SYSTEMATICS OF THE MEGADIVERSE SUPERFAMILY (INSECTA: LEPIDOPTEA)

DISSERTATION

Presented in Partial Fulfillment of the Requirements for The Degree of Doctor of Philosophy in the Graduate School of The Ohio State University

By

Sibyl Rae Bucheli, M.S. *****

The Ohio State University 2005

Dissertation Committee: Approved by Dr. John W. Wenzel, Advisor Dr. Daniel Herms Dr. Hans Klompen ______Dr. Steven C. Passoa Advisor Graduate Program in

ABSTRACT

The phylogenetics, systematics, , and biology of Gelechioidea (Insecta:

Lepidoptera) are investigated. This superfamily is probably the second largest in all of

Lepidoptera, and it remains one of the least well known. Taxonomy of Gelechioidea has

been unstable historically, and definitions vary at the and levels. In

Chapters Two and Three, I review the taxonomy of Gelechioidea and characters that have

been important, with attention to what characters or terms were used by different authors.

I revise the coding of characters that are already in the literature, and provide new data as

well. Chapter Four provides the first phylogenetic analysis of Gelechioidea to include

molecular data. I combine novel DNA sequence data from Cytochrome oxidase I and II

with morphological matrices for exemplar . The results challenge current concepts

of Gelechioidea, suggesting that traditional morphological characters that have united

taxa may not be homologous structures and are in need of further investigation.

Resolution of this problem will require more detailed analysis and more thorough

characterization of certain lineages. To begin this task, I conduct in Chapter Five an in-

depth study of morphological evolution, host- selection, and geographical

distribution of a medium-sized Haworth (), larvae of

ii

which generally feed on in the families and . Host-plant use is

commonly studied in this group because of physiological and behavioral responses

exhibited by Depressaria pastinacella to furanocoumarins produced by their host plants, yet no species level phylogeny is available. The phylogeny of Nearctic Depressaria is

constructed using a morphological data matrix analyzed under the parsimony criterion.

This study is the only modern phylogeny of the genus, and includes all North American

species but one, and about half the Old World species. I redescribe these species. In

Chapter Six I describe nine new species of Hübner (Scythridinae) from the

Galápagos Islands, Ecuador, and provide a key and illustration of genitalia and abdominal

modifications. Finally, Chapter Seven represents an application of taxonomy to

address questions of sampling protocols used for studies of and conservation.

I use Gelechioidea in eastern North America as indicators of diversity, with attention to

the effectiveness of different sampling protocols with respect to active versus passive

sampling, and plot-based versus plotless sampling. A list of Gelechioidea was produced

from trap sites from an Appalachian forest in southern Ohio. The composition and

diversity of Ohio Gelechioidea captured in a passive, plot-based protocol compares

favorably to more exhaustive sampling, and reinforces recent (and counterintuitive)

recommendations that it is more efficient and repeatable to focus surveys on target

groups in focal localities rather than to conduct extensive sampling programs.

iii

Dedicated to Randle, my best friend

And we don't notice any time pