3’ HybrpS5degR 5’ – CGGTTRGAYTTRGCAACACG – 3’

Reagents Volume Accuzyme Mix 12.5µl Taq Polymerase 0.5µl ddH2O 9.0µl Forward Primer [20pmol] 1.0µl Reverse Primer [20pmol] 1.0µl DNA template 1.0µl

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Table 3-3. Optimal partitioning schemes selected by PartitionFinder v1.0.1 (Lanfear et al. 2012) using the AIC, AICc, and BIC selection criteria. The model of evolution for each partition, the number of subsets, parameters, and the log likelihood for each scheme is included. Selection Model of Subset Parameter lnL Optimal Scheme Criterion Evolution # # AIC, AICc 28S = 1–805 GTR+I+G 10 168 -19363.032 COI nt1 = 806–1555\3 GTR+G COI nt2 = 807–1555\3 F81 COI nt3 = 808–1555\3 GTR+G CytB nt1 = 1556–2215\3 GTR+G CytB nt2 = 1557–2215\3 HKY CytB nt3 = 1558–2215\3 GTR+I+G RpS5 nt1 = 2216–2878\3 K80+I RpS5 nt2 = 2217–2878\3 JC+I RpS5 nt3 = 2218–2878\3 GTR+I+G

BIC 28S = 1–805 GTR+I+G 7 144 -19411.662 COI, CytB nt1 = 806–1555\3, 1556–2215\3 GTR+G COI, CytB nt2 = 807–1555\3, 1557–2215\3 HKY COI, CytB nt3 = 808–1555\3, 1558–2215\3 GTR+G RpS5 nt1 = 2216–2878\3 K80+I RpS5 nt2 = 2217–2878\3 JC+1 RpS5 nt3 = 2218–2878\3 GTR+I+G

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Table 3-4. Interpretation of the Bayes Factor (BF) adapted from Kass and Raftery (1995) BF1 Evidence for Hypothesis 1 1 – 3 Not worth mentioning 3 – 20 Positive 20 – 150 Strong >150 Very Strong

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Figure 3-1. Maximum likelihood tree from analysis of AIC, AICc optimal partitioning scheme. LnL = -18742.441364. Bootstrap branch support values ≥50 given above the branch. A – D = Clades A – D.

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Figure 3-2. Maximum likelihood tree from analysis of BIC optimal partitioning scheme. LnL = -18792.271033. Bootstrap branch support values ≥50 given above the branch. A – D = Clades A – D.

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Figure 3-3. Majority rule consensus tree of the Bayesian analysis using the AIC, AICc optimal partitioning scheme. Harmonic mean likelihood = -19870.87. Posterior probability branch support values given above the branch. A – D = Clades A – D.

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Figure 3-4. Majority rule consensus tree of the Bayesian analysis using the BIC optimal partitioning scheme. Harmonic mean likelihood = -19918.22. Posterior probability branch support values given above the branch. A – D = Clades A – D.

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CHAPTER 4 PHYLOGENY OF THE LICHEN MOTH GENUS LYCOMORPHA HARRIS (LEPIDOPTERA: EREBIDAE: ARCTIINAE)

Background Information

The lichen moth genus Lycomorpha Harris (Lepidoptera: Erebidae: Arctiinae) is currently composed of seven species of lichen moths that are restricted to North

American genus (Forbes 1960; Powell & Opler 2009). It is currently placed within the tribe Lithosiini, and Bendib & Minet (1999) placed Lycomorpha within the subtribe

Cisthenina. Although adults of several species within Lycomorpha exhibit both diurnal and nocturnal flight activity, L. pholus (Drury) is exclusively diurnal (Fullard &

Napoleone 2001), and L splendens Barnes and McDunnough is reported to be exclusively nocturnal (Powell & Opler 2009). The adults of the genus are aposematically colored. They are proposed to be Müllerian mimics of lycid beetles, other tiger moths

(Simmons 2009), and zygaenid moths. The wing color patterns of the adults have led to taxonomic confusion in the placement of Lycomorpha within a family as well as the placement of species within the genus. A phylogenetic study of the genus is necessary to elucidate and define the generic limits of Lycomorpha.

Lycomorpha is currently placed within the tribe Lithosiini. However, Harris (1839) originally described the genus as a member of Zygaenidae. Other authors treated the genus as a member of several other families (Table 4-1). It was not until Forbes (1960) noted the coloration of the adults and the loss of the Sc vein were the only characters that supported the placement of Lycomorpha within Euchromiidae that the genus was placed within the tribe Lithosiini. Although the wing color patterns caused the genus being misplaced, the lack of synapomorphies from the adult morphology of the lichen moth tribe also contributed to the taxonomic confusion. In a phylogenetic study of the

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tribes of Arctiinae, Jacobson & Weller (2002) identified eight synapomorphies for

Lithosiini. All of these characters were coded from the larval morphology. In addition,

Jacobson & Weller (2002) included L. pholus, the type species of Lycomorpha, in their analysis. It was recovered as sister to the genus Hypoprepia Hübner within the Lithosiini clade. Of the eight synapomorphies identified by Jacobson & Weller (2009), the presence of a mandibular mola (a structure that is hypothesized to aid the larvae in grinding the tougher fungal tissue of lichens (Gardner 1943)) is now used to definitively place taxa within the tribe. When known, the larvae of Lycomorpha possess the mandibular mola (Jacobson & Weller 2002) and are reported to feed on lichens

(Comstock & Henne 1967; Dyar 1897; Wagner et al. 2008).

Although Jacobson & Weller (2002) confirmed the classification of Lycomorpha as a member of the tribe Lithosiini, confusion remains about the evolutionary relationships among the species within the genus. The adult wing color patterns have been the most commonly used character to place species within Lycomorpha. The use of this character has resulted in an additional nineteen species being described as members of the genus (Fig. 4-1). However, all of these species are now placed within other genera of Arctiini, Lithosiini, and Zygaenidae. In addition, phenotypic variation present within adult Lycomorpha led to the description of subspecies: L. pholus miniata

Packard and L. grotei pulchra Dyar for the species, L. pholus and L. grotei (Packard), respectively. Within the southwestern United States, four species (L. fulgens (Henry

Edwards), L. grotei, L. grotei pulchra, and L. regulus (Grinnell)) that have red forewings and primarily black hindwings occur (Fig 4-2). Due to the similar coloration and the

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sympatric ranges, Comstock & Henne (1967) and Powell & Opler (2009) suggested that these species be studied to determine their taxonomic and systematic relationships.

Most recently while conducting a phylogenetic analysis of the tribe Lithosiini,

Scott & Branham (In Press) recovered all of the included species of Lycomorpha and the genus Propyria Hampson within a single, well-supported clade (Bremer Support = 3,

Posterior Probability (PP) ≥ 0.98). In the maximum parsimony analysis, the relationships among the species representing the two genera were completely unresolved. However, in the Bayesian Inference analysis Lycomorpha was found to be paraphyletic with respect to Propyria. Propyria is a North American genus composed of nine species.

Adults from the genus have been collected from Mexico and as far south as Costa Rica.

The adults are aposematically colored, and one species, P. schausi (Dyar), was originally described as a member of Lycomorpha. Using wing venation, one character separates these genera. In the hindwing of Propyria, the M3 and CuA1 are fused and stalk beyond the discal cell. These veins are not fused within Lycomorpha.

In this study, we used characters coded from the adult morphology to conduct the first phylogenetic analysis of the genera Lycomorpha (7 species) and Propyria (9 species). This phylogenetic analysis is used to assess their generic limits and form a hypothesis of the evolutionary relationships among the species within each genus.

Larval specimens were only available for one species, L. pholus. Therefore, it was not possible to include larval characters within the analysis even though the larval morphology of Lithosiini has been found to provide the strongest phylogenetic signal

(Bendib & Minet 1999; Jacobson & Weller 2002). Sixty-five morphological characters were coded for thirteen species representing Lycomorpha and Propyria. In addition, the

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phylogenetic usefulness of the adult morphological characters in resolving the evolutionary relationships within the genera was assessed.

Materials and Methods

Taxon Sampling

Thirteen species representing Lycomorpha (10 species) and Propyria (3 species) were sampled for this study (Table 4-2). The type species for each genus (L. pholus and

P. ptychoglene Hampson) was included in the analysis. The species representing

Lycomorpha included three species that have not yet been described, as well as both of the subspecies. The species status of each subspecies was revived. Each subspecies possesses morphological characters that are unique from the species with which it was synonymized. Male-female pairs were available for all but two of the species. Both internal and external outgroups were included in the analysis to assess the generic limits of Lycomorpha and Propyria. The genera Dolichesia Schaus, Hypermaepha

Hampson, Lycomorphodes Hampson, and Talara Walker were included as the internal outgroup. Scott & Branham (In Press) found that these four genera formed a strongly supported polytomy (PP ≥ 0.98) with the Lycomorpha + Propyria clade in the Bayesian

Inference analysis. Bendib & Minet (1999) included these genera in the subtribe

Cisthenina as well. To root the phylogenetic analysis, an external outgroup composed of

Hypoprepia and Ptychoglene Felder was included. Both of these genera are also members of Cisthenina. Forbes (1960) considered Lycomorpha to be a member of the

Hypoprepia generic group, and Jacobson & Weller (2002) found Hypoprepia and

Lycomorpha as sister taxa. However, in their analysis of Lithosiini, Scott & Branham (In

Press) found Hypoprepia to be more distantly related to Lycomorpha than the genera of the internal outgroup. The wing pattern colors of Ptychoglene have created confusion

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between it and Lycomorpha. Hampson (1914) synonymized the type species of

Ptychoglene, P. erythrophora Felder, with L. drucei Hampson, and the species P. coccinea (Henry Edwards) was originally described within Lycomorpha. Scott &

Branham (In Press) included both of these species in their analysis of Lithosiini. All species of Ptychoglene were found to be distantly related to the Lycomorpha + Propyria clade. P. erythrophora is included to assess the relationship between it and Lycomorpha and to determine the status of L. drucei.

Collections Consulted

The following is a list of collections consulted for this study. It includes the name of the individual(s) who prepared the loan as well as an acronym for each of the collections. The acronyms used follow Heppner & Lamas (1982): American Museum of

Natural History (AMNH) (D. Grimaldi), California Academy of Sciences (CAS) (N.D.

Penny), Carnegie Museum (CMNH) (J. Rawlins), Canadian National Collection,

Agriculture Canada (CNC) (C. Schmidt), Cornell University Insect Collection (CUIC) (J.

Liebherr, E.R. Hoebeke), Florida Museum of Natural History, University of Florida

(FLMNH) (J. Miller, D.M. Lott, A.D. Warren), Los Angeles County Museum of Natural

History (LACM) (W. Xie), Museum of Comparative Zoology, Harvard University (MCZ)

(R. Eastwood, P.D. Perkins), Museum of Southwestern Biology (MSB) (D. Lightfoot),

National Museum of Natural History, Smithsonian Institution (USNM) (D.G. Furth, D.

Harvey), Peabody Museum of Natural History, Yale University (YPM) (L.F. Gall), San

Diego Natural History Museum (SDNH) (M.A. Wall), A&M University Insect

Collection (TAMU) (E.G. Riley), Essig Museum of Entomology, University of California,

Berkeley (UCB) (J. Powell), University of Minnesota, St. Paul (UMSP) (R. Holzenthal),

University of Wisconsin – Oshkosh (UWO) (J.M. Zaspel).

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Morphology

Dissections of the genitalia were prepared after softening the abdomens in a warm 10% potassium hydroxide (KOH, Fisher Scientific, Pittsburg, PA) solution for 30 minutes to 1.5 hours using standard methods (Winter 2000). The scales and viscera were removed from the abdomen in several rinses of 20% ethanol using fine watercolor paintbrushes (# 000 – 2). Structures were stained with a solution of chlorazole black E

(Acrōs) dissolved in deionized water. Specimens were viewed in 20% ethanol and stored temporarily in 70% ethanol. Subsequently the abdominal pelts and genitalia were either stored in vials of glycerol or permanently slide mounted in Euparol (Bioquip,

Garden City, CA). The final preparation method used was dependent on the preference of the collection from which the material originated. The external morphological characters were coded from dried pinned specimens. All characters were scored using a Nikon SMZ800 light microscope. Pencil drawings were made using a camera lucida associated with a Leica MZ16 light microscope. These drawings were scanned and saved as PDFs that were imported into Adobe Illustrator CS5 and then inked using the pen tool. The scale bar included in each figure is equivalent to 1mm. The purpose of stippling in any figure is explained in the figure legend. The terminology for male and female genitalia follows Klots (1970) and Forbes (1939b, 1954). The Lithosiini specific terminology for the genitalia proposed by Birket-Smith (1965) was not used. This terminology can only be applied with confidence to members of the subtribe Lithosiina, and all species included in this analysis were placed in the subtribe Cisthenina (Bendib

& Minet 1999). The terminology for the wing venation follows Kristensen (2003).

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Characters Examined

Sixty-five morphological characters (35 binary and 31 multistate; 196 states) were scored for all of the species that were sampled (Appendix D). These characters were coded from the adult external morphology as well as the male and female genitalia. Non-genitalic and genitalic characters are as follows: head (5 characters, 12 states), thorax (1 characters, 2 states), wings (2 characters, 4 states), abdomen (1 character, 3 states), male abdomen and genitalia (38 characters, 113 states), and female abdomen and genitalia (18 characters and 62 states). Multistate characters were treated as unordered and non-additive. When characters were linked, e.g. the presence of androconia in the A7/A8 intersegmental membrane (Character 10) and the form of the androconia in the A7/A8 intersegmental membrane (Character 11), the resulting inapplicable characters were coded as missing “?” (Strong & Lipscomb 1999). By linking the characters, the presence of a morphological structure can contribute to the analysis, and the variation in the structure can be documented without oversimplifying characters

(Pogue & Mickevich 1990).

Phylogenetic Analysis

Phylogenetic trees were constructed using parsimony and Bayesian analyses.

Two data matrices were created and analyzed using the same set of morphological characters. One matrix contained all species (AS) from the ingroup including two species represented by a male specimen only (Appendix E). The other matrix contained only the eleven ingroup taxa that were represented by a male-female pairs. Both matrices were analyzed using a Maximum Parsimony (MP) analysis in Paup* 4.0b 10

(Swofford 2003). A heuristic search of 10,000 random–taxa additions was performed to identify all tree islands. The resulting cladograms of the AS matrix and the male-female

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(MF) matrix were compared to determine the effect of the missing data in the AS matrix.

Bremer support indices (BS: Bremer 1988, 1994) were calculated for the strict consensus of the MP trees obtained from the analysis of each dataset. The program

TreeRot.v3 (Sorenson & Franzosa 2007) was run in conjunction with Paup* to obtain the Bremer support indices. In the discussion of our results, we define Bremer support values as giving weak (BS 1-2), moderate (BS 3-5), good (BS 6-10), or strong (BS≥11) support (Wahlberg & Nylin 2003; Wahlberg et al. 2003, 2005). Based on the findings of the comparison between the AS matrix and the MF matrix, only the AS matrix was used to conduct the Bayesian Inference analysis and test the sensitivity of the ingroup topology to the removal of specific outgroup taxa. Additional nodal support values were calculated for the AS matrix from character jackknife analysis. Single taxon jackknife analysis (Lanyon 1985) was performed on each of the internal and external outgroup taxa using the AS matrix to determine if the ingroup topology was sensitive to taxon sampling. 10,000 random-taxa additions were performed for each outgroup manipulation. Further nodal support was calculated for the MP cladogram obtained from the analysis of the AS matrix using 1,000 character jackknife replicates (Kitching et al.

1998). Branch support was not calculated using bootstrap analysis (Felsenstein 1985) due to statistical objections that have been raised over its use (Sanderson 1995). The

Bayesian Inference (BI) analysis was conducted using MrBayes 3.1.2 (Ronquist &

Huelsenbeck 2003). The data set was input as standard and analyzed using the Mk+Γ model (Lewis 2001). The BI analysis was performed using four chains (one cold and three hot) using the default temperature settings. Five simultaneous, independent runs of 20,000,000 generations were conducted. The Markov chain Monte Carlo (MCMC)

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chains were started at random trees. The cold run was sampled every 1,000

generations. Five million generations were discarded as a ‘burn-in’. The potential scale

reduction factor (PSRF: Gelman & Rubin 1992) was calculated by summarizing the

probabilities of the five runs. The PSRF value is expected to approach 1 as the

independent runs converge. Posterior probability (PP) provided the clade credibility

values for the BI cladogram. The consensus trees from both types of analysis were

visualized using FigTree v1.3.1 (Rambaut 2010).

Results

Phylogenetic Analysis and Taxonomic Implications

The MP analyses of both the all species (AS) and the male-female (MF) data matrices recovered Lycomorpha as paraphyletic with respect to Propyria. The analysis of the AS matrix resulted in three most parsimonious trees (L=192, CI=0.68, RI=0.73) located within a single tree island. Despite the concerns of some authors (Kitching et al.

1998) that missing data within a dataset may confound the analysis, the strict consensus of the MP trees from this analysis was highly resolved (Fig. 4-4). The only unresolved portion of this tree was the pholus clade. The analysis of the MF matrix also resulted in three most parsimonious trees (L=192, CI=0.68, RI=0.71) within a single tree island. The strict consensus of these trees produced a cladogram (Fig. 4-3) with a topology identical to that found from the analysis of the AS matrix with the exception of the excluded taxa. However, the nodal support of the MF matrix MP cladogram was lower than that recovered for the AS cladogram.

Due to these preliminary findings, the Bayesian Inference (BI) analysis as well as the character and outgroup jackknife analyses were only conducted on the AS matrix.

The BI analysis also found Lycomorpha to be paraphyletic with respect to Propyria (Fig.

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4-5). However, the topology of the BI cladogram was less resolved than cladograms recovered through the MP analysis of both matrices. Although the BI analysis was unable to recover the deeper evolutionary relationships of Lycomorpha, the species groups that were recovered were identical to those found in the MP analysis.

In addition to the two types of phylogenetic analysis, an outgroup jackknife analysis was completed to determine whether the ingroup topology was sensitive to the exclusion of any outgroup taxon. This analysis revealed that Lycomorpha was recovered as paraphyletic to Propyria regardless of which outgroup taxon was removed

(Figs 4-6 to 4-11). In addition, all of topologies recovered from the outgroup jackknifing were shorter than the MP cladogram obtained from the analysis of the AS matrix (Table

4-4). Omission of Hypoprepia (Fig. 4-6) or Talara (Fig. 4-8) caused the deeper nodes of the ingroup to collapse. The exclusion of any other outgroup taxon produced no effect on the ingroup topology. However, the removal of Hypoprepia and Hypermaepha (Fig.

4-10) resulted in the loss of resolution among the internal outgroup taxa. In addition, the omission of Dolichesia (Fig. 4-11) resulted in the internal outgroup clade to forming a polytomy with the ingroup.

Both types of analysis recover the Lycomorpha + Propyria ingroup sister to the internal outgroup taxa with good support (BS = 5, Jackknife (JK) ≥ 92, PP ≥ 0.98). The relationship between Lycomorpha and Propyria is strongly supported (BS=9, JK ≥ 99,

PP = 1). An examination of the character support (Table 4-3) for this node reveals that there are twenty synapomorphies that define the ingroup clade. Four of the synapomorphies (L1, IV1, BP1, PG1) are unique and found in all members of the ingroup. The occurrence of a tibial spur formula of 0-2-3 (L1) is unique within the

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subfamily Arctiinae. Scoble (1995) describes Arctiinae as possessing a tibial spur formula of either 0-2-2 or 0-2-4. Two of the other unique states that occur in all members of the ingroup arise from the male genitalia. They include the presence of an androconial patch within the intravincular membrane (37:IV1) and the absence of a caecum on the proximal end of the basiphallus (38:BP1). The final unique state arises from the female genitalia and relates to the form of the dorsal pheromone gland (PG1;

Character 64, State 0). The states of another four characters defining the ingroup are unique to this clade (AB1, MA2, MA8, VS1). An anterolateral process occurring as a sclerotized bar (9:AB1) and an M-shaped vinculum/saccus (36:VS1) are unique character states that are reversed only once within the ingroup. In addition, the form of the androconia present in the A7/A8 intersegmental membrane (11:MA2), which is unique to the ingroup, is further modified to another unique state within the Propyria clade (Clade 4).

Within the ingroup clade (Clade 1), two main clades (Clade 2, Clade 3) are present. The pholus clade (Clade 2; BS = 2, JK ≥ 82, PP ≥ 0.81) is composed of three species including the type species of Lycomorpha. The other members of this clade include the former subspecies of L. pholus and a yet to be described species. The undescribed member of the clade represents a form of L. pholus that is only found in

Texas. Five characters support this clade. The rectangular sclerotization around the uncus base (23:UB2), which is unique to Clade 2, is present in all species. Although the evolutionary relationships among these three species are unresolved, differences are present in the female genitalia. However, they do not provide a strong enough phylogenetic signal to resolve the relationships. The remaining ingroup clade (Clade 3;

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BS = 2, PP ≥ 0.51) contains all of the Propyria and the remaining species of

Lycomorpha with the exception of L. splendens. There are ten synapomorphies that

support Clade 3. Unique states are present in three of these characters (33:J1, 41:PH2,

FA4). However, none of these states are present in all of the species, but each of these

states is further modified into another unique state.

All of the species representing Propyria are found within Clade 3, but these species do not form a monophyletic clade. The type species of Propyria, P. ptychoglene, is found sister to P. normani Schaus (Clade 4; BS = 2, JK ≥ 72, PP ≥

0.64). The Propyria clade is supported by six synapomorphies. The presence of a narrow pocket of sex scales (Character 11, State 2) and a trident shaped processus basalis (Character 29, State 2) in the male genitalia are unique and found in both members of this clade. Unique states are also present in two other characters that were coded from the female abdomen (FA3, FA4). However, a female specimen from P. normani was not available. Therefore, it is not possible to determine whether these states are present in all members of this clade. The other representative species of

Propyria, P. morelosia Schaus, is recovered within the clade that contains all of the species of Lycomorpha with red forewings and primarily black hindwings (Clade 5; BS =

2, PP ≥ 0.6). Six characters support this relationship. Two of these characters, an S- shaped uncus on the male genital capsule (Character 25, State 0) and continuous heavy sclerotization of the seventh female abdominal segment (Character 49, State 0), are unique to and present throughout Clade 5. The reduction of the A8 sternite to a sclerotized bar that is fused to the A8 tergite and A7 sternite (53:FA6) is also unique to this clade. However, the A8 sternite is lost within the grotei clade (Clade 6).

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The grotei clade receives moderate to strong support (BS = 4, JK ≥ 83, PP ≥

0.99). Three of the species whose wing color patterns led Comstock & Henne (1967) and Powell & Opler (2009) to suggest a phylogenetic study of the taxa was necessary are recovered within this clade. Seven synapomorphies define Clade 6. Three of these characters that arise from the female abdomen and genitalia (52:FA5, 57:DB2, 65:PG2) represent unique states that are present in all members of the clade.

The findings of our analysis demonstrate that Lycomorpha and Propyria are not reciprocally monophyletic. We propose to treat these two genera as congeners. As presently defined, the species of Lycomorpha are found within three clades (C1, C2,

C6). Constraining the monophyly of these species results in a tree that is six steps longer (Fig. 4-12, L = 198, CI = 0.66, RI = 0.71). The parsimony likelihood test (Kishino

& Hawsegawa 1989) does not find the two trees to be significantly different (P < 0.20).

However, when this constraint is enforced, Propyria is found to be paraphyletic with respect to Lycomorpha. When the monophyly of Propyria is constrained, a tree that is only two steps longer (Fig. 4-13; L = 194, CI = 0.67, RI = 0.72; P < 0.53) is found.

Constraining the monophyly of Propyria does not separate it from Lycomorpha. Instead,

Lycomorpha remains paraphyletic with respect to Propyria. Thus we propose to place

Propyria 1898 as a junior synonym of Lycomorpha 1839. A formal revision of

Lycomorpha will be provided elsewhere (Scott in prep.) Below we provide a species checklist of Lycomorpha. The species are listed in alphabetical order. Given the taxonomic confusion surrounding Lycomorpha, we also provide a checklist of the species originally described in the genus along with their current taxonomic treatment.

Several species are removed from Lycomorpha and placed in other genera. L. desertus

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is transferred to the lithosiine genus Eudesmia. Hampson (1914) and Jordan (1917) reported that the types of this species had been lost. However, examination of the type collection of the National Museum of Natural History, Smithsonian Institution revealed two unidentified Lithosiini type specimens that match the description of L. desertus

(pers. obs.). In addition, both specimens are missing abdomens, which is congruent with a male-female pair taken in copula. A new description of L. desertus will be included in the revision of Lycomorpha. L. nigridorsata Dognin is listed as incertae sedis. In his description of the genus, Dognin compares L. nigridorsata to three species that have been transferred to other genera. Two of these species are now members of

Zygaenidae. Dognin (1916) considered Lycomorpha to be a member of Zygaenidae.

Without examining the type specimen, it will not be possible to confirm if L. nigridorsata is a lithosiine or zygaenid. Finally, L. drucei is synonymized with P. erythrophora.

Species Checklist of Lycomorpha

LYCOMORPHA Harris, 1839: 317

Type species: Sphinx pholus Drury, by monotypy

Anatolmis Packard, 1864: 45

Prepodes Herrich-Schäffer, 1855: 100,101

Propyria Hampson, 1898: 521 New Synonym atroxantha (Schaus, 1906) (Propyria) Mexico New Combination criton (Druce, 1881-1900) (Cisthene) Guatemala New Combination

coatepeciensis (Strand, 1920) (Propyria) Mexico New Combination

a. hypoleuca (Draudt, 1917) (Propyria) Mexico New Combination

orizabae (Strand, 1920) (Propyria) Mexico New Combination flora (Schaus, 1911a) (Propyria) Costa Rica New Combination

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fridolinia (Schaus, 1925) (Propyria) Guatemala New Combination fulgens (Henry Edwards, 1881b) (Anatolmis) Arizona

tenuimargo (Holland, 1903) (Ptychoglene) Arizona, Mexico

tenumargo authors, misspelling grotei (Packard, 1864) (Anatolmis) Colorado

palmerii Packard, 1872 (Lycomorpha) Arizona miniata Packard, 1872 (Lycomorpha) Southern California Species Revised morelosia (Schaus, 1925) (Propyria) Mexico New Combination normani (Schaus, 1911b) (Propyria) Costa Rica New Combination pelopia (Druce, 1881-1900) (Talara) Panama New Combination pholus (Drury, 1773) (Sphinx) New England ptychoglene (Hampson, 1898) (Propyria) Mexico, Guatemala New Combination

aequalis (Walker, 1854a) (Lithosia) Guatemala New Combination

sinuata (Henry Edwards, 1885) (Lycomorpha) Mexico pulchra Dyar, 1898 (Lycomorpha) Texas Species Revised regulus (Grinnell, 1903) (Anatolmis) California schausi Dyar, 1898 (Lycomorpha) New Mexico, Arizona Combination Revised splendens Barnes & McDunnough, 1912 (Lycomorpha) Utah strigifera Gaede, 1926 (Lycomorpha) Mexico

Checklist of Species Described as Lycomorpha

Lithosiini

EUDESMIA Hübner, 1823: plate 187

Type species: Callimorpha ruficollis Donovan, by monotypy

Gerba Walker, 1865: 270

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Ruscina Walker, 1854: 563 desertus (Henry Edwards, 1881a) (Lycomorpha) Arizona

deserta Hampson, 1914 misspelling

LYCOMORPHODES Hampson, 1900: xvi, 84, 378

Type species: Trichromia strigosa Butler, by original designation. bipartita (Walker, 1866) (Lycomorpha) Brazil strigosa (Butler, 1877b) (Trichromia) Brazil

fumata (Möschler, 1890) (Lycomorpha) Puerto Rico

funesta Kirby, 1892, misspelling

PTYCHOGLENE Felder, 1874: plate 106

Type species: Ptychoglene erythrophora Felder, by monotypy. coccinea (Henry Edwards, 1886) (Lycomorpha) Arizona erythrophora Felder, 1874 (Ptychoglene) Mexico

drucei Hampson, 1901 (Lycomorpha) Mexico New Combination

Arctiini

CTENUCHA Kirby, In Richardson, 1837: 305

Type species: latreillana Kirby, by monotypy

Caralisa Walker, 1856: 1660

Compsoprium Blanchard, In Gay, 1852: 67

Erchia Walker, 1854a: 321

Euctenucha Grote, 1873: 33

Hoplarctia Butler, 1877a: 54 togata (Druce, 1881–1900) (Acreagris) Guatemala, Costa Rica, Panama

augusta (Henry Edwards, 1887) (Lycomorpha) Mexico

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obscurata Draudt, 1915 (Ctenucha)

Zygaenidae

MALTHACA Clemens, 1860: 540–541

Type species: Malthaca perlucidula Clemens, by monotypy

Coementa Druce, 1881–1900: 123

Gingla Walker, 1854b: 228

Felderia Kirby, 1892: 163 marginata (Henry Edwards, 1884) (Lycomorpha) Mexico

mexicana (Druce, 1881–1900) (Lycomorpha) Mexico

teos (Schaus, 1889) (Lycomorpha) Mexico

NEOALBERTIA Tarmann, 1984: 64

Type species: Lycomorpha constans Edwards

constans (Henry Edwards, 1881) (Lycomorpha) Arizona

sancta (Neumoegen & Dyar, 1894) (Seryda) Arizona

NEOFELDERIA Tarmann, 1984: 65

Type species: Acreagris correbioides Felder, by monotypy

Acreagris Felder, 1874: plate 83

rata (Henry Edwards, 1882) (Lycomorpha) Arizona

PYROMORPHA Herrich-Schäffer, 1850–1858: 6

Type species: dimidiata Herrich-Schäffer, by monotypy

Tetraclonia Jordan, 1917: 24

centralis (Walker, 1854a) (Lycomorpha)

notha (Henry Edwards, 1885) (Lycomorpha) Mexico

latercula (Henry Edwards, 1882) (Lycomorpha) Arizona

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flavescens Herig, 1924 (Tetraclonia) America

SERYDA Walker, 1856: 1598

Type species: Seryda cincta Walker, by monotypy anacreon (Druce 1881–1900) (Lycomorpha) Costa Rica basirei (Druce 1891–1900) (Lycomorpha) Mexico

semifulva (Druce, 1891–1900) (Lycomorpha) Mexico

Incertae sedis contermina Henry Edwards, 1884 (Lycomorpha) Mexico

Dyar (1898) does not consider contermina to be a true species of Lycomorpha.

No alternative placement is suggested. nigridorsata Dognin, 1916 (Lycomorpha) Panama regia Schaus, 1889 (Lycomorpha) Mexico

Dyar (1898) considers regia to be closely related to augusta

Characters and their Phylogenetic Usefulness

Here we present a list of the morphological characters used in this phylogenetic analysis. A discussion of the phylogenetic usefulness of each character is included. The individual character indices Consensus Index (CI) and Retention Index (RI) are included after each character. The indices are rounded up. Linked characters are indicated by a

“*”.

Head

1. Male flagellomere state. 0: simple and ciliate (Fig. 4-14A), 1: serrate (Fig. 4- 14B), 2: bipectinate (Fig. 4-14C). (CI = 0.67, RI = 0.75).

a) Most species possess serrate antennae. Clade 2 + Clade 3 is defined by the presence of state 1. State 0 occurs in all outgroup taxa except Hypoprepia. State 2 is unique to L. splendens.

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2. Female flagellomere state. 0: simple and ciliate (Fig. 4-15A), 1: serrate (Fig. 4- 15B). (CI = 0.33, RI = 0.50).

a) The females of most species possess simple and ciliate antennae. Serrate antennae (state 1) are a synapomorphy of Lycomorpha. They are lost once in Clade 3 but regained in L. fulgens.

3. Gena state. 0: well developed, can be seen as a continuous band around the eye that joins the frons when viewing the head in profile (Fig. 4-16A), 1: reduced, cannot be seen as a continuous band around the eye (Fig. 4-16B). (CI = 0.25, RI = 0.57).

a) The presence of a well-developed gena (State 0) is a synapomorphy for Clade 2 + Clade 3. Within this clade, it is reversed twice in New sp. A and L. grotei.

4. Proboscis length. 0: longer than the thorax, 1: shorter than thorax. (CI = 0.50, RI = 0.00).

a) The presence of a proboscis that is shorter than the thorax is autapomorphic for two outgroup taxa: Hypoprepia and Hypermaepha.

5. Labial palp segment fusion. 0: all segments fused (Fig. 4-17A), 1: 2nd and 3rd segments fused (Fig. 4-17B), 2: fusion absent (Fig. 4-17C). (CI = 0.67, RI = 0.00).

a) Fusion between any of the labial palp segments is uncommon in both Lycomorpha and the outgroup (OG) taxa. The fusion of all three segments is autapomorphic for Hypermaepha. State 1 arises independently in L. splendens and L. fulgens.

Thorax

6. Tibial spur formula. 0: 0-2-4, 1: 0-2-3. (CI = 1.00, RI = 1.00).

a) A tibial spur formula of 0-2-3 (state 1) is a synapomorphy of Lycomorpha. It unites all species of Lycomorpha and Propyria. This condition is unusual within Arctiinae. Scoble (1995) reported that the tibial spur formula for Arctiinae was either 0-2-2 or 0-2-4.

Wings

7. Forewing: R free to costal margin, not anastomosing with Sc. 0: present (Fig. 4- 18A), 1: absent (Fig. 4-18B). (CI = 0.50, RI = 0.50).

a) The absence of an R vein that is free to the costal margin (state 2) is a synapomorphy of the internal OG clade. It is reversed just once in Lycomorphodes.

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8. Hindwing: Sc + R. 0: present (Fig. 4-19A), 1: absent (Fig. 4-19B). (CI = 1.00, RI = 1.00).

a) The loss of the Sc + R (State 1) is a synapomorphy of the internal OG + Lycomorpha clade.

Abdomen

9. Form of the Anterolateral Process (ALP) on the A2 apodeme. 0: sclerotized bar (Fig. 4-20A), 1: flattened sclerotized lobe (Fig. 4-20B), 2: short nob, less than half the length of the apodeme (Fig. 4-20C). (CI = 0.50, RI = 0.60).

a) A synapomorphy of Lycomorpha is the sclerotized bar (state 0) form of the ALP. State 2 occurs only in the internal OG taxa Talara and Hypermaepha. All remaining members of the outgroup have the flattened sclerotized lobe (state 1) form of the ALP.

Male abdomen and genitalia

Male Abdomen

10. * A7/A8 ventral intersegmental membrane. 0: without androconia (Fig. 4-21A), 1: with androconia (Figs 4-21 B-D). (CI = 0.50, RI = 0.75).

a) The presence of androconia in the A7/A8 ventral intersegmental membrane (State 1) defines the ingroup clade and arises independently in Hypermaepha.

11. * Form of androconia in A7/A8 ventral intersegmental membrane. ?: does not apply (Fig. 4-21A), 0: 2 pockets with long hair-like scales (Fig. 4-21B), 1: shallow, broad pocket of sex scales one-third or more the width of A7 sternite (Fig. 4- 21C), 2: narrow pocket of sex scales less than one-third the width of A7 sternite (Fig. 4-21D). (CI = 1.00, RI = 1.00).

a) The presence of a shallow, broad pocket of sex scales androconia (State 1) is a synapomorphy of the Lycomorpha clade. It transforms into State 2 within the Propyria clade.

12. Cephalic margin of A8 sternite. 0: fused to cephalic margin of A8 tergite, 1: extending to the cephalic margin of A8 tergite, no fusion, 2: extending onto the A8 pleurites. (CI = 0.50, RI = 0.67).

a) Fusion of the cephalic margin of A8 sternite to the cephalic margin of A8 tergite (State 0) is diagnostic of the grotei clade. In all other members of the ingroup clade, the cephalic margin of the A8 sternite extends to the cephalic margin of the A8 tergite but no fusion occurs. State 2 is diagnostic of outgroup taxa Hypoprepia and Hypermaepha.

13. * Sclerotization of A8 tergite. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

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a) The absence of sclerotization on the A8 tergite is a synapomorphy of the internal OG clade.

14. * Pattern of sclerotization of A8 tergite. ?: does not apply, 0: T-shaped, 1: square, 2: heartshaped, 3: rectangular. (CI = 1.00, RI = 1.00).

a) The ingroup is defined by a rectangular A8 sclerite (State 3). This character transforms to a heartshaped plate (State 2) within New sp. A + grotei clade.

15. A8/A9 dorsal intersegmental membrane. 0: without androconia, 1: with 3 androconia, 2 pockets with long hair-like sex scales bracketing a shallow pocket with hair-like scales. (CI = 0.50, RI = 0.50).

a) The presence of androconia in the A8/A9 dorsal intersegmental membrane is diagnostic of the internal OG and is reversed once in Hypermaepha.

16. * A8/A9 ventral intersegmental membrane. 0: without androconia, 1: with androconia. (CI = 0.50, RI = 0.83).

a) Androconia in the A8/A8 ventral intersegmental membrane defines all members of Clade 5 except P. morelosia (the basal most species in the clade). It also arises independently within Dolichesia.

17. * Form of androconia in the A8/A9 ventral intersegmental membrane. ?: does not apply, 0: 2 shallow pockets of sex scales located on the outer edges of the membrane, 1: a small pocket of sex scales located in the center of the membrane. (CI = 1.00).

a) State 1 is a synapomorphy of all species of Lycomorpha within Clade 5. State 0 is unique to Dolichesia.

Genital Capsule

18. Pleural Sclerites. 0: present, 1: absent.

a) The presence of pleural sclerites is constant throughout all taxa included in the study. The character is included because pleural sclerites were thought to be absent throughout Arctiinae.

19. The two halves of the tegumen fused for their entire length. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

a) There is fusion between the two halves of the tegumen in all ingroup and outgroup taxa. However, fusion that extends the entire length of the two halves of the tegumen (state 0) is a synapomorphy of the internal OG+Lycomorpha clade.

20. * Sutures indicating the location of the fusion. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

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a) The absence of sutures that can be used to identify the two halves of the tegumen (state 1) is a synapomorphy of the internal OG clade. When present the form of the sutures helps to define species groups.

21. * Shape of the sutures on the tegumen. ?: does not apply, 0: inverted Y-shape, 1: V-shaped, 2: inverted U-shape, 3: Inverted T-shape, 4: straight line with an ovoid bulge medially. (CI = 0.80, RI = 0.67).

a) Variation in the shape of the sutures is useful for identifying species (e.g. State 3, L. splendens; State 2, L. fulgens). V-shaped sutures (State 1) are a synapomorphy of the ingroup.

22. Articulation of tegumen with uncus. 0: not fused, membranous break separating the tegumen and uncus, 1: fused. (CI = 0.33, RI = 0.33).

a) This character had low CI and RI values. Within both the external and internal outgroup taxa fusion between the tegumen and uncus (state 1) arose independently four times.

23. * Sclerotization around the uncus base. 0: present, 1: absent. (CI = 0.50, RI = 0.75).

a) The presence of sclerotization around the uncus base defines the Clade 2 + Clade 3 relationship. It also arises independently within both external OG taxa.

24. * Form of sclerotization around the base of the uncus. ?: does not apply, 0: reduced to narrow, strips occurring laterally on the uncus base, 1: triangular, 2: rectangular, 3: V-shaped, 4: U-shaped. (CI = 1.00, RI = 1.00).

a) A reduction of the sclerotization to narrow strips (State 0) is the most common state within the ingroup. However, the form of the sclerotization around the uncus is also useful for diagnosing species (e.g. State 4, L. grotei) and small species groups.

25. Curvature of the uncus. 0: S-shaped, emerges from the conjuctiva and undergoes two bends, 1: C-shaped, 2: knob, 3: straight. (CI = 0.75, RI = 0.86).

a) An S-shaped uncus (State 0) is unique to Clade 5 and is present in all species within the clade. All other members of the ingroup clade possess a C-shaped uncus (State 1). State 1 also evolves independently within the external OG taxa and Lycomorphodes. State 2 is autapomorphic for Hypermaepha.

26. Shape of the apex of the uncus. 0: fingerlike tapering to a point, 1: fingerlike not tapering to a point, 2: laterally compressed ridge, 3: ovoid, teardrop shaped, tapering to a point, 4: spade shaped, tapering to a point. (CI = 0.57, RI = 0.57).

a) The variation within this character is diagnostic of small species groups and species. States 0, 1, and 3 are found in both the ingroup and the outgroup taxa.

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State 4 is an autapomorphy of New sp. A. State 2 is an autapomorphy of Lycomorphodes.

27. Distal end of costal margin occurring as an identifiable break in the sclerotization. 0: present, 1: absent. (CI = 0.50, RI = 0.67).

a) An identifiable break at the distal end of the costal margin (State 0) defines the ingroup clade. It arises independently within the Lycomorphodes + Hypermaepha clade of the internal OG.

28. * Processus basalis of costa. 0: present, 1: absent. (CI = 0.33, RI = 0.60).

a) This character (State 0) defines clade 3. It is reversed in the grotei clade and arises independently within the internal OG Lycomorphodes.

29. * Shape of processus basalis of costa. ?: does not apply, 0: laterally compressed triangles, 1: elongate spines, tapering to a point, 2: dorso-ventrally compressed trident shaped extensions, tips are blunt, 3: short dorso-ventrally flattened lobe. (CI = 1.00, RI = 1.00).

a) The shape of the processus basalis of the costa is useful in diagnosing species (e.g. State 3, New sp. A) and species pairs (e.g. State 2, P. morelosia + P. normani).

30. Transtilla. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

a) The loss of the transtilla (State 1) defines the internal OG clade).

31. * Editum. 0: present, 1: absent. (CI = 0.33, RI = 0.60).

a) Most species in the study possess an editum (State 0). This state is reversed independently within the pholus clade, L. fulgens, and the internal OG clade of Lycomorphodes + Hypermaepha.

32. * Location of the editum. ?: does not apply, 0: proximally, occurs within the proximal ¼ of the costa, 1: processus basalis of the costa, 2: distally, at or beyond the proximal ¼ of the of the costa. (CI = 0.67, RI = 0.80).

a) The editum is located distally (State 2) in members of both the internal and external OG. State 1 arises twice in the ingroup (e.g. Propyria clade & New sp. A).

33. Shape of the juxta. 0: rectangular with length perpendicular to the body, 1: semicircular, 2: conical projection with lateral extensions, 3: square with elongate two pronged projection, 4: square with a concave indentation in distal margin, 5: triangular with a conical tip, 6: trapezoidal, 7: rectangular with a triangular indentation in distal margin. (CI = 0.86, RI = 0.83).

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a) The shape of the juxta is useful for diagnosing species (e.g. State 6, L. splendens) and uniting small species groups (e.g. State 4, pholus clade). States 5 is diagnostic for a smaller clade within Clade 5 (L. fulgens (New sp. A (L. grotei (L. regulus (L. pulchra + New sp. C))))). All former Propyria species possess State 7.

34. * Juxta ornamentation, transparent patch. 0: present, 1: absent. (CI = 0.20, RI = 0.50).

a) The presence of transparent patches ornamenting the juxta (State 0) is a synapomorphy of Clade 3. This state is reversed twice in L. pulchra and New sp. A.

35. * Shape of transparent patch. ?: does not apply, 0: triangular, 1: semicircular, 2: conical, 3: rectangular, 4: circular. (CI = 0.80, RI = 0.50).

a) A conical translucent patch in the juxta (State 2) is autapomorphic for P. ptychoglene. State 0 is diagnostic of a smaller clade within Clade 6 (L. regulus (L. pulchra + New sp. C)). It is lost once in L. pulchra. This state arises independently in P. morelosia and P. normani.

36. Shape of vinculum/saccus. 0: U-shaped, 1: V-shaped, 2: M-shaped. (CI = 0.67, RI = 0.80).

a) An M-shaped vinculum/saccus (State 2) is a synapomorphy of the ingroup. It is reversed to state 0 once in L. fulgens.

37. Ornamentation of intravincular membrane, single shallow pocket with sex scales. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

a) This character (State 0) is unique to the ingroup and present in all of the species of this clade.

Phallus

38. Proximal end of basiphallus (caecum). 0: well-developed, ductus ejaculatorious simplex (DES) located entirely dorsad of the caecum, 1: reduced, DES emerging from anterior, dorsal end of caecum, 2: absent, DES emerges from the anterior end of aedeagus. (CI = 1.00, RI = 1.00).

a) The absence of the caecum (State 2) is unique to the ingroup clade. This character unites all Lycomorpha and former Propyria species. A well-developed caecum (State 0) is a synapomorphy of the internal OG. State 1 is diagnostic of the external OG taxa.

39. Orientation of basiphallus. 0: straight, 1: inflected dorsally at distal end (curved away from venter), 2: inflected dorsally at midpoint (curved away from venter), 3: inflected ventrally at base (curved toward venter). (CI = 0.75, RI = 0.67).

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a) Most species possess a straight basiphallus (State 0). State 1 is unique to a smaller clade within Clade 5 (L. fulgens (New sp. A (L. grotei (L. regulus (L. pulchra + New sp. C))))). It is reversed once within the (L. regulus (L. pulchra + New sp. C))) species group of this clade. State 2 is autoapomorphic within Lycomorphodes. State 3 occurs in the internal OG taxa Talara and Hypermaepha.

40. * Phallic sclerite. 0: present, 1: absent, highly reduced. (CI = 0.25, RI = 0.50).

a) The presence of a phallic sclerite is a synapomorphy of the ingroup. It is lost twice in L. fulgens and P. ptychoglene. The sclerite arises independently within the internal OG Talara.

41. * Shape of phallic sclerite, left side. ?: does not apply, 0: flattened lobe, extending as a separate structure dorsad of the vesica, 1: rectangular, 2: narrowed, rounded triangle, 3: narrowed, pointed triangle. (CI = 1.00, RI = 1.00).

a) This character (State 2) is unique to clade 3. It is modified to state 3 in the L. pulchra + New sp. C species pair. A rectangular sclerite (State 1) is an autapomorphy of L. splendens. State 0 is found in both the ingroup (e.g. pholus clade) and the internal OG Talara.

42. Vesica bilobed. 0: absent, 1: present. (CI = 0.50, RI = 0.50).

a) A bilobed vesica is found in all external OG taxa and the internal OG taxon Hypermaepha.

43. * Membranous projections arising from the primary lobe(s) of vesica. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

a) This character (State 0) is unique to Clade 2 + Clade 3. This clade includes all members of the ingroup except L. splendens.

44. * Location of membranous projections. ?: does not apply, 0: apically on the left side, proximal to the distal margin of the aedeagus, 1: distal end of main lobe, upper right apice, 2: apically on the right side, 3: medially on the right side, 4: dorsal surface of the primary lobe. (CI = 1.00, RI = 1.00).

a) This character is useful for diagnosing small clades (e.g. State 1, grotei clade) and species (e.g. State 3, New sp. A).

45. * Ornamentation of the membranous projection located distally on the primary lobe. ?: does not apply, 0: peglike cornuti present the entire length, 1: peglike cornuti present on the distal third, remainder ruggose, 2: ruggose for the entire length. (CI = 1. 00).

a) A membranous projection located distally on the primary lobe is unique to the grotei clade. The different forms of ornamentation help to diagnose the species of

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this clade (e.g. State 2, L. grotei), which includes three of the species with red forewings and primarily black hindwings.

46. * Ornamentation of the vesica: heavily-sclerotized, spine like cornutus. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

a) The presence of a heavily-sclerotized, spine like cornutus (State 0) is unique to the ingroup. It is absent in L. splendens (the most basal species of the ingroup) only.

47. * Location of spine-like cornutus. ?: does not apply, 0: primary lobe, 1: distal point of the membranous projection off the vesica. (CI = 0.50, RI = 0.67).

a) The spine-like cornutus is located on the primary lobe of the vesica (State 0) in most species. State 1 is diagnostic of the pholus clade. This state arises independently within L. fulgens.

Female abdomen and genitalia

48. * A7 segment more heavily sclerotized than preceding segments. 0: present, 1: absent. (CI = 0.50, RI = 0.00).

a) Without Character 49, Character 48 is not phylogenetically informative. The females of most species have an A7 segment that is more heavily sclerotized than the preceding segments (State 0).

49. * Form of heavy sclerotization of A7. ?: does not apply, 0: continuous around the segment, 1: membranous breaks in the pleurites, 2: membranous break occurring on the sternite, 3: membranous break on the tergite, 4: membranous breaks on pleurites and sternite. (CI = 0.80, RI = 0.75).

a) A heavily sclerotized A7 segment with membranous breaks in the pleurites (State 1) is unique to the ingroup. This character is reversed in Clade 5 (State 0). States 2, 3, and 4 diagnose taxa within the external and internal outgroups.

50. Shape of A7 sternite. 0: sternite unsclerotized, no defined shape, 1: goblet- shaped, 2: M-shaped, 3: shield-shaped, 4: rectangular, length perpendicular to the body, 5: parallelogram, 6: ovoid with x-shaped posterior margin. (CI = 0.75, RI = 0.60).

a) The shape of the A7 sternite is useful for diagnosing species (e.g. State 4, L. pholus) and small species groups. This character will likely be helpful for future species level revisions.

51. Form of distal margin of the A7 sternite. 0: deep concave indentation, extends more than a quarter the length of the A7 sternite, 1: margin approximately horizontal, 2: distal margin membranous, not possible to distinguish from A7/A8 intersegmental membrane, 3: crenellated, 4: shallow concave indentation,

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extends less than a quarter the length of the A7 sternite, 5: convex, 6: A7 distal margin elongated into 2 projections that encircle the ostium bursa. (CI = 0.75, RI = 0.67).

a) A deep concave indentation on the distal margin of the A7 sternite (State 0) is unique to Clade 5. Within this clade, State 0 is modified once in L. fulgens (State 1). In addition, the variation in the form of the distal margin can be used to diagnose species (e.g. State 6, P. ptychoglene).

52. * A8 sternite. 0: present, 1: absent. (CI = 1.00, RI = 1.00).

a) The absence of the A8 sternite (State 1) is unique to the grotei clade and is not reversed in any of the taxa.

53. * Form of A8 sternite. ?: does not apply, 0: heavily sclerotized plate, 1: reduced to a bar fused with A8 tergite, ventral break present, 2: reduced to a continuous sclerotized bar, fused with A8 tergite, 3: semicircular lightly, sclerotized plate, anterior margin a sclerotized bar fused with A8 tergite, 4: sclerotized plate with W-shaped anterior margin, 5: sclerotized bar fused to A7 sternite and A8 tergite. (CI = 0.83, RI = 0.80).

a) The variation in the form of the A8 sternite can be used to diagnose the smaller species groups (e.g. State 0, pholus clade) and species (e.g. State 3, P. ptychoglene). State 5 is unique to the basal taxa within Clade 5. The A8 sternite is completely absent in the grotei clade, which is found in Clade 5.

54. * Location of the ostium bursa. 0: intersegmental membrane between A7/A8, 1: A8 sternite, 2: A7 sternite. (CI = 0.50, RI = 0.71).

a) The ostium bursa is either found in the intersegmental membrane between A7/A8 (State 0) or the A8 sternite (State 1) within the ingroup. State 1 is found within the pholus clade and P. ptychoglene. Both of these states along with state 2 can be found in the outgroup taxa.

55. * Placement of ostium bursa when it occurs in A8 sternite. ?: does not apply, 0: fused with anterior margin of A8 sternite, 1: broad horizontal opening, centered in A8 sternite, extends over half the width of A8 sternite, 2: circular opening occurring anteriorly in A8 sternite, not fused with margin of A8 sternite, 3: narrow horizontal opening, centered in A8 sternite, extends less half the width of A8 sternite. (CI = 1.00, RI = 1.00).

a) The placement of the ostium bursa in the A8 sternite can be used to diagnose species (e.g. State 2, P. ptychoglene) and species groups (e.g. State 1, pholus clade). State 0 defines the internal OG taxa Dolichesia and Talara. State 3 is autapomorphic for the internal OG taxon Lycomorphodes.

56. * Sclerotization of the ductus bursa. 0: present, 1: absent. (CI = 0.20, RI = 0.43).

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a) The presence of sclerotization of the ductus bursa is homoplastic and is not phylogenetically useful without Character 57.

57. * Location of the sclerotized portion of the ductus bursa. ?: does not apply, 0: proximal to the ostium bursa, 1: not proximal to the ostium bursa, membranous break between ostium bursa and sclerotized portion of ductus bursa. (CI = 1.00, RI = 1.00).

a) The presence of sclerotization of the ductus bursa that is not proximal to the ostium bursa (State 1) is unique to the grotei clade and is found in all members of the clade.

58. Corpus bursa. 0: wrinkled, 1: smooth. (CI = 0.33, RI = 0.00).

a) This character is not phylogenetically useful. Most species have a wrinkled corpus bursa (State 0). A smooth corpus bursa (State 0) evolve independently twice in the ingroup (L. fulgens and L. splendens) and once in the internal OG (Talara).

59. Corpus bursa. 0: single, 1: two bursae, connate, arising from ductus, 2: two separate bursae with common membranous duct. (CI = 1.00).

a) Most species possess a single corpus bursa (State 0). State 1 is an autapomorphy of P. ptychoglene. State 2 is unique to Talara.

60. Signa number. 0: one, 1: two, 2: three or more. (CI = 1.00, RI = 1.00).

a) Most of the species examined posses two signa (State 1). L. splendens is diagnosed by the presence of one signum. State 2 is limited to the external OG.

61. Signa form. 0: slightly sclerotized plates with heavily sclerotized bars running perpendicular to the length of the plate, 1: elongate heavily sclerotized strips with internal spines. (CI = 1.00, RI = 1.00).

a) This character (State 0) diagnoses the ingroup + internal OG clade. It is present in all members of the clade and undergoes no reversions.

62. * Origin of the ductus seminalis. 0: ductus bursa, 1: corpus bursa, 2: appendix bursa, 3: second corpus bursa. (CI = 0.60, RI = 0.00).

a) The ductus seminalis arises from the ductus bursa (State 0) in most species examined. Within the ingroup, this state is modified once in P. morelosia (State 1). All four states are present in the outgroup taxa. State 3 is an autapomorphy of Talara.

63. * Location of the ductus seminalis on the ductus bursa. ?: does not apply, 0: ventral, 1: lateral, right side, 2: lateral, left side, 3: dorsal. (CI = 0.60, RI = 0.50).

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a) The location of the ductus seminalis on the ductus bursa is useful for diagnosing species (e.g. State 3, L. fulgens) and small species groups (e.g. State 1, grotei clade). This character will likely be useful in future species level revisions.

64. * Dorsal pheromone gland shape. 0: two triangular projections arising from a single opening, 1: square with elongations from each apical corner. (CI = 1.00, RI = 1.00).

a) The presence of a dorsal pheromone gland that is composed of two triangular projections arising from a single opening (State 0) is unique to the ingroup and present in all members.

65. * Form of the two triangular projections of the dorsal pheromone gland. ?: does not apply, 0: broad triangles, width greater or equal to length of the triangle, 1: short, narrow triangles, length of triangle greater than the base but not more than two times greater. (CI = 1.00, RI = 1.00).

a) Most members of the ingroup clade are diagnosed by State 0. State 1 is unique to the grotei clade.

Characters examined but excluded from the analysis

Several morphological characters were examined but were excluded from the final analysis. These characters showed no variation among species, the range of variation could not be coded into discrete states, or the character was variable among individuals within a single brood of a species.

The two invariant characters arose from the thorax. They included the tarsal claw form and the length of the epiphysis. The tarsal claw was simple in all of the taxa. The epiphysis was found to be short (less than two-thirds the length of the tibia) in all species examined. These characters have been used in higher-level studies of Arctiinae

(Jacobson & Weller 2002; DaCosta & Weller 2005). The form of the tarsal claw was also included in a study of arctiine genera (Zaspel & Weller 2006). The form of the tarsal claw and the length of the epiphysis were found to be phylogenetically informative in the higher-level studies (Jacobson & Weller 2002; DaCosta & Weller 2005). However,

Zaspel & Weller (2006) found this the tarsal claw form to be highly homoplasious within

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the genus Virbia. These findings along with the absence of variation within the species

examined in this study suggest that these characters are informative for higher-level

phylogenetic studies.

Several characters examined on the male& female genitalia were found to have

a range of variation that could not be quantified into discrete states. On the male

genitalia, the shape of the dorsal tegumen was examined. Within both Lycomorpha and

Propyria, the fusion between the two halves of the tegumen results in this structure occurring as a contiguous plate. Although the fusion has occurred, the halves of the tegumen can still be identified in several species through the presence of sutures that are visible on the dorsal surface of the tegumen. In some species, a raised node is present between the two halves of the tegumen. However, no landmarks could be identified on the male genital capsule to accurately describe the location of this node.

Another character that was examined but excluded from the final analysis was the presence of a spine at the distal apex of the valvae. Within Lycomorpha and Propyria, the valvae either taper to a sclerotized spine or a sclerotized spine is present at the distal apex. However, the valvae of all the species examined are sclerotized for the entire length. It is difficult to identify the margins of structures such as the sacculus and costa. Therefore, it was not possible to determine whether the spines represented a homologous structure or several distinct structures. In the female genitalia, the shape of the ductus bursa was examined. Within some species the ductus bursa occurred as a narrow tube. However, in other species the width of the ductus bursa varied along its length. We were not able to quantify this variation within a single character state that was not continuous.

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Two characters were excluded because they were found to vary among individuals from a single brood. Comstock & Henne (1967) reared broods of the species

L. regulus. Through examination of this material it was found that within a single brood variation was present in form of the M3 and CuA1 vein in the hindwing and the number of frenular bristles present on the female hindwing. Members of this brood were found to have M3 and CuA1 arising separately from the discal cell or M3 and CuA1 were fused and stalk beyond the discal cell. Propyria has been distinguished from Lycomorpha by the latter form. The presence of both states within a single species of Lycomorpha suggests that this character can no longer be used to separate the two genera. Within this brood it was also found that the frenular bristle number varied between four and five. In addition, both states were present on some specimens. Although the frenular number has previously been used in higher-level studies (DaCosta & Weller 2005; Miller

1991), it would not be an informative character within this study unless a range of values was coded as a single state.

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Table 4-1. Historical family treatment of Lycomorpha including authors, year, and their placement of the genus. Author(s) Harris 1839 Edwards Neumogen & Dyar Hampson 1898, Forbes Hampson 1914 Packard 1864, 1886 1893 1901 1960 Comstock & 1872 Dyar 1898 Zerny 1912 Henne 1967 Druce 1881- Draudt 1917 1900 Edwards 1885, 1887 Schaus 1889 Dognin 1916 Family Zygaenidae Ctenuchidae Euchromiidae Syntomiidae Lithosiidae Amatidae

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Table 4-2. Species included in the phylogenetic analysis. CHS = specimen prepared by Clare H. Scott Genus species author Dissection Sex Collection Locality INGROUP

Lycomorpha New Mexico, L. fulgens (Henry Edwards) CHS084 Male AMNH USA CHS088 Female AMNH Arizona, USA L. grotei (Packard) CHS079 Male LACM Colorado, USA CHS093 Female LACM California, USA South Dakota, L. miniata Packard CHS071 Male AMNH USA CHS051 Female AMNH Wyoming, USA L. pholus (Drury) CHS136 Male USNM West Virginia, CHS074 Female CMNH USA L. pulchra Dyar CHS076 Male CAS California, USA CHS095 Female SDNH California, USA New Mexico, L. regulus (Grinnell) CHS099 Male AMNH USA CHS102 Female LACM California, USA L. splendens Barnes& CHS068 Male USNM Utah, USA McDunnough CHS061 Female LACM Arizona, USA New Mexico, New sp. A CHS086 Male AMNH USA CHS060 Female LACM Arizona, USA New sp. B CHS128 Male USNM Texas, USA CHS129 Female CAS Texas, USA New Mexico, New sp. C CHS253 Male CMNH USA

Propyria P. morelosia Schaus CHS196 Male USNM Mexico CHS247 Female AMNH Mexico P. normani Schaus CHS252 Male USNM Mexico P. ptychoglene Hampson CHS103 Male USNM Mexico CHS104 Female USNM Mexico

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Table 4-2. Continued Genus species author Dissection Sex Collection Locality OUTGROUP TAXA

Dolichesia D. falsimonia Schaus CHS158 Male MCZ Panama CHS213 Female MCZ Panama

Hypermaepha H. marionensis Schaus CHS163 Male CUIC Surinam CHS218 Female CUIC Surinam

Hypoprepia H. fucosa Hübner CHS002 Male UWO Florida, USA CHS004 Female FLMNH Florida, USA

Lycomorphodes L. correbiodes Schaus CHS052 Male CMNH Ecuador CHS053 Female CMNH Ecuador

Ptychoglene P. erythrophora Felder CHS121 Male USNM Mexico CHS122 Female USNM Mexico

Talara T. coccinea Butler CHS055 Male CMNH Brazil CHS056 Female CMNH French Guiana

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Table 4-3. Character support for major ingroup clades. ‘Abbreviation’ refers to the name given to the character in Appendix D. CI = consistency index. Clade Character #: Abbreviation State Character Change CI Clade 1. Ingroup 1: A1 0 to 1 0.67 2: A2 0 to 1 0.33 6: L1 0 to 1 1.00 9: AB1 1 to 0 0.50 10: MA1 0 to 1 0.50 11: MA2 0 to 1 1.00 17: MA8 0 to 1 1.00 27: CM1 1 to 0 0.50 29: PB2 0 to 1 1.00 32: ED2 2 to 0 0.67 33: J1 0 to 4 0.88 35: J3 3 to 0 0.80 36: VS1 0 to 2 0.67 37: IV1 1 to 0 1.00 38: BP1 0 to 2 1.00 40: PH1 1 to 0 0.25 49: FA2 0 to 1 0.80 51: FA4 2 to 1 0.75 55: OB2 0 to 1 1.00 64: PG1 1 to 0 1.00 Clade 2. pholus clade 24: UB2 0 to 2 1.00 26: US2 0 to 3 0.57 31: ED1 0 to 1 0.33 47: V6 0 to 1 0.50 50: FA3 2 to 1 0.75 Clade 3. Remaining Ingroup 2: A2 1 to 0 0.33 21: TF3 1 to 4 0.80 28: PB1 1 to 0 0.33 33: J1 4 to 7 0.88 34: J2 1 to 0 0.20 41: PH2 0 to 2 1.00 44: V3 0 to 4 1.00 51: FA4 1 to 0 0.75 53: FA6 0 to 3 0.83 55: OB2 1 to 2 1.00 Clade 4. Propyria 11: MA2 1 to 2 1.00 29: PB2 1 to 2 1.00 32: ED2 0 to 1 0.67 50: FA3 2 to 6 0.75 51: FA4 0 to 6 0.75 63: DS2 0 to 2 0.60

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Table 4-3. Continued Clade Character #: Abbreviation State Character Change CI Clade 5. morelosia + remaining 25: US1 1 to 0 0.75 Lycomorpha 26: US2 0 to 1 0.57 49: FA2 1 to 0 0.80 53: FA6 3 to 5 0.83 54: OB1 1 to 0 0.50 56: DB1 1 to 0 0.20 Clade 6. grotei clade 12: MA3 1 to 0 0.50 24: UB2 0 to 3 1.00 28: PB1 0 to 1 0.33 52: FA5 0 to 1 1.00 57: DB2 0 to 1 1.00 63: DS2 0 to 1 0.60 65: PG2 0 to 1 1.00

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Table 4-4. Results of Outgroup Jackknife Analysis. Consistency Index (CI) and Retention Index (RI) rounded Outgroup Excluded Number of Trees Tree Length CI RI None 3 192 0.68 0.73 Dolichesia 9 190 0.67 0.70 Hypermaepha 6 175 0.70 0.75 Hypoprepia 9 191 0.68 0.69 Lycomorphodes 3 179 0.70 0.74 Ptychoglene 3 184 0.70 0.72 Talara 6 186 0.68 0.70

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Figure 4-1. Species mistakenly described as members of Lycomorpha and their current taxonomic placement. A) Ctenucha augusta (Henry Edwards) (Lepidoptera: Erebidae: Arctiinae: Arctiini). B) Neoalbertia constans (Henry Edwards) (Lepidoptera: Zygaenidae). C) Ptychoglene coccinea (Henry Edwards) (Lepidoptera: Erebidae: Arctiinae: Lithosiini).

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Figure 4-2. Lycomorpha species with red forewings and primarily black hindwings. A) L. fulgens. B) L. grotei. C) L. pulchra. D) L. regulus.

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Figure 4-3. Strict consensus of 3 trees (L=192, CI=0.68, RI=0.71) resulting from the MP analysis of the MF matrix. Numbers above the branches are the Bremer support values for the nodes. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-4. Strict consensus of 3 trees (L=192, CI=0.68, RI=0.73) resulting from the MP analysis of the AS matrix. Numbers below the branches are the support values for the nodes: BS/JK. C1 – C6 = Clade 1 – Clade 6. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-5. Bayesian Inference consensus tree from the analysis of the AS matrix. Numbers listed on the branches are the posterior probability support values. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-6. Strict consensus of 9 trees (L=191, CI=0.68, RI=0.69) resulting from the exclusion of outgroup taxon Hypoprepia. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-7. Strict consensus of 3 trees (L=184, CI=0.70, RI=0.72) resulting from the exclusion of outgroup taxon Ptychoglene. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-8. Strict consensus of 6 trees (L=186, CI=0.68, RI=0.70) resulting from the exclusion of outgroup taxon Talara. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-9. Strict consensus of 3 trees (L=179, CI=0.70, RI=0.74) resulting from the exclusion of outgroup taxon Lycomorphodes. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-10. Strict consensus of 6 trees (L=175, CI=0.70, RI=0.75) resulting from the exclusion of outgroup taxon Hypermaepha. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-11. Strict consensus of 9 trees (L=190, CI=0.67, RI=0.70) resulting from the exclusion of outgroup taxon Dolichesia. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-12. Strict consensus of 24 trees (L=198, CI=0.66, RI=0.71) resulting from constraining the monophyly of Lycomorpha. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-13. Strict consensus of 30 trees (L=194, CI=0.67, RI=0.72) resulting from constraining the monophyly of Propyria. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 4-14. Male antennal flagellomeres. A) Lycomorphodes correbiodes; simple, ciliate flagellomeres. B) Lycomorpha fulgens; serrate flagellomeres. C) Lycomorpha splendens; bipectinate flagellomeres. The scale bar is equivalent to 1mm.

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Figure 4-15. Female antennal flagellomeres. A) Lycomorpha grotei; simple, ciliate flagellomeres. B) Lycomorpha splendens; serrate flagellomeres. The scale bar is equivalent to 1mm.

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Figure 4-16. State of gena. A) Lycomorpha pholus; G-gena well developed, can be seen as a continuous band that joins the F-frons. B) Lycomorpha splendens; G- gena reduced, cannot be seen as a continuous band. The scale bar is equivalent to 1mm.

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Figure 4-17. Labial palps. A) Hypermaepha maroniensis; all segments fused. B) Lycomorpha splendens; 2nd and 3rd segments fused. C) Lycomorpha pholus; each segment separate. The scale bar is equivalent to 1mm.

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Figure 4-18. Forewing venation. A) Lycomorpha grotei; R free to costal margin, does not anastomose with Sc. B) Dolichesia falsimonia; R anastomoses with Sc. The scale bar is equivalent to 1mm.

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Figure 4-19. Hindwing venation. A) Hypoprepia fucosa; Sc+R vein present. B) Propyria ptychoglene; Sc+R vein absent. The scale bar is equivalent to 1mm.

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Figure 4-20. Second abdominal sternite. A) Lycomorpha pholus; anterolateral process (ALP) that arises from A-apodeme present as a SB-sclerotized bar. B) Ptychoglene erythrophora; ALP that arises from A-apodeme occurs as FL- flattened lobe. C) Hypermaepha maroniensis; ALP that arises from A- apodeme present as SA-short knob ALP. The scale bar is equivalent to 1mm.

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Figure 4-21. A7/A8 androconia. A) Dolichesia falsimonis; androconia absent from A7/A8 intersegmental membrane. B) Hypermaepha maroniensis; TP-two pockets of long hair-like scales in A7/A8 intersegmental membrane. C) Lycomorpha splendens; SP-shallow pocket of scales that is greater than 1/3rd the width of 7S-seventh sternite. D) Propyria normani; NP-narrow pocket of scales less than 1/3rd the width of 7S-seventh sternite. The scale bar is equivalent to 1mm.

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CHAPTER 5 A REVISION OF THE GENUS LYCOMORPHA HARRIS (LEPIDOPTERA: EREBIDAE: ARCTIINAE: LITHOSIINI) INCLUDING SPECIES FORMERLY PLACED IN THE GENUS PROPYRIA HAMPSON (LEPIDOPTERA: EREBIDAE: ARCTIINAE: LITHOSIINI)

Background Information

The brightly colored species of the genus Lycomorpha Harris (Lepidoptera:

Erebidae: Arctiinae: Lithosiini), which includes species formerly placed in Propyria

Hampson, range from southeastern Canada to Panama. The adults typically possess either orange and black or red forewings. Species possessing either of these color patterns are hypothesized to form Müllerian mimicry complexes with other arctiine genera (e.g. Ctenucha Kirby, Dycladia Felder, and Correbia Herrich-Schäffer), zygaenid moths, and lycid beetles (Forbes 1960; Simmons 2009). In addition to sharing similar color patterns with these chemically defended, diurnal taxa, adults of several species of

Lycomorpha also exhibit diurnal flight activity. Although the species of Lycomorpha are well known for their bright color patterns, these patterns have also led to confusion over both the family placement and species composition of the genus.

In order to address the confusion over the species composition of Lycomorpha, a phylogenetic study of the genus was completed (see Chapter 4). This study included representatives of seven species of Lycomorpha and three undescribed species. In order to test the generic limits of Lycomorpha, species placed in the lithosiine genera

Propyria and Ptychoglene Felder were also included. Draudt (1917) noted a similarity of color pattern and body shape among these three genera. Furthermore, species originally described within Lycomorpha have been transferred to both Propyria and

Ptychoglene. The phylogeny that was recovered based on adult morphology found that

Lycomorpha was not reciprocally monophyletic with respect to the genus Propyria (Fig.

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5-1). However, Ptychoglene was found to be more closes related to the genus

Hypoprepia Hübner, the species used to root the analysis. As a result of this analysis,

Propyria (Hampson 1898) was placed as a junior synonym of Lycomorpha (Harris 1839)

(see Chapter 4).

Little is known of the biology of the species in the genus Lycomorpha. The flight activity Lycomorpha pholus (Drury), the most widely distributed species, has been studied. Fullard & Napoleone (2001) found that L. pholus was exclusively diurnal. Based on this finding, Muma & Fullard (2004) tested L. pholus to determine whether the tympanum was still functional. Since L. pholus is exclusively diurnal, the species is no longer under selection by bats to maintain functional tympana unless selected for by an alternative use. They found that L. pholus had low sensitivity to ultrasounds of the frequency produced by bats. However, L. pholus adults retained sensitivity to ultrasound frequencies in the range produced by the tymbal organ present on their metathorax.

The authors suggest that the tympanum of L. pholus may be used in short-range, social communication. This communication could occur during the courtship of female L. pholus. Males of both Lithosiini and Arctiini have been found to use ultrasonic noises produced by their tymbal organs as part of their courtship ritual (Černý 1990, Conner

1999, Sanderford 2009). In addition, the larvae of L. pholus have been reared on

Protococcus viridis and Physcia millegrana lichens (Wagner et al. 2008).

Lycomorpha regulus (Grinnell) is the only other species in the genus where biological information besides that present on collection labels is available. Comstock & Henne

(1967) reared several broods of this species on Parmelia lichens. They observed the adults flying during the day and feeding at flowers of Lepidospartum squamatum and

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Senecia douglasii. In addition, they found that the females oviposited in the crevices of

rocks.

Here a revision of the species of Lycomorpha is provided. A generic diagnosis and description of the genus is presented. In addition, a faunal treatment is given for fourteen of the twenty species currently placed in the genus. The inclusion of species was dictated by the availability of pinned material. Four of the species treated are formerly of the genus Propyria. Three new species (L. concolor Scott, L. neomexicanus

Scott, and L. texanus Scott) are also treated for the first time. Each faunal treatment includes a diagnosis, description, and illustrations of diagnostic characters. The adult habitus is figured for both sexes (Figs. 5-2 A-H, 5-3 A-H, 5-4 A-H, 5-5 A-H, 5-6A,B, and

5-7). A summary of life history information is also provided. This summary is based on the literature and the data available on the collection labels.

Taxonomic History of Lycomorpha and Propyria

The genus Lycomorpha was originally described as a subgenus of Glaucopis

Fabricius within Zygaenidae (Harris 1839). Sphinx pholus Drury was designated as the type species by monotypy. Harris (1839) also treated the arctiine genera Syntomeida

Harris, Cosmosoma Hübner, and Ctenucha Kirby as subgenera of Glaucopis. However, he noted that these four subgenera did not appear congeneric and might be raised to generic status when more was known of them. After this description, authors (Edwards

1881a, 1881b, 1882, 1884, 1885; Packard 1864, 1872; Druce 1881-1900) treated

Lycomorpha as a genus of Zygaenidae until Edwards (1886) transferred the genus to the family Ctenuchidae, which is now a subtribe of Arctiini. However, the following year

Edwards (1887) returned to treating Lycomorpha as a zygaenid genus. Neumogen &

Dyar (1893) then transferred Lycomorpha to Euchromiidae, which is now a subtribe of

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Arctiini. However, some authors (Dognin 1916) continued to treat the genus as a zygaenid. Later Hampson (1898) transferred Lycomorpha to Syntomiidae. This family was later synonymized with Ctenuchidae. This placement was generally accepted

(Zerny 1912; Draudt 1917) until Forbes (1960) transferred the genus to Lithosiini. In his description of Lycomorpha, Forbes noted that the loss of the Sc vein was the only character that supported the placement of Lycomorpha within Euchromiidae. Jacobson

& Weller (2002) confirmed the placement of the genus within Lithosiini in their phylogenetic analysis of Arctiinae.

Three genera have been synonymized with Lycomorpha: Anatolmis Packard,

Prepodes Herrich-Schäffer, and Propyria Hampson. In his description, Packard (1864) treated Anatolmis as member of Zygaenidae, and Anatolmis grotei Packard was designated as the type species by monotypy. Packard noted that Anatolmis was morphologically similar to Lycomorpha, which he considered to be the sister taxa to

Anatolmis. Furthermore, he stated that both genera superficially resemble Lithosiini.

Hampson (1898) later synonymized the two genera. However, no explanation was provided for this action. Herrich-Schäffer (1855) described Prepodes, and designated

Sphinx pholus as the type species by monotypy. However, this species was already designated as the type of Lycomorpha. The two genera were synonymized, and

Lycomorpha was given priority as the older name. Hampson (1898) described Propyria and designated Propyria ptychoglene Hampson as the type species. Propyria was treated as a member of Syntomiidae, and Draudt (1917) noted similarities between

Propyria, Ptychoglene, and Lycomorpha. A phylogenetic analysis of the three genera

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found that Lycomorpha and Propyria were not reciprocally monophyletic. Scott (Chapter

4) synonymized the two genera.

Throughout the shifts in family placement and the addition of synonyms, new species have continued to be described in Lycomorpha. However, nineteen of the species that were originally described in the genus have been transferred to other genera within Lithosiini, Arctiini, and Zygaenidae. Bryk (1936) transferred ten of these species to three genera of Zygaenidae. An additional three species are considered incertae sedis. Dyar (1898) did not consider two of these species, L, contermina

Edwards and L. regia Schaus, to be members of Lycomorpha. However, he did not suggest alternative placements. In addition, no further mentions can be found of these species in descriptions of the species composition of Lycomorpha. Scott (Chapter 4) transferred the third species, L. nigridorsata Dognin, based on the original species description. In his description of L. nigridorsata, Dognin (1916) only compared this species to other species of Lycomorpha that have been transferred into zygaenid genera.

The taxonomy of Lycomorpha has been confused by the wing color patterns of the adults. These patterns led to the placement of the genus in several families that are known for being composed of brightly colored individuals. Furthermore, the use of this color system instead morphological characters led to species being mistakenly described as members of Lycomorpha. Below a synonymic checklist is provided that includes all species currently placed within Lycomorpha. A list, which includes the species that have been transferred from the genus, is available in Chapter 4.

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Synonymic Checklist of the Genus Lycomorpha

In this checklist, the species names are arranged in alphabetical order. Valid species names are in bold and synonyms are in italics. If a species was originally described in a genus other than Lycomorpha, the original generic name follows the author’s name. The location where the type specimen of a species was collected is provided in parentheses following the original generic name. An asterisk (*) following a name indicates the type specimen of that species was examined.

Genus Lycomorpha Harris, 1839

Anatolmis Packard, 1864

Prepodes Herrich-Schäffer, 1855

Propyria Hampson, 1898 atroxantha* (Schaus, 1906) Propyria (Mexico) concolor Scott n. sp. (Arizona) criton (Druce, 1881-1900) Cisthene (Guatemala)

coatepeciensis* (Strand, 1920) Propyria (Mexico)

a. hypoleuca (Draudt, 1917) Propyria (Mexico)

orizabae* (Strand, 1920) Propyria (Mexico) flora (Schaus, 1911a) Propyria (Costa Rica) fridolinia* (Schaus, 1925) Propyria (Guatemala) fulgens* (Henry Edwards, 1881b) Anatolmis (Arizona)

tenuimargo* (Holland, 1903) Ptychoglene (Arizona, Mexico)

tenumargo authors, misspelling grotei (Packard, 1864) Anatolmis (Colorado)

palmerii* Packard, 1872 (Arizona)

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miniata* Packard, 1872 (Southern California) morelosia* (Schaus, 1925) Propyria (Mexico) neomexicanus Scott n. sp. (New Mexico) normani* (Schaus, 1911b) Propyria (Costa Rica) pelopia (Druce, 1881-1900) Talara (Panama) pholus (Drury, 1773) Sphinx (New England) ptychoglene (Hampson, 1898) Propyria (Mexico, Guatemala)

aequalis (Walker, 1854a) Lithosia (Guatemala)

sinuata (Henry Edwards, 1885) (Mexico) pulchra Dyar, 1898 (Texas) regulus (Grinnell, 1903) Anatolmis (California) schausi* Dyar, 1898 (New Mexico, Arizona) splendens Barnes & McDunnough, 1912 (Utah) strigifera Gaede, 1926 (Mexico) texanus Scott n. sp. (Texas)

Materials and Methods

Material Examined

This revision was based on the examination of 1,997 pinned specimens. To examine the species limits, 143 dissections of the genitalia of the fourteen species described below were completed. The label data was copied exactly for each specimen that was dissected. Each specimen was assigned a unique dissection voucher number.

This voucher number was placed on the pin with the remaining moth body. Another copy of the voucher number was kept with the genitalia being dissected. A voucher number beginning with either CHS or CHS-PC indicates that Clare H. Scott completed

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the dissection. In addition, specimens from the USNM were assigned a unique

dissection number, which is associated with the slide prepared for that dissection. This

number was also provided in the specimens examined section of each species

description.

The following is a list of institutions that were consulted for this study. The list

includes both the name of each collection and an acronym for each. The name of the

individual(s) that prepared the loan follows this information. Acronyms follow Heppner &

Lamas (1982): American Museum of Natural History (AMNH) (D. Grimaldi), California

Academy of Sciences (CAS) (N.D. Penny), Carnegie Museum (CMNH) (J. Rawlins),

Canadian National Collection, Agriculture Canada (CNC) (C. Schmidt), Cornell

University Insect Collection (CUIC) (J. Liebherr, E.R. Hoebeke), Florida Museum of

Natural History, University of Florida (FLMNH) (J. Miller, D.M. Lott, A.D. Warren), Los

Angeles County Museum of Natural History (LACM) (W. Xie), Museum of Comparative

Zoology, Harvard University (MCZ) (R. Eastwood, P.D. Perkins), Museum of

Southwestern Biology (MSB) (D. Lightfoot), National Museum of Natural History,

Smithsonian Institution (USNM) (D.G. Furth, D. Harvey), Peabody Museum of Natural

History, Yale University (YPM) (L.F. Gall), San Diego Natural History Museum (SDNH)

(M.A. Wall), Texas A&M University Insect Collection (TAMU) (E.G. Riley), Essig

Museum of Entomology, University of California, Berkeley (UCB) (J. Powell), University

of Minnesota, St. Paul (UMSP) (R. Holzenthal).

Morphology

Dissections of the genitalia were prepared after softening the abdomens in a warm 10% potassium hydroxide (KOH, Fisher Scientific, Pittsburg, PA) solution for 30 minutes to 1.5 hours using standard methods (Winter 2000). The scales and viscera

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were removed from the abdomen in several rinses of 20% ethanol using fine watercolor paintbrushes (# 000 – 2). Structures were stained with a solution of chlorazole black E

(Acrōs) dissolved in deionized water. Specimens were viewed in 20% ethanol and stored temporarily in 70% ethanol. Subsequently the abdominal pelts and genitalia were either stored in vials of glycerol or permanently slide mounted in Euparol (Bioquip,

Garden City, CA). The final preparation method used was dependent on the preference of the collection from which the material originated. The wing veins of Lycomorpha species are well developed. Therefore, it was not necessary to remove the wings and clear them using bleach. Instead, the venation was viewed by adding one to two drops of 95% ethanol to the wing. The external morphology of the dried, pinned specimens was examined using a Nikon SMZ800 light microscope. The terminology for male and female genitalia follows Klots (1970) and Forbes (1939b, 1954). The terminology for the wing venation follows Kristensen (2003).

Illustrations of the external morphology and male and female genitalia were completed using a camera lucida associated with a Leica MZ16 light microscope. The pencil drawings produced were scanned and saved as PDFs that were imported into

Adobe Illustrator CS5 and then inked using the pen tool. The purpose of stippling in any figure is explained in the figure legend. Illustrations are provided for all diagnostic characters. In addition, when a form of a morphological character was present in more than one species, a single illustration was produced and referenced more than once.

Species Descriptions

The species placed within Lycomorpha are not sexually dimorphic. There are slight size differences between males and females. Characters from the external morphology, as well as the male and female genitalia were used to diagnose species.

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Phenotypically similar species were primarily separated using male and female genitalia characters. However, it was difficult to separate some species using these characters.

In addition, variation was present in the wing color pattern of individuals reared from a single brood. However, studies of these broods made it possible to determine elements of the wing color pattern that were constant throughout a species. When describing elements of the wing color pattern the terminology for the typical noctuid ground plan

(Powell & Opler 2009) was used. Although Ferguson (1985) proposed a wing pattern terminology specific to Arctiinae, homologous pattern elements cannot be identified within Lycomorpha. The colors of the specimens were described using the Naturalist’s

Color Guide (Smith 1975). Images illustrating the adult habitus were taken using an

Olympus Camedia C-5500 Zoom digital camera.

Systematic Entomology of the Genus Lycomorpha

Order LEPIDOPTERA

Family EREBIDAE

Subfamily ARCTIINAE

Tribe LITHOSIINI

Genus Lycomorpha Harris

Lycomorpha Harris, 1839: p. 317. Type species: Sphinx pholus (Drury, 1773).

Anatolmis Packard, 1864: p. 45

Prepodes Herrich-Schäffer, 1855: pp. 100,101

Propyria Hampson, 1898: p. 521

Diagnosis (Figs. 5-2 A-H, 5-3 A-H, 5-4 A-H, 5-5 A-H, 5-6A,B, and 5-7). The adults of this genus are medium sized lithosiines with a forewing length of 11mm to

16.1mm (N = 13) in the males and 11.7mm to 15.5mm (N = 12) in the females. The

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male antennal flagellomeres are typically serrate (Fig. 5-8A). The female antennal

flagellomeres can be either simple and ciliate or serrate (Figs 5-9A,B). In both sexes,

the antennae are dilated in the middle. There are two primary wing color patterns within

the genus: 1) a red forewing with a primarily black hindwing and 2) a forewing with an

orange or red basal half and a black distal half and a black hindwing with the adbasal

region the same color as the basal half of the forewing. Individuals of both wing color

patterns form mimicry complexes with other genera of Arctiinae and beetles of the

family . Four autapomorphies can be used to separate Lycomorpha from other

Lithosiini: 1) a tibial spur formula of 0-2-3, 2) a shallow pocket of sex scales in the

intravincular membrane of the male genital capsule, 3) the absence of a caecum on the

basiphallus, and 4) a dorsal pheromone gland shaped like a square with elongations

arising from each of the cephalic corners.

Description. Adult Habitus (Figs. 5-2 A-H, 5-3 A-H, 5-4 A-H, 5-5 A-H, 5-6A,B, and 5-7): Head: The color is typically uniform and varies from dark gray brown (Fig. 5-

5C) to a dark brown that appears almost black (Fig. 5-4C). The scales covering the gena of some species are the same color as the patagia and forewing (Fig. 5-7).

Typically, none of the three labial palp segments are fused (Fig. 5-10A). Thorax: The

patagia are typically the same color as the red or orange color present on the forewings

(Figs. 5-2A,B). The color of the tegulae and dorsal mesothorax is either the same as the

head (Fig. 5-2C) or matches the color present on the patagia (Fig. 5-2B). The dorsal

metathorax is always the same color as the head. Tymbal organs are always present in

both the males and females. Forewing: The R and Rs1-4 veins are all present with a

branching pattern of R; Rs1; (Rs2 (Rs3, Rs4)). The M1-M3 veins are present. The M2 and

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M3 veins arise separately from the discal cell or are fused and stalk beyond the cell

(Figs 5-11A,B). Both states are found in some species. Hindwing: The Sc + R vein and the M2 vein are absent throughout the genus. The M3 and CuA1 veins arise separately

from the discal cell or are fused and stalk beyond the discal cell (Figs 5-12A,B). The

wing color pattern of the fore and hindwings is one of two main types discussed in the

diagnosis. Abdomen: The color of the abdomen is a dark brown that is the same as the

color of the head and dorsal metathorax (Figs 5-2 A-H). The anterolateral process is

typically present as a sclerotized bar (Fig. 5-13A). A single pouch of androconial scales

is present in the A7/A8 intersegmental membrane of males (Figs 5-14A,B). In addition,

a pouch of androconial scales is present in the A8/A9 intersegmental membrane of

some species. The variation in the sclerotization of the eighth tergite and the form of the

cephalic margin of the eighth sternite are specific to species groups. Male Genitalia:

Phallus: The caecum is always absent from the basiphallus, which is typically straight

without inflections. A phallic sclerite is present in most species and variation is present

in the shape: rectangular (L. splendens), narrow, rounded triangle (L. grotei), narrow,

pointed triangle (L. pulchra), and flattened lobe that extends dorsad over the vesica (L.

pholus). The membrane of the vesica is smooth, and the primary lobe is typically

ornamented with at least one membranous projection and a heavily sclerotized, spine

like cornutus. The placement of the projection(s) and cornutus are typically species

specific. Genital Capsule: The two halves of the tegumen are fused for their entire

length. A suture is present on the tegumen indicating the location of the fusion. The

shape and location of the suture is species specific. The uncus base is always

separated from the tegumen by membranous tissue. The curvature of the uncus is

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either C-shaped (L. splendens) or S-shaped (L. concolor). The distal end of the costa is

either defined by the presence of a processus basalis of the costa (L. morelosia) or a

membranous break in the dorsal edge of the valve (L. neomexicanus). The costa and

sacculus are fused into a contiguous structure. The distal end of the valve tapers to a

point (L. grotei) or a spine (L. ptychoglene) that is perpendicular to the length of the

valve and oriented toward the uncus. The shape of the juxta and its ornamentation with

transparent patches is generally species specific. The vinculum/saccus are typically M-

shaped. Female Genitalia: The seventh abdominal segment is heavily sclerotized. The

sclerotization is either continuous around the segment (L. concolor) or membranous

breaks occur on the pleurites (L. pholus). The shape of the seventh sternite is variable:

goblet shaped (L. pholus), inverted M-shaped (L. concolor), shield shaped (L.

splendens), rectangular, perpendicular to the length of the body (L. miniata), parallelogram shaped (L. fulgens), and ovoid with X-shaped posterior margin (L. ptychoglene). The ostium bursa is located in either the A7/A8 intersegmental membrane

(L. grotei) or the eighth sternite (L. texanus). The ductus bursa is typically unsclerotized.

The shape of the ductus bursa is species specific. Typically only a single corpus bursa is present. The membrane of the corpus bursa is typically wrinkled and ornamented with two signa, which occur on opposite sides of the corpus bursa. The signa are lightly sclerotized, ovoid plates with heavily sclerotized bars occurring perpendicular to the length.

Discussion. A wide variation of the color red is present on the fore and hindwing of species that possess the red forewing and primarily black hindwing phenotype (Figs.

5-2 F-J and 5-3A,B). The color can vary from a deep red to a light yellow orange. This

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variation appears to be due to the age of the specimen and cannot be associated with the location where the specimen was collected. In addition, a wide range of phenotypic variation is present within individuals from a single brood. The broods reared by

Comstock & Henne (1967) help to define the phenotypic variation present within L. regulus. In addition, this series was valuable in determining the species boundaries of L. grotei, L. pulchra, and L. regulus. These three species all possess the same wing color pattern phenotype. Comstock & Henne (1967) and Powell & Opler (2009) suggested these species be studied to determine their systematic and taxonomic relationships.

The male and female genitalia are useful for separating species that possess similar wing color patterns. In the male genitalia, the shape of suture on the tegumen and the shape of the valve are typically species specific. Within the female genitalia, the shape of the eighth sclerite and the ductus bursa are helpful in separating species with similar wing color phenotypes.

These moths are often found misplaced in uncurated Zygaenidae and Arctiini.

Adult Lycomorpha do not possess either ocelli or chaetostomata. Both of these structures are well developed in Zygaenidae, and their absence can be used to separate Lycomorpha from Zygaenidae. The absence of ocelli and tympanal hoods can be used to separate Lycomorpha from Arctiini. In addition, pleural sclerites are present on the male genital capsule of Lycomorpha. This structure is absent throughout Arctiini.

Lycomorpha atroxantha (Schaus)

Fig. 5-2A (female).

Propyria atroxantha Schaus 1906: 193-194 [type locality: Cuesta de Misantla,

Vera Cruz, Mexico]

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Material examined. Type material: Holotype of L. atroxantha: Propyria atroxantha Schaus, Cuesta de Misantla, Vera Cruz, Mexico, Cat. No. 8516, USNM.

Specimen preparations. All material from USNM unless otherwise noted:

GUATEMALA: Alta Verapaz: Tactic, viii, Schaus and Barnes collection (1 female,

CHS254, USNM 127,737).

Diagnosis. Lycomorpha atroxantha can be confused with L. pholus, L. pelopia, and L. texanus. The geographic distribution of L. atroxantha can be used to separate it from both L. pholus and L. texanus. Lycomorpha axtroxantha has been collected from

Mexico, Guatemala, and Costa Rica. Both L. pholus and L. texanus are restricted to

North America. The tibial spur formula and thoracic color pattern can be used to separate L. atroxantha from each of the previously mentioned species. The tibial spur formula of L. atroxantha is 0-2-2, whereas L. pholus, L. pelopia, and L. texanus have a tibial spur formula of 0-2-3. In addition, a stripe of orange yellow scales extends down the middle of the dorsal thorax in L. atroxantha. However, the dorsal surface of the thorax of L. pholus, L. pelopia, and L. texanus is composed only of dark brown scales.

Description. Female habitus (Fig. 5-2A): Head: Scales on the dorsal portion of the head, clypeus, antennae, and labial palps fuscous (21) colored. The antennae are serrate. No fusion occurs among the labial palp segments. The gena is well developed and can be seen as a continuous band around the eye that joins with the frons when the head is viewed in profile. The scales on the gena and frons are trogon yellow (153) colored. Thorax: The tegulae are covered with a mixture of spectrum orange (17), orange yellow (18), and trogon yellow (153) colored scales. The patagia are raw umber

(223) colored with orange yellow scales present around the margin. The dorsal thorax is

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covered with raw umber colored scales. A stripe of orange yellow scales is found in the center of the thorax. The stripe extends from the under the patagia to the posterior margin of the mesothorax. The ventral thorax and legs are fuscous (21) colored. The tibial spur formula is 0-2-2. Forewing: Length = 12mm (N = 1); M2 and M3 fused and stalk beyond the discal cell. Dorsal surface: The basal half of the wing is trogon yellow

(153) colored. In the basal half of the wing, mars brown (223A) and raw umber (223) colored scales are intermixed with the trogon yellow scales within the discal cell, between the CuA1 and A1 veins, and posterior to the A1 vein. The distal half of the wing is fuscous (21) colored. The internal margin of the distal half of the wing is curved convexly towards the wing base and serrate. The margin is indented by trogon yellow scales that line the A1 vein for one-third the width of the fuscous coloring, the CuA2 vein base, and the remainder of the posterior margin of the discal cell to the point where the

M2 + M3 and CuA1 veins arise. Dark grayish brown (20) colored scales line the posterior margin of the wing. These scales extend to the wing base. Ventral surface: The wing is warm buff (118) colored. A hair brown (119A) colored band (3 – 5mm wide) occurs along the outer margin of the wing. The interior margin of this band is curved convexly towards the wing base and smooth. Hindwing: CuA1 and M3 fused and stalk beyond the discal cell. Dorsal surface: The wing is orange yellow (18) colored. A raw umber colored marginal band is present. The interior margin of the band is sinuate. The band extends from the apex of the wing to the anal angle. Raw umber colored scales occur intermittently along the posterior margin of the wing. Ventral surface: The pattern is the same as the dorsal surface. The wing color is the same as the dorsal surface. The marginal band is hair brown colored. Hair brown colored scales do not occur along the

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posterior margin of the wing. Abdomen: Dorsal and lateral scales raw umber (223) colored. Ventral scales are fuscous (21) colored. Female genitalia: The heavy sclerotization of the seventh abdominal segment is continuous without any membranous breaks. The seventh sternite is shaped like an inverted M. On the distal margin of the seventh sternite, a deep, concave indentation, which extends more than three-quarters the length of the sternite, is present. The eighth sternite is present as an ovoid, heavily sclerotized plate whose surface is wrinkled. The distal margin of the plate extends ventrad of the papillae anales. The ostium bursa is located in the membrane between eighth sternite and the papillae anales. The opening is adjacent to the distal margin of the eighth sternite. The ductus bursa is equal to or longer than the length of the seventh tergite. The ductus bursa is tube shaped and straight. A rounded bulge is present at the midpoint of the ductus bursa on the left side. The ductus bursa is not sclerotized. The ductus seminalis arises from the lateral, left side of the ductus bursa. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa.

These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner. The width of the triangular projections is greater than or equal to the length of the projection.

Variation. Only one female specimen was available from this species. The specimen is congruent with the original holotype.

Biology. Unknown.

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Distribution. Based on label data, this species has been collected from Mexico,

Guatemala, and Costa Rica.

Lycomorpha concolor Scott New Species

Figs. 5-2B,C (female); Figs. 5-2D,E (male).

Material examined. Type material: Holotype: AZ: Cochise Co., Chiricahua

Mountains, East Turkey Creek, 6400’, 6.vii.1966, J.G. Franclemont, CUIC. Paratypes:

UNITED STATES: Arizona: Cochise Co., Chiricahua Mountains, East Turkey Creek,

6400’, 6.vii.1966, J.G. Franclemont, CUIC; Santa Cruz Co., Santa Rita Mts., Madera

Canyon, 4880’, 11.vii.1960, CUIC (1 female, CHS-PC082); Santa Cruz Co., Santa Rita

Mts., Madera Canyon, 5800’, 8.vii.1960, CUIC (4 females, 1 male); Santa Cruz Co.,

Santa Rita Mts., Madera Canyon, 4880’, 12.vii.1960, CUIC (1 female (CHS-PC081), 2 males); Santa Cruz Co., Santa Rita Mts., Madera Canyon, 4880’, 10.vii.1960, CUIC (1 female). Specimen preparations. All material from USNM unless otherwise noted:

UNITED STATES: Arizona: Cochise Co., Chiricahua Mt’s., Pinery Canyon, Upper

Camp, 4.vii.1956, LACM (1 male, CHS059); Cochise Co., Chiricahua Mt’s., Pinery

Canyon, Upper Camp, 6.vii.1956, Collected by Lloyd M. Martin, John A. Comstock, and

William A. Rees, LACM (1 female, CHS060); Cochise Co., Chiricahua Mts., 1-7.vii (1 female, CHS090, USNM 127,711); Chircahua Mts., July, A. Twomey, Carn. Mus. Acc.

12518, CMNH (1 male, CHS085); Santa Cruz Co., Madera Cyn., 15.vii.1972, Joseph

Cicero Coll. (1 female, CHS-PC076). New Mexico: Catron Co., Bursum Camp, 18 miles E. Alma, 9000ft., 11.vii.1961, F., P. & J. Rindge, AMNH (1 male, CHS086).

Etymology. The species is named in reference to the similarity between its wing color patterns and those of L. fulgens. In museum collection, these two species are generally found in the same drawer, and both will be labeled as L. fulgens.

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Diagnosis. The males and females of L. concolor can be confused with the L. ptychoglene and the darker color forms of L. fulgens and L. regulus. Lycomorpha concolor can be separated from L. ptychoglene based on size and geographic distribution. The forewing of male L. concolor is five millimeters longer than that of L. ptychoglene. The female forewing of L. concolor is two millimeters longer. Furthermore,

L. concolor and L. ptychoglene are geographically isolated from each other.

Lycomorpha concolor occurs in southeastern Arizona and is a stray into southwestern

New Mexico. Lycomorpha ptychoglene occurs from Mexico to Guatemala.

The geographic distributions of L. concolor, L. fulgens, and L. regulus overlap, and the size difference among the three species is not as pronounced. However, the gena of L. concolor is reduced unlike L. fulgens and L. regulus, which both possess a well-developed gena. In addition, characters from the male and female genitalia can be used to separate the L. concolor from L. fulgens and L. regulus. A narrow, rounded triangle shaped phallic sclerite is found in L. concolor. In L. fulgens, the phallic sclerite is absent or highly reduced. The heavily sclerotized, spine-like cornutus occurs at the distal end of the primary lobe of the vesica of L. concolor, but the spine-like cornutus of

L. fulgens occurs at the distal end of the membranous projection that arises from the primary lobe of the vesica. The processus basalis of the costa present in L. concolor occurs as a short dorso-ventrally flattened lobe, whereas in L. fulgens the processus basalis is present as an elongate spine that tapers to a point. Lycomorpha concolor has a spade-shaped apex of the uncus that tapers to a point. However, the apex of the uncus in L. fulgens is finger-like and does not taper to a point. From the female genitalia, a deep, concave indentation occurs in the distal margin of the seventh sternite

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of L. concolor, whereas the margin of the seventh sternite of L. fulgens is without any noticeable indentations.

The basiphallus of L. concolor is inflected dorsally at the distal end curving away from the venter, whereas the basiphallus of L. regulus is straight. On the vesica of L. concolor, the membranous projection arises from the right side of the primary lobe, midway down the length. However, in L. regulus the projection is found at the distal end of the primary lobe of the vesica at the right apex. A processus basalis of the costa is absent in L. regulus. In addition, the apex of the uncus in L. regulus is ovoid (teardrop shaped) not spade shaped as in L. concolor. Lycomorpha concolor retains an eighth sternite in the female genitalia. However, the eighth sternite is absent in L. regulus.

Description. Male habitus (Fig. 5-2D): Head: The head, antennal scales, and labial palps are sepia (119) to hair brown (119A) colored. The antennae are serrate.

There is no fusion among the labial palp segments. The gena is reduced and cannot be seen as a continuous band around the eye when viewing the head in profile. The genal scales are salmon (106) colored. Thorax: The tegulae, patagia, and dorsal mesothorax are burnt orange (116) to flesh ocher (132D) colored. The dorsal metathorax, all of the ventral thorax, and the legs are sepia (119) to hair brown (119A) colored. Forewing:

Length = 16.1mm (N = 10); M2 and M3 arise separately from the discal cell. Dorsal surface: The wing is Pratt’s rufous (140) to flesh ocher (132D) colored. The fringe of the wing is sepia to hair brown colored. Ventral surface: The wing is Pratt’s rufous (140) to salmon (106) colored. Below the A1 vein the scales are warm buff (118) colored. The fringe is sepia to hair brown colored. A few sepia to hair brown colored scales are scattered along the distal end of the A1 vein. Hindwing: CuA1 and M3 arise separately

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from the discal cell. Dorsal surface: The wing is fuscous (21) to hair brown colored. A warm buff (118) colored band costal band is present. The band arises at the wing and extends along the costal margin. This band tapers to a point adjacent to where the Rs vein terminates at the costal margin. The costal band is broadest in the proximal third of the wing where it stretches from the costal margin to the posterior margin of the discal cell. The posterior margin of the band is smooth except for a single serration that terminates in the proximal third along the posterior margin of the discal cell. Geranium pink (13) colored scales are present along the serration of the band. Ventral surface:

The pattern is identical to the dorsal surface. The wing is fuscous to hair brown colored.

The costal band is Pratt’s rufous (140) to salmon (106) colored. Abdomen: Dorsal, lateral, and ventral scales are sepia (119) to hair brown (119A) colored. A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The pocket is one-third or more the width of A7 sternite. The scales in the pocket are drab

(27) colored. A small, shallow pocket of androconial scales is located in the center of the A8/A9 intersegmental membrane. The scales in the pocket are hair brown (119A) colored. The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite. The pattern of sclerotization on the eighth tergite is heart-shaped. Female habitus (Fig. 5-

2B): Head: The color of the head, antennal scales, labial palps and gena the same as the male. The antennae are simple and ciliate. The fusion of the labial palp segments and the development of the gena are the same as the male. Thorax: The tegulae, patagia, and dorsal mesothorax are flesh ocher (132D) colored. The dorsal metathorax, all of the ventral thorax, and legs are sepia (119) to hair brown (119A) colored.

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Forewing: Length = 14.1 (N = 10); venation the same as the male. Dorsal surface: The

wings are Pratt’s rufous to flesh ocher colored. The fringe is sepia to hair brown colored.

The posterior margin is sometimes narrowly lined with sepia to hair brown colored

scales. Breaks may be present in these scales, or they may be absent entirely. Ventral

surface: The pattern is the same as the dorsal surface. The wing is Pratt’s rufous (140)

to salmon (106) colored. Below the A1 vein the scales are warm buff (118) colored. The

fringe is sepia to hair brown colored. Sepia to hair brown colored scales occur

intermittently along the posterior margin of the wing. Hindwing: Venation is the same as the male. Dorsal surface: The pattern and colors are the same as the male. Ventral surface: the pattern and colors are the same as the male. Abdomen: Color of the dorsal, lateral, and ventral scales the same as the male. Male genitalia: Phallus: The basiphallus is inflected dorsally at the distal end curving away from the venter. When the phallus is viewed from the left side, the phallic sclerite is present as a narrowed, rounded triangle. The vesica is composed of a single primary lobe with an unornamented, membranous projection arising medially on the right side of the vesica.

The distal end of the vesica is ornamented with a single heavily-sclerotized, spine-like cornutus. Genital capsule: The tegumen is square with concave indentations on the

proximal and distal margin. A V-shaped suture that indicates where fusion occurred

between the two halves of the tegumen extends from the proximal to distal margin of

the tegumen. A triangular shaped knob occurs between the two halves of the tegumen.

The knob is contained within the margins of the suture. The apex of the triangle is

oriented toward the proximal margin of the tegumen. Narrow, sclerotized strips of tissue

occur on the lateral margins of the uncus base. The uncus has an S-shaped curve. The

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base of the uncus teardrop shaped and narrows adjacent to the apex of the uncus. The apex of the uncus is spade shaped tapering to a point. The distal end of the costa of the valve is defined by a processus basalis of the costa. The processus basalis of the costa occurs as a short dorso-ventrally flattened lobe. An editum is found at the cephalic end of the costa but does not extend onto the processus basalis of the costa. The valve is triangular shaped with the ventral margin rounded not pointed. The distal end of the valve tapers to a sclerotized spine that is oriented inward and perpendicular to the length of the valve. The juxta is triangular shaped with a conical tip at the distal end.

The juxta is not ornamented with any transparent patches at the proximal margin.

Female genitalia: The heavy sclerotization of the seventh abdominal segment is continuous without any membranous breaks. The seventh sternite is shaped like an inverted M. On the distal margin of the seventh sternite, a deep, concave indentation, which extends more than a quarter the length of the sternite, is present. The eighth sternite is present as a sclerotized bar that fuses with the eighth tergite and the seventh sternite. The ostium bursa is located in the A7/A8 intersegmental membrane below the apex of the concave indention in the seventh sternite. The ductus bursa is shorter than the length of the seventh tergite. It is ribbon shaped proximal to the ostium bursa and the base of the corpus bursa. Between these areas it expends to form a triangular shaped bulb. The ductus bursa is sclerotized proximally to the ostium bursa. The ductus seminalis arises from the ventral side of the ductus bursa. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone glands are

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square shaped with two broad triangular projections arising from each apical corner.

The width of the triangular projections is greater than or equal to the length of the projection.

Variation. The color patterns on the head and thorax of male and female L. concolor collected in Madera canyon (Figs. 5-2C,E) differ from the holotype (Fig. 5-2D).

Both sexes have genal scales that are sepia (119) to hair brown (119A) colored. In both sexes, the only portion of the thorax that is red-orange colored is the tegulae. In the male, the tegulae are burnt orange (116) to flesh ocher (132) colored. In the female, the tegulae are Pratt’s rufous (140) to burnt orange (116) colored. The patagia and dorsal thorax of both sexes are sepia to hair brown colored. The coloring of the ventral thorax and legs is congruent with the holotype. A male L. concolor collected from Redington,

Arizona has a sepia colored terminal band with a hair brown colored fringe. The band is

2mm wide at the apex of the wing. No variation in size was observed for male or female specimens.

Biology. All specimens of L. concolor were collected in July. No other information is known about the biology of this species.

Distribution. With the exception of one specimen, this species is only found in

Arizona. It has been collected from two distinct locations within the state: Chiricahua

Mountain Range (Cochise Co.) and Madera Canyon (Santa Cruz Co.). Both locations occur within the Coronado National Forest. However, the two locations are in separate stands of the forest. One specimen was collected in Catron Co., New Mexico. This county is located on the southwest edge of the state adjacent to Arizona.

Lycomorpha fulgens (Henry Edwards)

Figs 5-2 F-H (female); Figs 5-2I,J and Figs 5-3A,B (male).

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Anatolmis fulgens Henry Edwards 1881b: 116 [type locality: Prescott, Arizona,

USA]

Ptychoglene tenuimargo Holland 1903: 110 [type locality: Arizona, USA]

Material examined. Type material: Holotype of L. fulgens: Anatolmis fulgens

Henry Edwards, Prescott, Arizona, AMNH. Holotype of a synonym of L. fulgens:

Ptychoglene tenuimargo, Type Holland, 3.ix, Arizona Moth Book, Plate XIII, Fig. 17,

CMNH. Specimen preparations. All material from USNM unless otherwise noted:

UNITED STATES: Arizona: Redington, 15.ix.1902, M. Chrissman, Holland Collection,

CMNH (1 female, CHS044); Graham Mt., 28.viii.1933, Parker Lot, LACM (1 male,

CHS083); Pima Co., Tucson, Mt. Lemmon, Elev. 8000’, 28.viii.1965, leg. J.H. Hessel,

AMNH (1 female, CHS088); Cochis Co., Huachuca Mts., Ramsey Canyon, 5400’,

9.vii.1959, LACM (1 female, CHS089). Colorado: Garcia, 12.ix.1898, Holland

Collection, CMNH (1 male, CHS034); Garcia, 3.ix.1898, Holland, CMNH (1 female,

CHS087). New Mexico: Socorro Co., W slope of San Mateo Mts., Wet Red Cyn.,

7500ft., uv, 14.ix.1985, R. Holland, AMNH (1 male, CHS084). UNLABELED: 1 male,

CHS050, USNM 127,706; 1 female, CHS057, USNM 127,708.

Diagnosis. Lycomorpha fulgens can be confused with L. concolor, L. ptychoglene, and L. regulus. As discussed in the diagnosis of L. concolor, characters from the head and male and female genitalia can be used to separate L. fulgens from L. concolor. Characters from the genitalia can be used to separate L. fulgens and L. ptychoglene. The basiphallus of L. fulgens is inflected dorsally at the distal end, whereas there is no curvature of the basiphallus in L. ptychoglene. Lycomorpha fulgens has a single membranous projection that arises from the right side of the primary lobe of

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the vesica. However, L. ptychoglene has three membranous projections that arise from the dorsal surface of the primary lobe of the vesica. On the male genital capsule, the uncus of L. fulgens has an S-shaped curve. The curvature of the uncus of L. ptychoglene is C-shaped. The processus basalis of the costa of L. fulgens is an elongate spine, whereas this structure has a dorso-ventrally flattened trident shape in L. ptychoglene. The juxta of L. fulgens is triangular with a conical tip. Lycomorpha ptychoglene has a rectangular juxta with a triangular indentation in the distal margin. In the female genitalia, the heavy sclerotization of the seventh abdominal is continuous around the segment in L. fulgens. However, membranous breaks in the sclerotization, which are located on the pleurites, occur in L. ptychoglene. The distal margin seventh sternite of L. fulgens is approximately horizontal, whereas in L. ptychoglene to elongate extensions arise from the distal margin and encircle the ostium bursa. The ostium bursa of L. fulgens is located in the A7/A8 intersegmental membrane, but is found in the eighth sternite of L. ptychoglene.

The wing color patterns of L. fulgens and L. regulus can be very similar in the darker specimens of each species. However, the thoracic coloration can be used to separate the two species. The only red colored scales that occur on the thorax of L. fulgens are restricted to the tegulae. Lycomorpha regulus always has red colored scales on the tegulae, patagia, and dorsal mesothorax.

Description. Male habitus (Figs. 5-2I,J and 5-3A,B): Head: Head, antennal scales, labial palps, and gena are Vandyke brown (121) to dark grayish brown (20) colored. The antennae are serrate. No fusion is present among the labial palp segments. The gena is well developed and can be seen as a continuous band around

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the eye that joins with the frons when the head is viewed in profile. Thorax: The tegulae

are geranium (13), flame scarlet (15), burnt orange (116), or warm buff (118) colored.

The anterior margin of the tegulae is Vandyke brown (121) to dark grayish brown (20)

colored. The patagia, dorsal and ventral thorax, and legs are Vandyke brown to dark

grayish brown colored. Forewing: Length = 13.4mm (N = 10); M2 and M3 arise

separately from the discal cell. Dorsal surface: The wing is geranium (12), flame scarlet

(15), burnt orange (116), orange rufous (132D), or warm buff (118) colored. A dusky

brown (19) to fuscous (21) colored terminal band is present. The band arises at the

distal third to quarter of the costal margin and extends to the anal angle. The posterior

margin of the wing is lined with dusky brown to fuscous colored scales. These scales

extend from the wing base to the terminal band. Ventral surface: The color and pattern

is the same as the dorsal surface. Hindwing: CuA1 and M3 arise separately from the

discal cell. Dorsal surface: The wing is dusky brown (19) to fuscous (21) colored. A

geranium (12), flame scarlet (15), burnt orange (116), orange rufous (132D), or warm

buff (118) colored band is present on the costal margin of the wing. The band extends

over three-quarters the length of the wing arising at the wing base. The band is widest at the wing base where it stretches from the costal margin to the posterior margin of the discal cell. The band then tapers to a point. The posterior margin of the band is serrate.

Ventral surface: The color and pattern is the same as the dorsal surface. Abdomen:

Dorsal, lateral, and ventral scales are Vandyke brown (121) to dark grayish brown (20)

colored. A shallow, broad pocket of androconial scales present in the A7/A8

intersegmental membrane. The pocket is one-third or more the width of A7 sternite. The

scales in the pocket are raw umber (23) colored. A small, shallow pocket of androconial

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scales is located in the center of the A8/A9 intersegmental membrane. The scales in the pocket are hair brown (119A) colored. The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite. The pattern of sclerotization on the eighth tergite is rectangular. Female habitus (Figs. 5-2 F-H): Head: Head, antennal scales, and labial palps dusky brown (19) colored. The antennae are serrate. The fusion of the labial palp segments and the development of the gena are the same as the male. The gena is covered with hair brown (119A) colored scales. Thorax: Tegulae geranium (12) to scarlet (14) or chrome orange (16) to ferruginous colored. The anterior margins of the tegulae are dusky brown (19) colored. The patagia, dorsal and ventral thorax, and legs are dusky brown colored. Forewing: Length = 11.9mm (N = 10); venation the same as the male. Dorsal surface: The pattern is the same as the male. The wing is geranium

(12) to scarlet (14) or chrome orange (16) to Pratt’s rufous (140) colored. The terminal band is sepia (119) to hair brown (119A) colored. The scales lining the posterior margin of the wing are also sepia to hair brown colored. Ventral surface: The pattern and color are the same as the dorsal surface except that yellow ocher (123C) to chamois (123D) colored scales are intermixed with the red scales posterior to the A1 vein. In addition, the retinaculum is covered in yellow ocher to chamois colored scales. Hindwing: The venation is the same as the male. Dorsal surface: The pattern is the same as the male.

The wing is dark grayish brown (20) to fuscous (21) colored. The costal band is geranium pink (13) to burnt orange (116) colored. The band is salmon (106) colored at the wing base. Ventral surface: The wing color is the same as the dorsal surface. The costal band is geranium pink colored for the entire length. The length of the band is the

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same as the dorsal surface. The posterior margin of the wing is smooth with no serrations. Abdomen: Scales on the dorsal, lateral, and ventral abdomen dusky brown

(19) colored. Male genitalia: Phallus: The basiphallus is inflected dorsally at the distal end curving away from the venter. When the phallus is viewed from the left side, the phallic sclerite is absent or highly reduced. The vesica is composed of a single primary lobe with a membranous projection arising on the left side of the vesica proximal to the distal margin of the phallus The distal end of the membranous projection of the vesica is ornamented with a single heavily-sclerotized, spine like cornutus. Genital capsule: The tegumen is rectangular with a concave indentation in the distal margin and a rectangular indentation in the proximal margin. An inverted U-shaped suture that indicates where fusion occurred between the two halves of the tegumen extends from the proximal margin of the tegumen adjacent to the distal margin. The apex of the U does not reach the distal margin of the tegumen. A triangular shaped knob occurs between the two halves of the tegumen. The knob is contained within the margins of the suture. The apex of the triangle is oriented toward the distal margin of the tegumen. Narrow, sclerotized strips of tissue occur on the lateral margins of the uncus base. The uncus has an S-shaped curve. The apex of the uncus is fingerlike but does not taper to a point. The distal end of the costa of the valve is defined by a processus basalis of the costa. The processus basalis of the costa occurs as an elongate spine that tapers to a point. No editum is present on the interior surface of the valve. The valve is rectangular shaped with a sinuate ventral margin. At the distal end, the valve tapers to a point that is oriented inward and perpendicular to the length of the valve. The juxta is triangular shaped with a conical tip at the distal end. The juxta is ornamented with a semicircular

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transparent patch at the proximal margin. The vinculum/saccus is U-shaped. Female genitalia: The heavy sclerotization of the seventh abdominal segment is continuous without any membranous breaks. The seventh sternite is parallelogram shaped. This sternite is widest at the cephalic end and narrows at the caudal end. The distal margin of the seventh sternite is approximately horizontal without any deep indentations. The eighth sternite is present as a sclerotized bar that fuses with the eighth tergite and the seventh sternite. The ostium bursa is located in the A7/A8 intersegmental membrane.

The ductus bursa is as long as the seventh sternite. The ductus bursa forms a ovoid bulb shape proximal to the ostium bursa. It tapers to a ribbon shape adjacent to the base of the corpus bursa. The ribbon shaped portion of the ductus bursa is approximately one third of the length of the ductus bursa. No sclerotization is present along the length of the ductus bursa. The ductus seminalis arises from the dorsal side of the ductus bursa. The membrane of the single corpus bursa is smooth and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner. The width of the triangular projections is greater than or equal to the length of the projection.

Variation. As noted in the description, a wide range of color variation is present in both males and females. This variation is thought to be due to fading of the specimens.

Specimens that were collected more recently are closer to the red color seen in the holotypes. The color of the forewings and tegulae lighten in older specimens. No

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correlation was found between the collection location and the color of the specimen. No variation in size was observed for male or female specimens.

Biology. This species has been collected during the months of May through

September. No other information is known about the biology of this species.

Distribution. Lycomorpha fulgens occurs in the southwestern United States in

Arizona and New Mexico. A single individual was collected in the Chihuahua state of

Mexico. This Mexican state is adjacent to the southwestern border of New Mexico.

Lycomorpha grotei (Packard)

Figs. 5-3C,D (female); Figs. 5-3E,F (male).

Anatolmis grotei Packard 1864: 47 [type locality: Pike’s Peak, Colorado, USA]

Lycomorpha palmerii Packard 1872: 84 [type locality: Arizona, USA]

Material examined. Type material: Holotype of L. grotei not examined.

Holotype of a synonym of L. grotei: Lycomorpha palmerii Packard, Arizona, MCZ.

Specimen preparations. All material from USNM unless otherwise noted: UNITED

STATES: Colorado: Denver, 8.viii.1915 (1 female, CHS043, USNM 127,704); Garfield

Co., Grizzly Creek, Glenwood Canyon, 11.viii.1970, O. Shields, S. Ellis, LACM (1 male,

CHS079). Montana: Stillwater Co., 8 road mi SW of Nye, Woodbine C.G., 5300ft.,

28.vii.1982, Julian P. Donahue, LACM (1 male, CHS094). New Mexico: Jemez Mts.,

6600 feet, 15.vii.1915, John Woodgate, CMNH (1 male, CHS080). Texas: Engel Coll’n,

C.M. Acc. 2436, CMNH (1 female, CHS092). Utah: Coll. No. 22, G.E. Wallace, Carn.

Mus. Acc. 11394, CMNH (1 male, CHS035); Provo, 7._.1937, H.P. Chandler, Col. No.

4235, CAS (1 female, CHS091).

Diagnosis. Lycomorpha grotei can be confused with L. pulchra and L. regulus.

Although there is overlap in the range of these three species, L. grotei does not occur in

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California unlike L. pulchra and L. regulus. The wing color pattern of each of these species varies, but certain color characters have been found to be constant within each species. Lycomorpha grotei does not have any dark brown scales lining the posterior margin of the forewing. Both L. pulchra and L. regulus have dark brown scales that extend at least half the length of the posterior margin of the forewing. On the hindwing, the dark brown scales of L. grotei occur intermittently along the posterior margin of the wing, whereas the dark brown scales are continuous to the wing base in L. regulus.

Characters from the male genitalia can also be used to separate these three species. The basiphallus of L. grotei is inflected dorsally at the distal end. No inflection is present on the basiphallus of either L. pulchra or L. regulus. Lycomorpha grotei has no peglike cornuti on the membranous projection arising from the primary lobe of its vesica. Peglike cornuti are present along the entire length of the projection in L. pulchra.

These cornuti only occur on the distal third of the projection in L. regulus. The uncus base of L. grotei occurs within a U-shaped sclerotized plate, whereas the uncus base of both L. pulchra and L. regulus occurs in a V-shaped sclerotized plate. Lycomorpha grotei has a juxta that is ornamented with a semicircular transparent patch at the proximal end. The juxta of L. regulus is ornamented with a triangular transparent patch.

However, no ornamentation is present on the juxta of L. pulchra.

Description. Male habitus (Figs. 5-3E,F): Head: The head and labial palps are cinnamon brown (33) to hair brown (119A) colored. Posterior to the antennae, the head is covered with orange rufous (132C) to salmon (106) to warm buff (118) colored scales.

The antennae are serrate. The antennal scales are dark grayish brown (20) colored. No fusion is present between the labial palp segments. The gena is reduced and cannot be

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seen as a continuous band around the eye when viewing the head in profile. The gena is covered with the same color of scales that are found posterior to the antennae on the head. Thorax: The tegulae, patagia, and dorsal mesothorax are orange rufous (132C) to salmon (106) to warm buff (118) colored. The color is the same as that found on the head posterior to the antennae. The dorsal metathorax is covered with a mixture of scales that are the same color as those found on the tegulae, patagia, and dorsal mesothorax and cinnamon brown (33) to hair brown (119A) colored scales. The entire ventral thorax and legs are covered with cinnamon brown to hair brown colored scales.

The color of the brown scales matches those found on the head. Forewing: Length =

15.2 (N = 10); M2 and M3 arise from the discal separately or are fused and stalk beyond the discal cell. Dorsal surface: The wing is orange rufous to salmon to warm buff colored. The color of the wing is the same as the color of the thorax. Cinnamon brown

(33) to russet (34) colored scales are intermixed. A dark grayish brown (20) to fuscous

(21) colored terminal band is present. The band arises in the distal quarter of the costal margin and extends to the anal angle of the wing. No brown scales extend from the terminal band along the posterior margin of the wing. Ventral surface: The wing is orange rufous to flesh ocher (132D) colored. The costal margin is lined with spectrum orange scales (17). Warm buff (118) colored scales occur posterior to the A1 vein. No cinnamon brown to russet colored scales are present. The color and pattern of the terminal band are the same as the dorsal surface. Hindwing: CuA1 and M3 arise separately from the discal cell. Dorsal surface: The wing is geranium pink (13) to peach red (94) to salmon (106) colored. Salmon (106) to warm buff (118) colored scales extend from the costal margin to the anterior margin of the discal cell. A fuscous (21) to

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dark drab (119B) colored marginal band is present. The band extends from the apex of the wing to the anal angle. The interior margin of the band is sinuate. Fuscous to dark drab colored scales extend from the band intermittently along the posterior margin of the wing to the wing base. Ventral surface: The wing is geranium pink to peach red to salmon colored. The color and pattern of the marginal band are the same as the dorsal surface. Abdomen: Dorsal, lateral, and ventral scales are dark grayish brown (20) colored. A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The pocket is one-third or more the width of A7 sternite. The scales in the pocket are drab gray (119D) colored. A small, shallow pocket of androconial scales is located in the center of the A8/A9 intersegmental membrane. The scales in the pocket are drab gray (119D) colored. The cephalic margin of the eighth sternite is fused to the cephalic margin of the eighth tergite. The pattern of sclerotization on the eighth tergite is heart-shaped. Female habitus (Figs. 5-3C,D): Head: The colors of the head, labial palps, and gena are the same as the male. The antennae are simple and ciliate. The antennal scales are natal brown (219A). The fusion of the labial palp segments and the development of the gena are the same as the male. Thorax: The tegulae, patagia, and dorsal mesothorax are orange rufous (132C) to salmon (106) to warm buff (118) colored. The mixture of colors on the dorsal metathorax is the same as the male. The colors of the entire ventral thorax and legs are the same as the male.

Forewing: Length = 15.5mm (N = 10); Venation is the same as the male. Dorsal surface: The colors and patterns are the same as those on the male. Ventral surface:

The colors and patterns are the same as those on the male. Hindwing: CuA1 and M3 arise from the discal separately or are fused and stalk beyond the discal cell. Dorsal

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surface: The colors and patterns are the same as those on the male. Ventral surface:

The colors and patterns are the same as those on the male. Abdomen: The scales on the dorsal, lateral, and ventral abdomen are natal brown (219) colored. Male genitalia:

Phallus: The basiphallus is inflected dorsally at the distal end curving away from the venter. When the phallus is viewed from the left side, a narrow, rounded triangular shaped phallic sclerite is present. The vesica is composed of a single primary lobe with a membranous projection present. The membranous projection arises from the distal end of the primary lobe at the upper right apices. The entire length of the membranous projection is ruggose. A heavily sclerotized, spine-like cornutus is present on the distal end of the primary lobe adjacent to the base of the membranous projection. Genital capsule: The tegumen is trapezoidal shaped. It is widest at the distal margin. A rectangular indentation is present in the proximal margin. A V-shaped suture that indicates where fusion occurred between the two halves of the tegumen arises from the midpoint of the tegumen and extends to the distal margin. A triangular knob is contained within the margins of the suture. The apex of the triangle is oriented toward the proximal margin of the tegumen. The uncus base occurs within a U-shaped sclerotized plate. The uncus has an S-shaped curve. The base of the uncus is spade shaped and tapers to a point before the first curve. The apex of the uncus is ovoid (tear drop shaped) tapering to a point. The distal end of the costa of the valve is defined by a membranous break that occurs at the midpoint of the dorsal edge of the valve. The break occurs as a concave indentation in the straight ventral margin of the valve. No processus basalis of the costa is present on the valve. An editum is located on the cephalic end of the costa.

The valve has a pentagon shape. The apex of the pentagon occurs on the ventral

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margin of the valve and is rounded. Distally, the valve tapers to a sclerotized point that is oriented inward and perpendicular to the length of the valve. The juxta is triangular shaped with a conical tip at the distal end. The juxta is ornamented with a semicircular transparent patch at the proximal margin. Female genitalia: The heavy sclerotization of the seventh abdominal segment is continuous without any membranous breaks. The seventh sternite is shaped like an inverted M. On the distal margin of the seventh sternite, a deep, concave indentation, which extends more than a quarter the length of the sternite, is present. The eighth sternite is absent. The ostium bursa is located in the

A7/A8 intersegmental membrane below the apex of the concave indention in the seventh sternite. The ductus bursa is shorter than the length of the seventh sternite.

The ductus bursa is tube shaped. The tube undergoes a helical curve before joining the base of the corpus bursa. Sclerotization is present on the ductus bursa but not proximal to the ostium bursa. The sclerotization is separated from the ostium bursa by a membranous break. The ductus seminalis arises from the lateral, right side of the ductus bursa. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone gland is square shaped with two, narrow triangular projections arising from each apical corner. The width of the triangular projections is less than the length of the projections.

Variation. As noted in the description, a wide range of color variation is present in both males and females. This variation is thought to be due to fading of the specimens.

Specimens that were collected more recently are a darker orange red color. The color of

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the forewings and the thorax lighten in older specimens. As the wing color lightens the light brown scales that are intermixed on the forewing become more apparent making the wings appear gray. No correlation was found between the collection location and the color of the specimen. No variation in size was observed for male or female specimens.

Biology. From the label data, this species flies from July through September.

Species have been collected from all of the months within this time period. It is unknown if this represents a single or multiple flights. No other information is known about the biology of L. grotei.

Distribution. This species is found in the western United States. It has been collected from the panhandle of Texas north to Montana. The western edge of its range is Arizona and Utah. The eastern edge of the range is Colorado and New Mexico. The range of this species does not extend into California. Specimens collected in California that have been labeled as L. grotei are representatives of either L. pulchra or L. regulus.

Lycomorpha miniata Packard

Fig. 5-3G (female); Fig. 5-3H (male).

Lycomorpha miniata Packard 1872: 84 [type locality: Southern California, USA]

Material examined. Type material: Holotype of L. miniata: Lycomorpha miniata

Packard, Southern California, 933, MCZ. of L. miniata: Lycomorpha miniata

Packard, Southern California, 933, MCZ. Specimen preparations. All material from

USNM unless otherwise noted: UNITED STATES: Colorado: Jeff. Co., Plainview,

7000-8000ft., 9-14.vii.1922, Brooklyn Museum (1 male, CHS069, USNM 127,710);

Larimer Co., 10mi. W Loveland, Big Thompson Cyn., 7.viii.1973, J. Powell Collr., UCB

(1 female, CHS138). New Mexico: Jemez Springs, Alt. 8500, 15.viii.1928, Marloff

Coll’n, Carn. Mus. Acc. 8992, CMNH (1 male, CHS070); Jemez Springs, 14.vii, Engel

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Coll., Carn. Mus. Acc. 13257, CMNH (1 female, CHS137); Colfax Co., Philmont Scout

Ranch, Lovers Leap, 18.vii.1973, Joseph Cicero Coll., LACM (1 female, CHS139).

South Dakota: Hill City, Black Hills, Joe Dollar Gulch, 3.viii.1964, D.C. Feruson (1 male, CHS038, USNM 127,702); Lawrence Co., Custer Peak, 6500-6794ft., 30.vii.1950,

F. and P. Rindge, AMNH (1 male, CHS071). Wyoming: Crook Co., Devils Tower N.M.,

4250ft., 6.vii.1962, F., P., and M. Rindge, AMNH (1 female, CHS051).

Diagnosis. L. miniata can be confused with L. pholus and L. texanus. Although there is overlap of the ranges of L. miniata and L. pholus, characters from the wing color patterns and the female genitalia can be used to separate these to species. On the dorsal surface of the forewing of L. miniata, the red-orange color of the basal half extends over half the length of the wing. This color extends less than half the length of the wing in L. pholus. In addition, the dark scales that extend from the distal half of the wing along the posterior margin cannot be seen without a microscope in L. miniata. This band of scales can be viewed with the naked eye in L. pholus. Furthermore, in L. miniata, these scales extend only a third of the length of the basal half of the wing. In L. pholus, they extend adjacent to the wing base. On the ventral surface of the forewing in

L. miniata, no dark colored scales extend along the posterior margin of the basal half of the wing unlike L. pholus. On the female genitalia, the seventh sternite of L. miniata is rectangular with the length of the rectangle perpendicular to the body, whereas this sternite is goblet shaped in L. pholus. In addition, the posterior margin of the seventh sternite has a shallow concave indentation in L. miniata. However, this margin is approximately horizontal in L. pholus.

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Although both L. miniata and L. texanus are restricted to the western United

States, their ranges are isolated. L. texanus occurs in the Edwards Plateau and Rolling

Plains regions of Texas. No records are available for L. miniata being found in Texas.

The same two characters of the female genitalia used to separate L. miniata and L. pholus can be used to separate L. miniata from L. texanus. The seventh sternite is goblet shaped in L. texanus, and the posterior margin of the seventh sternite is crenellated.

Description. Male habitus (Fig. 5-3H): Head: Scales on the head and labial palps are sepia (119) colored. The antennae are serrate. The antennal scales are sepia colored with cobalt (68) iridescence. No fusion is present between the labial palp segments. The gena is well developed and can be seen as a continuous band around the eye that joins with the frons when the head is viewed in profile. The gena is covered in sepia colored scales. A row of flame scarlet (14) to chrome orange (15) colored scales are present on the posterior margin of the gena. Thorax: The tegulae are flame scarlet (14) to chrome orange (15) colored. The patagia and dorsal and ventral thorax are sepia (119) colored. The leg scales are sepia colored with cobalt (68) iridescence.

Forewing: Length = 13.9mm (N = 10); M2 and M3 arise separately from the discal cell.

Dorsal surface: The basal half of the wing is flame scarlet (14) to chrome orange (15) colored. The external margin of the basal half of the wing tapers to a point where the

CuA1 vein arises from the discal cell. Anterior and posterior to this point, the external margin is bent convexly toward the wing margin. The distal half of the wing is dusky brown (19) colored. A narrow band of dusky brown colored scales extends from the distal half of the wing along the posterior margin. These scales are difficult to see

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without a microscope. The scales extend one-third the length of the posterior margin of the basal half of the wing. Ventral surface: The basal half of the wing is flame scarlet

(14) to orange rufous (123C) colored. Chrome orange (16) colored scales line the costal margin of the basal half of the wing. The retinaculum is also covered with chrome orange colored scales. Flesh ocher (132D) colored scales are found posterior to the A1 vein. The external margin of the basal half of the wing is serrate. The distal half of the wing is dusky brown colored. No dusky brown colored scales extend along the posterior margin of the basal half of the wing. Hindwing: CuA1 and M3 arise separately from the discal cell. Dorsal surface: The wing is dusky brown (19) to dark grayish brown (20) to fuscous (21) colored. The adbasal region is geranium pink (13) colored. Within this region, a mixture of burnt orange (116) and warm buff (118) colored scales are found between the costal margin and the anterior margin of the discal cell. The posterior margin of this region is sinuate. Ventral surface: The basal half of the wing is geranium pink colored. The external margin of the basal half of the wing is sinuate. The distal half of the wing is fuscous (21) colored. A narrow band of fuscous colored scales extend from the distal half of the wing along the posterior margin of wing to the wing base.

Abdomen: Dorsal, lateral, and ventral scales are sepia (119) colored. A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The pocket is one-third or more the width of A7 sternite. The scales in the pocket are fuscous (21) colored. The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite.

The pattern of sclerotization on the eighth tergite is rectangular. Female habitus (Fig.

5-3G): Head: The colors of the head, antennal scales, labial palps, and gena are the

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same as the male. The antennae are serrate. The fusion of the labial palp segments

and the development of the gena are the same as the male. Thorax: The tegulae are

flame scarlet (15) to chrome orange (16) colored. The colors of the patagia, dorsal and

ventral thorax, and legs are the same as the male. Forewing: Length = 13.5mm (N =

10); Venation the same as the male. Dorsal surface: The colors and patterns are the

same as those on the male. Ventral surface: The basal half of the wing is geranium pink

(13) to orange rufous (132C) colored. The costal margin of the basal half is lined with

chrome orange (16) colored scales. Flesh ocher (132D) colored scales occur posterior

to the A1 vein in the basal half of the wing. The external margin of the basal half of the wing is the same as the male. The distal half of the wing is dusky brown (19) colored.

No dusky brown scales occur on the posterior margin of the basal half of the wing.

Hindwing: The venation is the same as in the male. Dorsal surface: The colors and patterns are the same as those on the male. Ventral surface: The colors and patterns are the same as those on the male. Abdomen: The color of the scales on the dorsal, lateral, and ventral abdomen are the same as the male. Male genitalia: Phallus: The basiphallus is straight with no inflection. When the phallus is viewed from the left side, the phallic sclerite is present as a flattened lobe, which extends as a separate structure dorsad of the vesica. The vesica is composed of a single primary lobe with a membranous projection arising on the left side of the vesica proximal to the distal margin of the phallus. The distal end of the membranous projection of the vesica is ornamented with a single heavily-sclerotized, spine like cornutus. Genital capsule: The

tegumen is rectangular with a concave indentation that extends half the length of the

distal margin. The indentation is centered in the margin. The proximal margin of the

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tegumen is crenellated. A V-shaped suture that indicates where fusion occurred

between the two halves of the tegumen extends for three-quarters of the length of the

tegumen. The sutures extend from the distal margin anteriorly. A circular knob is

contained within the margins of the suture and arises adjacent to the distal margin of the

tegumen. The uncus base occurs within a rectangular sclerotized plate. The uncus has

a C-shaped curve. The apex of the uncus is ovoid (teardrop shaped) tapering to a point.

The distal end of the costa of the valve is defined by a triangular, membranous break

that occurs beyond the midpoint of the dorsal edge of the valve. Beyond the

membranous break, the dorsal edge of the valve occurs below the costa. No processus

basalis of the costa is present on the valve. No editum is present on the internal face of the valve. The valve is teardrop shaped. The ventral margin of the valve tapers distally and forms a lobe that is perpendicular to the length of the valve. The dorsal margin also tapers to a lobe distally. In addition, a sclerotized spine arises from the lobe formed from

the dorsal margin of the valve. The spine is oriented inward and perpendicular to the

length of the valve. The juxta is a flat square sclerotized plate with a concave

indentation in the distal margin. The juxta is not ornamented with any transparent

patches at the proximal margin. Female genitalia: The heavy sclerotization of the

seventh abdominal segment is interrupted by narrow membranous breaks that occur on

the pleurites. The seventh sternite is rectangular shaped with the length of the rectangle

perpendicular to the body. On the distal margin of the seventh sternite, a shallow,

concave indentation, which extends less than a quarter the length of the sternite, is

present. The eighth sternite is present as a heavily sclerotized plate. The ostium bursa

is located in the eighth sternite as a broad, horizontal slit that is centered in the sternite.

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The opening extends over half the length of the sternite. The ductus bursa is shorter than the seventh sternite and ribbon like. The ductus bursa is not curved but a bulge occurs on the right side of the ductus bursa. No sclerotization is present on the ductus bursae. The ductus seminalis arises from the ventral side of the ductus bursa. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner. The width of the triangular projections is greater than or equal to the length of the projection.

Variation. Almost no variation in size and color was observed among the male or female specimens examined.

Biology. This species has been collected in July and August based on the label data. No other information is available on its biology.

Distribution. This species occurs only in the western United States. It ranges from

Montana and South Dakota in the north to New Mexico in the south. Although the collection locality on the type specimen of L. miniata is Southern California, no other records are available for this species from California.

Lycomorpha morelosia (Schaus)

Fig. 5-3I (female); Fig. 5-3J (male).

Propyria morelosia Schaus 1925: 19 [type locality: Morelos, Mexico]

Material examined. Type material: Holotype of L. morelosia: Propyria morelosia

Schaus, Morelos, Mexico, USNM. Specimen preparations. All material from USNM unless otherwise noted: MEXICO: Nayarit: Zacualpan, ix.1914, Dognin Collection, (1

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male, CHS196, USNM 127,721). Federal District: Barranca, Mixcoac, 20.viii.1917,

C.C. Hoffman, AMNH (1 female, CHS247).

Diagnosis. This species can be confused with L. neomexicanus and L. normani.

Lycomorpha morelosia is geographically isolated from L. neomexicanus. Lycomorpha morelosia is only known to occur in Mexico, whereas L. neomexicanus has only been found in New Mexico. Although both L. morelosia and L. normani occur in Mexico, the color pattern of the hindwing can be used to separate the two species. The red scales on the hindwing of L. morelosia extend over three-quarters the length of the costal margin of the hindwing and arise adjacent to wing base. However, the pink scales present on the hindwing of L. normani do not extend more than half the length of the costal margin and arise from the base of the wing. Furthermore, the thorax of L. morelosia is entirely covered with dark brown scales, whereas the tegulae of L. normani are covered in red-orange scales. The remainder of the thorax in L. normani is dark brown.

Description. Male habitus (Fig. 5-3J): Head: Scales on the head, antennae, labial palps, and gena are Vandyke brown (221) colored. The antennae are serrate. No fusion is present between the labial palp segments. The gena is well developed and can be seen as a continuous band around the eye that joins with the frons when the head is viewed in profile. Thorax: Tegulae, patagia, dorsal and ventral thorax, and legs

Vandyke brown (221) colored. Forewing: Length = 12mm (N=2); M2 and M3 fused and stalk beyond the discal cell. Dorsal surface: The wing is sepia (119) to Vandyke brown

(121) colored. The costal vein is lined with sepia colored scales. A band of geranium

(12) and scarlet (14) colored scales is present parallel to the costal margin of the wing.

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The band extends over three-quarters the length of the wing beginning just beyond the wing base. In the proximal third of the wing, the band extends from directly posterior to the costal vein to just above the A1 vein. The band tapers to the width of the discal cell beyond the point where the R1 vein arises from the discal cell. Ventral surface: The pattern is identical to the dorsal surface. The costal vein is lined with sepia (119) colored scales. The anterior half of the wing is fuscous (21) colored. The posterior half of the wing is olive brown colored (28). The band that is parallel to the costal vein is geranium pink (13) colored. The shape of the band is identical to the dorsal surface. Hindwing:

CuA1 and M3 fused and stalk beyond the discal cell. Dorsal surface: The hindwing is primarily sepia (219) colored. A band of geranium pink (13) to scarlet (14) colored scales extends from just beyond the wing base to outside of the discal cell. The band extends from the posterior costal vein edge to the posterior edge of the discal cell.

Ventral surface: The pattern is identical to the dorsal surface. The ventral surface is primarily fuscous (21) colored. The band parallel to the costal margin is geranium pink

(13) colored. Abdomen: Dorsal, ventral, and lateral scales Vandyke brown (221). The anterolateral process is present as a flattened sclerotized lobe (Fig. 5-13B). The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite. The pattern of sclerotization on the eighth tergite is rectangular. Female habitus (Fig. 5-3I): Head:

Scales on the head, antennae, labial palps, and gena are dusky brown (19) colored.

The antennae are simple and ciliate. The fusion of the labial palp segments and the development of the gena are the same as the male. Thorax: Tegulae, patagia, dorsal and ventral thorax, and legs dusky brown (19) colored. Forewing: Length = 12mm

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(N=3); venation the same as in the male. Dorsal surface: The dorsal surface of the

forewing is sepia (119) colored. A geranium (12) and scarlet (14) to crimson (108) band

runs parallel to the costal vein. The band occurs from just beyond the wing base to

outside of the discal cell where the Rs5 and Rs4 veins begin to fork. In the basal half of

the wing, the band extends from directly below the costal vein to above the A1 vein. In the distal half of the wing, the band narrows to the width of the discal cell. Ventral surface: The ventral surface is primarily geranium pink (13) colored. The costal vein is

lined with Vandyke brown (221) colored scales. A hair brown (119A) colored terminal

band is present. The hair brown colored scales extend from the terminal band along the

posterior margin of the wing to the wing base. The terminal band is 1.5-2mm wide

proximal to the apex and expands to 3mm wide proximal to the anal angle. The band

does not extend above the A1 vein along the posterior margin of the wing. Geranium

pink (13) scales are also found below the proximal half of the A1 vein. The A1 vein is

lined with spectrum orange (17) colored scale. A spectrum orange discal spot is

present. The Sc vein is lined with chrome orange (16) colored scales. Hindwing:

Venation is the same as in the male. Dorsal surface: The wing is crimson (108) to

geranium pink (13) colored. A sepia (219) colored marginal band extends from the apex

of the wing to the wing base on the posterior margin of the wing. The band is 2mm wide.

The internal margin of the band is sinuate. Sepia colored scales are found intermixed

with the crimson to geranium pink colored scales on the CuA2 vein and between the

internal margin of the marginal band and the posterior margin of the discal cell. Ventral

surface: The pattern is the same as the dorsal surface. The wing is geranium pink (13)

colored. The marginal band is hair brown (119A) colored. Hair brown colored scales are

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found intermixed with geranium pink colored scales between the internal margin of the marginal band and the posterior margin of the discal cell. Spectrum orange (17) colored scales line the R1 vein. Abdomen: Dorsal, lateral, and ventral scales dusky brown (20) colored. The anterolateral process is the same as the male. Male genitalia: Phallus:

The basiphallus is straight with no inflection. When the phallus is viewed from the left side, a narrowed, rounded triangular shaped phallic sclerite is present. The vesica is composed of a single primary lobe with two membranous projections arising from the dorsal surface of the vesica. A heavily sclerotized, spine-like cornutus is present on the distal end of the primary lobe Genital capsule: The tegumen has an elongate, rectangular shape. The lateral margins of the tegumen are bent concavely. A triangular indentation is present in the proximal margin. A straight suture with an ovoid bulge medially that indicates where fusion occurred between the two halves of the tegumen runs from the cephalic to caudal end of the tegumen. Narrow, sclerotized strips of tissue occur laterally on the uncus base. The uncus has an S-shaped curve. The apex of the uncus is fingerlike but does not taper to a point. The distal end of the costa of the valve is defined by a processus basalis of the costa. The processus basalis of the costa occurs as an elongate spine that tapers to a point. An editum is located on the cephalic end of the costa and does not extend onto the processus basalis of the costa. The valve is rectangular shaped and all of the edges are rounded. The dorsal margin of the valve is curved convexly. The juxta occurs as a flat, sclerotized plate that is rectangular with a triangular indentation in the distal margin. The juxta is ornamented with a triangular transparent patch at the proximal margin. Female genitalia: The heavy sclerotization of the seventh abdominal segment is continuous without any membranous breaks. The

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seventh sternite is shaped like an inverted M. On the distal margin of the seventh sternite, a deep, concave indentation, which extends more than a quarter the length of the sternite, is present. The eighth sternite is present as a sclerotized bar that fuses with the eighth tergite and the seventh sternite. The ostium bursa is located in the A7/A8 intersegmental membrane below the apex of the concave indention in the seventh sternite. The ductus bursa is longer than the seventh sternite. The ductus bursa is ribbon like, sclerotized for its entire length, and is not curved. The ductus bursa is sclerotized proximally to the ostium bursa. Two corpus bursae that are connate and arise from the ductus bursa are present. The membrane of the corpus bursae is smooth. The larger of the two bursae is ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa.

These two signa occur on opposite sides of this bursa. The ductus seminalis arises from the larger corpus bursa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner. The width of the triangular projections is greater than or equal to the length of the projection.

Variation. Almost no variation in size and color was observed among the male or female specimens examined.

Biology. Unknown.

Distribution. This species occurs within Mexico. Based on the label data, this species has been collected from the Federal District and the Mexican states of Nayarit and Morelos.

Lycomorpha neomexicanus Scott New Species

Fig. 5-4A (male).

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Material examined. Type material: Holotype: NM: Union Co., Johnson Mesa,

10mi N Folsom, Sta. 448, 21.vi., leg. H.K. and M.A. Clench, 1977, C.M. Acc. 29258,

CMNH, CHS253, male. Specimen preparations. This species is represented by one specimen from CMNH.

Etymology. This species is named for the state where the only representative was collected.

Diagnosis. Lycomorpha neomexicanus can be confused with L. morelosia and L. normani. This species is geographically isolated from L. morelosia as noted in the diagnosis of that species. The locality of L. neomexicanus can also be used to separate it from L. normani. Lycomorpha neomexicanus occurs in New Mexico, whereas L. normani is found in Mexico and Costa Rica. The color pattern of the hindwing can also be used to separate L. neomexicanus and L. normani. Although the orange-red costal band on the hindwing of L. neomexicanus extends three-quarters the length of the costal margin, the geranium pink scales in the adbasal region of L. normani extend only half the length of the costal margin.

Description. Male habitus (Fig. 5-4A): Head: The head and labial palps are hair brown (119A) colored. The antennae are serrate. The antennal scales are dark grayish brown (20) colored with turquoise green (64) iridescence. There is no fusion among the labial palp segments. The gena is well developed and can be seen as a continuous band around the eye that joins with the frons when the head is viewed in profile. The gena is covered with light drab (119C) colored scales. Thorax: The tegulae are spectrum orange (17) colored. The patagia are hair brown (119A) colored with spectrum orange (17) and orange yellow (18) colored scales occurring on the outer third of the

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patagia. The dorsal mesothorax is sepia (119) colored with hair brown (119A) colored

scales around the margins. The dorsal metathorax is sepia colored. The ventral thorax

and legs are hair brown (119A) colored. Forewing: Length = 13mm (N = 1); M2 and M3

are fused and stalk beyond the discal cell. Dorsal surface: The wings are dark grayish brown (20) colored. A warm buff (118) colored band arise at the wing base and extends along the costal margin. The band tapers to a point beyond the discal cell where the

Rs3+Rs4 fork from Rs2. At the wing base the band stretches from the costal margin to

the posterior margin of the wing. It tapers continuously from the wing base. The

posterior margin of the band is burnt orange (116) colored and smooth. Burnt orange

scales are intermixed with dark grayish brown scales between the A1 vein and the

posterior margin of the discal cell. Ventral surface: The wing is orange rufous (123C)

colored. The costal vein is lined with spectrum orange (17) colored scales. Posterior to

the A1 vein, the wing scales are buff (124) colored. A dark grayish brown (20) terminal

band is present. The band is 2mm wide. Hindwing: CuA1 and M2 arise separately from

the discal cell. Dorsal surface: The wing is dark grayish brown (20) colored. A costal

band extends from the wing base to the distal margin of the discal cell. The anterior half

of the band is warm buff (118) colored. The posterior half of the band is orange-rufous

(123C) colored. The band tapers to a point. The posterior margin of the band is serrate.

Ventral surface: The pattern is the same as the dorsal surface. The wing is dark grayish

brown colored. The costal band is burnt orange (116) colored. Abdomen: Dorsal,

lateral, and ventral scales are sepia (119) colored. A shallow, broad pocket of

androconial scales present in the A7/A8 intersegmental membrane. The pocket is one-

third or more the width of A7 sternite. The scales in the pocket are dark drab (119B)

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colored. A small, shallow pocket of androconial scales is located in the center of the

A8/A9 intersegmental membrane. The scales in the pocket are hair brown (119A)

colored. The cephalic margin of the eighth sternite is fused to the cephalic margin of the

eighth tergite. The pattern of sclerotization on the eighth tergite is heart-shaped. Male

genitalis: Phallus: The basiphallus is straight with no inflection. When the phallus is

viewed from the left side, a narrow, pointed triangular shaped phallic sclerite is present.

The vesica is composed of a single primary lobe with a membranous projection present.

The membranous projection arises from the distal end of the primary lobe at the upper

right apices. The distal third of the projection is ornamented with peglike cornuti. The

proximal two-thirds of the projection are ruggose. A heavily sclerotized, spine-like

cornutus is present on the distal end of the primary lobe adjacent to the base of the

membranous projection. Genital capsule: The tegumen is rectangular shaped. The

length of the rectangle is perpendicular to the length of the body. A rectangular

indentation is present in the proximal margin. A V-shaped suture that indicates where

fusion occurred between the two halves of the tegumen arises adjacent to the proximal

margin of the tegumen and extends to the distal margin of the valve. A triangular knob is

contained within the margins of the suture. The apex of the triangle is oriented toward

the proximal margin of the tegumen. The uncus base occurs within a V-shaped

sclerotized plate. The uncus has an S-shaped curve. The base of the uncus is teardrop shaped and tapers to a point proximal to the first curve of the uncus. The apex of the uncus is ovoid (tear drop shaped) tapering to a point. The distal end of the costa of the valve is defined by a membranous break that occurs at the midpoint of the dorsal edge of the valve. No processus basalis of the costa is present on the valve. An editum is

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located on the cephalic end of the costa. The valve is rectangular shaped with a ventral margin that is curved convexly. Distally, the valve tapes to a sclerotized point that is oriented inward and perpendicular to the length of the valve. The juxta is triangular shaped with a conical tip at the distal end. The juxta is ornamented with a triangular transparent patch at the proximal margin.

Variation. Only one specimen was available for this species.

Biology. Unknown.

Distribution. The specimen representing this species was collected in New

Mexico in Union County.

Lycomorpha normani (Schaus)

Fig. 5-4B (male).

Propyria normani Schaus 1911: 613 [type locality: Alajuela, Costa Rica]

Material examined. Type material: Holotype of L. normani: Propyria normani

Schaus, Alajuela, Costa Rica, USNM. Specimen preparations. All material from

USNM unless otherwise noted: MEXICO: Morelos: Cuernavaca, vii.1906, Collection

Wm. Schaus (1 male, CHS252, USNM 127,736).

Diagnosis. L. normani can be confused with L. morelosia and L. neomexicanus.

Lycomorpha normani can be separated from L. morelosia based on color and male genitalia characters as described in the diagnosis of L. morelosia. Characters separating L. normani and L. neomexicanus are provided in the diagnosis of L. neomexicanus.

Description. Male habitus (Fig. 5-4B): Head: Scales on the head, antennal, labial palp, and gena are fuscous (21) colored. The antennae are serrate. No fusion is present between the labial palp segments. The gena is well developed and can be seen

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as a continuous band around the eye that joins with the frons when the head is viewed

in profile. Thorax: The tegulae are chrome orange (16) colored with flame scarlet (15)

colored scales intermixed. The patagia, dorsal and ventral thorax, and legs are fuscous

(21) colored. Forewing: Length = 12mm (N = 1); M2 and M3 are fused and stalk beyond

the discal cell. Dorsal surface: The wing is fuscous (21) colored. The distal half of the

costal vein is lined with sepia (119) colored scales. Chrome orange (16) to flame scarlet

(15) colored scales form a band that is parallel to the costal margin. The proximal half of

the costal vein is lined with scales from this band. The band extends from the wing base

to within 1mm of the wing apex. The band is thickest at the wing base. At the wing base,

it stretches from the costal vein towards the posterior margin of the wing. The band is

separated from the posterior margin of the wing by a narrow band of fuscous colored

scales. Beyond this point the band narrows and tapers to a point. The posterior margin

of the band is smooth and angled toward the apex of the wing. Ventral surface: The wing is hair brown (119A) colored. A flame scarlet (15) colored band occurs parallel to the costal margin. The length of this band is the same as the band on the dorsal surface. The width of the band at the wing base is the same as the dorsal surface. The posterior margin of the band is sinuate and angled toward the M2 + M3. Hair brown

scales are intermixed along the posterior margin of the band. Hindwing: CuA1 and M3

fused and stalk beyond the discal cell. Dorsal surface: The wing is burnt umber (22)

colored. The fringe is fuscous (21) colored. A patch of geranium pink (13) colored

scales is present in the adbasal region. The patch extends half the length of the costal

margin. At the wing base, the patch stretches from the costal margin to the posterior

margin of the discal cell. The posterior margin of the geranium pink patch curves

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convexly. Ventral surface: The pattern is identical to the dorsal surface. The wing is fuscous (21) colored. The adbasal patch is flame scarlet (15) colored. Abdomen:

Dorsal, ventral and lateral scales fuscous (21). A narrow pocket of hair-like androconial scales is found in the A7/A8 ventral intersegmental membrane. The pocket is less than one-third the width of the A7 sternite. The androconial scales are cinnamon brown (33) colored. The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite. The pattern of sclerotization on the eighth tergite is rectangular. Male genitalis: Phallus:

The basiphallus is straight with no inflection. When the phallus is viewed from the left side, a narrowed, rounded triangular shaped phallic sclerite is present. The vesica is composed of a single primary lobe with three membranous projections arising from the dorsal surface of the vesica. A heavily sclerotized, spine-like cornutus is present on the distal end of the primary lobe. Genital capsule: The tegumen has an elongate, rectangular shape. The lateral margins of the tegumen are bent concavely. A straight suture with an ovoid bulge medially that indicates where fusion occurred between the two halves of the tegumen runs from the cephalic to caudal end of the tegumen.

Narrow, sclerotized strips of tissue occur laterally on the uncus base. The uncus has an

C-shaped curve. The apex of the uncus is fingerlike and tapers to a point. The distal end of the costa of the valve is defined by a processus basalis. The processus basalis is dorso-ventrally compressed and trident shaped. The tips of the processus basalis are blunt. An editum is located on the base of the processus basalis. The valve is curved at an obtuse angle. Proximally to the genital capsule the valve is rectangular shaped with smooth, uncurved edges. Distal to the curve, the valve is rectangular shaped with

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curved edges. The valve tapers to a sclerotized spine that is oriented inward and perpendicular to the valve. In addition, a sclerotized lobe occurs adjacent to the ventral margin of the spine. The juxta occurs as a flat, sclerotized plate that is rectangular with a triangular indentation in the distal margin. The juxta is ornamented with a triangular transparent patch at the proximal margin.

Variation. Only one male specimen was available from this species. The specimen is congruent with the original holotype.

Biology. Unknown.

Distribution. This species has been collected in Mexico and Costa Rica.

Lycomorpha pholus (Drury)

Figs. 5-4C,D (female); Figs. 5-4E,F (male).

Sphinx pholus Drury 1773: 49 [type locality: New England, USA]

Material examined. Type material: Holotype not examined. Specimen preparations. All material from USNM unless otherwise noted: CANADA: Ontario:

24mi. SE Kenora, ex flowers Meliotus alba, 5.viii.1967, J.R. Powers Collr., UCB (1 male,

CHS134); 20mi. SE Kenora, 30.vii.1967, J.R. Powers Collr., UCB (1 female, CHS125).

Quebec: Ottawa Co., Meach Lake, 1-7.viii (1 male, CHS136, USNM 127,720). UNITED

STATES: : St. Louis Co., 24.ix.1944, Rockwood Res., T.B. Bleulusle Collecter,

SDMNH (1 male, CHS064); St. Louis, 20.ix.1931, Meiners, Collection of Grace H. and

John L. Sperry, AMNH (1 female, CHS072). New York: Big Indian V’y, Catskill Mt’s,

10.viii.1907, R.F. Pearsall (1 female, CHS045, USNM 127,705); Oliveria, 18.vi.1927,

LACM (1 male, CHS133). Virginia: Page Co., Pinnacle, Shenandoah Nat. Park, 3400’,

5.viii.1979, Leg. D.C. Ferguson (1 male, CHS036, USNM 127,701). West Virginia:

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Pendleton Co., Spruce Knob, Elev. 4890’, 5.viii.1960, Milliron and Wallace, CMNH (1 female, CHS074).

Diagnosis. Lycomorpha pholus may be confused with L. atroxantha, L. miniata,

L. pelopia, and L. texanus. Color patterns of the thorax, tibial spur formula, and the geographic distribution can be used to separate L. pholus and L. atroxantha as noted in the diagnosis of the latter species. The geographic distribution and color patterns of the head are useful in separating L. pholus and L. pelopia. Lycomorpha pholus is restricted to North America, whereas L. pelopia has been collected from Mexico and Panama.

The scales on the head and gena of L. pholus are all sepia colored. However, the genal scales of L. pelopia are orange yellow colored. Characters from the forewing color patterns and the female genitalia help to identify L. pholus and L. miniata. Full descriptions of these characters are provided in the diagnosis of L. miniata. Lycomorpha pholus and L. texanus are geographically isolated. Lycomorpha texanus is restricted to the Edwards Plateau and Rolling Plains regions of Texas. The range of L. pholus does not extend into Texas. In addition, a narrow band of dark brown colored scales is present on the posterior margin of L. pholus. These scales extend adjacent to the wing base. A narrow band of dark brown scales also occurs in L. texanus. However, the band does not extend more than one-third the length of the posterior margin of the basal half of the wing.

Description. Male habitus (Fig. 5-4E): Head: Scales on the head, labial palps, and gena are sepia (119) colored. The antennae are serrate. The antennal scales are sepia colored with cobalt (68) iridescence. No fusion is present between the labial palp segments. The gena is well developed and can be seen as a continuous band around

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the eye that joins with the frons when the head is viewed in profile. Thorax: The tegulae are spectrum orange (17) to orange yellow (18) colored. The patagia and dorsal and ventral thorax are sepia (119) colored. The leg scales are sepia colored with cobalt (68) iridescence. Forewing: Length = 13.1mm (N = 10); M2 and M3 arise separately from the discal cell. Dorsal surface: The basal half of the wing is spectrum orange (17) to orange yellow (18) colored. The external margin is sinuate and tapers to a point where the

CuA1 vein arises from the discal cell. The distal half of the wing is sepia (119) colored. A narrow band of sepia colored scales extends along the posterior margin of the basal half of the wing. These scales are adjacent to but do not reach the wing base. Ventral surface: The basal half of the wing is orange yellow colored. The external margin tapers to a point where the CuA1 vein arises. Anterior and posterior to the point, the margin is bent convexly toward the wing base. From the costal margin to the CuA2 vein, the distal half of the wing is sepia to dusky brown (19) colored. Posterior to the CuA2 vein the distal half of the wing is hair brown (119A) colored. Hair brown scales line the posterior margin of the wing and end adjacent to the wing base. Hindwing: The CuA1 and M3 veins arise separate from the discal cell. Dorsal surface: The wing is dusky brown (19) to fuscous (21) colored. The adbasal region is trogon yellow (153) colored. The posterior margin of the adbasal region is orange yellow (18) colored and sinuate.

Ventral surface: The pattern is the same as the dorsal surface. The wing is dark grayish brown (20) to fuscous (21) colored. The adbasal region is orange yellow colored.

Abdomen: Dorsal, lateral, and ventral scales are sepia (119) colored. A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The pocket is one-third or more the width of A7 sternite. The scales in the pocket are

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cinnamon brown (33) colored. The cephalic margin of the eighth sternite extends

adjacent to the cephalic margin of the eighth tergite. There is no fusion between the

sternite and tergite. The pattern of sclerotization on the eighth tergite is rectangular.

Female habitus (Fig. 5-4C): Head: Color of head, antennal scales, labial palps, and

gena the same as the male. Antennae are serrate. The fusion of the labial palp

segments and the development of the gena are the same as the male. Thorax: Color of the tegulae, patagia, dorsal and ventral thorax, and legs same as the males. Forewing:

Length = 14.2mm (N = 10); M2 and M3 arise separately from the discal cell or M2 and M3

are fused and stalk beyond the discal cell. Dorsal surface: The pattern of the wing is the same as the male. The basal half of the wing is spectrum orange (17) to orange yellow

(18) colored. The distal half of the wing and the scales lining the posterior margin of the wing are sepia (119) to dusky brown (19) colored. Ventral surface: The color is the same as in the ventral forewing surface of the male. The external edge of the basal half of the wing is either bent concavely towards the wing base above and below the posterior edge of the discal cell or almost smooth with no distinctive indentations.

Hindwing: Venation is the same as the male. Dorsal surface: The pattern is the same

as the male. The wing is sepia (119) to fuscous (21) colored. The color of the adbasal

region is the same as the male. Ventral surface: The pattern is the same as the male.

The wing is either dusky brown (19) or fuscous colored. The color of the adbasal region

is the same as the male. Abdomen: The color of the scales on the dorsal, lateral, and

ventral abdomen are the same as the male. Male genitalia: Phallus: The basiphallus is

straight with no inflection. When the phallus is viewed from the left side, the phallic

sclerite is present as a flattened lobe, which extends as a separate structure dorsad of

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the vesica. The vesica is composed of a single primary lobe with a membranous projection arising on the left side of the vesica proximal to the distal margin of the phallus. The distal end of the membranous projection of the vesica is ornamented with a single heavily-sclerotized, spine like cornutus. Genital capsule: The tegumen is rectangular with a concave indentation that extends a third of the length of the distal margin. The indentation is centered in the margin. The proximal margin of the tegumen is crenellated. A V-shaped suture that indicates where fusion occurred between the two halves of the tegumen extends for the distal two-thirds the length of the tegumen. A circular knob is contained within the margins of the suture and arises adjacent to the distal margin of the tegumen. The uncus base occurs within a rectangular sclerotized plate. The uncus has a C-shaped curve. The apex of the uncus is ovoid (teardrop shaped) tapering to a point. The distal end of the costa of the valve is defined by a triangular, membranous break that occurs beyond the midpoint of the dorsal edge of the valve. No processus basalis of the costa is present on the valve. No editum is present on the internal face of the valve. The valve is teardrop shaped. The ventral margin of the valve tapers distally and forms a lobe that is perpendicular to the length of the valve.

The dorsal margin also tapers to a lobe distally. In addition, a sclerotized spine arises from the lobe formed from the dorsal margin of the valve. The spine is oriented inward and perpendicular to the length of the valve. The juxta is a flat square sclerotized plate with a concave indentation in the distal margin. The juxta is not ornamented with any transparent patches at the proximal margin. Female genitalia: The heavy sclerotization of the seventh abdominal segment is interrupted by narrow membranous breaks that occur on the pleurites. The seventh sternite is goblet shaped. The distal margin of the

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seventh sternite is approximately horizontal with no significant indentations or crenellations. The eighth sternite is present as a heavily sclerotized plate. The ostium bursa is located in the eighth sternite as a broad, horizontal slit that is centered in the sternite. The opening extends over half the length of the sternite. The ductus bursa is shorter than the seventh sternite. The ductus bursa is ribbon shaped and undergoes a

C-shaped curve. No sclerotization is present on the ductus bursae. The ductus seminalis arises from the ventral side of the ductus bursa. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner.

The width of the triangular projections is greater than or equal to the length of the projection.

Variation. Males of L. pholus collected from have burnt orange (116) colored scales between the anterior margin of the discal cell and the A1 vein on the basal half of the ventral surface of the forewing (Fig. 5-4F). A mixture of geranium pink

(13), chrome orange (16), and peach red (94) colored scales are present on the dorsal surface of the adbasal region of the hindwing of some males collected from Oklahoma and Missouri. On the ventral surface of the hindwing of some males collected from

Oklahoma and Missouri, the adbasal region is spectrum orange (17) and geranium pink colored. Some female L. pholus collected from Oklahoma and Missouri have a mixture of peach red (94) and orange yellow (18) colored scales on the posterior margin of the dorsal surface of the adbasal region of the hindwing (Fig. 5-4D). On the ventral surface

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of the hindwing, the adbasal region of these individuals is composed of a mixture of orange yellow and peach red colored scales. No size variation was observed for males or females.

Biology. The larvae of L. pholus possess a mandibular mola. In addition, they have been raised on the algae Protococcus viridis and the lichen Physcia millegrana

(Wagner et al. 2008). The adults of L. pholus are entirely diurnal (Fullard & Napoleone

2001). The adults have been reported nectar feeding on flowers of Apocynum,

Ericameria nauseosa, and Solidago (Powell & Opler 2009). Based on label data, the adults fly from from June through September.

Distribution. Lycomorpha pholus has the broadest distribution of any of the species within Lycomorpha. This species occurs as far north as Ontario and Quebec in

Canada. The range of L. pholus extends south along the eastern coast of the United

States to . Based on the label data a few specimens have been collected from Florida. However, no other information besides the state name was provided. From the east coast, the range of L. pholus extends west to Colorado. Within Western North

America, L. pholus occurs from Montana south to New Mexico.

Lycomorpha ptychoglene (Hampson)

Fig. 5-4G (female); Fig. 5-4H (male).

Propyria ptychoglene Hampson 1898: 521 [type locality: Vera Cruz, Mexico]

Lithosia aequalis Walker 1854: 511 [type locality: Guatemala]

Lycomorpha sinuata Henry Edwards 1885: 128 [type locality: Jalapa, Mexico]

Material examined. Type material: Holotype not examined. Specimen preparations. All material from USNM unless otherwise noted: MEXICO: Coatepec:

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Collection Wm. Schaus (1 female, CHS104, USNM 127,713). Vera Cruz: Jalapa,

Collection Wm. Schaus (1 male, CHS103, USNM 127,712).

Diagnosis. Lycomorpha ptychoglene may be confused with L. concolor, L. fulgens, and L. regulus. Lycomorpha ptychoglene is geographically isolated from L. concolor as noted in the diagnosis of that species. Characters from the male and female genitalia that are described in the diagnosis of L. fulgens can be used to separate L. ptychoglene and L. fulgens. The geographic distribution of L. ptychoglene and L. regulus are disjunct. Lycomorpha ptychoglene has been collected in Mexico and

Guatemala, whereas L. regulus is found in the southwestern United States from

California east to the panhandle of Texas. In addition, the coloration of the thorax can be used to separate these two species. The thorax of L. ptychoglene has red tegulae and the remainder is dark brown. In L. regulus, the tegulae, patagia and dorsal mesothorax are all red. The metathorax and entire ventral thorax are dark brown.

Description. Male habitus (Fig. 5-4H): Head: Scales on the head, antennae, labial palps, and gena are fuscous (21) colored. The antennae are serrate. No fusion is present among the labial palp segments. The gena is well developed and can be seen as a continuous band around the eye that joins with the frons when the head is viewed in profile. Thorax: Scales on the tegulae are spectrum orange (17). The patagia and the remaining dorsal and ventral thoracic scales are fuscous (21). The leg scales are fuscous with a small patch of chrome orange (16) scales present on the profemur.

Forewing: Length = 11mm (N=3); M2 and M3 are fused and stalk beyond the discal cell.

Dorsal surface: The wing is chrome orange (16). The proximal half of the costal vein is covered in spectrum orange (17) scales. Fuscous (21) scales line the distal half of the

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costal vein. The fuscous scales form a terminal band. A narrow band of fuscous scales extends from the terminal band along the posterior margin of the wing. They do not extend to the wing base. The fringe is fuscous. Ventral surface: The pattern is identical to the dorsal surface. The scales on the ventral surface are burnt orange (116) at the wing base and transition to salmon (106) colored distally. Salmon colored scales are found along the posterior margin of the wing. The scale colors on the costal vein and marginal band are identical to dorsal surface. Hindwing: CuA1 and M3 are fused and stalk distally beyond the discal cell. Dorsal surface: The hindwing is raw umber (223) colored. A band of salmon (106) colored scales is present along the basal two-thirds of the costal margin. The band begins at the wing base. The salmon colored scales line the costal vein. The posterior margin of the band occurs proximal to the R1 vein. No salmon colored scales are found on or posterior to the R1 vein. Ventral surface: The pattern and primary wing color are identical to the dorsal surface. The costal vein is lined with spectrum orange (17) colored scales. On the ventral surface, the band is burnt orange (116). Abdomen: Dorsal, ventral and lateral scales fuscous. A narrow pocket of hair-like androconial scales is found in the A7/A8 ventral intersegmental membrane. The pocket is less than one-third the width of the A7 sternite. The androconial scales are hair brown (119A) colored. The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite. The pattern of sclerotization on the eighth tergite is rectangular. Female habitus (Fig. 5-4G): Head: Color of head, labial palp, and genal scales the same as in the male. Antennae are simple and ciliate. The antennal scales are dark grayish brown (20) to fuscous (21) colored. The fusion of the labial palp

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segments and the development of the gena are the same as the male. Thorax: The tegulae are chrome orange (16). The color of the patagia, dorsal and ventral thorax, and legs is the same as in the male. Forewing: Length = 12mm (N=3); Venation the same as in the male. Dorsal surface: Color and pattern the same as in the male. Ventral surface: Color and pattern the same as in the male. Hindwing: The venation is the same as in the male. Dorsal surface: Color and pattern the same as in the male. Ventral surface: Color and pattern the same as in the male. Abdomen: Color the same as in the male. Male genitalia: Phallus: The basiphallus is straight with no inflection. When the phallus is viewed from the left side, the phallic sclerite is absent or highly reduced.

The vesica is composed of a single primary lobe with three membranous projections arising from the dorsal surface of the vesica. A heavily sclerotized, spine-like cornutus is present on the distal end of the primary lobe. Genital capsule: The tegumen has an elongate, rectangular shape. The lateral margins of the tegumen are curved concavely.

A triangular indentation is present in the proximal margin of the tegumen. A straight suture with an ovoid bulge medially that indicates where fusion occurred between the two halves of the tegumen runs from the cephalic to caudal end of the tegumen.

Narrow, sclerotized strips of tissue occur laterally on the uncus base. The uncus has a

C-shaped curve. The apex of the uncus is fingerlike and tapers to a point. The distal end of the costa of the valve is defined by a processus basalis of the costa. The processus basalis of the costa is dorso-ventrally compressed and trident shaped. The tips of the processus basalis of the costa are blunt. An editum is located on the base of the processus basalis of the costa. The valve is curved at an obtuse angle. Proximally to the genital capsule the valve is rectangular shaped with smooth, uncurved edges.

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Distal to the curve, the valve is rectangular shaped with curved edges. The valve tapers to a sclerotized spine that is oriented inward and perpendicular to the valve. In addition, a sclerotized lobe occurs adjacent to the ventral margin of the spine. The juxta occurs as a flat, sclerotized plate that is rectangular with a triangular indentation in the distal margin. The juxta is ornamented with a conical transparent patch at the proximal margin. Female genitalia: The heavy sclerotization of the seventh abdominal segment is interrupted by narrow membranous breaks that occur on the pleurites. The seventh sternite is ovoid with an X-shaped posterior margin. On the distal margin of the seventh sternite two elongated projections arise that encircle the ostium bursa and overlap forming an X-shape. The eighth sternite is present as a semicircular, lightly sclerotized plate. The anterior margin of the sternite is a sclerotized bar that fuses with the eighth tergite. The ostium bursa is located in the eighth sternite as a circular opening that occurs anteriorly in the plate but is not fused with the anterior margin of the plate. The ductus bursa is equal to or longer than the length of the seventh tergite. The ductus bursa is ribbon shaped proximal to the ostium bursa and the base of the corpus bursa.

Between these two regions the ductus bursa expands to form a circular shaped bulb.

The ductus bursa is not sclerotized. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner. The width of the triangular projections is greater than or equal to the length of the projection.

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Variation. Almost no variation in size and color was observed among the male or

female specimens examined.

Biology. Unknown.

Distribution. This species is found in the Mexican states of Coatepec and Vera

Cruz. The type of a synonym of L. ptychoglene, L. aequalis, was collected in

Guatemala.

Lycomorpha pulchra Dyar

Figs. 5-4I,J and 5-5A (female); Figs. 5-5 B-D (male).

Lycomorpha pulchra Dyar 1898: 34 [type locality: Belfrage, Texas, USA]

Material examined. Type material: Holotype not examined. Specimen

preparations. All material from USNM unless otherwise noted: UNITED STATES:

California: Mt. Home Can., 19.ix.1949, A.L. Melander, AMNH (1 male, CHS040);

Forest Home, 4.vii.1927, Engel Coll., CMNH (1 female, CHS049); Forest Home,

4.vii.1927, LACM (1 male, CHS077); San Diego Co., Flinn Springs, 18.vi.1981, leg:

Faulkner, Brown, SDNH (1 male, CHS075; 1 female, CHS095); Corona, 19.x.1930,

CAS (1 male, CHS076); Baja del Norte Mexico, Sierra Juarez Mtns., El Tajo Canyon,

12.ix.1958, F.S. Truxal, LACM (1 female, CHS096); Banner, Chariot Canyon,

11.x.1946, R.F. Allen Collector, CAS (1 female, CHS097); Los Angeles Co., S. Fork, Big

Rock Cr., 4500’, 27.ix.1967, Chris Henne Collection, LACM (1 female, CHS098).

Diagnosis. Lycomorpha pulchra can be confused with L. grotei and L. regulus.

Although there is partial overlap of the ranges of L. pulchra and L. grotei, only L. pulchra occurs in California. In addition, characters from the forewing color pattern and the male genitalia can be used to separate L. pulchra and L. grotei. These characters are fully described in the diagnosis of L. grotei. Characters of the wing color pattern and the

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male genitalia can be used to separate L. pulchra and L. regulus. The dark brown scales on the hindwing of L. pulchra occur intermittently along the posterior margin of the wing. The posterior margin of the hindwing in L. regulus is continuously lined with dark brown scales. Lycomorpha pulchra possesses a phallic sclerite with a narrow, pointed triangular shape. The phallic sclerite of L. regulus has a narrow, rounded triangular shape. Peglike cornuti are present the entire length of the membranous projection from the primary lobe of the vesica in L. pulchra. These cornuti are only found on the distal third of the projection in L. regulus. Furthermore, the juxta of L. pulchra is not ornamented with any transparent patches, whereas a triangular shaped transparent patch is present on the proximal edge of the juxta in L. regulus.

Description. Male habitus (Figs. 5-5 B-D): Head: The head and labial palps are sepia (119) to hair brown (119A) colored. The antennae are serrate. The antennal scales are sepia colored. There is no fusion among the labial palp segments. The gena is well developed and can be seen as a continuous band around the eye that joins with the frons when the head is viewed in profile. The genal scales are light drab (119C) colored. Thorax: The tegulae, patagia, and dorsal mesothorax Pratt’s ruby (210), flame scarlet (15), or burnt orange (116) colored. The dorsal metathorax is covered with sepia colored scales. The entire ventral thorax and legs are sepia (119) to hair brown (119A) colored. Forewing: Length = 12.3mm (N = 10); M2 and M3 arise separately from the discal cell. Dorsal surface: The wing is Pratt’s ruby, flame scarlet, or burnt orange colored. The color of the wing is the same as the color of the tegulae, patagia, and dorsal mesothorax. A sepia (119) colored terminal band is present. The band arises in the distal third of the costal margin and extends to the posterior margin of the wing. The

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fringe on the wing is hair brown (119A) colored. The posterior margin of the wing is lined with sepia colored scales. These scales extend over half the length of the posterior margin but do not reach the wing base. Ventral surface: The pattern is the same as the dorsal surface. The wing is geranium pink (13) to peach red (94) colored. Salmon (106) colored scales are present below the A1 vein. The retinaculum is covered in salmon colored scales. The terminal band is sepia colored. The fringe is hair brown colored.

Hindwing: CuA1 and M3 arise separately from the discal cell. Dorsal surface: The wing is geranium (12) to geranium pink (13) or orange rufous (123C) colored. Salmon (106) colored scales stretch from the costal margin to the anterior edge of the discal cell. A narrow dusky brown (19) marginal band is present. The band is less than 1mm wide except at the anal angle of the wing. The interior margin of the band is sinuate. At the anal angle, the band is bent convexly towards the costal margin. The dusky brown scales continue intermittently from the marginal band along the posterior margin of the wing to the wing base. The distal ends of the M1, M3, CuA1, CuA2, and A1 veins are lined with dusky brown scales. The hair-like scales that occur between the A1 and A2 veins may be dusky brown colored. Ventral surface: The wing is geranium (12) to geranium pink (13) or orange rufous (123C) colored. No salmon (106) colored scales are present.

The marginal band is dusky brown. The pattern of the marginal band is the same as the dorsal surface. The dusky brown colored scales do not extend along the posterior margin of the wing. The distal end of the M3 vein is lined with dusky brown colored scales. The dusky brown colored scales form a discal spot. Abdomen: Dorsal, lateral, and ventral scales are sepia (119) to hair brown (119A) colored. A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The

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pocket is one-third or more the width of A7 sternite. The scales in the pocket are drab gray (119D) colored. A small, shallow pocket of androconial scales is located in the center of the A8/A9 intersegmental membrane. The scales in the pocket are drab (27) colored. The cephalic margin of the eighth sternite is fused to the cephalic margin of the eighth tergite. The pattern of sclerotization on the eighth tergite is heart-shaped. Female habitus (Figs. 5-4I,J and 5-5A): Head: The head is sepia (119) to hair brown (119A) colored. Posterior to the antennae, scarlet (14) to chrome orange (16) or burnt orange

(116) colored scales intermixed on the head posterior to the head. The antennae are simple and ciliate. The color of the antennal scales, labial palps, and gena is the same as the male. The fusion of the labial palp segments and the development of the gena are the same as the male. Thorax: The tegulae, patagia, and dorsal mesothorax are scarlet (14) to chrome orange (16) or burnt orange (116) colored. This color is the same as the scales found posterior to the antennae on the head. The dorsal metathorax, all of the ventral thorax, and the legs are sepia to hair brown colored. Forewing: Length =

13.2mm (N = 10); M2 and M3 arise separately from the discal cell or are fused and stalk beyond the discal cell. Dorsal surface: The wing is the pattern is the same as the male.

The wings are scarlet (14) to chrome orange (16) or burnt orange (116) colored. As in the males, the color of the forewing is the same as the color on the thorax. The terminal band is sepia (119) colored with a hair brown (119A) colored fringe. The terminal band extends the same length as the males. Ventral surface: The pattern is the same as the dorsal surface. The wing is geranium pink (13) to peach red (94) colored. Posterior to the A1 vein the scales are salmon (106) colored. The retinaculum is covered in salmon colored scales. The colors of the terminal band and fringe are the same as the dorsal

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surface. Hindwing: CuA2 and M3 arise separately from the discal cell or are fused and

stalk beyond the discal cell. Dorsal surface: The wing is geranium pink (13) to scarlet

(14) colored. A band of salmon (106) colored scales extends along the costal margin.

The band stretches from the costal margin to the anterior margin of the discal cell. A

dusky brown (19) marginal band is present. This band is broader (1mm) than the

marginal band found on the male hindwing (<1mm). The interior margin of the band is

sinuate. The band is bent convexly toward the costal margin at the anal angle. The

dusky brown scales continue intermittently from the marginal band along the posterior

margin of the wing to the wing base. The distal end of the M3, CuA1, CuA2, and A1 veins

are lined with dusky brown scales. The long hair-like scales found between A1 and A2

may be dusky brown colored. Ventral surface: The wing color is the same as the dorsal

surface. No salmon colored scales are present along the costal margin. The pattern and

color of the marginal band are the same as the dorsal surface. No dusky brown scales

extend from the marginal band along the posterior margin of the wing. Dusky brown

colored scales line the distal end of the CuA1 vein. A discal spot is formed from the dusky brown colored scales. Abdomen: The color of the dorsal, lateral, and ventral

scales the same as the male. Male genitalia: Phallus: The basiphallus is straight with

no inflection. When the phallus is viewed from the left side, a narrow, pointed triangular

shaped phallic sclerite is present. The vesica is composed of a single primary lobe with

a membranous projection present. The membranous projection arises from the distal

end of the primary lobe at the upper right apex. The entire length of the projection is

ornamented with peglike cornuti. A heavily sclerotized, spine-like cornutus is present on

the distal end of the primary lobe at the upper left apex. Genital capsule: The tegumen

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is trapezoidal shaped. It is widest at the distal margin. A rectangular indentation is present in the proximal margin. A V-shaped suture that indicates where fusion occurred between the two halves of the tegumen extends for the distal two-thirds the length of the tegumen. A triangular knob is contained within the margins of the suture. The apex of the triangle is oriented toward the proximal margin of the tegumen. The uncus base occurs within a V-shaped sclerotized plate. The uncus has an S-shaped curve. The base of the uncus is teardrop shaped and tapers to a point proximal to the first curve of the uncus. The apex of the uncus is ovoid (tear drop shaped) tapering to a point. The distal end of the costa of the valve is defined by a membranous break that occurs beyond the midpoint of the dorsal edge of the valve. No processus basalis of the costa is present on the valve. An editum is located on the cephalic end of the costa. The valve is parallelogram shaped with rounded edges. Distally the valve tapers to a point that is oriented inward and perpendicular to the length of the valve. The juxta is triangular shaped with a conical tip at the distal end. The juxta is not ornamented with any transparent patches at the proximal margin. Female genitalia: The heavy sclerotization of the seventh abdominal segment is continuous without any membranous breaks. The seventh sternite is shaped like an inverted M. On the distal margin of the seventh sternite, a deep, concave indentation, which extends more than a quarter the length of the sternite, is present. The eighth sternite is absent. The ostium bursa is located in the

A7/A8 intersegmental membrane below the apex of the concave indention in the seventh sternite. The ductus bursa is shorter than the length of the seventh sternite.

The ductus bursa is tubed shaped. Sclerotization is present on the ductus bursa but not proximal to the ostium bursa. The sclerotized portion of the ductus bursa forms a C-

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shaped curve. The sclerotization is separated from the ostium bursa by a membranous break. The ductus seminalis arises from the lateral, right side of the ductus bursa. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone gland is square shaped with two, narrow triangular projections arising from each apical corner. The width of the triangular projections is less than the length of the projections.

Variation. The wing and thoracic color of male and female L. pulchra can vary significantly (Figs. 5-4I,J and 5-5 A-D). As in L. fulgens, this color variation appears associated with the age of the specimen not the collection locality. Older specimens have a lighter, faded coloration. No size variation was observed for males or females.

Biology. Based on the label data, the flight of this species occurs from June to

October. Nothing else is known about the biology of L. pulchra.

Distribution. Lycomorpha pulchra is found in the southwestern United States. Its range extends from California to Colorado. This species is found as far south as Baja

California. The furthest north L. pulchra has been collected is in Teller County,

Colorado.

Lycomorpha regulus (Grinnell)

Figs. 5-5E,F (female); Figs. 5-5G,H (male).

Anatolmis regulus Grinnell 1903: 11 [type locality: Mt. Wilson, California, USA]

Material examined. Type material: Holotype not examined. Specimen preparations. All material from USNM unless otherwise noted: UNITED STATES:

Arizona: Cochise Co., Chiricahua Mts., Barfoot Ridge, 8500 ft., 6.viii.1927, CAS (1

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male, CHS081). California: Emgd. 4.ii.1968, ova ex. cont. female, Los Angeles Co.,

San Gabriel Mts., So. Fork Big Rock Cy., 4500’, 27.ix.1967, Coll. C. Henne, CAS (1 male, CHS046); Emgd. 3.ii.1968, ova ex. cont. female, Los Angeles Co., San Gabriel

Mts., So. Fork Big Rock Cy., 4500’, 27.ix.1967, Coll. C. Henne, CAS (1 female,

CHS047); Emgd. 17.ii.1968, ova ex. cont. female, Los Angeles Co., San Gabriel Mts.,

So. Fork Big Rock Cy., 4500’, 8.x.1967, Coll. C. Henne, CAS (1 male, CHS101); Emgd.

20.ii.1968, ova ex. cont. female, Los Angeles Co., San Gabriel Mts., So. Fork Big Rock

Cy., 4500’, 18.x.1967, Coll. C. Henne, CAS (1 female, CHS102); Ventura Co., Sespe

Creek, 15.vi.1976, leg. M. Bell and T. Haglund, E.C. Olson Collection, FLMNH (1 male,

CHS042); Los Angeles Co., 6.vi.1922, K.R Coolidge, Harold M. Bower, Coll. Access’d

LACM 1964, LACM (1 female, CHS093). New Mexico: Catron Co., Bursum Camp, 18 miles E Alma, 9000ft., 14.vii.1961, F., P., and J. Rindge, AMNH (1 male, CHS099);

Grant Co., McMillan Camp, 14 miles N Silver City, 7000ft., 13.vii.1964, F., P., and J.

Rindge, AMNH (1 female, CHS100).

Diagnosis. Lycomorpha regulus can be confused with L. concolor, L. fulgens, L. grotei, L. ptychoglene, and L. pulchra. Differences in the development of the gena, as well as characters from the male and female genitalia can be used to separate L. regulus from L. concolor. These characters are fully described in the diagnosis of the latter species. The color pattern of the thorax can be used to separate L. regulus from L. fulgens. The tegulae, patagia, and dorsal mesothorax of L. regulus are covered with red scales, whereas only the tegulae of L. fulgens have red scales present. The wing color pattern and male genitalia help to separate L. regulus and L. grotei. These characters are described in the diagnosis of L. grotei. In addition, the gena of L. regulus is well

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developed, whereas it is reduced in L. grotei. Lycomorpha regulus is geographically isolated from L. ptychoglene. In addition, the color pattern of the thorax separates L. regulus and L. ptychoglene. Only the tegulae are red in L. ptychoglene. Wing color patterns and characters of the male genitalia are also used to separate L. regulus from

L. pulchra. These characters are discussed in the diagnosis of L. pulchra.

Description. Male habitus (Figs. 5-5G,H): Head: The head, labial palps, and gena are dusky brown (19) colored. The antennae are serrate. The antennal scales are sepia (119) colored. No fusion is present between the labial palp segments. The gena is well developed and can be seen as a continuous band around the eye that joins with the frons when the head is viewed in profile. Thorax: The tegulae, patagia, and dorsal mesothorax are scarlet (14) to flame scarlet (15) to chrome orange (16) colored. The dorsal metathorax, the entire ventral thorax, and the legs are sepia (119) colored.

Forewing: Length = 13.2mm (N = 10); M2 and M3 arise from the discal separately or are fused and stalk beyond the discal cell. Dorsal surface: The wing is scarlet (14) to flame scarlet (15) to chrome orange (16) colored. The color of the wing matches the color of the tegulae, patagia, and dorsal mesothorax. A sepia (119) colored terminal band is present. The terminal band arises in the distal third of the costal margin of the wing and extends to the anal angle of the wing. The fringe of the wing is dusky brown (19) colored. A narrow band of sepia (119) colored scales extends from the terminal band along the posterior margin of the wing. They extend adjacent to but do not reach the wing base. Ventral surface: The pattern of the wing is the same as the dorsal surface.

The wing is scarlet to flame scarlet colored. Peach red (94) to salmon (106) colored scales occur posterior to the A1 vein. The retinaculum is covered with salmon colored

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scales. The colors of the terminal band, fringe, and scales lining the posterior margin of

the wing are the same as the dorsal surface. Hindwing: CuA1 and M3 arise separately

from the discal cell or are fused and stalk beyond the discal cell. Dorsal surface: The

wing is dark grayish brown (20) colored with a fuscous (21) colored fringe. A poppy red

(210) to geranium pink (13) band is present along the costal margin. The band extends

three-quarters the length of the costal margin and tapers to a point. At the wing base,

the band stretches from the costal margin to the A1 vein. Flesh ocher (132D) to salmon

(106) colored scales occur within the anterior half of the band. The posterior margin of

the band is serrate. Ventral surface: The color of the wing is the same as the dorsal

surface. The costal band is entirely poppy red to geranium pink colored. The costal

band is broader stretching from the costal margin to the A2 vein. The posterior margin of

the band is serrate and bent convexly toward the anal angle of the wing. The band does

not taper to a point. Abdomen: Dorsal, lateral, and ventral scales are sepia (119) colored. A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The pocket is one-third or more the width of A7 sternite. The scales in the pocket are hair brown (119A) colored. A small, shallow pocket of androconial scales is located in the center of the A8/A9 intersegmental membrane. The

scales in the pocket are sayal brown (223C) colored. The cephalic margin of the eighth

sternite is fused to the cephalic margin of the eighth tergite. The pattern of sclerotization

on the eighth tergite is heart-shaped. Female habitus (Figs. 5-5E,F): Head: The colors

of the head, antennal scales, labial palps, and gena are the same as the male. The

antennae are simple and ciliate. The fusion of the labial palp segments and the

development of the gena are the same as the male. Thorax: The tegulae, patagia, and

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dorsal mesothorax are scarlet (14) to flame scarlet (15) to chrome orange (16) colored.

The colors of the dorsal metathorax, the entire ventral thorax, and legs are the same as the male. Forewing: Length = 13mm (N = 10); Venation is the same as the male.

Dorsal surface: The colors and patterns are the same as those on the male. Ventral surface: The colors and patterns are the same as those on the male. Hindwing: The venation is the same as in the male. Dorsal surface: The colors and patterns are the same as those on the male. Ventral surface: The colors and patterns are the same as those on the male. Abdomen: The color of the scales on the dorsal, lateral, and ventral abdomen are the same as the male. Male genitalia: Phallus: The basiphallus is straight with no inflection. When the phallus is viewed from the left side, a narrow, rounded triangular shaped phallic sclerite is present. The vesica is composed of a single primary lobe with a membranous projection present. The membranous projection arises from the distal end of the primary lobe at the upper right apices. The distal third of the projection is ornamented with peglike cornuti. The proximal two-thirds of the projection are ruggose. A heavily sclerotized, spine-like cornutus is present on the distal end of the primary lobe adjacent to the base of the membranous projection. Genital capsule: The tegumen is trapezoidal shaped. It is widest at the distal margin. A rectangular indentation is present in the proximal margin. A V-shaped suture that indicates where fusion occurred between the two halves of the tegumen arises adjacent to the proximal margin of the tegumen and extends to the distal margin of the valve. A triangular knob is contained within the margins of the suture. The apex of the triangle is oriented toward the proximal margin of the tegumen. The uncus base occurs within a V-shaped sclerotized plate. The uncus has an S-shaped curve. The base of the uncus is teardrop

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shaped and tapers to a point proximal to the first curve of the uncus. The apex of the uncus is ovoid (tear drop shaped) tapering to a point. The distal end of the costa of the valve is defined by a membranous break that occurs posterior to the midpoint of the valve. No processus basalis of the costa is present on the valve. An editum is located on the cephalic end of the costa. The valve is rectangular shaped with a sinuate ventral margin. The apex of the pentagon occurs on the ventral margin of the valve and is rounded. Distally, the valve tapers to a sclerotized point that is oriented inward and perpendicular to the length of the valve. The juxta is triangular shaped with a conical tip at the distal end. The juxta is ornamented with a triangular transparent patch at the proximal margin. Female genitalia: The heavy sclerotization of the seventh abdominal segment is continuous without any membranous breaks. The seventh sternite is shaped like an inverted M. On the distal margin of the seventh sternite, a deep, concave indentation, which extends more than a quarter the length of the sternite, is present.

The eighth sternite is absent. The ostium bursa is located in the A7/A8 intersegmental membrane below the apex of the concave indention in the seventh sternite. The ductus bursa is equal to or longer than the length of the seventh sternite. The ductus bursa is tube shaped and sinuate. Sclerotization is present on the ductus bursa but not proximal to the ostium bursa. The sclerotization is separated from the ostium bursa by a membranous break. The ductus seminalis arises from the lateral, right side of the ductus bursa. The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone gland is square shaped with two, narrow triangular

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projections arising from each apical corner. The width of the triangular projections is less than the length of the projections.

Variation. The wing and thoracic color of male and female L. regulus can vary significantly (Figs. 5-5 E-F). This color variation appears associated with the age of the specimen not the collection locality. Older specimens have a lighter, faded coloration.

This is similar to the color variation seen in other species of Lycomorpha with red forewings and primarily black hindwings. No size variation was observed for males or females.

Biology. Based on the label data, there appear to be two flights of L. regulus. The first flight occurs from May through July. The second flight begins in late August and continues through October. Comstock & Henne (1967) reared the larvae of L. regulus on the lichen Parmelia plittii. Their series of L. regulus have proved valuable in determining the phenotypic variation present within the wing color pattern of a single species. This information has aided in the delimitation of the species boundaries of

Lycomorpha species with red forewings and primarily black hindwings.

Distribution. Lycomorpha regulus is another species whose range occurs within the southwestern United States. This species is found from California east to the panhandle of Texas. The farthest north L. regulus has been found is Kane County,

Utah, which occurs on the border of Utah and Arizona.

Lycomorpha splendens Barnes and McDunnough

Fig. 5-5I (female); Fig. 5-5J (male).

Lycomorpha splendens Barnes and McDunnough 1912: 3 [type locality: Eureka,

Utah, USA]

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Material examined. Type material: Holotype not examined. Specimen

preparations. All material from USNM unless otherwise noted: UNITED STATES:

Arizona: Yavapai Co., Montezuma Well, 4.x.1955, Lloyd M. Martin, LACM (1 female,

CHS061). California: San Bernadino Co., Burns Pinyon Ridge Res., 2.5 air mi. N Yucca

Valley, T1N, R5E, NW ¼ sec. 27, el. 4100ft, 31.v.1992, Jerry Freilich, LACM (1 male,

CHS066). Texas: Jeff Davis Co., Fort Davis, 13.x.1966, A. & M.E. Blanchard (1 male,

CHS039, USNM 127,703); Presidio Co., Big Bend Ranch St. Nat. Ar., Sauceda Rnch.,

26-1.x-xi.1989, E.G. Riley, TAMU (1 male, CHS067); Armstrong Co., Palo Duro Canyon

St. Pk., 25.ix.1968, A. & M.E. Blanchard, AMNH (1 male, CHS065). Utah: St. George,

1-7.vi (1 male, CHS068, USNM 127,709).

Diagnosis. The coloration of L. splendens is unique within Lycomorpha. It has not been confused with any other members of the genus.

Description. Male habitus (Fig. 5-5J): Head: Scales on the head, labial palps, and gena are dusky brown (19) colored. The antennae are bipectinate with comblike rami. The antennal scales are dusky brown with and iridescent cobalt (68) sheen. The second and third segments of the labial palps are fused together. The gena is reduced and cannot be see as a continuous band around the eye when the head is viewed in profile. Thorax: The tegulae are geranium pink (13) to scarlet (14) colored. The fringe emerging from the posterior margin of the tegulae is dusky brown (19) colored. The scales on the patagia are dusky brown with cobalt (68) iridescence. The scales on the ventral thorax and remaining dorsal thorax are fuscous (21) colored. The scales on the legs are the same color as those on the patagia. Forewing: Length = 14.1mm (N = 10);

M2 and M3 arise separately from the discal cell. Dorsal surface: The wing is Vandyke

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brown (121) colored. The scales lining the costal vein have cobalt (68) iridescence.

Ventral surface: The wing is sepia (219) to fuscous (21) colored. No iridescence is

present. Hindwing: CuA1 and M3 arise separately from the discal cell. Dorsal surface:

The wing is geranium pink (13) colored. A narrow hair brown (119A) band extends from the wing base on the costal margin of the wing to the anal angle. No hair brown colored scales occur along the posterior margin of the wing. Along the costal margin of the wing, the band stretches from the costal margin to the center of the discal cell. Ventral surface: The pattern and colors are the same as the dorsal surface. Abdomen: Dorsal, lateral, and ventral scales are fuscous (21) colored. A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The pocket is one- third or more the width of A7 sternite. The scales in the pocket are raw umber (23) colored. The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite. The pattern of sclerotization on the eighth tergite is rectangular. Female habitus (Fig. 5-5I):

Head: Scales on the head, labial palps, and gena are fuscous (21) colored. The

antennae are serrate. The antennal scales are fuscous colored with a cobalt (68)

iridescence. The fusion of the labial palp segments and the development of the gena are the same as the male. Thorax: The tegulae are geranium pink (13) to scarlet (14)

colored. The fringe emerging from the posterior margin is fuscous (21). Patagia, dorsal

and ventral thorax, and legs are fuscous colored. Forewing: Length = 11.7mm (N = 3);

venation the same as the male. Dorsal surface: The wing is Vandyke brown (121)

colored. No iridescence is present. Ventral surface: The color is the same as the dorsal

surface. Hindwing: The venation is the same as the male. Dorsal surface: The colors of

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the wing are the same as the male. The hair brown (119A) band is narrower than in the

males. It does not stretch to the center of the discal cell. On the outer margin of the

wing, the hair brown colored scales are restricted to the fringe. Ventral surface: The

colors and pattern are the same as the dorsal surface. Abdomen: The dorsal scales on

the abdomen are fuscous (21) colored. The lateral and ventral scales are sepia (219)

colored. Male genitalia: Phallus: The basiphallus is straight with no inflection. When

the phallus is viewed from the left side, a rectangular phallic sclerite is present. The

vesica is composed of a single primary lobe. No membranous projections arise from the

vesica. A heavily sclerotized, spine-like cornutus is not present. Genital capsule: The

tegumen is trapezoidal with a concave indentation on the proximal margin. The lateral

margins of the tegumen are curved convexly. The tegumen is widest at the proximal

margin. An inverted T-shaped suture that indicates where fusion occurred between the

two halves of the tegumen extends from cephalic to the caudal margin of the tegumen.

No sclerotization extends around the base of the uncus. The uncus has a C-shaped

curve. The apex of the uncus is fingerlike tapering to a point. The distal end of the costa of the valve is defined by a triangular, membranous break that occurs at the midpoint of the dorsal edge of the valve. No processus basalis of the costa is present on the valve.

An editum is found at the cephalic end of the costa. The valve is ovoid shaped. Distally,

the valve tapers to a sclerotized point that is oriented inward and perpendicular to the

length of the valve. The juxta is a trapezoidal shaped plate. The juxta is not ornamented

with any transparent patches at the proximal margin. Female genitalia: The heavy

sclerotization of the seventh abdominal segment is interrupted by narrow membranous

breaks that occur on the pleurites. The seventh sternite is shaped like a shield. The

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distal margin of the seventh sternite is approximately horizontal without any deep indentations. The eighth sternite is present. However, it is reduced to two bars that are fused to the eighth tergite. The bars are not continuous across the venter. The ostium bursa is located in the A7/A8 intersegmental membrane. The ductus bursa is shorter than the length of the seventh sternite. The ductus bursa is tube shaped and does not curve. No sclerotization is present on the ductus bursa. The ductus seminalis arises from the lateral, left side of the ductus bursa. The membrane of the single corpus bursa is smooth and ornamented with a single slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner. The width of the triangular projections is greater than or equal to the length of the projection.

Variation. Female L. splendens are rare in collections. Almost no variation in size and color was observed among the male or female specimens examined.

Biology. Based on the collection label data, there appear to be two flights of L. splendens. The first flight occurs from May to June. The second flight lasts from August to October. No specimens have been collected in July. Lycomorpha splendens is reported to be nocturnal and attracted to lights (Powell & Opler 2009). No other information is known about the biology of this species.

Distribution. This species is found in the Southwestern United States. Its range extends from Texas west to eastern California. L. splendens is found as far north as

Colorado and Utah.

Lycomorpha texanus Scott New Species

Fig. 5-6A (female); Fig. 5-6B (male).

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Material examined. Type material: Holotype: Mt. View Acres, Bexar Co.,

Texas, 11.x.1970, Leg. R.O. & C.A Kendall, USNM (1 male, CHS128, USNM 127,719).

Paratypes: UNITED STATES: Texas: Bexar Co., Mt. View Acres, 11.ix.1970, Leg.

R.O., USNM (1 male, CHS-PC049, USNM 127,729); Bexar Co., Mt. View Acres,

11.x.1970, Leg. R.O. & C.A Kendall, USNM (2 males); Tiger Hill USNM (1 female, 1 male). Specimen preparations. All material from TAMU unless otherwise noted:

UNITED STATES: Texas: 10 miles south of Kerrville, 8.v.1983, W.F. Chamberlain (2 females, CHS073, CHS-PC048; 1 male CHS062); 28 miles N of Del Rio, 10.x.1993,

W.F. Chamberlain (1 male, CHS131); Palo Pinto, 24.v.1941, O. Buchholz Collection,

AMNH (1 male, CHS063); Palo Pinto, 1.v.1954, W.J. Reinthal, AMNH (1 female,

CHS130); Bexar Co., 14mi N.W. San Antonio, 14.x.1970, J.W. Tilden Collector, CAS (1 female, CHS129); Comfort, Lucock, 13.x.1899, Holland Collection, CMNH (1 female,

CHS132); Comal Co., New Braunfels, 16.x.1977, leg. E.C. Knudson, 16.x.1977, FLMNH

(1 male, CHS-PC038); Bexar Co., 12mi N.W. San Antonio, 14.x.1970, J.W. Tilden

Collecter, UCB (1 male, CHS-PC047).

Etymology. The name of this species is derived from the only state that it has been collected: Texas.

Diagnosis. Lycomorpha texanus can be confused with L. atroxantha, L. miniata,

L. pelopia and L. pholus. The geographic distribution of L. texanus can be used to separate it from each of these species. The ranges of L. texanus and L. atroxantha are fully detailed in the diagnosis of L. atroxantha. Characters of the female genitalia also help to separate L. texanus and L. miniata. These characters are fully described in the previous diagnosis of L. miniata. Lycomorpha texanus is restricted to Texas, whereas L.

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pelopia has been collected from Panama and Mexico. The color of the scales on the

head and gena are sepia colored in L. texanus. However, the scales found on the gena

of L. pelopia are orange yellow colored. The color patterns of the forewing noted in the

diagnosis of L. pholus, as well as the geographic distribution of the species, help to

separate this species from L. texanus.

Description. Male habitus (Fig. 5-6B): Head: Scales on the head, labial palps,

and gena are sepia (119) colored. The antennae are serrate. The antennal scales are

sepia colored with cobalt (68) iridescence. No fusion is present between the labial palp

segments. The gena is well developed and can be seen as a continuous band around

the eye that joins with the frons when the head is viewed in profile. Thorax: The tegulae

are chrome orange (16) to spectrum orange (17) colored. The patagia and dorsal and

ventral thorax are sepia (119) colored. The leg scales are sepia colored with cobalt (68)

iridescence. Forewing: Length = 13.4mm (N = 10); M2 and M3 arise separately from the

discal cell. Dorsal surface: The basal half of the wing is spectrum orange (17) colored.

Geranium pink (13) colored scales occur on the basal third of the posterior margin of the

basal half. The external margin of the basal half of the wing tapers to a point where the

CuA1 vein arises from the discal cell. Anterior and posterior to this point, the external

margin is bent convexly toward the wing margin. The distal half of the wing is dusky

brown (19) colored with cobalt (68) iridescence. No dusky brown scales extend along

the posterior margin of the basal half of the wing. Ventral surface: The basal half of the

wing is spectrum orange (17) colored. Burnt orange (116) colored scales are intermixed

in the basal half of the wing posterior to the anterior margin of the discal cell. The

external margin of the basal half of the wing is serrate. The distal half of the wing is

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dusky brown (19) colored. No dusky brown cells extend along the posterior margin of

the basal half of the wing. Hindwing: CuA1 and M3 arise separately from the discal cell.

Dorsal surface: The wing is dusky brown (19) to dark grayish brown (20) to fuscous (21)

colored. The adbasal region is chrome orange (16) to spectrum orange (17) colored.

Geranium pink (13) colored scales are intermixed posterior to the middle of the discal

cell. The geranium pink colored scales continue to the posterior margin of the adbasal

region. The posterior margin of the adbasal region is sinuate. Ventral surface: The basal

half of the wing is composed of a mixture of chrome orange, spectrum orange, and geranium pink colored scales. The external margin of the basal half of the wing is

sinuate. The distal half of the wing is fuscous (21) colored. A narrow band of fuscous

colored scales extend from the distal half of the wing along the posterior margin of wing

to the wing base. Abdomen: Dorsal, lateral, and ventral scales are sepia (119) colored.

A shallow, broad pocket of androconial scales present in the A7/A8 intersegmental membrane. The pocket is one-third or more the width of A7 sternite. The scales in the pocket are burnt umber (22) colored. The cephalic margin of the eighth sternite extends adjacent to the cephalic margin of the eighth tergite. There is no fusion between the sternite and tergite. The pattern of sclerotization on the eighth tergite is rectangular.

Female habitus (Fig. 5-6A): Head: The colors of the head, antennal scales, labial palps, and gena are the same as the male. The antennae are serrate. The fusion of the labial palp segments and the development of the gena are the same as the male.

Thorax: The tegulae are chrome orange (16) to spectrum orange (17) colored. The colors of the patagia, dorsal and ventral thorax, and legs are the same as the male.

Forewing: Length = 13mm (N = 10); Venation is the same as the male. Dorsal surface:

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The colors and patterns are the same as those on the male. Ventral surface: The colors

and patterns are the same as those on the male. Hindwing: The venation is the same

as in the male. Dorsal surface: The colors and patterns are the same as those on the

male. Ventral surface: The colors and patterns are the same as those on the male.

Abdomen: The color of the scales on the dorsal, lateral, and ventral abdomen are the

same as the male. Male genitalia: Phallus: The basiphallus is straight with no

inflection. When the phallus is viewed from the left side, the phallic sclerite is present as

a flattened lobe, which extends as a separate structure dorsad of the vesica. The vesica

is composed of a single primary lobe with a membranous projection arising on the left

side of the vesica proximal to the distal margin of the phallus. The distal end of the

membranous projection of the vesica is ornamented with a single heavily-sclerotized,

spine like cornutus. Genital capsule: The tegumen is rectangular with a concave

indentation that extends half the length of the distal margin. The indentation is centered

in the margin. Two half-crescent shaped sclerotized extensions arise from the corners

of the distal margin. The proximal margin of the tegumen is crenellated. A V-shaped

suture that indicates where fusion occurred between the two halves of the tegumen

arises adjacent to the proximal margin of the tegumen and extends to the distal margin

of the valve. A circular knob is contained within the margins of the suture and arises

adjacent to the distal margin of the tegumen. The uncus base occurs within a

rectangular sclerotized plate. The uncus has a C-shaped curve. The apex of the uncus is ovoid (teardrop shaped) tapering to a point. The distal end of the costa of the valve is defined by a triangular, membranous break that occurs beyond the midpoint of the dorsal edge of the valve. Beyond the membranous break, the dorsal edge of the valve

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occurs below the costa. No processus basalis of the costa is present on the valve. No editum is present on the internal face of the valve. The valve is teardrop shape. The ventral margin of the valve tapers to a point that is perpendicular to the length of the valve distally. The dorsal margin tapers to a lobe distally. In addition, a sclerotized spine arises from the lobe formed from the dorsal margin of the valve. The spine is oriented inward and perpendicular to the length of the valve. The juxta is a flat square sclerotized plate with a concave indentation in the distal margin. The juxta is not ornamented with any transparent patches at the proximal margin. Female genitalia: The heavy sclerotization of the seventh abdominal segment is interrupted by narrow membranous breaks that occur on the pleurites. The seventh sternite is goblet shaped. The distal margin of the seventh sternite is crenellated. The eighth sternite is present as a heavily sclerotized plate. The ostium bursa is located in the eighth sternite as a broad, horizontal slit that is centered in the sternite. The opening extends over half the length of the sternite. The ductus bursa is shorter than the length of the seventh sternite. The ductus bursa is ribbon like with no curves or bulges. No sclerotization is present on the ductus bursae. The ductus seminalis arises from the ventral side of the ductus bursa.

The membrane of the single corpus bursa is wrinkled and ornamented with two, slightly sclerotized ovoid signa with heavily sclerotized bars running perpendicular to the length of the signa. These two signa occur on opposite sides of the corpus bursa. The dorsal pheromone glands are square shaped with two broad triangular projections arising from each apical corner. The width of the triangular projections is greater than or equal to the length of the projection.

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Variation. In some individuals of L. texanus, a narrow band of dusky brown scales extends from the distal half of the forewing along the distal third of the posterior margin of the basal half of the dorsal surface of the forewing. On the ventral surface of the forewing, the pattern of the external margin of the basal half of the wing is the same as the dorsal surface. Almost no variation in size was observed among the male or female specimens examined.

Biology. There appear to be two flights of L. texanus based on collection label data. The first flight occurs in April and May. The second flight is in September and

November. No other information is known about the biology of this species.

Distribution. L. texanus is restricted to Central Texas. Specimens of L. texanus have been collected in the Edwards Plateau region and the Rolling Plains region. The

Rolling Plains region is contiguous with the northern edge of the Edwards Plateau.

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Figure 5-1. Strict consensus of 3 most parsimonious trees (L=192, CI=0.68, RI=0.73) resulting from the maximum parsimony analysis of the all species (AS) dataset. Numbers below the branches are the support values for the nodes: Bremer Support (BS)/Jackknife (JK). C1 – C6 = Clade 1 – Clade 6. Solid line = species placed in Lycomorpha, Dashed line = species placed in Propyria, Dotted line = outgroup taxa.

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Figure 5-2. Images of the adult habitus of the genus Lycomorpha, Plate 1. A) L. atroxantha (Schaus), Female. B) L. concolor Scott, Female. C) L. concolor Scott, Female. D) L. concolor Scott, Male. E) L. concolor Scott, Male. F) L. fulgens (Edwards), Female. G) L. fulgens (Edwards), Female. H) L. fulgens (Edwards), Female. I) L. fulgens (Edwards), Male. J) L. fulgens (Edwards), Male. Collection locality data for each specimen photographed can be found in Appendix F.

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Figure 5-3. Images of the adult habitus of the genus Lycomorpha, Plate 2. A) L. fulgens (Edwards), Male. B) L. fulgens (Edwards), Male. C) L. grotei (Packard), Female. D) L. grotei (Packard), Female. E) L. grotei (Packard), Male. F) L. grotei (Packard), Male. G) L. miniata Packard, Female. H) L. minata Packard, Male. I) L. morelosia (Schaus), Female. J) L. morelosia (Schaus), Male. Collection locality data for each specimen photographed can be found in Appendix F.

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Figure 5-4. Images of the adult habitus of the genus Lycomorpha, Plate 3. A) L. neomexicanus Scott, Male. B) L. normani (Schaus), Male. C) L. pholus (Drury), Female. D) L. pholus (Drury), Female. E) L. pholus (Drury), Male. F) L. pholus (Drury), Male. G) L. ptychoglene (Hampson), Female. H) L. ptychoglene (Hampson), Male. I) L. pulchra Dyar, Female. J) L. pulchra Dyar, Female. Collection locality data for each specimen photographed can be found in Appendix F.

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Figure 5-5. Images of the adult habitus of the genus Lycomorpha, Plate 4. A) L. pulchra Dyar, Female. B) L. pulchra Dyar, Male. C) L. pulchra Dyar, Male. D) L. pulchra Dyar, Male. E) L. regulus (Grinnell), Female. F) L. regulus (Grinnell), Female. G) L. regulus (Grinnell), Male. H) L. regulus (Grinnell), Male. I) L. splendens Barnes and McDunnough, Female. J) L. splendens Barnes and McDunnough, Male. Collection locality data for each specimen photographed can be found in Appendix F.

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Figure 5-6. Images of the adult habitus of the genus Lycomorpha, Plate 5. A) L. texanus Scott, Female. B) L. texanus Scott, Male. Collection locality data for each specimen photographed can be found in Appendix F.

Figure 5-7. Adult habitus of female L. pelopia. Collection locality data for the specimen can be found in Appendix F.

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Figure 5-8. Male flagellomere state. A) Lycomorpha fulgens; serrate flagellomeres. B) Lycomorpha splendens; bipectinate flagellomeres. The scale bar is equivalent to 1mm.

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Figure 5-9. Female flagellomere state. A) Lycomorpha grotei; simple, ciliate flagellomeres. B) Lycomorpha splendens; serrate flagellomeres. The scale bar is equivalent to 1mm.

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Figure 5-10. Labial palp segment fusion. A) Lycomorpha pholus; No fusion of segments. B) Lycomorpha splendens; 2nd and 3rd segments fused. The scale bar is equivalent to 1mm.

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Figure 5-11. Medial veins of the forewing. A) Lycomorpha pholus; M2 and M3 arise separately from the discal cell. B) Lycomorpha grotei; M2 and M3 are fused and stalk beyond the discal cell. The scale bar is equivalent to 1mm.

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Figure 5-12. Medial and cubital veins of hindwing. A) Lycomorpha grotei; M3 and CuA1 arise separately from the discal cell. B) Lycomorpha ptychoglene; M3 and CuA1 are fused and stalk beyond the discal cell. The scale bar is equivalent to 1mm.

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Figure 5-13. Anterolateral process (ALP) form. A) Lycomorpha pholus; ALP present as a SB-sclerotized bar. B) Lycomorpha morelosia; ALP present as a FL- flattened lobe. The scale bar is equivalent to 1mm.

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Figure 5-14. Androconia of the A7/A8 intersegmental membrane. A) Lycomorpha splendens; SP-shallow pocket that is more than 1/3rd the width of 7S-seventh sternite. B) Lycomorpha normani; NP-narrow pocket that is less than 1/3rd the width of 7S-seventh sternite. The scale bar is equivalent to 1mm.

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CHAPTER 6 DISCUSSION AND FUTURE DIRECTIONS

Tribal Study

Phylogenies have been constructed for the tribe Lithosiini (Lepidoptera:

Erebidae: Arctiinae) based on either a morphological or a molecular dataset. The

phylogenies were used to assess the monophyly of both the tribe and the subtribes

proposed by Bendib & Minet (1999). Although overlap was present in the taxon

sampling of the two phylogenetic analyses, the datasets were not combined for a total

evidence analysis (De Queiroz 1993; Kluge 1989; Wiens 1998a). Only seventeen of the

species sampled for Lithosiini were present in both datasets. In addition, each of the

four gene fragments (two mitochondrial and two nuclear) included in the molecular

dataset were not successfully amplified for these seventeen species. Although Whiting

et al. (1997) recovered a resolved phylogeny despite the inclusion of taxa that were not

represented by all of the data, Wiens & Reeder (1995) and Wilkinson (1995) noted that

the inclusion of taxa that were missing portions of the data could detrimentally impact

the accuracy of the phylogeny that was obtained. Furthermore, Wiens (1998b) found

that as the proportion of missing data increased within a dataset, the accuracy of the

phylogeny decreased. Given the percentage of data that would have been missing by

combining the two datasets, a total evidence analysis was not appropriate at this time.

For the morphological study, seventy-six species of Lithosiini that included representatives of each of the seven subtribes were sampled for eighty-two morphological characters. This dataset was analyzed using maximum parsimony (MP) and Bayesian inference (BI) analysis. The phylogenies that were recovered were almost entirely unresolved. The BI phylogram recovered the tribe Lithosiini as monophyletic;

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however, none of the subtribes were recovered as monophyletic by either analysis.

These findings are not unusual since all synapomorphies that have been identified for

the tribe, to date, arise from the larval morphology. In addition, Bendib & Minet (1999)

noted that the larval morphology was the most accurate for placing taxa within Lithosiini.

Although the results of this phylogenetic study do not provide resolution to the

evolutionary relationships among the genera, they do provide confirmation that the

phylogenetic signal present within the adult lithosiine morphology is not informative at

the tribal level.

Phylogenetic analyses of datasets composed of several gene fragments have

been used to resolve the higher-level relationships within superfamilies, families, and

subfamilies of Lepidoptera (Regier et al. 2012; Ugelvig et al. 2011; Zahiri et al. 2011,

2012). A molecular dataset composed of four gene fragments (two nuclear and two

mitochondrial) was constructed for forty-six species of Lithosiini representing four of the lithosiine subtribes. The molecular dataset was analyzed using maximum likelihood

(ML) and BI analysis. In addition, the optimal partitioning strategies and models of evolution identified by two different selection criteria were compared. The same species groups were found with both partitioning strategies and both methods of analysis. The relationships recovered among the species groups varied based on the type of phylogenetic analysis. Both the type of analysis and the partitioning strategy affected the nodal support of these groups. Although the phylogenies recovered using the molecular dataset were more resolved than those obtained from the morphological dataset, nodal support was low or absent for the deeper nodes of the molecular phylogeny. Each of the analyses found the tribe Lithosiini to be paraphyletic with

269

respect to the other tribes of Arctiinae, Syntomiini and Arctiini. However, in the most

recent phylogenetic analysis of the subfamily Arctiinae (S. Weller pers. comm.),

Lithosiini was recovered as monophyletic. As in the morphological study of the tribe,

none of the subtribes included in the analysis were recovered as monophyletic.

Although the use of molecular data led to an increase in the resolution of the

evolutionary relationships of Lithosiini, further phylogenetic studies are needed to

resolve the higher-level relationships.

Review of Lycomorpha

A review of the lithosiine genus Lycomorpha Harris was completed as part of this

study. The wing color patterns of the adults of this genus have led to confusion in both

the family placement of the genus and the species composition. In their phylogenetic

study of the tribes of Arctiinae, Jacobson & Weller (2002) confirmed the placement of

Lycomorpha within Lithosiini. However, no study had been completed that examined the

relationships of the species within the genus or the generic limits of Lycomorpha. A

similarity of wing color pattern and body shape had been noted between Lycomorpha

and the lithosiine genera Propyria Hampson and Ptychoglene Felder. This study used sixty-five morphological characters coded for seven species of Lycomorpha, which included three new species. Also included in the analysis were three species of Propyria and a single species of Ptychoglene. The species sampling included the type species of each genus. The dataset was analyzed using MP and BI analysis. Both types of analysis, recovered Lycomorpha as paraphyletic with respect to Propyria. Twenty synapomorphies were identified that supported this relationship. In addition,

Ptychoglene was more closely related to the genus Hypoprepia Hübner than either

Lycomorpha or Propyria. Further MP analyses were conducted in which the monophyly

270

of either Lycomorpha or Propyria was constrained. Although the lengths of the two phylogenies recovered from these analyses were not significantly greater, the two genera remained paraphyletic. Based on these findings, Lycomorpha and Propyria were synonymized. The older generic name Lycomorpha was given priority. Based on the results of the morphological phylogeny of the genus, a revision of Lycomorpha has been completed. A description and diagnosis have been provided for both the genus and fourteen of the twenty species placed within Lycomorpha.

Future Directions

Tribal Systematics

Studies of the evolution of behaviors within the tribe and assessments of the monophyly of the subtribes of Lithosiini require a resolved phylogeny. As part of this study, two phylogenies were constructed for the tribe; however, neither phylogeny was able to resolve the higher-level relationships within Lithosiini. Of the two datasets that were analyzed, the phylogenies obtained using molecular data were more resolved.

However, only four of the seven subtribes were represented in the taxon sampling of the molecular phylogeny. In addition, the Neotropics, an area of high biodiversity for the tribe Lithosiini, was represented by only four species. Future studies must attempt a denser taxon sampling that includes species from each of the subtribes and provides better representation for areas of high biodiversity. The lack of higher-level resolution might be addressed by including more nuclear gene fragments in the data analysis.

Nuclear DNA evolves more slowly than mitochondrial DNA, and this property has made it useful in resolving the higher-level relationships of the order Lepidoptera (Regier et al.

2009; Mutanen et al. 2010). This process of selecting, amplifying, and sequencing gene fragments to construct a dataset for a phylogenetic analysis is reliant on Sanger

271

sequencing. Although other sequencing methods were available, the cost of these

methods has been prohibitive in comparison to Sanger sequencing. However,

sequencing protocols collectively referred to as “Next-Generation Sequencing” that

produce large volumes of data cheaply have become available (Shendure & Ji 2008;

Metzker 2010). The greater quantities of information obtained are equivalent to

amplifying a large number of loci using Sanger sequencing. In addition, Next-

Generation sequencing methods allow for genomic data to be obtained for non-model

organisms. This data can then be used both to construct a phylogeny and to study the evolution of behavior (Nygaard et al. 2011). Next-Generation sequencing methods could also be used to obtain a dataset for future studies to examine the evolutionary relationships of the tribe and obtain resolution at the deeper nodes.

Chemical Defense

Lichen phenolics are compounds produced by the fungal component of the lichen to defend against herbivores. Hesbacher et al. (1995) examined wild-caught

adults of Lithosiini for the presence of these phenolics. They examined 103 adults of

sixteen species representing eight lithosiine genera. Twenty-four individuals were found to have known lichen phenolic compounds stored in their bodies. In addition, forty- seven of the individuals examined possessed lichen phenolic-like compounds. They found both inter- and intraspecific variation in the presence of these compounds. This is the only the study that has been completed that examined adults of Lithosiini for the presence of lichen based compounds. However, the sampling of the study was too small to provide a clear idea of how common this behavior is within Lithosiini. Future studies could examine a broader sampling of lithosiine taxa from around the world for the presence of lichen phenolics. HPLC analysis could be used to determine which taxa

272

possess lichen phenolics. This data could then be correlated with wing color pattern and ultrasonic noise production in order to produce hypotheses as to which species that appear distasteful possess defensive chemicals.

Courtship Behavior of Lycomorpha

The courtship rituals of some species within the tribe Arctiini involve both pheromones released from androconia and ultrasonic clicks produced by tymbal organs

(Weller et al. 1999; Sanderford 2009). Males of Lycomorpha species possess both of these structures. The males of all species of Lycomorpha possess at least one pocket of androconial scales on their abdomen and another pocket of androconial scales within the intravincular membrane of the genital capsule. However, no studies have been completed to examine the courtship behavior of Lycomorpha species. Lycomorpha pholus, the most widespread species within the genus, can be used as a model for this research. The tympanum of L. pholus has been found to be most sensitive to the ultrasound frequency produced by its tymbal organs. Furthermore, Muma & Fullard

(2004) hypothesize that the ability of L. pholus to hear in this range could be selected for by short range, social communications, which are the types of cues used in courtship rituals. Studies of L. pholus courtship could determine whether the males use either of these systems. If the results of these studies demonstrate that these structures are used in mating, the role of androconia, ultrasound, or both in mating success could also be examined.

273

APPENDIX A MORPHOLOGICAL DATA MATRIX FOR TRIBAL PHYLOGENY

1 2 3 4 5 6 7 8 0 0 0 0 0 0 0 0 0 Acsala anomala 1021010201 1001?00001 0101000111 000211?100 1101?110?1 ??11?11??? 0111?1?12? 0001?000?1 1? Aemene altaica 1010011201 1010100001 000101?100 ??01100101 1101?10211 ??00010111 1110103110 0101?100?1 00 Barsinella 001???1201 1110100001 0020000011 1002100101 0101?101?0 1001?11??? 10001????? ?????????? ?? mirabilis Bruceia 11?01?1001 1000000001 0101000211 101??1?001 00101110?1 ??01?11??? 111011?111 00000000?1 1? hubbardi Bruceia 11?01?1101 1000000001 0001000011 101??1?001 00101110?1 ??01?11??? 2110102111 00000000?1 00 pulverina Cisthene 0010011201 1000100001 0001001001 000111?101 01000110?1 ??11?01??? 2010002111 1001?00311 00 plumbea Cisthene 0010011201 1000100001 0001001211 000211?101 01000110?1 ??11?01??? 2010002111 1000110301 1? subjecta Clemensia 0010011211 1100000001 000101?101 0100100101 1100110231 ??11?10011 200001?110 0000110310 00 albata Clemensia 0010011211 1100000001 000101?101 0100100101 1100010231 ??11?10011 200001?110 0000110310 1? leopardina Dolichesia 0010011201 1110100010 1?01000001 0001103101 11100110?1 ??11?01??? 0??0100111 0000010101 1? falsimonia Euthyone 01?01?1001 1000000001 0011000211 000101?101 11001110?1 ??01?01??? 1010000010 11000000?1 01 grisescens Euthyone 11?11?1001 1000000001 0011000211 0000002100 1111?110?1 ??01?11??? 101001?110 00000000?1 1? simplex Gaudeator 1010011211 1000000001 0011000201 100111?101 11100110?1 ??01?01??? 0110101111 1100110110 1? paidicus Haematomis 11?01?1101 1000000001 00010??211 101??1?001 00101110?1 ??01?11??? 2110102111 11000000?1 1? mexicana

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Hypermaepha 0010011201 1011?00010 1?111??211 1100100100 1101?110?1 ??00011??? 0110102010 0100000101 00 maroniensis Hypoprepia 1011011211 1000000001 1?11000201 0001100101 1101?110?0 1001?11??? 201011?110 0000000201 00 cadaverosa Hypoprepia 1010011211 1000000001 1?11000201 0001100101 1101?110?0 1001?11??? 201011??10 1??0000201 1? fucosa Hypoprepia 1011011201 1000000001 1?11000201 0001100101 0101?110?0 1001?11??? 2010101110 0100000201 00 inculta Lycomorpha 1011010101 1000000010 1?01001001 0000100101 0001?110?1 ??01?11??? 2110000110 1101?10101 1? fulgens Lycomorpha 1010011101 1000000010 1?01001001 0000100101 00000110?1 ??01?01??? 2110000111 1100010101 1? fulgens nr. Lycomorpha 1010011101 1000000010 1?01001001 0000100101 11000110?1 ??01?01??? 2100000111 1100010101 1? grotei Lycomorpha 1011010101 1000000010 1?01001001 0000100101 0101?110?1 ??01?01??? 211000312? 1100010101 1? pholus Lycomorpha 1021011101 1000000010 1?01001001 0000100101 01000110?1 ??11?11??? 2111?03111 1101?000?1 1? splendens Lycomorphodes 0010011201 1000100010 1?11000301 0100100001 0011?110?1 ??01?01??? 1111?1?111 0101?10101 1? correbioides Metalobosia elis 0010011211 1010000001 1?20000301 101??1?000 0100110221 ??01?10100 011011?110 0001?10300 01 Nodozana 0010011?01 1000000001 0000001211 0001100001 1101?110?0 1011?01??? 0001?03111 1101?10310 1? jucunda Odozana 0010011211 1110000001 1?20000001 111??1?001 11100110?1 ??01?01??? 0011?03110 0000010310 00 floccosa Prepiella aurea 0010011201 1000100001 0011000201 010011?101 11101110?0 1011?11??? 110001?110 0001?10300 00 Prepiella 1021021211 1100100001 0011000210 ??1??00101 11100101?0 1011?10101 0111?00010 00000100?1 1? radicans Propyria 1010010101 1000000010 1?11000001 0001100101 00000110?1 ??01?11??? 2010000111 1100010101 01 morelosia Propyria 1010010101 1000000010 1?11001001 0000100101 00000110?1 ??01?11??? 2110003111 1101?10101 1?

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pytchoglene Pytchoglene 0010011201 1100000001 1?11000201 000111?101 11000110?0 1011?11??? 111011?010 1001?10211 00 coccinea Pytchoglene 0010010211 1000000001 1?11000001 0000100101 11000010?0 1001?11??? 100010310? 1100110200 00 erythrophora Pytchoglene 0010010211 1000000001 1?11000001 0000100101 110??010?0 1001?11??? 100010310? 1100110200 00 sanguineola Rhabdatomis 0010011101 1000000001 0011001301 100211?100 1101?110?1 ??00110100 2011?1?110 0101?000?1 1? coroides Rhabdatomis 0010011101 1000000001 0011001311 100211?100 11000110?0 0101?10101 0011?00110 0101?100?1 1? pueblae Talara coccinea 0010011201 1010100010 1?01001301 0100100101 11100110?1 ??01?01??? 0?11?01010 0101?10101 00 Talara 001???1201 1010100010 1?01001301 0100100101 11101110?1 ??11?01??? 0011?????? ?????????? ?? megaspila Setina aurita 101???0201 1000100001 010101?111 1000100100 1100000221 ??11?10101 11101????? ?????????? ?? Setina irrorella 1010011201 1010100001 010101?111 1000100100 1100000221 ??11?10101 1110103111 0000000301 00 Stigmatophora 1010011211 1100100001 010101?111 1000100100 1100000220 1010110100 1000100011 1100000301 00 flava Eudesmia arida 1011011211 1000100001 0001000201 000011?111 11001110?1 ??00111??? 011011?011 00000100?1 01 Eudesmia 1011011211 1000100001 0001000201 100011?111 11001110?1 ??00111??? 011011?011 10000100?1 01 praxis Eudesmia 1010011211 1000100001 0001000201 0000100111 10001110?1 ??00111??? 0101?0112? 0101?000?1 1? ruficollis Euryptidia ira 0010011201 1100100001 0001001201 000011?101 11001110?1 ??11?11??? 111011?110 00000100?1 01 Euryptidia 1011011211 1000100001 0001000201 0000100101 11001110?1 ??00111??? 1110101111 01000100?1 01 univitta Josioides 0010011211 1000100001 0001001211 1000100111 11001110?1 ??00111??? 1101?01111 00000100?1 01 celena Josioides 0010011211 1000100001 0001001211 1000100111 11001110?1 ??00111??? 1101?01110 00000100?1 01 myrrha

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Apistosia judas 1021011210 1100100001 1?0101?110 ??0211?100 0101?00210 0010011??? 0110101010 1101?100?1 1? Crambidia 0010011201 1110101101 1?11???011 0002100101 1101?00201 ??11?00001 0?11?1?110 1101?01??? 1? pallida Cybosia 0010011211 1010000001 010101?211 000111?101 1111?00201 ??11?00100 401011??11 0000010101 00 mesomella Eilema bicolor 001???1201 1110101001 1?11???211 0002000101 1110100221 ??01?00100 00101????? ?????????? ?? Eilema bipuncta 0010011211 1110001001 1?111??211 0001100101 1111?00220 1111?00111 001001?111 11001100?1 1? Eilema heimi 0010011211 1110101001 1?11000001 0002000101 1100010201 ??11?10100 0011?02111 0101?100?1 01 Gardinia 1021021211 1010100001 001101?311 100211?100 01000110?0 1111?11??? 1011?0310? 1101?10101 1? anoploa Gnamptonychia 1011011210 0100100001 1?11???111 100211?001 0101?00211 ??00011??? 0010103011 00000100?1 1? flavicollis Inopsis 1021020210 0100100001 010101?111 0002101001 0110100210 0010011??? 0010103011 11000100?1 1? modulata Lithosia quadra 1010011211 1000100001 1?1101?011 000201?101 1101?00201 ??11?00100 011000012? 0000101??? 1? Lithosia 0010011211 1010101001 1?1101?001 0002000101 1101?00201 ??11?00100 0010102011 1000111??? 01 sororcula Pelosia 0010011211 1110111001 1?1101?201 1002000101 1101?10210 1011?00001 1011?1?02? 0001?000?1 01 muscerda Asura cervicalis 1021020210 1010100001 0001001201 100211?100 11000101?1 ??11?00101 1000100011 0100011??? 00 Asura 0010011211 1110100001 0001000211 001??1?100 1101?101?1 ??11?00110 0000102111 1100010300 00 strigipennis Cyana 0010011211 1000110001 0011000101 000211?100 1100000221 ??11?10100 101011?011 0101?000?1 00 interrogationis Hemipsilia coa- 0010011211 1000110001 00011??211 101??1?100 11101101?1 ??01?00101 1010101010 0100000300 00 vestis Miltochrista 0010011211 1000100001 0001000101 000011?100 11001101?1 ??01?10110 1000101111 1100010300 00 flexuosa Miltochrista 0010011211 1000100001 0101000201 000011?111 11001101?1 ??01?10010 1000103111 11000100?1 00 gratiosa

277

Paidia murina 101???1211 1001?00001 0011000111 1002101100 1100000220 1111?10110 00001????? ?????????? ?? Phryganopteryx 0010011201 1?00100001 0101???201 010111?101 0101?10210 1110001??? 0001?03111 00000000?1 00 convergens Ardonea moria 1021011211 1010100001 1?01000311 1001100100 1101?110?0 1001?01??? 1111?03111 1100110101 1? Ardonea 0010011211 1010100001 1?00000311 001??1?100 10100110?0 1111?01??? 2011?0010? 0001?10101 00 peculiaris Balbura 0010011201 1?10100001 0000000311 011??1?101 0101?110?0 1111?11??? 1011?1?110 0001?11??? 00 intervenata Epeiromulona 1010011211 1000000001 1?11000101 100111?101 11101110?0 0001?00111 001011?110 00000100?1 1? icterinus Eurylomia 01?01?1201 0000100001 1?0101?110 ??02100100 0100000210 0010011??? 0110100110 11000100?1 1? cordula Heliosia 0010011201 1000000001 1?0101?110 ??01100100 11101101?0 1111?10111 1011?1?110 00000100?1 01 jacunda Oeonistis 1020011211 1100100001 010101?311 100011?101 1111?00221 ??10100101 0011?0010? 11000000?1 01 entella Paramulona 1021021201 1?00100001 1?11000101 000211?101 1101?010?1 ??01?10111 101011?110 0100110310 01 nephalistis Pagara simplex 1021021200 1100100001 0001???111 100211?101 0011?10221 ??11?10000 111001?111 0110101??? 01 Asota heliconia 1001001211 1100100000 0001002111 1101100101 1100?00201 ??11?10011 001001?02? 1110001??? 00

278

APPENDIX B CHARACTER DIAGNOSTICS FOR STRICT CONSENSUS TREE

Character Number Best Fit Worst Fit Tree Steps CI RI 0 1 38 22 0.045 0.432 1 1 6 2 0.500 0.800 2 2 12 11 0.182 0.100 3 1 20 16 0.062 0.211 4 1 6 2 0.500 0.800 5 2 7 7 0.286 0.000 6 1 10 9 0.111 0.111 7 2 14 4 0.500 0.833 8 1 39 24 0.042 0.395 9 1 5 4 0.250 0.250 10 1 3 2 0.500 0.500 11 1 21 16 0.62 0.250 12 1 22 16 0.062 0.286 13 1 3 3 0.333 0.000 14 1 30 14 0.071 0.552 15 1 3 3 0.333 0.000 16 1 6 2 0.500 0.800 17 1 1 1 1.000 0/0 18 1 12 6 0.167 0.545 19 1 13 7 0.143 0.500 20 1 35 18 0.056 0.500 21 1 10 9 0.111 0.111 22 2 33 22 0.091 0.355 23 1 6 5 0.200 0.200 24 1 3 3 0.333 0.000 25 1 16 9 0.111 0.467 26 2 18 12 0.167 0.375 27 3 41 38 0.097 0.378 28 1 37 25 0.040 0.333 29 1 5 4 0.250 0.250 30 1 27 19 0.053 0.308 31 1 11 10 0.100 0.100 32 1 10 6 0.167 0.444 33 2 39 27 0.074 0.324 34 1 7 4 0.250 0.500 35 1 35 24 0.042 0.324 36 3 4 4 0.750 0.000 37 1 9 6 0.167 0.375 38 1 6 3 0.333 0.600 39 1 21 16 0.062 0.250 40 1 23 14 0.071 0.409 41 1 11 9 0.111 0.200 42 1 22 20 0.050 0.095

279

43 1 28 24 0.042 0.148 44 1 22 12 0.083 0.476 45 1 20 13 0.077 0.368 46 1 33 16 0.062 0.531 47 2 33 17 0.118 0.516 48 3 15 10 0.300 0.417 49 1 25 20 0.050 0.208 50 1 5 3 0.333 0.500 51 1 8 8 0.125 0.000 52 1 38 21 0.048 0.459 53 1 16 8 0.125 0.533 54 1 7 4 0.250 0.500 55 1 28 23 0.043 0.185 56 1 31 16 0.062 0.500 57 1 7 5 0.200 0.333 58 1 12 8 0.125 0.364 59 1 14 13 0.077 0.077 60 3 48 30 0.100 0.400 61 1 32 20 0.050 0.387 62 1 18 13 0.077 0.294 63 1 23 19 0.053 0.182 64 1 17 10 0.100 0.438 65 1 26 23 0.043 0.120 66 3 32 26 0.115 0.207 67 1 17 15 0.067 0.125 68 2 11 10 0.200 0.111 69 1 28 21 0.048 0.259 70 1 32 22 0.045 0.323 71 1 31 27 0.037 0.133 72 1 2 1 1.000 1.000 73 1 23 22 0.045 0.045 74 1 12 8 0.125 0.364 75 1 25 18 0.056 0.292 76 1 7 6 0.167 0.167 77 3 35 23 0.130 0.375 78 1 8 7 0.143 0.143 79 1 13 8 0.125 0.417 80 1 31 22 0.045 0.300 81 1 16 10 0.100 0.400

280

APPENDIX C DATA MATRIX USED TO PRODUCE TREES BASED ON MOLECULAR DATA

[28S] Acsala anomala ------AAAATAGGACCAGGCATAGTTCACGTTTTTTCGGGTCCCAGCAGCGCGGCTCAGAGTG---CGACTACATT-CAC GGAGCAGAAACGAGACGCCTCGGGAGTGCGAGAGATCGACCTAGATCGACGCTCCATCCTCCCTGAGCGCGCGGC AAGCGCACGCCTTCACTTTCGTTGCGCCTTTCAGTTTTATCATCTCAATGACTCGCAT-ACATGCTAGACTCCTTGGTC CGTGTTTCAAGACGGGTCCTGCGAGTGCCCGAA------ACTGAATCATC-GCAGACAGAGA---CGCGCACAGTCCG A-GACTGCACGGCTGCAACAAC-AGACGCGT-CGCAC-TCACGTCCGCGCCAAGGCGGTACAAAGGATACGACGTTGA ---CTTGCGTCG--GGCCGGACGCGTTATGAACGCGTGCGCGTTTCGTCAGA-----ACTACCGTCCGACGGCCGGTCGGC CACCGTCGCGGTCCCACAGACCCCGTGAAGGATCCGTG--GGGCTCCCGGACGGC---GCTTAGACCGACATCGAACG GGTCGCG-ATGTAT-----TTACTAAGAGA-----GAAGTGCACGCCGTCCGCGGACGTCGACGGACGCGCGTCGTGCACG CCG------Agylla septentrionalis ------Asura cervicalis ----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCCG CGGCGTACTGGC-GCGCGCTTCGATGTCG------TCGGCCTCGGTCGGC------GCGCACGACGCGCGTCTGTC GACGTCCGCG-GTGGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCCATCGGACCCC--TCACCGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTG C-CGCCAAAGCGCG-----GACGCGAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTC TGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTA TGCGAGTCATTGAGATTATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGCGCGCGCTAGCCGCGCGCTCAGGGAG GATGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGCAGGCGC GCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Asura polyspila

281

------CGGGGTCTTT-TGGGACCG CGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG- ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC-CGCCTAAGCGCG-----GACGTGAGTGCGGCGCGTCTGT-TGTTGCCGC CGTGTCAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTG AAACACGGACCAAGGAGTCTAGCATGTATGCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGG CGCGCGCTCGCCGCGCGCTCAGGGAGGATGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGT CTCGTTTCCAATCCGTGAATGTAGGCGCGCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTA TGCCTGGTCAGCTTTA ----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCCT CGGCGTACGAAC-GCGCGCTTCGATGTCG------TCGGCTTCGGTCGGC------GTGCACGACGCGCGTCCGTCG ACGTCCGAG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GC CGTTCGGAAGCCCCAC-TGACCCT--TCACGGGGTC-TC-TGGGACCGGTATTGCGGTGGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTACC GCCTAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGC GAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGAT GGAG-CGTCGATCTCGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGCT CTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Bruceia hubbardi ------TAAAGCTGACCAGGCATAGTTCACCATCTTTCGGGTCCCAGCATTTATGCTCAGAGCG---CGCCTACATT-CAC GGATTGGAAACGAGACGCCTCGGGAGTGCGAGAGATCGACCTAGATCGACGCTCCATCCTCCCTGAGCGCGCGGC GAGCGCGCGCCTTCACTTTCGTTACGCCTTTCAGTTTTATTATCTCAATGACTCGCAT-ACATGCTAGACTCCTTGGTC CGTGTTTCAAGACGGGTCCTGCGAGTGCCCGAA------ACTGAATCATC-GCAGACAGAGA---CGCGCACAGTCCG A-GACTGCACGGCTGCAACAAC-AGACGCGC-CGCAC-CTGCGTCCGCTCTTGGGCGGTACGCAGGATACGACGTTGA ---CTTGCGTCG--GGCCGGACGCGTTGTAAACGCGTGCGCGTTTCGTCAGA-----ACTACCGTCCGACGGCCGGTCGGC CACCGTCGCGGTCCCACAGACCCCGTGAAGGATCCGTG--GGGCTCCCGAACGGC---GCTTAGACCGACATCGAACG GGTCGCG-ATGTAT-----TTACTAAGAGA-----GAAGTGCACGCCGTCCGCGGACGTCGACGGAAGAGCGTCGTGCACG CCGACCGAGATCGACGACATCGA-----AACGCGCTGCCGTACGCCGCGGAGTGACGATGAATCTC------TCCGTTCGT TCATTCGAGTTTCG------CAGGTTTACCCCTGAACG------GTTTCACGTACTCCCCTA------Bruceia pulverina

282

------ATTCATCGTCATTCCGCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCGCGTACGTCGACGTCCGCG-GACGGCGTGCACTTCT---CT CTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GCCGTTCGGGAGCCCCAT-TGTGCCT--TCACG GGTAT-AG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGCGCACGCGT-TTAC A-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTCACGTAC-CGCCTCAGCGCG-----GACGCGGGTGCGGCG CGTCTGCGTATCGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCAC TCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCGAGTCATTGAGACAATA--AAACTGAAAGG CG-CAACGAAAGTGAAGGCGCGCGCTAGCCGCGCGCTCAGGGAGGATGGAG-CGTCGATCTCGGTCGATCTCTCGC ACTCC------CGAGGCGTCTCGTTTCCAATC------Calamidia hirta ----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCCG CGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCGCGTACGTC GACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCCAT-TGATCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCACG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC- CGCCTAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTT-CAGTCTCGGACTGTGCGCGTCTCTGT ------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGC GAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGAT GGAG-CTTCGATCTAGGTCGTACTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGCT CTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Cisthene juanita GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CGCGGCGTACGGAC-GCGCGCTTCGATGTAG------TCGGCCTCGGTCGGC------TTGCACGACGCGCGTCCGT CGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGATCCT--TCACGGGATC-TT-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTAAA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTTCGTAC -CGCCTAAGCGCG-----GACGCAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTTGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGTCAGCTT— Cisthene plumbea

283

------TAAAGCTGACCAGGCATAGTTCACCATCTTTCGGGTCCCAGCATTTATGCTCAGAGCG---CGCCTACATT-CAC GGATTGGAAACGAGACGCCTCGGGAGTGCGAGAGATCGACCTAGATCGACGCTCCATCCTCCCTGAGCGCGCGGC AAGCGCGCGCCTTCACTTTCGTTACGCCTTTCAGTTTTATTATCTCAATGACTCGCAT-ACATGCTAGACTCCTTGGTC CGTGTTTCAAGACGGGTCCTGCGAGTGCCCGAA------ACTGAATCATC-GCAGACAGAGA---CGCGCACAGTCCG A-GACTGCACGGCAGCAACAAC-AGACGCGC-CGCAC-TTACGTCCGCGCTTAGGCGGTACGCAAGATACGACGTTGA ---CTTGCGTCG--GGCCGGACGCGTTATAAACGCGTGCGCGTTTCGTCAGA-----ACTACCGTCCGACGGCCGGTCGGC CACCGTCGCGGTCCCAAAGATCCCGTGAAGGATCCATG--GGGCTCCCGAACGGC---GCTTAGACCGACATCGAACG GGTCGCG-ATGTAT-----TTACTAAGAGA-----GAAGTGCACGCCGTCCGCGGACGTCGACGGACGCGCGTCGTGCGAG CCGACCGAGGCCGACTACATCGA-----AGCGCGCGTCCGTACGCCGCGGAATGACGATGAATCTC------TCCGTTCGT TCATTCGAGTTTCG------CAGGTTTACCCCTGAACG------GTTTCACGTACTCCCCTAT------ GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CGCGGCGTACGGAC-GCGCGGCTCGATGCCC------GCCGGTCTCGATCGGC------GGGCACGACGCGCGTCC GTCGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAT-----ACGACAT-CGCGACCCGTTCGATGTCGGTCTAAG C---GCCGCGCGGGAGTCCCGT-CGCCCCC--CCTCGGGGTGGTGATTGGACCGCTG---CGGTCGCCGACCGGCGGTC GGACGGTAGCTTCGACGAATCGCGCACGCGT-TTACACACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CCAA CGTCTACGCCCGAGTGCG-----GACGTAGGTGCGGCGCGTCTGT-CGTCGCAGCCGTG-CTGTCTCGGACTGTGCGCG TCTCTGT------CTGCGATGATTCTGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCA TGTATGCGAGTCATTGAGATGACAACAAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTAGCCGCGCGCTCA GGGAGGATGGAA-ATGCGGTCTAGGTCGCACTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGC AGGCGCGCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGTCAGCTT— Cisthene tenuifascia ------TTAAGCTGACCAGGCATAGTTCACCATCTTTCGGGTCCCAGCATTTATGCTCAGAGCG---CGCCTACATT-CAC GGATTGGAAACGAGACGCCTCGGGAGTGCGAGAGATCGACCTAGATCGACGCTCCATCCTCCCTGAGCGCGCGGC AAGCGCGCGCCTTCACTTTCGTTACGCCTTTCAGTTTTATTATCTCAATGACTCGCAT-ACATGCTAGACTCCTTGGTC CGTGTTTCAAGACGGGTCCTGCGAGTGCCCGAA------ACTGAATCATC-GCAGACAGAGA---CGCGCACAGTCCG A-GACTGCACGGCAGCAACAAC-AGACGCGC-CGCAC-TTACGTCCGCGCTTAGGCGGTACGCAAGATACGACGTTGA ---CTTGCGTCG--GGCCGGACGCGTTATAAACGCGTGCGCGTTTCGTCAGA-----ACTACCGTCCGACGGCCGGTCGGC CACCGTCGCGGTCCCACAGAACCCGTGAAGGATCCATG--GGGCTCCCGAACGGC---GCTTAGACCGACATCGAACG GGTCACG-ATGTAT-----TTACTAAGAGA-----GAAGTGCACGCCGTCCGCGGACGTCGACGGACGCGCGTCGTGCGAG CCGACCGAGGCCGACTACATCGA-----AGCGCGCGTCCGTACGCCGCGGAATGACGATGAATCTC------TCCGTTCGT TCATTCGAGTTTCG------CAGGTTTACCCCTGAACGGTTTCACGTACTCCCCTATAGTGAGTCGTATTAAA------Crambidia lithosioides

284

GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCACTC CCCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTACGT CGACGTCCGGG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGACCCT--TCACGGGGTA-CG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC -CGCCTCAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTTGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGTCAGCTT— Cyana meyricki ----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTTATTCCG CGGCGTACTGAC-GCGCGATTCGATGTCG------TCGGCTTAGGTCGGC------GCGCACGACGCGTGTCTGTCG ACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GC CGTTCGGGAGCCCCAT-GGATCCT--TCACGGGGTC-TA-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACGG TATTACTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAAAGTCGTAT-CTTACGTAC-CG CCTAAGCGCG-----GACGTGAGTGCGGCTCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCGA GTCATTGAGATTATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGCGTGCGCTCGTCGCGCGCTCAGGGAGGATGGA G-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATTCGTGAATGTAGGCGCGCTCTG AGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Cybosia mesomella ----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCCA CGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTCCGTCG ACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GC CGTTCGGGAGCCCCAT-GGACCCT--TCGCGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC- CGCCTAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Eilema bicolor

285

---AGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCACTCCT CGGCGTACGGAC-GCGCGGTTCGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTACGTCG ACGTCCGAG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GC CGTTCGGGAGCCCCAT-GGACCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACGG TAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC-C GCCTTAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCG AGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGATG GAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGCTC TGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Eilema complana ------GAACGAACGGAGAGATTCATCGTCACTCCTCGGCGTACGGAC-GCGCGCTT CGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTACGTCGACGTCCGTG-GACGGCGTGCA CTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GCCGTTCGGGAGCCCCAT-TGACC CT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGTGCACG CGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGTGTAC-CGCCTCAGCGCG-----GACGTAAGT GCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCG GGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCGAGTCATTGAGATAATA-- AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGATGGAG-CGTCGATCTAGGT CGACCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGCTCTGAGCATAAATGCTGG GA------Eilema dorsalis GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCACTC CTCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTCCGT CGACGTCCGGG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGACCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC -CGCCTAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGTCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Eilema griseola

286

GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCACTC CTCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTCCGT CGACGTCCGAG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAC-GGTCCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC -CGCCTAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Eilema plana ---AGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCACTCCT CGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGATCTCGGTCGGC------GTGCACGACGCGCGTCCGTCG ACGTCCTTG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GC CGTTCGGGAGCCCCAC-GGACCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TYATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC- CGCCCAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Eilema sp.1 GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CTCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCGCGTCCGT CGACGTCCGGG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGACCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC -CGCCTAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Eilema sp.2

287

GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCACCGTCATTC CGCGGCGTACTGGC-GCGCGTATCGATGTCG------TCGGCTTCGGTCGGC------GTGCACGACGCGCGTCTGA CGATGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCATTGGGCCCT--TCACGGGGTCTTT-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGA CGGTAGTTCTGACGAAACGCGCACGCGT-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTA C-CGCCTAAGCGCG-----GACGTGAGTGCGGCGCGTCTGT-TGTTGCTGCCGTG-CAGTCTCGGACTGTGCGCGTCTCT GT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTAT GCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGG ATGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCG CTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Eilema sp.3 GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCACTC CTCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTCCGT CGACGTCCGAG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGACCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC -CGCCTAAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCTGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTTGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Eilema sp.4 ------GAACGAACGGAGAGATTCATCGTCACTCCTCGGCGTACGGAC-GCGCGCTT CGATGTCG------TCGATCTCGATCGGC------GTGCACGACGCGCGTCCGTCGACGTCCGAG-GACGGCGTGCA CTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GCCGTTCGGGAGCCCCAT-GGAC CCT--TCGCGGGGTC-TA-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGCGCAC GCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC-CGCCTAAGCGCG-----GACGTAAG TGCGTCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTC GGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCGAGTCATTGAGATAATA--AAACT GAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGATGGAG-CGTCGATCTAGGTCGATC TCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGCTCTGAGCATAAATGCTGGGACCC GAAAG------ATGGTGAACTATGCCTGGT------Gardinia anoploa

288

GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CGCGGCGTACGTAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GTGCACGACGCGCGTCCGC CGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGATCCT--TCGCGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAGCGCGCACGCGT-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTCGCGTA C-CGCCTCAGCGCG-----GACGCGAGTGCGGCGCGTCTGC-TGTTGCTGCCGTG-CAGTCTCGGACTGTGCGCGTCTC TGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTA TGCGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAG GATGGAG-CGTCGATCTCGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCTGTGAATGTAGGCGC GCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Gnamptonychia flavicollis ------GAACGGAGAGATTCACCGTCATTCCTCGGCGTACGGAC-GCGCGCTTCGA TGTCG------TCGGCTTAGGTCGGC------GTGCACGACGCGCGTACGTCGACGTCCGAG-GACGGCGTGCACTT CT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GCCGTTCGGGAGCCCCAT-GGACCCT- -TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGCGCACGCG T-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC-CGCCTAAGCGCG-----GACGTAGGTG CGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGG GCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCGAGTCATTGAGATAATA—AAACTGA AAGGCG-CAACGAAAGTGAAGGCGCGCGCTTGCCGCGCGCTCAGGGAGGATGGAG-CGTCGATCTAGATCGATCTC TCGCACTCC------CGAGGCGTCTCGTTTCCAATC------Heliosia jucunda GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CGCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCCACCTCGGTGGGC------GTGCACGACGCGCGTACGT CGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTCCGGGAGTCCCAT-GGTTTTCCTTCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGA CGGTAGTTCTGACGAAGCGCGCACGCGT-TCCAA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTAACGT AC-CGCCTCAGCGCG-----GACGTTGGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTC TGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTA TGCGAGTCATTGAGAATATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGTGCGCGCTTGCCGCGCGCTCAGGGAGG ATGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCTGTGAATGTAGGCGCG CTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Hiera gyge

289

------CGGGGTC-TG-TGGGACCG CGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG- ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC-CGCCTAAGCGCG-----GCCGTAAGTGCGGCGCGTCTGT-TGTTGCAGC CGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTG AAACACGGACCAAGGAGTCTAGCATGTGTGCGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAG GCGCGCGCTCGCCGCGCGCTCAGGGAGGATGGAG-CGTCGATCTAGGTCGATCTTTCGCACTCC------CGAGG------Hypoprepia cadaverosa ------CATTCCGCGGCGTACGGAC-GCGCGCTTCGATGTCG------T CGGCCTCGGTCGGC------GTGCACGACGCGCGTACGTCGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTA GTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---GCCGTTCGGGAGCCCCAT-TGTGCCT--TCACGGGT AT-AG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGCGCACGCGT-TTACA-AC GCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTCACGTAC-CGCCTCAGCGCG-----GACGCGGGTGCGGCGCGT CTGCGTATCGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCACTCG CAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCGAGTCATTGAGACAATA--AAACTGAAAGGCG- CAACGAAAGTGAAGGCGCGCGCTAGCCGCGCGCTCAGGGAGGATGGAG-CGTCGATCTCGGTCGATCTCTCGCACT CC------CGAGGCGTCTCGTTTCCAA------Hypoprepia fucosa GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CTCGGCGTACGTAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GCGCACGACGCGCGTCCGT CTACGTCCGCG-GACGGCGTGCACTTCT---CTCTCAGTAT-----GT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAC-GGATCCT--TTACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAATCGCGCACGCGT-TTACG-ACGCGTCCGGCCCG-ACGCAAG-TCCAACGTCGTAT-CTTTCGTA C-CGCCCAAGCGCG-----GACGAGAGTGCGGCGCGCCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTC TGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTA TGCGAGTCATTGAGACAATT--AAACTGAAAGGCG-TAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAG GATGGAG-CGTCGATCTCGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCTGTGAATGTAGGCGC GCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGTCAGCTT— Hypoprepia fucosa tricolor GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CTCGGCGTACGTAC-GCGCGCTTCGATGTCG------TCGACCTCGGTCGGC------GCGCACGACGCGCGTCCGT CTACGTCCGCG-GACGGCGTGCACTTCT---CTCTCAGTAT-----GT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---

290

GCCGTTCGGGAGCCCCAC-GGATCCT--TTACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAATCGCGCACGCGT-TTACG-ACGCGTCCGGCCCG-ACGCAAG-TCCAACGTCGTAT-CTTTCGTA C-CGCCCAAGCGCG-----GACGAGAGTGCGGCGCGCCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTC TGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTA TGCGAGTCATTGAGACAATT--AAACTGAAAGGCG-TAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAG GATGGAG-CGTCGATCTCGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCTGTGAATGTAGGCGC GCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Inopsis modulata GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCACCGTCATTC CTCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCTGCTTAGGTCGGC------GTGCACGACGCGCGTCCGT CGACGTCCGAG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGACCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC -CGCCTAAGCGCG-----GACGTAGGTGCGGCGCGTCTGT-TGTTGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGTCAGCTT— Lithosia quadra --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCC TCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCGCGTCCGTC GACGTCCGGG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCAAC-GGACCCT--TCACGGGGTC-TG-TTGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC- CGCCTCAGCGCG-----GACGTAAGTGCGGCGCGTCTGT-TGTTGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Lyclene pyraula --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCC GCGGCGTACTGGC-GTGCGATTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCACGTCTGTC GACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAT-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G

291

CCGTCCGGGAGCCCCATTGGACCCC--TCACGGGGTCTGG-TGGGACCGAGA---CGGTGGCCGACCGGCCGTCGGA CGGTAGTTCTGACGAAACGCGCACGCGT-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTACGTA C-CGCCTTAGCGCG-----GACGTGAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCT GT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTAT GCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGTGTGCGCTTGCCGCACGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Lyclene reticulata ------Lyclene sp.1 --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCC GCGGCGTACTGGC-GTGCGATTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCACGTCTGTC GACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAT-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTCCGGGAGCCCCATTGGACCCC--TCACGGGGTCTGG-TGGGACCGAGA---CGGTGGCCGACCGGCCGTCGGA CGGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTACGTA C-CGCCTTAGCGCG-----GACGTGAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCT GT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTAT GCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGTGTGCGCTTGCCGCACGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Lyclene sp.1 nr. --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCC GCGGCGTACTGGC-GTGCGATTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCACGTCTGTC GACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAT-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTCCGGGAGCCCCATTGGACCCC--TCACGGGGTCTGG-TGGGACCGAGA---CGGTGGCCGACCGGCCGTCGGA CGGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTACGTA C-CGCCTTAGCGCG-----GACGTGAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCT GT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTAT GCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGTGTGCGCTTGCCGCACGCTCAGGGAGGA

292

TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCC------Lycomorpha pholus GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CGCGGCGTACGGAC-GCGCGCTTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCGTTTCCGT CGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCAT-GGATCCT--TCACGGGATC-TT-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC -CGCCAAAGCGCG-----GACGCAAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGATGATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGCGCGCGCTTGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGTCAGCTT— Lycomorphodes sordida ------CGGGATC-TT-TGGGACCG CGA---CGGTGGCCGACCGGCCGTCGGACGGTAGTTCTGACGAAACGCGCACGCGT-TTACA-ACGCGTCCGGCCCG- ACGCAAG-T-CAACGTCGTAT-CTTGCGTAC-CGCCAAAGCGCG-----GACGCAAGTGCGGCGCGTCTGT-TGTTGCAGC CGTG-CAGTCTCGGACTGTGCGCGTCTCTGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTG AAACACGGACCAAGGAGTCTAGCATGTATGCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGG CGCGCGCTTGCCGTGCGCTCAGGGAGGATGGAG-CTTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTC TCGTTTCCAATCCGTGAATGTAGGCGCGCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTAT GCCTGGTCAGCTTTA Meterhythosia sangala ------TAAAACGGACCAAGCATAGTTCACCATCTTTCGGGTCCCAGCATTTATGATCAGAGCG---CGCCTACATT-CATG GATTGGAAACGAGACGCCTCGGGAGTGCGAGAGATCGACCTAGATCGACGCTCCATCCTCCCTGAGCGCGCGGCA AGCGCGCGCCTTCACTTTCGTTACGCCTTTCAGTTTTATCATCTCAATGACTCGCAT-ACATGCTAGACTCCTTGGTCC GTGTTTCAAGACGGGTCCTGCGAGTGCCCGAA------ACTGAATCATC-GCAGACAGAKA---MRCGCACAGTCCGA- GACTGCACGGCTGCAACAAC-AGACGCGC-CGCAC-TTGCGTCCGCGCTTTGGCGGTACGCAAGATACGACGTTGA--- CTTGCGTCG--GGCCGGACGCGTTGTAAACGCGTGCGCGTTTCGTCAGA-----ACTACCGTCCGACGGCCGGTCGGCC ACCGTCGCGGTCCCAAAGATCCCGNGAAGGATCCATG--GGGCTCCCGAACGGC---GCTTAGACCGACATCGAACGG GTCGCG-ATGTAT-----TTACTAAGAGA-----GAAGTGCACGCCGTCCGCGGACGTCGACGGAAACGCGTCGTGCACGCC

293

GACCGAGGCCGACGACATCGA-----AGCGCGCGTCCGTACGCCGCGGAATGACGATGAATCTC------TCCGTTCGTTC ATTCGAGTTTCG------CAGGTTTACCCCTGAACG------GTTTCACGTACTCCCCTATAGTGA--- Miltochrista miniata ------Miltochrista sp.1 --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCC GCGGCGTACTGGC-GCGCGTTTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGGCGCGCGTCTGG AGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC--- GCCGTTCGGGAGCCCCATTGGGCCCT--TCACGGGGTCTTT-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGA CGGTAGTTCTGACGAAACGCGCACGCGT-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTGCGTA C-CGCCTTAGCGCG-----GACGTGAGTGCGGCGCGTCTGT-TGTTGCTGCCGTG-CAGTCTCGGACTGTGCGCGTCTCT GT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTAT GCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGCGTGCGCTTGCCGCGCGCTCAGGGAGG ATGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCG CTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Prepiella aurea nr. ------TAAAGCTGACCAGGCATAGTTCACCATCTTTCGGGTCCCAGCATTTATGCTCAGAGCG---CGCCTACATT-CAC GGATTGGAAACGAGACGCCTCGGGAGTGCGAGAGATCGACCTAGATCGATGCTCCATCCTCCCTGAGCGCGCGGC GAGCGCGCGCCTTCACTTTCGTTGCGCCTTTCAGTTTTATTATCTCAATGACTCGCAT-ACATGCTAGACTCCTTGGTC CGTGTTTCAAGACGGGTCCTGCGAGTGCCCGAA------ACTGAATCATC-GCAGACAGAGA---CGCGCACAGTCCG A-GACTGCACGGCTGCAACAAC-AGACGCGC-CGCAC-CTRCGTCCGCRCTKRGGCGGTACGCAAGATACGACGTTGA ---CTTGCGTCG--GGCCGGACGCGTTGTAAACGCGTGCGCGTTTCGTCAGA-----ACTACCGTCCGACGGCCGGTCGGC CACCGTCGCGGTCCCAMAGAYCCCGNGAAGGATCCGTG--GGGCTCCCGAACGGC---GCTTAGACCGACATCGAACG GGTCGCG-ATGTAT-----TTACTAAGAGA-----GAAGTGCACGCCGTCCGCGGACGTCGACGGAAGAGCGTCGTGCACG CCKACCGAGATCGACGACATCGA-----AACGCGCTGCCGTACGCCGCGGAGTGACGATGAATCTC------TCCGTTCGT TCATTCGAGTTTCG------CAGGTTTACCCCTGAACG------GTTTCACGTACTCCCCTATAGTGAG— Prinasura quadrilineata --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCC GCGGCGTACTGGC-GTGCGATTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCACGTCTGTC

294

GACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAT-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTCCGGGAGCCCCATTGGACCCC--TCACGGGGTCTGG-TGGGACCGAGA---CGGTGGCCGACCGGCCGTCGGA CGGTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CTTACGTA C-CGCCTTAGCGCG-----GACGTGAGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCT GT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTAT GCGAGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGTGTGCGCTTGCCGCACGCTCAGGGAGGA TGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Ptychoglene coccinea ------TAAAGGTGACCCGGCATAGTTCACCATTTTTCGGGTCCCAGCATTTATGGTCAGAGCG---GGCCTACATT-CAC AGATTGGAAACGAGACGCCTCGGGAGTGCGAGAGATCGACCGAGATCGACGCTCCATCCTCCCTGAGCGCGCGAC TAGCGCACGCCTTCACTTTCGTTACGCCTTTCAGTTTTATWATCTCAATGACTCGCAT-ACATGCTAGACTCCTTGGTC CGTGTTTCAAGACGGGTCCTGCGAGTGCCCGAA------ACTGAATCATC-GCTGACAGAGA---CGCGCACAGTCCGA -GACTGCACGGCTGCAACAAC-AGGCGCGC-CGCACTTTGCGTCCGCGCTGAGGGGGTATAAAGGATACGACGTTGA- --CTTGCGTCG--GGCCGGACGCGTCGTAAACGCGTGCGCGATTCGTCAGA-----ACTACCGTCCGACGGCCGGTCGGC CACCGTCGCGGTCCCACAGATCCCGTGAAGGATCCRTG--GGGCTCCCGAACGGC---GCTTAGACCGACATCGAACG GGTCGCG-ATGTAY-----WTACTAAGAGA-----GAAGTGCACGCCGTCCGCGGACGCMGACGGACGCGCGTCGTGCGC GCCGACCGAGGCCGACGACATCGA-----AGCGCGCGTCCGTACGCCGCGGAATGACGATGAATCTC------TCCGTTC GTTCATTCGAGTTTCG------CAGGTTTACCCCTGAACG------GTTTCACGTACTCCCCTATAGTGAG-- Schistophleps albida --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCC GCGGCGTACGGAC-GGGCGGTTCGATGTCG------TCGGCCTCGGTCGGC------GTGCACGACGCTCGTACGTC GACGTCCACG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCCAT-GGACCCT--TCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCACG-ACGCAAG-T-CAACGTCGTAT-CTTACGTAC-C GCCTAAGTGCG-----GACGTGAGTGCCGCGCGTCTGT-TGTCGCCGCCGTG-CAGTCTCGGACTGTGCGTGTCTCTGT------CAGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATGCG AGTCATTGAGATAATA--AAACTGAAAGGCG-TAACGAAAGTGAAGGTTCGCGCTTGCCGCGTACTCAGGGAGGATGG AG-TGTCGATCTAGGTCGACCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCTGTGAATGTAGGCGCGCTCT GAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------Amata aperta ----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTCCG CGGCGTACGGGCTGCGCGGTTCGATGTCG------TCGACCTCGGTCGTC------AGGCACGACGCGCGTCCGTC

295

GACGTCCGCG-GAGGGCGTGCACTTCT---CTCTTAGTAA-----AT-GCAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCCGC-GGTCCCT--TCACGGGGTC-CG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGATGAAGCGCGCACGCGT-TTACA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CCAACGTA C-CGCCTCAGCGCG-----GACGTAGGTGCGGCGCGTCTGC-TGTTGCCGCCGTG-CAGTCTCGGACTGTGCGCGTCTC TGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTA TGCGAGTCATTGAGACAATA--AAACTGAAAGGCG- CAACGAAAGTGAAGGTGCGCGCTAGACGCGCGCTCAGGGAGGATGGAG- CTTCGATCTAGGTCGTACTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGC GCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Pagara simplex GGGAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCGTCATTC CTCGGCGTACGGGCTGCGCGATTCGATGTCG------TCGACCTCGGTCGTC------AGGCACGACGCGCGTCCG TCGACGTCCGCG-GACGGCGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC-- -GCCGTTCGGGAGCCCCAT-AGTCCCCGGTCACGGGGTC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGG ACGGTAGTTCTGACGAAACGCGCACGCGT-TTTTA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CCAACGT AC-CGCCTCAGCGCG-----GACGTAGGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTC TGT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTA TGCGAGTCATTGAGATAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAG GATGGAG-CATCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGC GCTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGTCTGCTT-- Asota heliconia ----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCTTCATTCCC CGGCGTACGGAC-GCGCGGTTCGATGTCG------TCGGCCTCGGTCGGC------CGGCACGACGCGCGCACGTC GACGTCCGTG-GAGGTTGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCCGC-TCACCCT--TCGCGGGGTG-TG-TGGGACCGCGA---CGGTTGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CCAACGTAC- CGCCTCAGCGCG-----GACGTAGGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGACAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTCGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTCGATCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Asota orbona

296

----GTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCTTCATTCCC CGGCGTACGGGC-GCGCGGTACGATGTCG------TCGGCCTCGGTCGGC------CGGCACGACGCGCGCACGTC GACGTCCGGG-GAGGTTGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCCGCTTACCCTT--TCACGGGGTA-TG-TGGGACCGCGA---CGGTTGCCGACCGGCCGTCGGACG GTAGTTCTGACGAAACGCGCACGCGT-TTATA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTAT-CCAACGTAC- CGCCTCAGCGCG-----GACGTAKGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCTG T------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTATG CGAGTCATTGAGACAATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTTGCCGCGCGCTCAGGGAGGA TGGAG-CGTCGATCTCGATCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCGC TCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGC------Neochera dominia --GAGTACGTGAAACCGTTCAGGGGTA------AACCTGCGAAACTC-GAATGAACGAACGGAGAGATTCATCTTCATTCC GCGGCGTACGGAC-GCGCGATTCGATGTCG------TCTACTTCGGTCGGC------CGGCACGACGTGCGTACGTC GACGTCCGCG-GAGGTAGTGCACTTCT---CTCTTAGTAA-----AT-ACAT-CGCGACCCGTTCGATGTCGGTCTAAGC---G CCGTTCGGGAGCCCCAC-GGACCCC--TTACGGGGCC-TG-TGGGACCGCGA---CGGTGGCCGACCGGCCGTCGGAC GGTAGTTCTGACGAAACGCGCACGCGT-TTTCA-ACGCGTCCGGCCCG-ACGCAAG-T-CAACGTCGTATCCTTGCGTA C-CGCCTCAGCGCG-----GACGTAGGTGCGGCGCGTCTGT-TGTTGCAGCCGTG-CAGTCTCGGACTGTGCGCGTCTCT GT------CTGCGATGATTCAGTTTCGGGCACTCGCAGGACCCGTCTTGAAACACGGACCAAGGAGTCTAGCATGTAT GCGAGTCATTGAGATTATA--AAACTGAAAGGCG-CAACGAAAGTGAAGGCGCGCGCTTGCCGCACGCTCAGGGAGG ATGGAG-CGTCGATCTAGGTCGATCTCTCGCACTCC------CGAGGCGTCTCGTTTCCAATCCGTGAATGTAGGCGCG CTCTGAGCATAAATGCTGGGACCCGAAAG------ATGGTGAACTATGCCTGGT------

[COI] Acsala anomala ------Agylla septentrionalis ------TCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCTGGAATAGTAGGAAC ATCCTTAAGATTATTAATTCGAGCTGAATTAGGAAATCCAGGATCTTTAATTGGAGACGATCAAATTTATAATACTATTG TAACTGCCCATGCTTTTATTATAATTTTTTTTATAGTTATACCCATTATAATTGGAGGATTTGGTAATTGATTAGTACCTC

297

TTATATTAGGAGCCCCAGATATAGCATTCCCACGAATAAATAATATAAGTTTTTGACTACTTCCCCCCTCTTTAACTCTT CTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGTACAGGATGAACAGTTTATCCCCCACTTTCATCAAATATTGCCC ATAGAGGTAGTTCAGTAGACTTAGCTATTTTCTCATTACATCTAGCAGGTATTTCTTCAATTTTAGGAGCTATTAATTTT ATTACCACAATTATTAACATACGATTAAATAAATTAATATTTGATCAAATACCTTTATTTGTATGAGCAGTTGGTATTACA GCATTTTTACTTCTTCTTTCACTACCAGTTTTAGCTGGAGCAATTACTATATTATTAACTGATCGAAATCTTAATACTTCA TTTTTTGATCCTGCAGGAGGGGGAGATCCCATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTTACTT T------Asura cervicalis ------TATAGGGATTCAACCAATCATAAAGATATTGGAACTTTATATTTTATTTTTGGAATTTGAGCTGGAATAGTAG GAACCTCTTTAAGTTTACTAATTCGAGCTGAATTAGGAAACCCAGGATCTTTAATTGGGGATGATCAAATTTATAATAC TATTGTTACTGCACATGCTTTTATTATAATTTTTTTTATAGTAATACCAATTATAATCGGAGGATTTGGAAATTGGTTGGT ACCCCTTATATTAGGAGCTCCTGATATAGCTTTCCCCCGAATAAATAATATAAGATTTTGATTATTACCCCCCTCATTAA CATTGCTAATTTCAAGAAGAGTTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTACCCCCCACTTTCATCTAATA TTGCTCATAGTGGAAGATCAGTAGATTTAGCTATTTTTTCTCTTCATTTAGCGGGTATTTCATCAATTTTAGGAGCAATT AATTTTATTACTACCATTATTAACATACGATTAAATTCACTATCATTCGATCAAATACCATTATTTGTTTGAGCTGTAGGT ATTACAGCATTTTTATTACTTTTATCTTTACCTGTTTTAGCAGGAGCTATTACTATATTACTAACTGATCGAAATTTAAAT ACTTCTTTTTTTGATCCAGCTGGAGGAGGTGATCCAATTTTATATCAACATTTATTTTGATTTTTT------Asura polyspila ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCTGG AATAGTAGGAACTTCATTAAGTTTATTAATTCGAGCCGAATTAGGTAATCCAGGTTCTTTAATTGGAGATGATCAAATTT ATAATACTATTGTAACTGCTCATGCCTTTATTATAATTTTTTTTATAGTAATACCAATTATAATTGGAGGATTTGGAAATT GATTAGTACCTTTAATACTTGGAGCCCCTGATATAGCTTTCCCCCGAATAAATAATATAAGTTTTTGACTCCTACCTCCT TCTCTAACTTTATTGATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACTGGATGAACAGTTTATCCACCTTTATCAT CTAATATTGCTCATAGAGGTAGTTCCGTAGATTTAGCTATTTTTTCATTACATTTAGCTGGAATTTCATCAATTTTAGGA GCAATTAATTTTATTACAACTATTATTAATATACGACTTAATAGATTAACCTTTGATCAAATACCTTTATTTGTTTGAGCT GTAGGAATTACAGCATTTCTTCTTCTTCTTTCATTACCTGTTCTAGCAGGAGCTATTACTATATTATTAACTGATCGAAA TTTAAATACTTCATTTTTTGATCCTGCAGGTGGAGGTGATCCAATTTTATATCAACATTTATTTTGATTTTTTGGACATCC AGAAGTTTACTTT------Atolmis rubricollis ------

298

------Bruceia hubbardi ------TTAAGATTATTAATTCGAGCTGAATTAGGAAA TCCGGGATCTTTAATTGGAGATGATCAAATTTATAATACTATTGTAACTGCACACGCTTTTATTATAATTTTTTTTATAGT TATACCTATTATAATTGGAGGATTTGGTAATTGATTAGTACCTCTTATACTAGGTGCTCCTGATATGGCATTCCCACGA ATAAATAACATAAGTTTTTGATTATTACCACCATCTCTTATTCTTTTAATCTCAAGAAGAATTGTAGAAAATGGAGCAGG AACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATTGCTCATAGTGGAAGATCAGTAGATTTAGCTATTTTTTCTT TACACTTAGCTGGAATTTCATCAATCCTAGGAGCTATTAATTTCATTACAACAATTATTAATATACGATTAAATAATTTAT CATTTGATCAAATACCTTTATTTGTATGAGCTGTAGGAATTACTGCATTTTTATTATTACTTTCATTACCTGTTCTAGCTG GAGCTATTACTATACTTTTAACAGACCGAAACTTAAATACTTCATTTTTAAATCCTTTA------ATACGAC TCACTATAGGGATCAACCAATCA------Bruceia pulverina ------TATTTTATTTTTGGTATCTGAGCGGGAATAGTAGGAACATCTTTAAGTTTATT AATTCGAGCTGAATTAGGAAATCCAGGATCTTTAATTGGAGATGATCAAATTTATAATACTATTGTAACTGCTCATGCTT TTATCATAATTTTTTTTATGGTTATACCTATTATAATTGGAGGATTTGGTAATTGATTAGTACCTCTCATATTAGGTGCCC CTGATATAGCATTTCCACGAATAAATAATATAAGTTTTTGATTATTACCACCATCTCTTATTCTTTTAATTTCAAGAAGAA TCGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATTGCCCATAGAGGTAGATCAG TAGATTTAGCTATTTTTTCATTGCATTTAGCCGGAATTTCATCAATCTTAGGTGCTATTAATTTTATTACAACAATTATTA ATATACGATTAAATAATTTATCATTTGATCAAATACCTTTATTTGTATGAGCTGTAGGGATTACTGCATTTTTACTACTAC TCTCATTACCTGTTTTAGCGGGAGCTATCACTATACTTTTAACAGATCGAAAT------Calamidia hirta ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGTATTTGAGCAGGTATAGTAG GAACTTCATTAAGTTTATTAATTCGAGCTGAATTAGGAAATCCAGGATCTTTAATTGGAGATGATCAAATTTATAATACT ATTGTAACTGCCCATGCTTTTATTATAATTTTCTTTATAGTTATACCTATTATAATTGGAGGATTTGGAAATTGATTAGTT CCCCTAATATTAGGTGCTCCCGATATAGCATTCCCTCGAATAAATAATATAAGATTTTGATTGCTTCCCCCTTCATTAAC CCTTTTAATTTCAAGAAGAATCGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTATCCCCCACTTTCATCAAATATC GCTCATAGTGGTAGTTCTGTTGATTTAGCTATTTTTTCTTTACATTTAGCTGGTATTTCTTCTATTTTAGGAGCTATTAAT TTTATTACAACAATTATTAATATACGATTAAATAAATTAATATTTGATCAAATACCTTTATTCGTTTGAGCTGTCGGAATT ACAGCATTTTTACTATTACTCTCCCTACCTGTTTTAGCAGGAGCTATTACCATATTATTAACAGATCGAAATCTTAATAC TTCCTTTTTTGACCCCGCAGGTGGAGGT------Cisthene juanita

299

------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGTATTTGAGCTGGAATAGTAG GAACTTCTCTAAGATTATTAATTCGAGCTGAATTAGGTAATCCTGGTTCTTTAATTGGTGATGATCAAATTTATAATACT ATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATAGTAATACCTATTATAATTGGAGGATTTGGTAACTGATTAGTA CCATTAATATTAGGGGCCCCTGATATAGCTTTCCCACGAATAAATAATATAAGTTTTTGACTTCTTCCCCCGTCTTTAAC TTTATTAATTTCAAGAAGAATTGTAGAAAATGGTGCAGGAACAGGATGAACAGTTTATCCCCCACTTTCATCTAATATT GCTCATGGAGGTAGATCAGTTGATCTAGCTATTTTTTCCCTACATTTAGCTGGAATTTCTTCAATTTTAGGAGCTATTAA TTTCATTACCACTATTATTAATATACGATTAAATAGACTATCATTTGATCAAATACCTTTATTCGTTTGAGCTGTAGGAAT TACAGCATTTTTATTATTACTTTCTTTACCAGTTTTAGCCGGAGCTATTACTATATTATTAACAGATCGAAATTTAAATAC ATCTTTCTTTGACCCTGCGGGAGGAGGAGATCCAATCCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTT TAC------Cisthene plumbea ------TCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGTATTTGAGCTGGTATAGTAGGAAC TTCCTTAAGACTATTAATTCGAGCTGAATTAGGTAATCCAGGGTCTTTAATTGGTGATGATCAAATTTATAACACAATC GTAACTGCTCACGCTTTTATTATAATTTTTTTTATAGTAATACCTATTATAATTGGAGGATTTGGTAATTGATTAGTACCA TTAATATTAGGAGCCCCTGATATAGCTTTCCCACGAATAAATAATATAAGTTTTTGACTTTTACCACCATCTTTAACTTT ATTAATTTCAAGAAGAATTGTAGAAAACGGTGCAGGAACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATTGCT CATGGAGGTAGATCAGTTGATCTCGCTATTTTTTCCCTTCATTTAGCCGGTATTTCATCAATTTTAGGAGCTATTAATTT TATTACTACAATTATCAATATACGATTAAATAAATTATCATTTGATCAAATACCTTTATTTATTTGAGCTGTGGGAATTAC TGCATTTTTATTATTACTTTCCTTACCAGTATTAGCAGGAGCTATTACCATACTTCTAACAGATCGAAATTTAAATACAT CTTTTTTTGAT------Cisthene subjecta ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTCGGTATTTGATCAGGAATAGTAG GAACTTCTCTTAGATTATTAATTCGAGCAGAATTAGGTAATCCTGGGTCATTAATTGGAGATGATCAAATTTATAATACT ATTGTAACAGCTCATGCTTTTATTATAATTTTTTTTATAGTAATACCAATTATAATTGGTGGATTTGGAAATTGATTAGTA CCTTTAATATTAGGAGCTCCTGATATAGCTTTCCCCCGAATAAATAATATAAGATTTTGATTATTACCACCATCATTAAC TTTACTTATTTCCAGAAGAATTGTAGAAAATGGAGCTGGAACAGGATGAACAGTTTATCCCCCACTTTCATCTAATATT GCTCATGGAGGAAGATCAGTCGATTTAGCTATTTTTTCTTTACATTTAGCTGGTATTTCATCAATTTTAGGAGCAATCAA TTTCATTACTACAATTATTAATATACGATTAAATAATTTATCTTTTGATCAAATACCTCTATTTGTTTGAGCTGTTGGTATT ACAGCATTTTTATTATTACTTTCTTTACCAGTATTAGCAGGAGCTATTACTATATTATTAACAGATCGTAACTTAAATACA TCATTTTTTGATCCTGCAGGAGGAGGAGATCCAATCTTATATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTTA C------Cisthene tenuifascia

300

------Crambidia lithosioides ------Cyana meyricki ------AGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCAGGAATAGTAGG TACTTCTTTAAGATTATTAATTCGAGCAGAATTAGGAAATCCTGGATCTTTAATTGGAGATGACCAAATTTATAATACTA TTGTTACAGCCCATGCTTTTATTATAATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTTGGTAATTGATTAGTAC CTTTAATATTAGGAGCTCCTGATATAGCTTTCCCCCGTATAAATAATATAAGTTTTTGATTATTACCCCCTTCATTAACT CTTCTTATTTCAAGAAGAATTGTAGAAAATGGAGCAGGTACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATCG CTCATAGAGGAAGATCAGTAGATTTAGCTATTTTTTCTCTTCATTTAGCTGGAATTTCATCAATTTTAGGAGCTATTAAT TTTATTACCACAATTATTAATATACGACTTAATAACTTATCTTTTGATCAAATACCTTTATTTGTTTGAGCAGTTGGAATT ACTGCATTTTTATTACTTTTATCTTTACCAGTTTTAGCAGGTGCTATTACTATATTACTAACTGATCGAAATTTAAATACA TCCTTCTTTGATCCTGCAGGAGGAGGAGATCCTATTCTCTACCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTT AC------Cybosia mesomella ------Eilema bicolor ------TCAACCAATCATAAAGATATTGGAACACTATATTTTATTTTTGGAATTTGGGCAGGTATAGTAGGAAC TTCCCTTAGATTATTAATTCGAGCAGAATTGGGTAATCCTGGATCATTAATTGGAGATGATCAAATTTATAATACTATTG TAACTGCTCATGCTTTTATTATAATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTTGGAAATTGATTAGTTCCCC TTATATTAGGAGCCCCTGACATAGCTTTCCCCCGAATAAATAACATAAGTTTTTGACTACTCCCCCCCTCTTTAACATT

301

ACTCATCTCAAGTAGAATCGTAGAAAATGGGGCAGGAACAGGATGAACAGTTTATCCCCCACTTTCATCTAATATTGC TCATAGAGGTAGTTCTGTAGACTTAGCTATTTTTTCTCTACATTTAGCAGGTATTTCTTCTATTTTAGGAGCTATTAATTT TATTACAACAATTATTAATATACGATTAAATAAATTAATATTTGATCAAATACCTTTATTTGTTTGAGCTGTAGGTATTAC AGCATTTTTATTGCTTCTTTCATTGCCTGTTTTAGCAGGAGCTATTACTATATTATTAACTGATCGAAATCTTAATACTTC TTTTTTTGACCCTGCAGGAGGGGGAGATCCTATCCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTTACT TT------Eilema complana ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTCATTTTTGGAATTTGAGCTGGAATAGTA GGAACTTCACTTAGATTATTAATTCGAGCAGAATTAGGTAATCCTGGATCTTTAATTGGTGATGATCAAATTTATAATAC TATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTTGGAAATTGATTGAT CCCCCTTATATTAGGGGCCCCTGATATAGCTTTCCCCCGAATAAATAATATAAGTTTTTGACTACTCCCCCCCTCTTTA ACATTACTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTATCCCCCACTTTCATCTAATA TTGCTCATAGAGGTAGTTCTGTAGACTTAGCTATTTTTTCTTTGCATTTAGCAGGTATTTCCTCTATTCTAGGAGCTATT AATTTTATTACAACAATTATTAATATACGATTAAATAAACTAATATTTGATCAAATACCTTTATTTGTATGAGCTGTAGGT ATTACAGCATTCTTATTACTTCTTTCACTACCTGTTTTAGCAGGAGCTATCACCATATTATTAACCGATCGAAATCTTAA CACTTCTTTCTTTGATCCTGCAGGAGGAGGAGATCCAATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAG TTTACTTT------Eilema dorsalis ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCTGGTATAGTAG GAACATCTTTAAGATTATTAATTCGAGCTGAATTAGGAAATCCAGGTTCTTTAATTGGAGATGATCAAATTTATAATACC ATTGTAACTGCTCATGCTTTTATTATAATTTTTTTCATAGTTATACCTATTATAATTGGAGGATTTGGTAATTGATTAGTC CCCCTTATATTAGGAGCCCCAGATATAGCCTTCCCTCGAATAAATAATATAAGTTTTTGATTACTTCCCCCCTCATTAA CTCTTTTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGTACAGGATGAACAGTTTATCCCCCACTTTCATCTAATAT TGCTCATAGAGGTAGTTCTGTAGATTTAGCTATTTTTTCCCTTCATTTAGCTGGTATTTCTTCTATTTTAGGAGCTATCA ATTTTATCACAACAATTATTAATATACGACTTAATAAATTAATATTTGATCAAATACCTTTATTTGTTTGAGCAGTGGGAA TTACAGCATTTTTATTACTCCTTTCATTACCTGTTTTAGCAGGAGCAATTACTATGTTACTAACAGACCGAAATCTTAAT ACTTCTTTCTTTGACCCTGCAGGAGGAGGAGATCCAATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGT TTACTTT------Eilema griseola ------YYAAGATTATTAATTCGAGCTGAATTAGGTAA TCCTGGTTCATTAATTGGAGATGATCAAATTTATAATACTATTGTCACTGCACATGCTTTTATTATAATTTTTTTTATAGT AATACCAATTATAATTGGAGGATTTGGAAATTGATTAGTACCTTTAATACTTGGAGCCCCTGATATAGCTTTCCCCCGA ATAAATAATATAAGTTTTTGACTTCTTCCCCCATCTCTAACATTATTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGG

302

AACTGGATGAACAGTTTACCCCCCTTTATCCTCTAATATTGCCCATAGAGGAAGCTCTGTAGATTTAGCTATTTTCTCC CTTCATTTAGCTGGAATTTCATCAATTTTAGGAGCAATTAATTTTATTACTACTATTATTAATATACGTCTTAATAGTTTA ACTTTCGATCAAATACCTTTATTTGTTTGAGCTGTAGGAATTACAGCATTTTTACTTCTTTTATCATTACCTGTTTTAGCT GGAGCTATTACTATATTATTAACAGATCGAAATTTAAATACTTCTTTT------Eilema plana ------TCAACCAATCATAAAGATATTGGAACAATATATTTTATTTTTGGAATTTGAGCTGGAATAGTAGGAAC TTCTTTAAGACTTTTAATTCGAGCTGAATTAGGAAATCCAGGATCTTTAATTGGTGATGATCAAATTTATAATACTATTG TAACTGCTCATGCTTTTATTATAATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTTGGAAATTGATTAGTTCCTC TTATATTAGGAGCCCCTGATATAGCTTTCCCACGAATAAATAATATAAGTTTTTGATTACTTCCCCCTTCCCTAACCTTA YTTATTTCAAGTAGAATTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTATCCCCCACTTTCATCTAATATCGCTC ATAGAGGTAGATCTGTAGATCTAGCTATTTTCTCCTTACATTTAGCAGGTATTTCTTCTATTTTAGGAGCTATTAATTTT ATTACAACAATTATTAATATACGATTAAATAAATTAATATTTGATCAAATACCATTATTTGTTTGAGCAGTAGGTATTACA GCATTTTTATTACTTCTTTCTCTACCAGTTTTAGCGGGAGCTATTACTATATTATTAACTGATCGAAATCTTAATACTTCT TTTTTTGACCCTGCAGGAGGGGGTGATCCAATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTTACTT T------Eilema sp.1 TTTTATACGACTCACTATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCTGG AATAGTAGGAACATCATTAAGATTATTAATTCGAGCTGAATTAGGAAATCCTGGATCCTTAATTGGAGATGATCAAATT TATAATACTATTGTAACTGCCCATGCTTTTATTATAATTTTTTTTATAGTTATACCAATTATAATTGGAGGATTTGGAAAT TGATTAATTCCTCTAATATTAGGAGCTCCTGATATAGCATTCCCCCGAATAAATAATATAAGTTTTTGATTACTTCCCCC CTCATTAACTCTTTTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTACCCCCCACTTTCA TCAAATATTGCTCATAGAGGTAGATCTGTAGATTTAGCTATTTTTTCATTACATTTAGCGGGTATTTCCTCAATCCTAGG AGCTATTAATTTCATTACCACAATCATTAATATACGATTAAATAAATTAATATTTGATCAAATACCATTATTTGTATGAGC TGTAGGAATCACAGCATTTTTATTACTTCTTTCTTTACCTGTTCTAGCGGGAGCTATTACTATATTATTAACTGATCGAA ATCTTAATACTTCTTTTTTTGACCCTGCAGGAGGGGGAGATCCTATCTTATATCAACATTTATTTTGATTTTTTGGACAT CCAGAAGTTTACTTTAGTGAAGGGTAAATA Eilema sp.2 ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATCTTTGGAATTTGAGCTGGAATAGTA GGAACTTCTTTAAGATTATTAATTCGAGCTGAATTAGGTAATCCTGGCTCACTAATTGGAGACGACCAAATTTATAATA CTATTGTAACCGCTCACGCTTTTATTATAATTTTTTTTATAGTAATACCAATCATAATTGGTGGATTTGGAAATTGATTAG TACCCCTTATACTTGGAGCTCCTGATATAGCTTTCCCTCGGATAAATAATATAAGTTTTTGACTTCTTCCACCTTCTTTA ACTTTATTAATCTCAAGAAGAATTGTTGAAAATGGAGCTGGAACTGGATGAACAGTCTACCCGCCTTTATCATCTAATA

303

TTGCCCATGGAGGAAGCTCCGTAGATTTAGCTATTTTTTCTTTACATTTAGCTGGAATTTCATCAATTTTAGGAGCAATT AATTTTATTACTACTATTATTAATATACGATTAAATAGACTATCCTTTGATCAAATACCCCTATTTGTATGAGCAGTCGG AATTACAGCATTTTTACTTCTTCTTTCATTACCTGTATTAGCAGGAGCTATTACAATATTATTAACAGATCGAAATTTAAA TACTTCTTTTTTTGATCCTGCAGGAGGAGGAGACCCAATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAGAAG TTTACTTT------Eilema sp.3 ------CTTAGATTATTAATTCGAGCTGAATTAGGAAA TCCAGGATCTTTAATTGGTGATGATCAAATTTATAATACTATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATAGT TATACCTATTATAATTGGAGGATTTGGAAATTGATTAGTTCCCCTTATATTAGGAGCCCCAGATATAGCTTTCCCCCGA ATAAATAATATAAGTTTTTGATTACTCCCCCCCTCCTTAACCCTTTTAATTTCAAGAAGAATTGTAGAAAACGGAGCAG GAACAGGATGAACAGTCTATCCCCCACTTTCATCTAATATTGCTCATAGAGGTAGTTCTGTAGATTTAGCTATTTTTTC CTTACATTTAGCTGGTATTTCTTCTATTTTAGGAGCTATTAATTTCATTACAACAATTATTAACATACGACTAAATAAATT AATATTTGATCAAATACCTTTATTTGTATGAGCTGTAGGTATTACAGCATTTTTATTACTTCTCTCATTACCTGTTCTAGC AGGAGCTATTACTATATTATTAACTGATCGAAATCTCAACACTTCTTTT------Eilema sp.4 ------TATAGGGATTCAACCAATCATAAAGATATTGGTACATTATATTTTATTTTTGGAATTTGGGCAGGTATAGTA GGAACATCATTAAGACTTCTAATTCGAGCTGAATTAGGAAACCCTGGATCTTTAATTGGTGATGATCAAATTTACAATA CCATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTTGGTAACTGATTAA TCCCTCTTATACTAGGAGCCCCTGATATAGCTTTCCCCCGAATAAATAATATAAGTTTTTGATTACTCCCCCCTTCCTTA ACCCTTTTAATCTCAAGAAGAATTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTATCCCCCACTTTCATCTAATA TTGCTCATAGAGGTAGATCAGTGGATCTAGCTATTTTTTCACTTCATTTAGCTGGTATTTCTTCTATTTTAGGAGCTATT AACTTTATTACTACAATTATTAATATACGATTAAATAGATTAATATTTGATCAAATACCATTATTTGTTTGAGCTGTAGGA ATTACAGCATTTTTATTACTTCTTTCATTACCTGTTTTAGCAGGAGCTATTACTATACTATTAACTGATCGAAATTTAAAT ACATCTTTTTTTGATCCTGCCGGAGGAGGTGATCCTATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGT TTACTTT------Gardinia anoploa ------TTAAGATTATTAATTCGAGCTGAATTAGGTAA TCCTGGATCTTTAATTGGAGACGATCAAATTTATAATACTATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATAGT TATACCCATTATAATTGGAGGATTTGGTAATTGATTAGTTCCTTTAATATTAGGAGCTCCAGATATAGCATTCCCTCGAA TAAATAATATAAGTTTTTGACTTTTACCTCCTTCTTTAATACTTTTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGA ACAGGATGAACAGTTTATCCTCCACTTTCTTCTAATATAGCTCATAGAGGAAGATCTGTAGATTTAGCTATTTTTTCTTT ACATTTAGCTGGTATTTCATCAATCTTAGGTGCTATTAATTTTATTACTACAATTATTAATATACGTTTAAATAAATTATCA

304

TTTGATCAAATACCTTTATTTATTTGAGCTGTAGGAATTACTGCTTTTTTACTTCTCCTTTCATTACCAGTATTAGCAGGA GCTATTACTATACTTTTAACTGATCGAAATTTAAATACTTCATTT------Gnamptonychia flavicollis ------TTAAGATTATTAATTCGAGCTGAATTAGGAAA TCCTGGATCTTTAATTGGAGATGATCAAATTTATAATACTATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATAGT TATACCTATTATAATTGGAGGATTTGGAAATTGATTAGTCCCTCTTATATTAGGAGCTCCTGATATAGCATTTCCACGAA TAAATAATATAAGTTTTTGACTTCTTCCCCCCTCATTAACTTTATTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGT ACAGGATGAACAGTTTATCCCCCACTTTCATCTAATATTGCTCATGGAGGTAGTTCCGTAGATTTAGCTATTTTTTCATT ACATTTAGCAGGAATTTCTTCAATTTTAGGAGCTATTAATTTTATTACTACAATTATTAATATGCGATTAAATAGATTAAT ATTTGATCAAATACCTTTATTTGTATGAGCTGTTGGTATTACAGCATTTTTATTACTTCTTTCATTACCAGTTTTAGCTGG AGCAATTACTATATTATTAACAGATCGAAATCTTAATACATCATTT------Heliosia jucunda ------Hiera gyge ------Hypoprepia cadaverosa ------TTAAGTTTATTAATTCGAGCTGAATTAGGTAAT CCTGGATCATTAATTGGTGATGATCAAATTTACAATACTATTGTAACTGCCCATGCTTTTATTATAATTTTTTTTATAGTA ATACCTATTATAATTGGAGGATTTGGTAATTGATTAGTCCCTTTAATATTAGGTGCCCCAGATATAGCTTTCCCTCGAAT AAATAACATAAGTTTTTGACTCCTACCCCCATCATTAACTTTATTAATCTCAAGAAGAATTGTAGAAAATGGAGCAGGA ACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATTGCCCATAGAGGTAGATCAGTAGATTTAGCTATTTTTTCCC TACATTTAGCTGGAATTTCTTCTATTTTAGGGGCTATTAATTTTATTACTACAATTATTAATATACGTCTTAACAAATTAT CATTTGATCAAATACCTTTATTTGTTTGAGCTGTAGGAATTACAGCATTTCTTTTATTACTTTCTCTTCCAGTTTTAGCTG

305

GAGCTATTACTATATTACTTACTGATCGAAATTTAAATACATCTTTC------Hypoprepia fucosa ------Hypoprepia fucosa tricolor ------TATAGGGATTCAACCAATCATAAAGATATTGGAACACTATATTTTATTTTTGGTATCTGAGCCGGAATAGTA GGAACCTCTTTAAGTTTATTAATTCGAGCTGAATTAGGTAATCCTGGATCATTAATTGGTGATGATCAAATTTATAATAC TATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATAGTAATACCTATTATAATTGGAGGATTTGGTAATTGATTAGT CCCTTTAATATTAGGTGCCCCAGATATAGCTTTTCCTCGAATAAATAACATAAGTTTTTGACTCCTACCCCCATCATTAA CTTTATTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTACCCTCCACTTTCATCCAATAT TGCTCACAGAGGTAGATCAGTAGATTTAGCTATTTTTTCCCTACATTTAGCTGGAATTTCTTCTATTTTAGGAGCTATTA ATTTTATTACTACAATTATTAATATACGTCTTAACAAATTATCATTTGATCAAATACCTTTATTTGTTTGAGCTGTAGGAA TTACAGCATTTCTTCTATTACTTTCTCTTCCAGTTTTAGCTGGAGCTATTACTATATTACTTACTGATCGAAATTTAAATA CATCTTTCTTTGATCCTGCTGGAGGAGGAGATCCTATTCTTTACCAACACTTATTTTGATTTTTTGGACATCCAGAAGTT TACTTT------Inopsis modulata ------AATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCTGGAATAGTAGGAACTTC TTTAAGATTATTAATTCGAGCTGAATTAGGAAACCCTGGATCTTTAATYGGAGACGATCAAATTTATAATACTATTGTAA CTGCTCATGCCTTTATTATAATTTTTTTTATAGTTATACCCATTATAATTGGAGGATTTGGAAATTGATTAGTTCCTCTTA TATTAGGAGCTCCTGATATAGCATTCCCMCGAATAAATAATATAAGTTTTTGACTTCTTCCCCCCTCTCTAACTCTTTTA ATTTCAAGAAGAATTGTAGAAAACGGAGCAGGTACAGGATGAACAGTTTATCCCCCACTTTCATCTAATATTGCCCATA GAGGTAGTTCTGTAGATTTAGCTATTTTTTCATTACATTTAGCAGGTATTTCCTCTATTTTAGGAGCTATTAACTTTATTA CTACAATTATCAATATACGATTAAATAGATTAATATTTGATCAAATACCTTTATTTGTATGAGCTGTTGGTATTACAGCAT TTTTATTACTTCTTTCATTACCTGTTCTAGCTGGAGCTATTACTATATTACTAACTGATCGAAATCTTAATACATCATTTT TTGACCCAGCTGGAGGAGGAGATCCTATTCTTTACCAACATTTATTTTGATTTTTTGGACAT------Lithosia quadra ------TTAAGATTATTAATTCGAGCAGAATTAGGAAA TCCAGGATCCTTAATTGGAGATGATCAAATTTATAATACTATTGTAACTGCTCATGCTTTTATTATAATTTTTTTTATGGT TATACCTATTATAATTGGAGGATTTGGAAATTGATTAGTTCCTTTAATATTAGGAGCTCCTGATATAGCATTCCCTCGAA

306

TAAATAATATAAGTTTTTGATTATTACCCCCCTCTTTAACTCTTTTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGA ACAGGATGAACAGTTTATCCCCCACTCTCATCAAATATTGCCCATAGAGGTAGTTCCGTAGATTTAGCCATTTTTTCTT TACATTTAGCAGGTATTTCTTCTATTTTAGGAGCTATTAATTTTATTACCACAATCATTAATATACGATTAAATAAATTAA TATTTGATCAAATACCTCTATTTGTATGAGCTGTAGGAATTACAGCATTTTTATTACTTTTATCATTACCTGTATTAGCTG GAGCTATTACTATACTTCTAACAGATCGAAACCTCAATACTTCATTT------Lyclene pyraula ------TATAGGGATTCAACCAATCATAAAGATATTGGAACTCTATATTTTATTTTTGGTATTTGAGCTGGAATAATTG GAACTTCTTTAAGTTTATTAATTCGAGCTGAATTAGGTAACCCTGGGTCTTTAATTGGTGATGACCAAATTTATAATACT ATTGTCACTGCCCATGCTTTCATTATAATTTTTTTTATAGTTATACCAATTATAATTGGAGGATTTGGAAATTGATTAGTC CCTTTAATATTAGGAGCTCCTGATATAGCTTTCCCTCGAATAAATAATATAAGTTTTTGACTTCTCCCCCCTTCTTTAAC TTTATTAATTTCAAGAAGAATTGTTGAAAATGGAGCTGGAACAGGATGAACTGTTTATCCTCCTTTATCTTCTAATATTG CCCATGGTGGAAGTTCTGTAGATTTAGCTATTTTTTCCCTTCATTTAGCAGGAATTTCATCAATTTTAGGAGCAATTAAT TTTATTACAACTATTATTAATATACGTTTAAATAATTTATCTTTTGATCAAATACCTTTATTTGTTTGAGCTGTAGGTATTA CAGCATTCTTATTACTTCTTTCTTTACCAGTTTTAGCAGGAGCTATTACTATATTATTAACTGATCGAAATTTAAATACTT CTTTTTTTGATCCTGCTGGAGGTGGAGATCCTATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTTAC TTT------Lyclene reticulata ------TATTTTATTTTTGGAATTTGAGCTGGAATAGTAGGAACCTCTTTAAGTTTACT AATTCGAGCTGAATTAGGAAACCCAGGATCTTTAATTGGGGATGATCAAATTTATAATACTATTGTTACTGCACATGCT TTTATTATAATTTTTTTTATAGTAATACCAATTATAATCGGAGGATTTGGAAATTGGTTGGTACCCCTTATATTAGGAGC TCCTGATATAGCTTTCCCCCGAATAAATAATATAAGATTTTGATTATTACCCCCCTCATTAACATTGCTAATTTCAAGAA GAGTTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATTGCTCATAGTGGAAGAT CAGTAGATTTAGCTATTTTTTCTCTTCATTTAGCGGGTATTTCATCAATTTTAGGAGCAATTAATTTTATTACTACCATTA TTAACATACGATTAAATTCACTATCATTCGATCAAATACCATTATTTGTTTGAGCTGTAGGTATTACAGCATTTTTATTAC TTTTATCTTTACCTGTTTTAGCAGGAGCTATTACTATATTACTAACTGATCGAAATTTAAATACTTCTTTTTTTGATCCAG CTGGAGGAGGTGATCCAATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTTACTTT------Lyclene sp.1 ------TCAACCAATCATAAAGATATTGGAACTCTATATTTTATTTTTGGTATTTGAGCTGGAATAATTGGAAC TTCTTTAAGTTTATTAATTCGAGCTGAATTAGGTAATCCTGGGTCTCTAATTGGTGATGACCAAATTTATAATACTATTG TTACTGCTCATGCTTTCATTATAATTTTTTTTATAGTTATACCAATTATAATTGGAGGATTTGGAAATTGATTAGTTCCTT TAATATTAGGAGCTCCTGATATAGCTTTCCCTCGAATAAATAATATAAGTTTTTGACTTCTCCCCCCTTCTTTAACTTTA TTAATTTCAAGAAGAATTGTTGAAAATGGAGCTGGAACAGGATGAACTGTTTATCCTCCTTTATCTTCTAATATTGCTCA

307

CGGTGGAAGTTCTGTAGATTTAGCTATTTTTTCTCTTCATTTAGCAGGAATTTCATCAATTTTAGGAGCAATTAATTTTA TTACAACTATTATTAATATACGTTTAAATAATTTATCTTTTGATCAAATACCTTTATTTGTTTGAGCTGTAGGTATTACAG CATTCTTATTACTTCTTTCTTTACCAGTTTTAGCAGGAGCTATTACTATATTATTAACTGATCGAAATTTAAATACTTCTT TTTTTGATCCTGCTGGAGGTGGAGATCCTATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTTTACTTT------Lyclene sp.1 nr. ---AATACGACTCACTATAGGGATTCAACCAATCATAAAGATATTGGAACTCTATATTTTATTTTTGGTATTTGAGCTGGA ATAATTGGAACTTCTTTAAGTTTATTAATTCGAGCTGAATTAGGTAATCCTGGGTCTCTAATTGGTGATGACCAAATTTA TAATACTATTGTTACTGCTCATGCTTTCATTATAATTTTTTTTATAGTTATACCAATTATAATTGGAGGATTTGGAAATTG ATTAGTTCCTTTAATATTAGGAGCTCCTGATATAGCTTTCCCTCGAATAAATAATATAAGTTTTTGACTTCTCCCCCCTT CTTTAACTTTATTAATTTCAAGAAGAATTGTTGAAAATGGAGCTGGAACAGGATGAACTGTTTATCCTCCTTTATCTTCT AATATTGCTCACGGTGGAAGTTCTGTAGATTTAGCTATTTTTTCTCTTCATTTAGCAGGAATTTCATCAATTTTAGGAGC AATTAATTTTATTACAACTATTATTAATATACGTTTAAATAATTTATCTTTTGATCAAATACCTTTATTTGTTTGAGCTGTA GGTATTACAGCATTCTTATTACTTCTTTCTTTACCAGTTTTAGCAGGAGCTATTACTATATTATTAACTGATCGAAATTTA AATACTTCTTTTTTTGATCCTGCTGGAGGTGGAGATCCTATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAGA AGTTTACTTTAGTGAGGGGATAATA Lycomorpha pholus ------Lycomorphodes sordida ------Meterhythosia sangala ------

308

------Miltochrista miniata ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCTGGTATAGTTG GAACTTCTTTAAGATTATTAATTCGAGCTGAATTAGGTAATCCTGGTTCATTAATTGGAGATGATCAAATTTATAATACT ATTGTCACTGCACATGCTTTTATTATAATTTTTTTTATAGTAATACCAATTATAATTGGAGGATTTGGAAATTGATTAGTA CCTTTAATACTTGGAGCCCCTGATATAGCTTTCCCCCGAATAAATAATATAAGTTTTTGACTTCTTCCCCCATCTCTAAC ATTATTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACTGGATGAACAGTTTACCCCCCTTTATCCTCTAATATT GCCCATAGAGGAAGCTCTGTAGATTTAGCTATTTTCTCCCTTCATTTAGCTGGAATTTCATCAATTTTAGGAGCAATTA ATTTTATTACTACTATTATTAATATACGTCTTAATAGTTTAACTTTCGATCAAATACCTTTATTTGTTTGAGCTGTAGGAA TTACAGCATTTTTACTTCTTTTATCATTACCTGTTTTAGCTGGAGCTATTACTATATTATTAACAGATCGAAATTTAAATA CTTCTTTTTTTGATCCTGCTGGTGGGGGAGATCCAATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAGAAGTT TACTTT------Miltochrista sp.1 ---TATACGACTCACTATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATCTGAGCTGGT ATAGTTGGAACTTCTTTAAGATTATTAATTCGAGCTGAATTAGGTAATCCTGGTTCATTAATTGGTGATGATCAAATTTA TAATACTATTGTTACTGCACATGCTTTTATTATAATTTTTTTCATAGTAATACCAATTATAATTGGAGGATTTGGTAATTG ATTAGTACCTTTAATACTTGGAGCTCCTGATATAGCTTTCCCTCGAATAAATAATATAAGTTTTTGACTTCTCCCTCCAT CTTTAACATTATTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACTGGATGAACAGTTTACCCCCCTTTATCCTC TAATATTGCTCATAGAGGAAGCTCTGTAGATTTAGCTATTTTTTCCCTTCATTTAGCTGGAATTTCATCAATTTTAGGAG CAATTAATTTTATTACCACTATTATTAATATGCGTCTTAATAGTTTAACTTTTGATCAAATACCTTTATTTGTTTGAGCTGT AGGAATTACCGCATTTTTACTTCTTTTATCTTTACCTGTTTTAGCTGGAGCTATTACTATATTATTAACAGATCGAAATTT AAATACTTCTTTTTTTGATCCTGCTGGAGGAGGAGATCCAATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAG AAGTTTACTTTAGTGAGGGGTTAATA Prepiella aurea nr. ------Prinasura quadrilineata TTTTATACGACTCACTATAGGGATTCAACCAATCATAAAGATATTGGAACTCTATATTTTATTTTTGGTATTTGAGCTGG AATAATTGGAACTTCTTTAAGTTTATTAATTCGAGCTGAATTAGGTAATCCTGGGTCTCTAATTGGTGATGACCAAATTT

309

ATAATACTATTGTTACTGCTCATGCTTTCATTATAATTTTTTTTATAGTTATACCAATTATAATTGGAGGATTTGGAAATT GATTAGTTCCTTTAATATTAGGAGCTCCTGATATAGCTTTCCCTCGAATAAATAATATAAGTTTTTGACTTCTCCCCCCT TCTTTAACTTTATTAATTTCAAGAAGAATTGTTGAAAATGGAGCTGGAACAGGATGAACTGTTTATCCTCCTTTATCTTC TAATATTGCTCACGGTGGAAGTTCTGTAGATTTAGCTATTTTTTCTCTTCATTTAGCAGGAATTTCATCAATTTTAGGAG CAATTAATTTTATTACAACTATTATTAATATACGTTTAAATAATTTATCTTTTGATCAAATACCTTTATTTGTTTGAGCTGT AGGTATTACAGCATTCTTATTACTTCTTTCTTTACCAGTTTTAGCAGGAGCTATTACTATATTATTAACTGATCGAAATTT AAATACTTCTTTTTTTGATCCTGCTGGAGGTGGAGATCCTATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAG AAGTTTACTTTAGTGAAGGAAAAATA Ptychoglene coccinea ------Schistophleps albida ------TATAGGGATTCAACCAATCATAAAGATATTGGAACTTTATATTTTATTTTTGGAATTTGAGCAGGAATAGTA GGAACCTCACTAAGTTTACTAATTCGAGCTGAATTAGGAAACCCCGGATCTCTAATTGGAGATGATCAAATTTATAATA CTATTGTAACAGCTCATGCCTTTATTATAATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTTGGGAATTGATTAG TACCTTTAATATTAGGAGCACCTGATATAGCATTTCCTCGAATAAATAACATAAGTTTTTGACTTTTACCACCATCTTTA ACCTTATTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACAGGATGAACAGTTTATCCCCCACTTTCATCTAACA TTGCCCATAGAGGCAGATCCGTTGACTTAGCCATTTTTTCTTTACATCTTGCAGGTATTTCATCAATTTTAGGAGCTATT AATTTTATTACTACTATTATTAATATACGATTAAATAATTTATCATTTGATCAAATACCATTATTCGTTTGAGCAGTAGGA ATTACAGCTTTCTTATTATTATTATCATTACCGGTTTTAGCTGGAGCTATTACTATATTATTAACTGATCGAAATTTAAAT ACCTCTTTTTTTGACCCTGCTGGAGGAGGAGATCCTATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGT TTACTTT------Amata aperta ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGTATTTGAGCAGGAATAGTA GGAACTTCTTTAAGATTATTAATTCGAGCTGAATTAGGAACTCCTGGTTCCATAATTGGAGACGATCAAATCTATAATA CTATTGTTACAGCTCATGCTTTTATTATAATTTTTTTTATAGTCATACCTATTATAATTGGAGGATTTGGTAACTGATTAG TACCTTTAATATTAGGAGCCCCTGATATAGCTTTTCCCCGAATAAATAATATAAGTTTTTGACTTTTACCCCCTTCTTTA ACCCTTTTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGAACTGGATGAACAGTTTATCCCCCACTTTCATCTAATA TTGCTCATAGAGGAAGTTCAGTTGATTTAGCTATTTTTTCCCTACATTTAGCTGGAATTTCTTCAATTCTAGGAGCTATT AATTTTATTACAACAATTATTAATATACGATTAAATAATTTATTTTTTGATCAAATACCTTTATTTGTATGAGCAGTAGGA

310

ATTACAGCTTTTTTATTACTTCTTTCCTTACCTGTTTTAGCTGGTGCTATTACTATATTATTAACAGACCGTAATCTCAAT ACATCTTTTTTTGACCCCGCTGGAGGAGGAGATCCAATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGT TTAC------Pagara simplex ------Asota heliconia ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCAGGAATAGTA GGAACATCTTTAAGATTATTAATTCGAGCTGAATTAGGTAACCCTGGATCTTTAATTGGGGATGATCAAATTTATAACA CTATTGTTACAGCTCATGCCTTTATTATAATTTTTTTTATAGTTATACCTATTATAATTGGAGGATTTGGTAATTGATTAG TCCCTCTTATATTAGGAGCCCCTGATATAGCTTTCCCCCGAATAAATAATATAAGTTTTTGACTTCTTCCCCCCTCATTA ACACTCTTAATTTCAAGAAGAATTGTTGAAAACGGAGCAGGTACCGGATGAACAGTTTACCCCCCACTTTCATCTAATA TTGCTCATGGAGGAAGATCAGTTGATTTAGCTATTTTTTCTTTACATTTAGCTGGAATTTCTTCAATTTTAGGAGCTATT AACTTCATTACCACAATTATTAATATACGATTAAATAATTTATCATTTGATCAAATACCTTTATTTGTATGAGCTGTAGGA ATTACAGCATTTTTATTACTTTTATCTTTACCAGTATTAGCTGGAGCTATTACCATACTTCTCACTGATCGAAATTTAAMT ACWTCTTTTTTTGATCCAGSTGGAGGAGGAGATCCAATTTTA------Asota orbona ------TATAGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCAGGAATAGTA GGAACATCTTTAAGATTGCTAATTCGAGCTGAATTAGGTACCCCCGGATCTTTAATTGGAGATGATCAAATTTATAATA CTATTGTTACAGCTCATGCTTTTATTATAATCTTTTTCATAGTTATACCTATTATAATTGGAGGATTTGGTAATTGATTAA TCCCTCTTATATTAGGAGCCCCCGATATAGCTTTCCCCCGAATAAATAATATAAGTTTTTGACTTCTTCCCCCCTCACT AACTTTATTAATCTCAAGAAGAATTGTTGAAAATGGAGCAGGTACCGGATGAACAGTTTACCCCCCACTTTCATCTAAT ATTGCTCACGGAGGAAGATCAGTTGATTTAGCCATTTTTTCATTACATTTAGCTGGAATTTCTTCAATCTTAGGAGCTAT TAATTTTATTACTACAATTATCAATATACGATTAAATAATTTATCATTTGATCAAATACCTTTATTTGTATGAGCTGTAGG AATTACAGCATTTTTATTACTCTTATCTTTACCAGTATTAGCTGGAGCTATTACTATACTTCTTACTGATCGAAATTTAAA CACATCTTTTTTTGATCCTGCTGGAGGAGGAGATCCAATTTTATATCAACATTTATTTTGATTTTTTGGACATCCAGAAG TTTAC------Neochera dominia ------AGGGATTCAACCAATCATAAAGATATTGGAACATTATATTTTATTTTTGGAATTTGAGCAGGTATAGTAGG AACTTCCCTAAGATTATTAATTCGAGCTGAACTAGGAAATCCTGGCTCTTTAATTGGAGATGATCAAATTTATAATACTA

311

TTGTAACAGCTCATGCTTTTATTATAATTTTTTTTATAGTTATGCCAATTATAATTGGAGGATTTGGAAATTGATTAGTAC CTCTAATATTAGGAGCACCTGATATAGCATTCCCTCGAATAAATAATATAAGTTTTTGACTTCTACCCCCATCATTAACT TTATTAATTTCAAGAAGAATTGTAGAAAATGGAGCAGGTACAGGATGAACAGTTTACCCCCCACTTTCATCTAATATTG CTCACGGAGGAAGCTCAGTTGATTTAGCTATTTTTTCATTGCATCTAGCAGGTATTTCCTCAATTTTAGGAGCTATTAA TTTTATTACTACAATTATTAATATACGATTAAATAGCTTATCATTTGATCAAATACCTTTATTTGTTTGAGCTGTAGGTAT TACTGCATTTCTTTTACTCCTTTCTTTACCAGTTCTAGCAGGAGCTATTACCATACTTTTAACAGACCGAAATTTAAATA CATCTTTCTTTGACCCAGCTGGAGGGGGAGATCCAATTCTTTATCAACATTTATTTTGATTTTTTGGACATCCAGAAGT TTAC------

[CytB] Acsala anomala ------Agylla septentrionalis ------Asura cervicalis ------Asura polyspila ------TCAATATTAGTTAATTGAATTTGAGGGGGTTTTGCAGTAGATAATGCTACATTAA CACGTTTCTACACATTTCATTTCTTATTACCATTTATTATTTTAATAATAACTATAATTCACTTAMTATTTMTACATCAAAC AGGATCCAATAACCCATTAGGATTAAATAGAAATTATGATAAAATTCCTTTTCACCCATTTTTTACTTTCAAAGATTTAAT TGGAGCTATTATTTTAATATTCCTATTAATTATAATTACTTTAACTAATCCCTATTTATTAGGAGATCCAGATAATTTTATT

312

CCAGCAAATCCTTTAGTAACTCCAGTTCATATTCAACCTGAATGATATTTTTTATTTGCTTATGCCATTTTACGATCAATT CCAAATAAATTAGGTGGAGTAATTGCTTTAATTATATCTATTTTAATTTTAATTATTTTACCTTTTACATTTAATAAAAAAA TTCAAGGAATTCAATTTTATCCAATTAATCAAATTATATTT------TTAATACGACTCACT ATAGGGATGGCAGAAGAGAATTGGAACGACGAAGCCGTGG Atolmis rubricolis ------TTAGGGTGAGGACAAATATCATTTTGAGGAGCTACAGTTATTACAAATCTTTTATCTGCAATCCCTTATTTA GGATCAATATTAGTAAATTGAATTTGAGGAGGATTTGCCGTAGATAATGCTACATTAACACGATTTTATACTTTTCATTT TTTACTACCCTTTATTATTTTAATAATAACTATAATTCATTTATTATTTCTTCATCAAACGGGATCAAATAATCCATTAGGA TTAAATAGAAATTTTGATAAAATTCCTTTCCATCCATTTTTCACTTATAAAGATCTTATTGGAGCTATTATATTATTAACAA TATTAATTATATTAACTCTTACAAATCCCTATTTATTAGGTGATCCTGATAATTTTATTCCTGCTAATCCCTTAGTCACAC CAGTTCATATTCAACCTGAATGATATTTTTTATTTGCATATGCTATTTTACGATCTATCCCTAATAAATTAGGAGGAGTT ATTGCTTTAATTATATCAATTTTAATTTTAATTATTTTACCTTTCACATTTAATAAAAAAATTCAAGGAATTCAATTCTACC CAATCAATCAAGTATTATTT------Bruceia hubbardi ------Bruceia pulverina ------Calamidia hirta ---ATACGACTCACATTAGGGTGAGGACAAATATCATTTTGAGGAGCTACAGTAATTACAAATCTTTTATCTGCTATTCCC TATTTAGGATCCATATTAGTAAATTGAATTTGAGGAGGATTTGCTGTAGATAATGCAACATTAACACGATTCTATACATT TCACTTTTTATTACCTTTTATTATTTTAATAATAACAATAATTCATTTATTATTTCTTCATCAAACAGGATCTAATAATCCA CTAGGACTAAATAGAAATTTAGATAAAATTCCTTTTCATCCATTTTTTACCTTTAAAGATTTAATTGGAGCAATTATATTA ATAATATTTTTAATTATATTAACACTTACAAATCCTTATTTACTTGGAGATCCAGATAATTTTATCCCAGCCAATCCTTTA GTCACTCCAGTTCATATTCAACCTGAATGATATTTTTTATTTGCTTATGCTATTTTACGATCTATTCCCAATAAATTAGGA

313

GGTGTAATTGCTTTAATTATATCAATTTTAATTTTAATTATCTTACCTTTTACATTCAATAAAAAAATTCAAGGAATTCAAT TTTACCCAATTAATCAAATTTTATTT------Cisthene juanita ------Cisthene plumbea ------GGACAAATATCATTTTGAGGAGCTACTGTTATTACTAACTTATTGTCAGCCATCCCTTATTTAGGAAC TATATTAGTAAATTGAATTTGAGGAGGATTTGCAGTAGATAACGCTACATTAACTCGATTTTACACATTTCATTTTCTAC TACCATTTATTATTGCATTATTAATTATAATTCATTTATTATTTTTACATCAGACAGGGTCAAATAATCCTTTAGGATTAAA TAGAAATTATGATAAAATCCCCTTTCATCCATTTTTTTCTTACAAAGATTTAATTGGAATCATTATTATACTATCAATTCTA ATCTTATTAAATCTCACTAACCCTAACCTACTTGGAGACCCAGATAATTTTATTCCAGCTAACCCATTAGTAACCCCTG TACACATTCAACCTGAATGATATTTTTTATTCGCCTATGCAATTTTACGATCAATTCCTAATAAATTAGGAGGAGTAATT GCTTTAATTATATCCATTTTAATTCTAATTATTTTACCTTTTACATTTAATAAGAAAATCCAAGGAATTCAATTCTACCCAA TTAAT------Cisthene subjecta ------Cisthene tenuifascia ------Crambidia lithosioides ------

314

------Cyana meyricki ------GGGTGAGGACAAATATCATTTTGAGGAGCTACTGTAATTACTAATTTACTATCAGCAATCCCTTATTTAG GATCCATATTAGTAAATTGAATTTGAGGGGGATTTGCAGTTGAAAATGCTACTTTAACTCGATTTTACACTTTCCATTTT TTATTACCTTTTATCATTTTAATAATAACTATAATTCATTTATTATTTTTACATCAAACTGGATCTAATAATCCCCTAGGAT TAAATAGTAACTATGATAAAATTCCTTTCCATCCATTTTTTACTTATAAAGATTTAATTGGAGCTATTTTTATATTATTTTT ATTAATTATGTTAACTCTAACTAATCCTTACTTATTAGGAGACCCTGATAATTTTATTCCCGCTAATCCCTTAGTTACTC CCGTTCATATTCAACCTGAATGATATTTTCTATTCGCTTATGCTATTTTACGATCGATTCCAAATAAATTAGGAGGAGTT ATTGCTCTAATTATATCAATTCTTATTTTAATTATTTTACCATTTACTTTTAATAAAAAAATTCAAGGAATTCAATTTTAT------CTCTAT------Cybosia mesomella ------Eilema bicolor TTAATACGACTCACATTAGGGTGAGGACAAATATCATTTTGAGGAGCTACAGTTATTACAAATCTTCTATCAGCAATCC CCTATCTTGGATCTATATTAGTAAATTGAATTTGAGGGGGATTTGCTGTAGATAATGCTACATTAACACGATTTTATACA TTTCATTTTTTATTACCTTTTATTGTATTATTACTAGTTATTATTCATCTTTTATTTCTCCATCAAACAGGATCTAATAATC CTTTAGGACTTAATAGAAATCTAGATAAAATTCCATTTCATCCATTTTTTACTTATAAAGATTTAATTGGAGCTATTATTA TAATAATAATCTTAATTATGTTAACTTTTACAAATCCTTATTTATTGGGAGACCCTGATAATTTCATTCCTGCTAACCCAT TAGTTACTCCTGTCCATATCCAACCTGAATGATACTTTTTATTTGCATATGCTATTTTACGATCTATCCCTAATAAATTAG GAGGTGTAATTGCTTTAATTATATCAATTTTAATTTTAATTATTTTACCTTTTACATTTAATAAAAAAATTCAAGGAATTCA A------Eilema complana ------Eilema dorsalis

315

------CTCACATTAGGGTGAGGACAAATATCATTTTGAGGAGCTACAGTCATTACTAATCTTTTATCGGCAATTCCATAT TTAGGATCCATATTAGTAAATTGAATTTGAGGAGGATTTGCTGTAGATAATGCCACATTAACACGATTTTACACTTTTCA TTTTTTATTACCATTTATTGTATTAATATTAACTATTATTCACTTACTATTTTTACACCAAACAGGATCTAATAATCCCTTA GGTATTAATAGAAATTTTGATAAAATTCCTTTCCACCCATTCTTTACTTATAAGGATTTAATTGGGGCTATTATTATATTA ACAATTTTAATTATATTAACCTTTACAAATCCTTATTTATTGGGAGATCCTGATAATTTTATTCCCGCTAACCCTTTAGTC ACCCCCGTCCATATTCAACCTGAATGATATTTTTTATTTGCCTATGCTATTTTACGCTCTATTCCTAATAAATTAGGAGG TGTAATTGCTTTAGTTATATCAATTCTAATTTTAATTATTTTACCATTTACATTTAATAAAAAAATACAAGGAATTCAATTT TATCCCATTAATCAAATTTTATTTTGATTTTTAGTAGTAATAATCATTTTATTAACATGAATTGGAGCTCGACCAGTCTTT------Eilema griseola ------TTAGGGTGAGGACAAATATCATTTTGAGGAGCTACAGTCATTACTAATCTCCTATCAGCAATCCCCTATCTT GGATCTATATTAGTAAATTGAATTTGAGGAGGATTTGCTGTAGATAATGCTACATTAACACGATTTTATACATTTCATTT TTTATTACCATTTGTTGTATTAATATTAACTATTATTCATCTACTATTTCTACATCAAACAGGATCTAATAATCCATTAGG AATTAATAGAAACTTTGATAAAATTCCTTTCCACCCCTTTTTCACGTATAAAGACTTAATTGGAGCTATTATTATATTAAC AATTTTAATTATATTAACTTTTACAAATCCTTATTTATTGGGAGACCCTGATAATTTTATTCCTGCAAATCCCTTAGTTAC TCCTGTACACATTCAACCTGAATGATATTTTCTATTTGCATATGCCATTTTACGATCTATCCCAAATAAATTAGGAGGTG TAATCGCTTTAATTATATCAATTTTAATTTTAATTATTCTACCATTTACATTTAATAAAAAAATCCAAGGAATTCAATTTTA TCCAATTAATCAAATCCTTTTT------Eilema plana ------Eilema sp.1 ------ACATTAGGGTGAGGACAAATATCATTTTGAGGAGCTACAGTTATTACCAATCTTTTATCTGCTATTCCCTATCT TGGATCTATATTAGTAAATTGAATTTGAGGAGGTTTTGCTGTAGATAATGCAACATTAACACGATTTTACACATTTCATT TTTTACTACCATTTATTATCTTAATAATAGTAATAATTCACTTATTATTTCTCCATCAAACAGGATCTAATAATCCCCTAG GACTAAACAGAAATTTAGATAAAATCCCATTTCACCCATTTTTCACATTTAAAGATCTAATTGGAGCTATTATATTATTAA TATTATTAACTATACTAACTCTCACAAACCCAAATTTACTTGGGGATCCTGATAATTTTATTCCAGCTAATCCACTGGTT ACACCAGTACACATTCAACCTGAGTGATATTTTTTATTTGCTTACGCTATTTTACGATCTATCCCTAATAAATTAGGAGG TGTAATTGCTTTAATTATATCAATTCTAATTTTAATTATTCTACCTTTTACATTTAATAAAAAAATTCAAGGAATTCAATTT TACCCCATTAATCAAATCTTATTTTGATTTTTAGTAATAATAATTATTTTACTAACATGA------

316

Eilema sp.2 ------Eilema sp.3 ------GGGTGAGGACAAATATCATTTTGAGGAGCTACAGTTATCACTAATCTCTTATCAGCAATTCCTTACCTTG GATCTATATTAGTTAATTGAATCTGAGGAGGATTTGCTGTTGATAATGCAACTCTAACTCGATTTTACACATTTCATTTT TTATTACCTTTTATTGTATTAATATTAACTATTATTCATTTATTATTTTTACATCAAACAGGATCAAATAATCCCTTAGGAC TTAATAGAAATTTAGATAAAATTCCCTTTCATCCCTTTTTCACATATAAGGATTTAATTGGAGGTATTATCATATTAATAA TTTTAATTATATTAACTTTTACAAATCCCTATTTATTAGGAGATCCTGATAACTTTATCCCCGCAAATCCCTTAGTTACTC CAGTTCATATTCAACCTGAATGATATTTTTTATTTGCTTATGCTATTTTACGATCAATCCCTAATAAATTAGGAGGTGTA ATTGCTTTAATCATATCAATCTTAATTTTAATTATTTTACCTTTTACATTTAATAAAAAAATTCAAGGAATTCAATTCTATC CCATTAATCAAATTCTATTTTGATTCCTAGTA------Eilema sp.4 ------Gardinia anoploa ------Gnamptonychia flavicollis ------ACATTAGGGTGAGGACAAATATCATTTTGAGGAGCTACTGTCATTACAAAT CTTTTATCAGCTATTCCTTATTTAGGATCAATATTAGTAAATTGAATTTGAGGGGGATTCGCCGTTGATAATGCAACATT AACTCGTTTTTATAGTTTTCACTTTTTATTACCTTTTATTATTTTAATAATAGTTATAATTCATTTATTATTTCTCCATCAAA CAGGATCTAATAACCCTTTAGGATTAAATAGAAATTATGATAAAATTCCCTTTCATCCATTTTTTACTTATAAAGATTTAA TTGGAAGTATTATATTATTATTTTTATTAATTATATTAACTCTTACTAATCCCTACCTATTAGGAGATCCTGATAATTTTAT

317

TCCTGCTAATCCTTTAGTTACCCCTGTACATATTCAACCTGAATGATATTTTTTATTTGCTTATGCTATTTTACGATCAAT TCCTAATAAACTTGGAGGAGTAATTGCTCTTATCATATCAATTTTAATTTTAATTATTTTACCTTTTACATTTAATAAAAAA ATTCAAGGAATTCAATTTTAT------Heliosia jucunda ------Hiera gyge ------Hypoprepia cadaverosa ------TTAGGGTGAGGACAAATATCATTTTGAGGTGCAACAGTAATTACTAATTTATTATCAGCTATTCCTTATTTA GGAACTATAATAGTAAATTGAATTTGAGGGGGATTCGCAGTAGATAATGCTACTTTAATTCGATTTTATACTTTCCATTT TCTTTTTCCATTTATTATTTTAATAATAACTATAATTCATTTATTATTCTTACATCAAACAGGATCTAATAATCCTTTAGGA TTAAATAGAAATTATGATAAAATTCCATTTCATCCATTTTTTACATACAAAGATTTAATTGGAGCTATCATATTATTATTTA TATTAATTTTATTAACTTTAACTAACCCATATCTATTAGGAGATCCAGATAATTTTATCCCTGCAAATCCATTAGTTACTC CTATTCATATTCAACCAGAATGATATTTTTTATTTGCTTATGCCATTCTTCGATCCATTCCCAATAAATTAGGAGGAGTT ATTGCATTAATTATATCAATTTTAATTTTAATTATCTTACCATTTACTTTTAATAAAAAAATTCAAGGAATTCAATTCTATC CCATTAATCAAATC------Hypoprepia fucosa ------Hypoprepia fucosa tricolor ------TTAGGGTGAGGACAAATATCATTTTGAGGTGCAACAGTAATTACTAATTTATTATCAGCTATTCCTTATTTA GGAACTATAATAGTAAATTGAATTTGAGGAGGATTCGCAGTAGATAATGCTACTTTAATTCGATTTTATACTTTCCATTT

318

TCTTTTCCCATTTATTATTTTAATAATAACTATAATTCATTTATTATTTTTACATCAAACAGGATCTAATAATCCTTTAGGA TTAAATAGAAATTATGATAAAATCCCATTTCATCCATTTTTTACATACAAAGATTTAATTGGAGCTATCATATTATTATTT ATATTAATTTTATTAACTTTAACTAATCCATATCTATTAGGAGATCCAGATAATTTTATCCCTGCAAATCCATTAGTCACT CCTATTCATATTCAACCAGAATGATATTTTTTATTTGCTTATGCCATCCTTCGATCTATCCCCAATAAACTAGGAGGAGT TATTGCATTAATCATATCAATTTTAATTTTAATTATCTTACCATTTACTTTTAATAAAAAAATTCAAGGAATTCAATTCTAT CCCATTAATCAAATC------Inopsis modulata ------Lithosia quadra ------Lyclene pyraula ---ATACGACTCACATTAGGGTGAGGACAAATATCATTTTGAGGAGCAACAGTAATTACTAATTTACTTTCAGCTATTCCT TATTTAGGTTCAATATTAGTTAATTGAATTTGAGGAGGATTTGCCGTAGATAATGCTACATTAACACGATTTTATACCTT TCATTTTTTATTACCTTTTATTATTTTAATAATAACTATAATTCATTTATTATTTTTACATCAAACAGGATCAAATAATCCTT TAGGTTTAAATAGAAACTATGATAAAATTCCTTTCCATCCATTTTTTACTTATAAAGATTTAATTGGAGCTATTATTATAT TATTTATTTTAATTATACTTACTCTTACTAACCCTTATATATTAGGAGATCCTGATAATTTTATTCCAGCTAACCCTTTAG TTACCCCTGTTCATATTCAACCAGAATGATATTTTTTATTTGCTTATGCAATTTTACGATCAATTCCTAATAAATTAGGAG GTGTTATTGCATTAATTATATCAATTTTAATTTTAATTATTTTACCTTTTACTTTCAACAAAAAAATTCAAGGAATTCAATT TTATCCTATTAATCAAATTATATTTTGATTT------Lyclene reticulata ---ATACGACTCACATTAGGGTGAGGACAAATATCATTTTGAGGGGCAACAGTTATTACTAATTTACTTTCAGCAATTCCT TATTTAGGGTCAATATTAGTTAATTGAATCTGAGGGGGGTTTGCTGTAGATAATGCTACATTAACACGATTTTATACTTT CCACTTTCTTTTACCCTTTATCATTTTAATAATAACTATAATTCACTTACTTTTTCTTCACCAAACAGGATCTAATAACCC ATTAGGATTAAATAGAAACTATGATAAAATTCCTTTCCATCCATTCTTTACTTATAAGGATTTAATTGGAGCAATTATTAT AATATTTATTTTAATTATACTCACTCTTACTAATCCTTACCTATTAGGAGATCCTGATAATTTTATTCCAGCCAATCCTTT

319

AGTCACCCCTGTTCATATTCAACCAGAATGATACTTTTTATTTGCTTATGCAATTTTACGATCAATTCCTAATAAATTAG GAGGTGTAATTGCATTAATTATATCAATTTTAATTTTAATTATTTTACCTTTTACTTTTAATAAAAAAATTCAAGGAATCCA ATTTTATCCTATTAATCAAATTATATTT------Lyclene sp.1 ------Lyclene sp.1 nr. ------Lycomorpha pholus ------Lycomorphodes sordida ------Meterhythosia sangala ------

320

Miltochrista miniata ------CGACTCACATTAGGGTGAGGACAAATATCATTTTGAGGGGCAACAGTCATTACTAATCTACTTTMTGYTATTCCTT ATTTAGGATCTATATTAGTAAATTGAATTTGAGGAGGATTTGCAGTAGATAATGYTACATTAACACGTTTTTATACTTTT CATTTTTTATTACCATTTATTATTATAATAATAACTATAATTCATTTATTATTTTTACATCAAACAGGATCTAATAACCCAT TAGGATTAAATAGAAATTATGACAAAATCCCATTTCATCCATTTTTTACATATAAAGATTTAATTGGAGCTATTATTTTAA TATTTATATTAATTATTCTTACTCTTACTAATCCTTATTTATTAGGAGACCCTGATAATTTTATTCCCGCTAATCCTTTAGT TACCCCAGTTCACATTCAACCAGAATGATATTTTTTATTTGCTTATGCTATTTTACGATCAATTCCTAATAAATTAGGAG GAGTAATTGCTTTAATCATATCAATTTTAATTTTAATTATTTTACCTTTCACTTTTAATAAAAAAATTCAAGGAATTCAATT TTACCCAATTAACCAAATTATATTTTGATTT------Miltochrista sp.1 ------Prepiella aurea nr. ------Prinasura quadrilineata ------Ptychoglene coccinea ------

321

Schistophleps albida ------Amata aperta ------Pagara simplex ------Asota heliconia ------GGGTGAGGACAAATATCATTTTGAGGAGCCACTGTTATTACCAATCTTTTATCTGCAATTCCTTATTTAG GATCAATATTAGTAAATTGAATTTGAGGAGGATTTGCAGTTGATAATGCAACCCTAACTCGATTTTATACTTTCCACTTT CTCCTTCCCTTTATTATTATAATAATAACCATAATTCATCTGCTATTTTTACACCAAACTGGATCTAATAATCCTCTTGGA TTAAATAGTAATTATGATAAAATCCCTTTTCATCCATTTTTTTCTTATAAAGATCTTATTGGAGCAATTATTATAATTTTTA TCTTAATTATATTAACCTTAACTAATCCTTATTTATTAGGAGATCCTGATAATTTTATCCCAGCTAATCCTCTTGTTACAC CAGTTCACATTCAACCAGAATGATACTTTTTATTCGCCTATGCAATTTTACGTTCAATTCCCAATAAATTAGGAGGAGTA ATTGCTTTAGTACTTTCAATTTTAATTTTAATTATTTTACCATTTACTTTTAATAAAAAAATCCAAGGAATTCAATTTTATC CTTTAAATCAAATTTTATTTTGATCTTTAGTAACTATA------Asota orbona ------ACATTAGGGTGAGGACAAATATCATTTTGAGGGGCTACTGTTATTACTAATCTTTTATCCGCAATTCCCTATTT AGGATCAATATTAGTAAATTGAATTTGAGGAGGATTTGCAGTTGATAACGCAACATTAACTCGATTTTACACTTTCCATT TTCTTTTACCTTTTATTATTTTAATAATAACTATAATTCATTTATTATTTCTACATCAAACTGGTTCTAATAATCCCCTTGG ATTAAATAGTAACTATGATAAAATCCCCTTTCATCCATTTTTCTCCTATAAAGATCTTATTGGAGCAATTATTTTAATTTT TATCTTAATTATATTAACTTTAACTAATCCTTATCTATTAGGAGATCCTGATAATTTTATCCCGGCTAACCCCCTTGTTAC

322

ACCAGTTCATATCCAACCAGAATGATATTTTTTATTTGCTTATGCAATTTTACGTTCAATTCCTAATAAATTAGGAGGAG TAATTGCTCTAGTTCTTTCAATTTTAATTTTAATTATTCTACCTTTTACTTTTAATAAAAAAATTCAAGGAATTCAATTCTA TCCATTAAATCAAATTTTATTTTGATCTTTAGTAACAATAATTATTTTATTAACATGAATTGGAGCTCGACCAGTCTTTAG TGAGGGT Neochera dominia ------ACATTAGGGTGAGGACAAATATCATTTTGAGGGGCAACTGTTATTACTAATCTTTTATCTGCAATCCCTTACTT AGGATCAATATTAGTTAATTGAATTTGAGGGGGATTTGCAGTAGATAACGCTACATTAACTCGATTCTATACCTTTCAT TTTTTATTACCATTTATTATTTTAATAATAACTATAATTCACTTATTATTCCTACATCAAACAGGATCCAATAATCCACTAG GATTAAATAGTAATTATGATAAAATTCCTTTCCATCCTTTTTTTACTTATAAAGACATTGTCGGAGCTATTTTATTAATAT TTTTATTAATTATATTAACCCTAACAAACCCATATCTATTAGGAGATCCTGATAATTTTATCCCCGCTAATCCCCTTGTA ACTCCAGTCCATATTCAACCAGAATGATATTTTTTATTTGCTTATGCAATTCTACGCTCTATTCCCAATAAACTAGGAGG AGTAATTGCATTAGTTCTTTCAATCCTTATTTTAATTATTTTACCATTTACATTTAATAAAAAAATCCAAGGAATTCAATTC TACCCTATTAATCAAATCTTATTTTGATCTCTAGTTACT------

[RpS5] Acsala anomala ------CGACTCACTATAGGGATGGCGGAGGAGAATTGGAATGATGATGCCGTAGATGCAGGCA GCATGGCTGTTGACA GCATGCCACTGCCTCAGCCGGCTGATATCCCCGAAATTAAACTGTTCGGAAGATGGAGTTGTTACGACGTTCAAGTG TCTGACATGTCTCTGCAGGATTACATCTCCGTTAAGGAAAAGTACGCCAAGTATTTACCTCACTCGGCTGGCAGGTAT GCCCACAAACGTTTCCGTAAAGCTCAGTGCCCCATCGTCGAGCGTTTGACCAACTCTCTTATGATGCACGGTCGCAA CAATGGCAAAAAACTGATGGCCGTCAGGATCGTGAAGCACGCATTTGAAATTATTCATTTGCTCACTGGTGAGAACCC TCTGCAGGTTCTTGTGACAGCCATCATTAACTCAGGCCCCCGTGAAGACTCCACCAGAATTGGTCGCGCCGGTACGG TGCGGCGGCAGGCTGTTGACGTTTCTCCTCTGCGTCGTGTTAACCAAGCCATCTGGTTATTATGCACAGGTGCACGT GAAGCTGCATTCCGTAACATCAAGACTATTGCTGAGTGTGTAGCTGATGAACTTATTAATGCAGCGAAGGGTTCATCA AACTCTTATGCCATCAAGAAGAAAGATGAGTTGGAGCGTGTT Agylla septentrionalis ------Asura cervicalis

323

------Asura polyspila ATGCTGGCAGCATGGCTGTTGAT---ATGCCTTTGCCCCCGGCAGCCGATATTCCCGAAATAAAACTGTTCGGAAGATG GAGCTGTTTTGATGTGCAGGTTTCTGATATGTCTCTGCAAGATTACATTTCAGTCAAAGAGAAGTACGCCAAATACTTA CCTCACTCGGCTGGCAGGTATGCGCACAAACGTTTCCGTAAAGCTCAGTGCCCCATCGTTGAGCGTTTAGCTAACTC TCTTATGATGCACGGCCGCAATAACGGCAAGAAACTGATGTCCGTTAGAATCGTGAAGCACGCATTCGAAATTATACA TTTGCTCACTGGTGAGAACCCTCTGCAAGTGTTGGTGACAGCTATCATTAACTCTGGGCCCCGGGAAGACTCAACTA GGATTGGTCGTGCTGGTACAGTACGTCGTCAGGCCGTCGACGTTTCCCCTCTTCGCCGTGTAAACCAAGCTATTTGG CTATTGTGCACAGGTGCACGTGAAGCTGCATTCCGTAACATCAAGACTATTGCTGAATGTGTAGCTGATGAACTGATC AATGCAGCAAAGGGATCTTCAAATTCCTATGCCATCAAGAAGAAGGACGAATTAGAACGTGTT Atolmis rubricollis TTAATACGACTCACTATAGGGATGGCGGAGGAGAATTGGAATGACGATGCCGTGGATACAGGCAGCATGGCTGTTGA CAGCATGCCATTGCCACAGGCGGCTGATATCCCCGAAATTAAACTTTTCGGAAGATGGAGTTGCTATGATGTTCAAGT GTCTGACATGTCCCTGCAGGATTACATCTCCGTTAAAGAGAAGTACGCCAAGTATTTACCTCACTCGGCTGGCAGGT ACGCGCACAAACGTTTCCGTAAAGCTCAGTGTCCCATCGTTGAACGTTTGACCAACTCTCTTATGATGCACGGCCGC AACAATGGCAAAAAGCTGATGGCCGTAAGAATCGTGAAGCACGCTTTTGAAATTATTCATTTGCTCACTGGTGAGAAC CCTCTGCAGGTTCTTGTGACAGCTATCATAAACTCCGGTCCCCGTGAAGACTCAACTAGAATTGGTCGCGCTGGTAC AGTGCGCCGGCAGGCTGTTGACGTCTCTCCTCTGCGTCGTGTGAACCAGGCAATCTGGTTACTATGCACAGGTGCA CGTGAGGCTGCATTCCGTAACATCAAGACTATTGCTGAGTGTGTAGCTGATGAACTCATCAACGCAGCAAAGGGTTC ATCAAACTCTTACGCCATTAAGAAGAAGGATGAGTTGGAACGTGTT Bruceia hubbardi ------Bruceia pulverina ------

324

------Calamidia hirta ------ATCCCCGAAATTAAGTTATTCGGCAGA TGGAGTTGTTATGATGTCCAAGTGTCTGATATGTCCCTGCAGGATTACATCTCTGTTAAAGAGAAGTACGCCAAGTAT TTACCACACTCGGCTGGCAGGTACGCGCACAAACGTTTCCGTAAAGCTCAATGTCCCATCGTTGAGCGTTTGACCAA CTCTCTTATGATGCACGGCCGCAACAATGGCAAGAAACTGATGGCCGTAAGGATCGTGAAGCACGCCTTTGAAATCA TCCATTTGCTCACTGGTGAGAACCCTCTGCAGGTTCTTGTGACTGCTATCATTAACTCTGGCCCCCGTGARGACTCAA CTAGGATCGGTCGCGCTGGTACAGTGCGTCGTCAGGCTGTGGACGTTTCTCCACTCCGT------Cisthene juanita ------Cisthene plumbea ------Cisthene subjecta ------Cisthene tenuifascia ------

325

------Crambidia lithosioides ------Cyana meyricki ------GCGGCTGATATTCCTGAAATCAAACTTTTTGG CAGATGGAGTTGTTATGATGTCCAAGTGTCAGATATGTCTCTACAGGATTACATTTCTGTTAAGGAAAAATACGCTAAG TATTTACCCCACTCTGCTGGCAGGTATGCACACAAACGTTTCCGTAAAGCTCAATGCCCCATCGTTGAGCGTTTGACC AACTCACTTATGATGCACGGCCGCAACAATGGCAAGAAGCTAATGGCTGTCAGGATTGTCAAGCACGCGTTTGAAAT CATTCATTTATTGACTGGTGAAAATCCTTTACAAGTTCTTGTAACGGCTGTCATTAATTCTGGCCCTCGTGAAGATTCA ACTAGAATTGGTCGTGCTGGTACAGTTCGTCGCCAAGCTGTTGATGTCTCTCCTCTGCGTCGTGTGAACCAGGCTAT CTGGCTATTGTGCACTGGTGCTCGGGAAGCAGCATTCCGTAATATCAAGACCATTGCTGAGTGTGTGGCC------Cybosia mesomella ------Eilema bicolor ------CGACTCACTATAGGGATGGCGGAGGAGAATTGGAACGATGATGCCGTGGAGTCAGGCAGCATGGCTGTTGACA ACATGCCATTGCCGCAGGCGGCAGATATCCCCGAAATTAAACTCTTCGGCAGATGGAGTTGTTATGATGTTCAAGTG TCTGACATGTCCCTACAGGATTACATCTCCGTTAAAGAGAAGTACGCCAAGTATTTACCACACTCGGCTGGCAGGTAT GCGCACAAACGTTTCCGTAAAGCTCAGTGTCCCATCGTTGAGCGTTTGACCAACTCTCTTATGATGCACGGCCGCAA CAATGGCAAGAAACTGATGGCCGTAAGGATCGTGAAACACGCATTTGAAATTATTCATTTGCTGACTGGTGAGAACCC TCTGCAGGTTCTTGTAACAGCTATCATAAACTCTGGGCCCCGTGAAGACTCTACTAGAATTGGTCGTGCTGGTACTGT GCGTCGTCAGGCTGTTGACGTTTCACCCCTGCGTCGTGTAAACCAGGCAATCTGGCTATTGTGCACAGGCGCACGT GAGGCTGCATTCCGTAACATCAAGACTATTGCTGAGTGTGTAGCTGATGAACTTATCAACGCAGCTAAGGGTTCATCA AACTCTTATGCCATCAAGAAGAAGGATGAGTTGGAACGT---

326

Eilema complana TTAATACGACTCACTATAGGGATGGCGGAGGAAAATTGGAACGATGAAGCCGTGGAGGCAGGCAGCATGGCTGTTG ACAGCATGCCACTGCCGCAGGCGGCAGATATCCCCGAAATTAAACTCTTCGGCAGATGGAGTTGTTATGATGTTCAG GTGTCTGACATGTCCCTGCAGGATTACATCTCCGTTAAAGAGAAGTACGCCAAGTATTTACCACACTCGGCTGGCAG GTATGCGCACAAACGTTTCCGTAAAGCTCAGTGCCCCATCGTTGAGCGTTTGACCAACTCTCTTATGATGCACGGCC GCAACAATGGCAAGAAACTGATGGCCGTAAGGATCGTGAAACACGCATTTGAAATTATTCATTTGCTGACTGGTGAGA ACCCTCTGCAGGTTCTTGTGACAGCTATCATTAACTCTGGCCCGCGTGAAGACTCTACTAGAATTGGTCGCGCTGGT ACTGTGCGTCGTCAGGCTGTTGACGTTTCACCCCTACGTCGTGTAAACCAGGCAATCTGGTTATTGTGCACAGGCGC ACGTGAGGCTGCATTCCGTAACATCAAGACTATTGCTGAGTGTGTAGCTGATGAACTTATCAACGCAGCGAAGGGTT CATCAAACTCTTATGCCATCAAGAAGAAGGATGAGTTGGAACGTGTT Eilema_dorsalis ------Eilema griseola ------TGGAATGACGATGCCGTGGAGGCAGGCAGCATGGCTGTTGACAGCATGCCACTGCCGCA GGCGGCAGATATCCCTGAAATTAAACTCTTCGGCAGATGGAGTTGTTATGATGTTCAAGTGTCTGACATGTCCCTGCA GGATTACATCTCCGTAAAAGAGAAGTACGCCAAGTATTTACCACACTCTGCTGGCAGGTATGCGCACAAACGTTTCC GTAAAGCTCAGTGTCCCATCGTTGAGCGTTTGACCAACTCTCTTATGATGCACGGCCGCAACAACGGCAAGAAACTG ATGGCCGTAAGGATCGTGAAACACGCATTTGAAATTATTCATTTGCTTACTGGCGAGAACCCTCTGCAGGTTCTAGTG ACGGCTATCATTAACTCTGGCCCCCGTGAAGACTCTACTAGGATTGGTCGCGCTGGTACCGTGCGTCGTCAGGCTGT TGACGTTTCACCCCTGCGTCGTGTGAACCAGGCTATCTGGTTATTGTGCACAGGAGCACGAGAGGCTGCATTCCGCA ACATCAAGACTATCGCTGAATGTGTAGCTGATGAGCTTATCAACGCAGCCAAGGGTTCATCAAACTCTTACGCCATCA AGAAGAAGGATGAGTTGGAACGTGTT Eilema plana ------Eilema sp.1

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------CGACTCACTATAGGGATGGCGGAGGAGAATTGGAACGACGATGCCGCCGATGCAGGCAGCATGGCTGTTGACA ACATGCCACTGCCGCAGGCGGCAGATATCCCCGAAATCAAACTCTTCGGCAGATGGAGTTGTTATGATGTGCAAGTG TCTGACATGTCCCTGCAGGATTACATCTCCGTTAAAGAAAAGTACGCTAAATACTTACCACACTCGGCTGGCAGGTAC GCGCACAAACGTTTCCGTAAGGCTCAGTGTCCCATCGTTGAGCGTTTGACCAACTCTCTTATGATGCACGGCCGCAA CAATGGCAAGAAACTGATGGCCGTGAGAATCGTGAAGCACGCCTTTGAAATCATTCATTTGCTCACTGGTGAGAACC CTCTGCAGGTTCTTGTGACGGCTATCATTAACTCTGGCCCCCGTGAAGACTCAACTAGAATCGGTCGCGCTGGTACA GTGCGTCGTCAGGCTGTTGACGTTTCTCCTCTTCGTCGTGTGAACCAGGCAATCTGGTTATTGTGCACAGGTGCACG TGAGGCTGCATTCCGTAACATCAAGACTATTGCTGAGTGTGTAGCTGATGAACTTATCAACGCAGCCAAGGGTTCTTC AAACTCTTATGCCATCAAGAAGAAGGATGAGTTGGAACGTGTT Eilema sp.2 ------CGACTCACTATAGGGATGGCGGAGGAGAATTGGAACGAAGAATCCGCGGATGCTGGCAGCATGGCTGTTGAC--- ATGCCTTTGCCCCCGGCGGCCGATATCCCCGAAATAAAACTGTTCGGAAGATGGAGCTGTTTTGATGTGCAGGTTTC TGATATGTCTCTGCAGGATTACATCTCAGTCAAGGAGAAATACGCTAAGTATTTACCACACTCGGCTGGCAGGTATGC GCATAAACGTTTCCGTAAAGCTCAGTGCCCCATTGTTGAGCGTTTAGCCAACTCTCTTATGATGCACGGCCGCAATAA TGGCAAGAAACTTATGTCCGTAAGAATTGTGAAGCACGCCTTTGAAATCATTCATTTACTCACTGGTGAGAACCCCCT TCAAGTCTTGGTGACAGCTATCATCAACTCTGGGCCCCGTGAAGACTCAACTAGGATTGGTCGTGCTGGTACAGTAC GTCGTCAAGCCGTCGACGTTTCCCCGCTGCGCCGTGTAAACCAAGCTATCTGGCTATTGTGCACAGGTGCACGTGAA GCTGCATTCCGTAACATCAAGACTATTGCTGAATGTGTAGCCGATGAACTGATTAATGCAGCAAAGGGATCATCAAAT TCCTATGCCATCAAGAAGAAGGACGAATTAGAACGTGTT Eilema sp.3 ------Eilema sp.4 ------Gardinia anoploa

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------GATATCCCAGAAATTAAGCTTTTTGGCAG ATGGAGTTGTTATGATGTTCAAGTATCTGACATGTCACTGCAGGATTATATTTCCGTTAAAGAGAAGTACGCCAAGTAT TTACCTCACTCGGCTGGCAGGTATGCTCATAAACGTTTCCGCAAAGCTCAGTGCCCCATCGTTGAGCGTTTGACCAA CTCTCTTATGATGCACGGCCGTAACAATGGCAAAAAGCTGATGGCTGTCAGGATCGTGAAACACGCCTTTGAAATTAT TCACCTGCTCACTGGTGAGAACCCTCTGCAGGTTCTTGTGACCGCTATCATAAACTCTGGACCCCGTGAAGACTCGA CCAGAATTGGCCGTGCTGGTACAGTGCGTCGTCAGGCAGTTGACGTTTCTCCTCTGCGTCGTGTGAACCAGGCTATC TGGTTATTGTGCACAGGTGCACGTGAAGCTGCATTCCGCAACATCAAGACTATTGCTGAATGTGTTGCTGATGAACTT ATCAATGCAGCAAAGGGTTCATCAAACTCTTATGCCATTAAAAAGAAGGATGAGTTGGAACGT--- Gnamptonychia flavicollis ------ATACGACTCACTATAGGGATGGCGGAGGAGAATTGGAACGATGATGCCGTGGA C---GGCAGCATGGCTGTCGACAACATGCCACTGCCCCAGGCCGCTGATATCCCCGAAATCAAACTTTTTGGCAGATG GAGTTGTTACGACGTTCAAGTGTCTGATATGTCTCTGCAGGATTACATTTCTGTCAAAGAGAAGTACGCCAAGTATTTA CCTCACTCGGCTGGCAGGTATGCGCATAAACGTTTCCGTAAAGCTCAGTGCCCCATCGTTGAGCGTTTGACCAACTC TCTTATGATGCACGGCCGCAACAATGGCAAAAAGCTGATGGCCGTAAGGATCGTGAAGCACGCTTTTGAAATTATTCA TTTGCTCACCGGTGAGAACCCTCTGCAGGTTCTAGTGACAGCTATCATTAACTCTGGGCCCCGTGAAGATTCAACTAG AATTGGTCGTGCTGGTACAGTGCGCAGACAGGCTGTCGACGTTTCTCCTCTGCGTCGTGTGAACCAGGCAATCTGGT TATTGTGCACAGGTGCACGTGAGGCTGCATTCCGTAACATCAAGACAATTGCTGAGTGTGTAGCTGATGAACTCATCA ATGCAGCAAAGGGTTCATCAAACTCTTATGCCATTAAGAAGAAGGATGAGTTGGAACGTGTT Heliosia jucunda ------Hiera gyge ------Hypoprepia cadaverosa ------

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------Hypoprepia fucosa ------Hypoprepia fucosa tricolor ------GATGCAGGCAGCATGGCTGTTGACAGCATGCCACTGCCCCAAGCGGCTGA CATCCCCGAAATTAAACTTTTCGGCAGATGGAGTTGTTATGATGTTCAAGTATCTGACATGTCACTGCAGGATTACATC TCCGTAAAAGAGAAGTACGCCAAGTATTTACCTCACTCGGCTGGCAGGTACGCTCACAAACGTTTCCGTAAAGCTCA GTGCCCCATCGTTGAGCGTTTGACCAACTCTCTTATGATGCACGGCCGCAACAATGGCAAAAARCTGATGGCCGTCA GGATCGTAAAGCACGCGTTTGAAATTATTCACTTGCTCACTGGTGAGAACCCACTGCAGGTTCTTGTAACAGCTATCA TAAACTCTGGACCCCGTGAAGACTCTACGAGAATTGGTCGTGCTGGTACAGTGCGTCGCCAGGCTGTTGACGTCTCT CCTCTGCGCCGTGTTAACCAGGCTATCTGGTTATTGTGCACAGGTGCACGTGAAGTTGCATTCCGTAATATCAAGACC ATTGCTGAATGTGTAGCTGATGAACTTATCAATGCAGCAAAGGGTTCTTCC------Inopsis modulata ------Lithosia quadra ------Lyclene pyraula ------

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------Lyclene reticulata ------CCTCCGGCGGCCGATATCCCAGAAATCAAGCTTT TCGGAAGATGGAGCTGTTTTGATGTGCAGGTGTCGGATATGTCCCTGCAAGATTACATCTCGGTGAAAGAGAAATAT GCCAAGTATTTACCGCACTCCGCTGGCAGGTATGCACACAAACGTTTCCGTAAAGCTCAGTGCCCCATCGTTGAACG TTTGACTAACTCTCTTATGATGCATGGCCGCAACAATGGTAAGAAGCTGATGGCCGTGAGAATCGTCAAACATGCGTT TGAAATAATCCATTTACTCACGGGCGAAAACCCTCTGCAAGTGTTGGTGACAGCTATCATCAATTCGGGACCTCGTGA AGACTCCACTAGGATCGGTCGTGCTGGTACCGTTCGTCGTCAAGCTGTTGACGTCTCTCCTCTGCGCCGTGTGAACC AGGCTGTATGGTTATTGTGCACAGGAGCCCGTGAAGCTGCATTCCGTAATATCAAAACTATCGCCGAGTGTGTAGCT GATGAGCTCATTAATGCAGCCAAAGGATCTTCTAATTCCTATGCAATTAAGAAGAAGGATGAGTTAGAACGTGTT Lyclene sp.1 ------Lyclene sp.1 nr. ------CGACTCACTATAGGGATGGCGGAGGAGAATTGGAACGACGATGCAGTGGAGGCTGGCAGCATGGCTGTCGATT CCATGCCCCTTCCTCCGGCGGCCGATATTCCAGAAATTAAGCTTTTCGGAAGATGGAGCTGTTTTGATGTGCAGGTTT CTGATATGTCCCTGCAAGATTACATCTCGGTGAAAGAGAAATACGCCAAGTATTTACCTCACTCCGCTGGCAGGTATG CTCATAAACGTTTCCGTAAAGCTCAGTGCCCTATTGTTGAGCGTTTGACTAACTCCCTTATGATGCATGGCCGCAACA ATGGCAAGAAGCTGATGGCCGTCAGAATCGTCAAACATGCGTTTGAAATAATCCATTTACTCACTGGCGAGAACCCTC TGCAAGTGTTGGTGACAGCTATCATCAATTCGGGACCTCGTGAAGACTCCACTAGGATCGGTCGTGCYGGTACTGTT CGTCGTCAGGCTGTTGACGTTTCTCCTCTGCGCCGTGTGAACCAGGCTATATGGTTATTGTGCACAGGAGCCCGTGA AGCTGCATTCCGTAACATCAAAACTATTGCTGAATGTGTAGCCGATGAGCTCATTAACGCAGCCAAAGGATCTTCTAA TTCCTATGCAATTAAGAAGAAGGATGAGTTAGAACGT--- Lycomorpha pholus ------

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------Lycomorphodes sordida ------Meterhythosia sangala ------Miltochrista miniata ------GACCTCACTATAGGGATGGCGGAGGAGAATTGGAATGACGATGCCGTGGACGCTGGCGGCATGGCTATTGACA ACATGCCCCTGCCCCTGGCGGCCGATATTCCCGAAATAAAACTTTTCGGAAGGTGGAGCTGTTTTGATGTACAGGTT TCTGATATGTCCCTGCAGGATTATATATCAGTCAAAGAGAAGTACGCTAAATATTTACCTCACTCAGCTGGCAGGTAT GCGCATAAACGTTTCCGTAAAGCACAGTGCCCCATCGTTGAGCGTTTGGCCAACTCTCTTATGATGCATGGCCGCAA CAACGGCAAGAAACTGATGTCCGTGAGAATCGTGAAGCATGCTTTCGAAATCATCCATTTACTCACTGGTGAGAACCC CCTGCAAGTGCTGGTCACTGCTATCATCAACTCTGGGCCCCGTGAAGACTCAACTAGGATTGGCCGTGCTGGTACTG TACGTCGTCAGGCCGTCGACGTTTCCCCTCTGCGCCGTGTGAACCAGGCTATCTGGTTATTGTGCACAGGTGCACGT GAAGCTGCATTCCGTAACATCAAGACTATTGCTGAATGTGTAGCTGATGAACTGATCAACGCAGCAAAGGGATCCTCA AATTCCTATGCCATCAAGAAGAAGGACGAGCTGGAACGTGTT Miltochrista sp.1 ------Prepiella aurea nr. ------

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------Prinasura quadrilineata ------Ptychoglene coccinea ------Schistophleps albida ------Amata aperta ------CCCCCGGCGGCCGATATCCCCGAAATAAAACTGT TCGGAAGATGGAGCTGTTTTGATGTGCAGGTTTCTGATATGTCTCTGCAGGATTACATCTCAGTCAAGGAGAAATACG CTAAGTATTTACCACACTCGGCTGGCAGGTATGCGCATAAACGTTTCCGTAAAGCTCAGTGCCCCATTGTTGAGCGTT TAGCCAACTCTCTTATGATGCACGGCCGCAATAATGGCAAGAAACTTATGTCCGTAAGAATTGTGAAGCACGCCTTTG AAATCATTCATTTACTCACTGGTGAGAACCCCCTTCAAGTCTTGGTGACAGCTATCATCAACTCTGGGCCCCGTGAAG ACTCAACTAGGATTGGTCGTGCTGGTACAGTACGTCGTCAAGCCGTCGACGTTTCCCCGCTGCGCCGTGTAAACCAA GCTATCTGGCTATTGTGCACAGGTGCACGTGAAGCTGCATTCCGTAACATCAAGACTATTGCTGAATGTGTAGCCGAT GAACTGATTAATGCAGCAAAGGGATCATCAAATTCCTATGCCATCAAGAAGAAGGACGAATTAGAACGTGTT Pagara simplex ------

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------Asota heliconia ------Asota orbona ------Neochera dominia ------

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APPENDIX D LIST OF CHARACTERS AND STATES IN PHYLOGENY OF LYCOMORPHA

List of Characters and States used in the Phylogenetic Analysis. Characters that are linked are marked with *. A unique abbreviation is provided to identify each character. Illustrations of the characters and their phylogenetic distribution are provided in the Chapter 4.

Head

1. A1: Male flagellomere state. 0) simple and ciliate, 1) serrate, 2) bipectinate.

2. A2: Female flagellomere state. 0) simple and ciliate, 1) serrate.

3. E1: Gena state. 0) well developed, can be seen as a continuous band around the eye that joins the frons when viewing the head in profile, 1) reduced, cannot be seen as a continuous band around the eye.

4. P1: Proboscis length. 0) longer than the thorax, 1) shorter than thorax but longer than the head.

5. P2: Labial palp segment fusion. 0) all segments fused, 1) 2nd and 3rd segments fused, 2) fusion absent.

Thorax

6. L1: Tibial spur formula. 0) 0-2-4, 1) 0-2-3.

Wings

7. FW1: Forewing: R free to costal margin, not anastomosing with Sc. 0) Present, 1) Absent.

8. HW1: Hindwing: Sc + R. 0) present, 1) absent.

Abdomen

9. AB1: Form of the Anterolateral Process (ALP) on the A2 apodeme. 0) sclerotized bar, 1) flattened sclerotized lobe, 2) short nob, less than half the length of the apodeme.

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Male Abdomen and Genitalia

Male Abdomen

10. * MA1: A7/A8 ventral intersegmental membrane. 0) without androconia, 1) with androconia.

11. * MA2: Form of androconia in A7/A8 ventral intersegmental membrane. ?) does not apply, 0) 2 pockets with long hair-like scales, 1) shallow, broad pocket of sex scales one-third or more the width of A7 sternite, 2) narrow pocket of sex scales less than one-third the width of A7 sternite.

12. MA3: Cephalic margin of A8 sternite. 0) fused to cephalic margin of A8 tergite, 1) extending to the cephalic margin of A8 tergite, no fusion, 2) extending onto the A8 pleurites.

13. * MA4: Sclerotization of A8 tergite. 0) present, 1) absent.

14. * MA5: Pattern of sclerotization of A8 tergite. ?) does not apply, 0) T-shaped, 1) square, 2) heartshaped, 3) rectangular.

15. MA6: A8/A9 dorsal intersegmental membrane. 0) without androconia, 1) with 3 androconia, 2 pockets with long hair-like sex scales bracketing a shallow pocket with hair-like scales.

16. * MA7: A8/A9 ventral intersegmental membrane. 0) without androconia, 1) with androconia.

17. * MA8: Form of androconia in the A8/A9 ventral intersegmental membrane. ?) does not apply, 0) 2 shallow pockets of sex scales located on the outer edges of the membrane, 1) a small pocket of sex scales located in the center of the membrane.

Genital Capsule

18. PS1: Pleural Sclerites. 0) present, 1) absent. (The presence of pleural sclerites is constant throughout all taxa included in the study.)

19. TF1: The two halves of the tegumen fused for their entire length. 0) present, 1) absent.

20. * TF2: Sutures indicating the location of the fusion. 0) present, 1) absent.

21. * TF3: Shape of the sutures on the tegumen. ?) does not apply, 0) inverted Y- shape, 1) V-shaped, 2) inverted U-shape, 3) Inverted T-shape, 4) straight line with an ovoid bulge medially.

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22. TU1: Articulation of tegumen with uncus. 0) not fused, membranous break separating the tegumen and uncus, 1) fused.

23. * UB1: Sclerotization around the uncus base. 0) present, 1) absent.

24. * UB2: Form of sclerotization around the base of the uncus. ?) does not apply, 0) reduced to narrow, strips occurring laterally on the uncus base, 1) triangular, 2) rectangular, 3) V-shaped, 4) U-shaped.

25. US1: Curvature of the uncus. 0) S-shaped, emerges from the conjuctiva and undergoes two bends, 1) C-shaped, 2) knob, 3) straight.

26. US2: Shape of the apex of the uncus. 0) fingerlike tapering to a point, 1) fingerlike not tapering to a point, 2) laterally compressed ridge, 3) ovoid, teardrop shaped, tapering to a point, 4) spade shaped, tapering to a point.

27. CM1: Distal end of costal margin occurring as an identifiable break in the sclerotization. 0) present, 1) absent.

28. * PB1: Processus basalis of costa. 0) present, 1) absent.

29. * PB2: Shape of processus basalis of costa. ?) does not apply, 0) laterally compressed triangles, 1) elongate spines, tapering to a point, 2) dorso-ventrally compressed trident shaped extensions, tips are blunt, 3) short dorso-ventrally flattened lobe.

30. TR1: Transtilla. 0) present, 1) absent.

31. * ED1: Editum. 0) present, 1) absent.

32. * ED2: Location of the editum. ?) does not apply, 0) proximally, occurs within the proximal ¼ of the costa, 1) processus basalis of the costa, 2) distally, at or beyond the proximal ¼ of the of the costa.

33. J1: Shape of the juxta. 0) rectangular with length perpendicular to the body, 1) semicircular, 2) conical projection with lateral extensions, 3) square with elongate two pronged projection, 4) square with a concave indentation in distal margin, 5) triangular with a conical tip, 6) trapezoidal, 7) rectangular with a triangular indentation in distal margin.

34. * J2: Juxta ornamentation, transparent patch. 0) present, 1) absent.

35. * J3: Shape of transparent patch. ?) does not apply, 0) triangular, 1) semicircular, 2) conical, 3) rectangular, 4) circular.

36. VS1: Shape of vinculum/saccus. 0) U-shaped, 1) V-shaped, 2) M-shaped.

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37. IV1: Ornamentation of intravincular membrane, single shallow pocket with sex scales. 0) present, 1) absent.

Phallus

38. BP1: Proximal end of basiphallus (caecum). 0) well developed, ductus ejaculatorious simplex (DES) located entirely dorsad of the caecum, 1) reduced, DES emerging from anterior, dorsal end of caecum, 2) absent, DES emerges from the anterior end of aedeagus.

39. BP2: Orientation of basiphallus. 0) straight, 1) inflected dorsally at distal end (curved away from venter), 2) inflected dorsally at midpoint (curved away from venter), 3) inflected ventrally at base (curved toward venter).

40. * PH1: Phallic sclerite. 0) present, 1) absent, highly reduced.

41. * PH2: Shape of phallic sclerite, left side. ?) does not apply, 0) flattened lobe, extending as a separate structure dorsad of the vesica, 1) rectangular, 2) narrowed, rounded triangle, 3) narrowed, pointed triangle.

42. V1: Vesica bilobed. 0) absent, 1) present.

43. * V2: Membranous projections arising from the primary lobe(s) of vesica. 0) present, 1) absent.

44. * V3: Location of membranous projections. ?) does not apply, 0) apically on the left side, proximal to the distal margin of the aedeagus, 1) distal end of main lobe, upper right apice, 2) apically on the right side, 3) medially on the right side, 4) dorsal surface of the primary lobe.

45. * V4: Ornamentation of the membranous projection located distally on the primary lobe. ?) does not apply, 0) peglike cornuti present the entire length, 1) peglike cornuti present on the distal third, remainder ruggose, 2) ruggose for the entire length.

46. * V5: Ornamentation of the vesica: heavily-sclerotized, spine like cornutus. 0) present, 1) absent.

47. * V6: Location of spine-like cornutus. ?) does not apply, 0) primary lobe, 1) distal point of the membranous projection off the vesica.

Female Abdomen and Genitalia

48. * FA1: A7 segment more heavily sclerotized than preceding segments. 0) present, 1) absent.

49. * FA2: Form of heavy sclerotization of A7.?) does not apply, 0) continuous around the segment, 1) membranous breaks in the pleurites, 2) membranous

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break occurring on the sternite, 3) membranous break on the tergite, 4) membranous breaks on pleurites and sternite.

50. FA3: Shape of A7 sternite. 0) sternite unsclerotized, no defined shape, 1) goblet- shaped, 2) M-shaped, 3) shield-shaped, 4) rectangular, length perpendicular to the body, 5) parallelogram, 6) ovoid with x-shaped posterior margin.

51. FA4: Form of distal margin of the A7 sternite. 0) deep concave indentation, extends more than a quarter the length of the A7 sternite, 1) margin approximately horizontal, 2) posterior margin membranous, not possible to distinguish from A7/A8 intersegmental membrane, 3) crenellated, 4) shallow concave indentation, extends less tha a quarter the length of the A7 sternite, 5) convex, 6) A7 distal margin elongated into 2 projections that encircle the ostium bursa.

52. * FA5: A8 sternite. 0) present, 1) absent.

53. * FA6: Form of A8 sternite. ?) does not apply, 0) heavily sclerotized plate, 1) reduced to a bar fused with A8 tergite, ventral break present, 2) reduced to a continuous sclerotized bar, fused with A8 tergite, 3) semicircular lightly, sclerotized plate, anterior margin a sclerotized bar fused with A8 tergite, 4) sclerotized plate with W-shaped anterior margin, 5) sclerotized bar fused to A7 sternite and A8 tergite.

54. * OB1: Location of the ostium bursa. 0) intersegmental membrane between A7/A8, 1) A8 sternite, 2) A7 sternite.

55. * OB2: Placement of ostium bursa when it occurs in A8 sternite. ?) does not apply, 0) fused with anterior margin of A8 sternite, 1) broad horizontal opening, centered in A8 sternite, extends over half the width of A8 sternite, 2) circular opening occurring anteriorly in A8 sternite, not fused with margin of A8 sternite, 3) narrow horizontal opening, centered in A8 sternite, extends less half the width of A8 sternite.

56. * DB1: Sclerotization of the ductus bursa. 0) present, 1) absent.

57. * DB2: Location of the sclerotized portion of the ductus bursa. ?) does not apply, 0) proximal to the ostium bursa, 1) not proximal to the ostium bursa, membranous break between ostium bursa and sclerotized portion of ductus bursa.

58. CB1: Corpus bursa. 0) wrinkled, 1) smooth.

59. CB2: Corpus bursa. 0) single, 1) two bursae, connate, arising from ductus, 2) two separate bursae with common membranous duct.

60. CB3: Signa number. 0) one, 1) two, 2) three or more.

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61. CB4: Signa form. 0) slightly sclerotized plates with heavily sclerotized bars running perpendicular to the length of the plate, 1) elongate heavily sclerotized strips with internal spines.

62. * DS1: Origin of the ductus seminalis. 0) ductus bursa, 1) corpus bursa, 2) appendix bursa, 3) second corpus bursa.

63. * DS2: Location of the ductus seminalis on the ductus bursa. ?) does not apply, 0) ventral, 1) lateral, right side, 2) lateral, left side, 3) dorsal.

64. * PG1: Dorsal pheromone gland shape. 0) two triangular projections arising from a single opening, 1) square with elongations from each apical corner.

65. * PG2: Form of the two triangular projections of the dorsal pheromone gland. ?) does not apply, 0) broad triangles, width greater or equal to length of the triangle, 1) short, narrow triangles, length of triangle greater than the base but not more than two times greater.

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APPENDIX E ALL SPECIES DATA MATRIX FOR LYCOMORPHA PHYLOGENY

0 0 1 1 2 2 3 3 4 4 5 5 6 1 6 1 6 1 6 1 6 1 6 1 6 1 Lycomorpha fulgens 11001 10101 11030 11000 20000 10010 1?501 00211 ?002? 01005 1050? 1?101 00300 Lycomorpha grotei 10102 10101 10020 11000 10040 301?0 00501 20210 20012 00002 01?0? 01001 00101 Lycomorpha miniata 11002 10101 11030 0?000 10021 301?0 1?41? 20200 0000? 01014 40011 1?001 00000 Lycomorpha pholus 11002 10101 11030 0?000 10021 301?0 1?41? 20200 0000? 01011 10011 1?001 00000 Lycomorpha pulchra 10002 10101 10020 11000 10030 301?0 0051? 20200 30010 00002 01?0? 01001 00101 Lycomorpha regulus 10002 10101 10020 11000 10030 301?0 00500 20200 20011 00002 01?0? 01001 00101 Lycomorpha splendens 21101 10101 11030 0?000 301?1 001?0 0061? 20200 101?? 1?013 1010? 1?100 00200 New Species A 10102 10101 11020 11000 10000 40030 0151? 20210 2003? 00002 0050? 00001 00000 New Species B 11002 10101 11030 0?000 10021 301?0 1?41? 20200 0000? 01011 30011 1?001 00000 New Species C 1?002 10101 10020 11000 10030 301?0 00500 20200 30011 00??? ????? ????? ????? Propyria morelosia 10002 1011? ?1030 0?000 40000 10010 00700 20200 2004? 00002 0050? 00011 01?00 Propyria normani 1?002 101?1 21030 0?000 40001 00020 01700 20200 2004? 00??? ????? ????? ????? Propyria ptychoglene 10002 10101 21030 0?000 40001 00020 01702 20201 ?004? 00016 60312 1?001 00200 Dolichesia falsimonia 00102 01110 ?11?1 10001 ?11?3 011?1 0231? 01001 ????? 1?002 40210 00001 0011? Hypermaepha 00110 01121 021?0 0?001 ?11?2 301?1 1?204 11031 ?11?? 1?035 5020? 00001 02?1? maroniensis Hypoprepia fucosa 10112 00010 ?2000 0?010 01011 011?0 0211? 01101 ?11?? 1?1?0 2022? 00002 11?1? Lycomorphodes 00102 00110 ?01?1 0?001 ?01?1 20001 1?003 11021 ?01?? 1?1?0 20413 1?001 0011? correbiodes Ptychoglene erythrophora 00002 00010 ?1010 0?010 01011 011?0 02003 01101 ?11?? 1?040 2012? 00002 12?1? Talara coccinea 00102 01120 ?01?1 0?001 ?01?3 111?1 0211? 11030 001?? 1?020 20210 1?121 03?1?

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APPENDIX F COLLECTION LOCALITY DATA FROM SPECIMENS USED TO ILLUSTRATE THE ADULT HABITUS OF LYCOMORPHA

Figure 5-2 A. L. atroxantha (Schaus), Female: Tactic, Guatemala, August, Schaus and Barnes Collection, USNM. B. L. concolor Scott, Female: Cave Ck. Canyon, Portal, Arizona, Cochise Co., 13.vii.1948, C. & P. Vaurie, AMNH. C. L. concolor Scott, Female: Madera Canyon 5800’, Santa Rita Mts., Santa Cruz Co., Arizona, 8.vii.1960, J.G. Franclemont, CUIC. D. L. concolor Scott, Male: East Turkey Creek 6400’, Chiricahua Mountains, Cochise Co., Arizona, 6.vii.1966, J.G. Franclemont, CUIC. E. L. concolor Scott, Male: Madera Canyon 5800’, Santa Rita Mts., Santa Cruz Co., Arizona, 8.vii.1960, J.G. Franclemont, CUIC. F. L. fulgens (Edwards), Female: Palmerlee, Arizona, USNM. G. L. fulgens (Edwards), Female: Huachuca Mts., Arizona, USNM. H. L. fulgens (Edwards), Female: Wet Canyon, Pinaleño Mts., Arizona, 8.vi.1967, J.H. Hessel, AMNH. I. L. fulgens (Edwards), Male: Ramsey Canyon, Huachuca Mts., Cochise Co., Arizona, 28.v.1964, R.F. Sternitzky, Collection of Fred H. Rindge, AMNH. J. L. fulgens (Edwards), Male: Redington, Arizona, USNM.

Figure 5-3 A. L. fulgens (Edwards), Male: Redington, Arizona, M. Chrissman, 15.ix.1902, Holland Collection, CMNH. B. L. fulgens (Edwards), Male: Chihuahua Townsend, 16.ix, Holland Collection, CMNH. C. L. grotei (Packard), Female: Black Canon Cimarron, Colorado, 13-15.ix.1917, R.C. Shannon, CUIC. D. L. grotei (Packard), Female: So. Utah, Poling, 1.vii.1910, USMN. E. L. grotei (Packard), Male: Sylvan Lakes, South Dakota, 13.viii.1912, R.A. Leussler, AMNH. F. L. grotei (Packard), Male: Eureka, Utah, 27.viii.1911, Tom Spalding, USNM. G. L. miniata Packard, Female: Joe Dollar Gulch, Hill City, Black Hills, South Dakota, 2.viii.1964, D.C. Ferguson, USNM. H. L. miniata Packard, Male: Joe Dollar Gulch, Hill City, Black Hills, South Dakota, 3.viii.1964, D.C. Ferguson, YPM. I. L. morelosia (Schaus), Female: Barranca, Mixcoac, Mexico, 20.viii.1917, C.C. Hoffman, AMNH. J. L. morelosia (Schaus), Male: Zacualpan, Mexico, ix.1914, Dognin Collection, USNM.

Figure 5-4 A. L. neomexicanus Scott, Male: New Mexico: Union Co., Johnson Mesa, 10mi N Folsom, 21.vi, Sta. 448, leg. H.K. & M.A. Clench, 1977, C.M. Acc. 29258, CMNH.

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B. L. normani (Schaus), Male: Cuernavaca, Mexico, vii.1906, Collection Wm. Schaus, USNM. C. L. pholus (Drury), Female: Big Indian V’y, Catskill Mt’s, New York, 3.vii.1906, R.F. Pearsall, Accession No. 11291, Collection Brklyn Mus., USNM. D. L. pholus (Drury), Female: Oklahoma: Sequo. Co., Lake Tenkiller, 2mi NW Blackgum, 11-14.vi.1981, D. & M. Davis, USNM. E. L. pholus (Drury), Male: USA: VA: Page Co., Pinnacles, Shenandoah Nat. Park 3400’, 5.viii.1979, Leg. D.C. Feguson, USNM. F. L. pholus (Drury), Male: Oklahoma: Kiowa Co., 7mi SW Lone Wolf, 3.iv.1979, R.J. McGinley, USNM. G. L. ptychoglene (Hampson), Female: Jalapa, Mexico, Collection Wm. Schaus, USNM. H. L. ptychoglene (Hampson), Male: Mexico, Dognin Collection, USNM. I. L. pulchra Dyar, Female: Cuyamaca State Pk., California, 17.vi.1943, S.S. Nicolay, LACM. J. L. pulchra Dyar, Female: Chariot Canyon, Banner, California, 11.x.1946, R.P. Allen Collector, CAS.

Figure 5-5 A. L. pulchra Dyar, Female: Forest Home, California, 7.iv.1927, Engel Coll., Carn. M. Acc. 13257, CMNH. B. L. pulchra Dyar, Male: Mill Cr. Cn., San Bernadino Co., California, 21.ix.1923, E.P. Van Duzce Collector, CAS. C. L. pulchra Dyar, Male: Chariot Canyon, Banner, California, 11.x.1946, R.P. Allen Collector, UCB. D. L. pulchra Dyar, Male: Forest Home, California, 7.iv.1927, LACM. E. L. regulus (Grinnell), Female: S. Fork Big Rock Cr., San Gabriel Mts., L.A. Co., California, 23.ix.1971, el. 4500, coll. C. Henne, Chris Henne Collection purchased 1978 LACM Foundation, LACM. F. L. regulus (Grinnell), Female: Upper Camp., Pinery Canyon, Chiricahua Mts., Cochise Co., Arizona, 5.vii.1956, Collected by Lloyd M. Martin, John A. Comstock, & William A. Rees, LACM. G. L. regulus (Grinnell), Male: Emgd. 18.ii.1968, ova ex. conf. female, So. Fork Big Rock Cn., L.A. Co., California, 18.x.1967 4500’, coll. C. Henne, Chris Henne Collection purchased 1978 LACM Foundation, LACM. H. L. regulus (Grinnell), Male: Diamond Rock, White Mts., Arizona, 30.viii.1947, Collection of Grace H. and John L. Sperry, AMNH. I. L. splendens Barnes and McDunnough, Female: Utah: Kane Co., Fivemile Valley, 5000’, (Hwy. 89, 5 road mi. W of Paria R.), 4.ix.1968, leg. J.F. Emmel & A.O. Shields, YPM. J. L. splendens Barnes and McDunnough, Male: Fort Davis, Texas, Jeff Davis Co., 5.x.1969, A. & M.E. Blanchard, USNM.

Figure 5-6 A. L. texanus Scott, Female: Texas, USNM. B. L. texanus Scott, Male: Texas, Collection Brklyn Museum, USNM.

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Figure 5-7 L. pelopia (Druce), Female: Mexico: Guerrero, 5-7km NW Taxco, 1850-1900M, 14.ix.1982, J.A. Powell and J.A. Chemsak Collectors, UCB.

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BIOGRAPHICAL SKETCH

Clare Hilary Scott was born in Columbia, . She graduated from

D.W. Daniel High School in May 2003. Clare began her undergraduate studies in

August 2003 at the University of . During her time at the university, Clare was able to complete undergraduate research in Dr. John Wares’ population genetics lab and Dr. Joe McHugh’s insect systematics lab. In May 2007, she graduated with high honors and received a Bachelor of Science degree in genetics and a Bachelor of

Sciences in Environmental Sciences degree in entomology. In August 2007, Clare enrolled in a Ph.D program in the University of Florida’s Department of Entomology and

Nematology. Her major advisor was Dr. Marc Branham. While at the University of

Florida, Clare had the opportunity to mentor to undergraduate students, Pablo Chialvo and Liset Perez. In addition, she was able to participate in the Science Partners in

Inquiry-based Collaborative Education (SPICE) GK-12 fellowship program, which allowed her to obtain teaching experience at a Mebane Middle School in Alachua,

Florida. Upon completeing her doctoral degree, Clare will begin a postdoctoral position at Purdue.

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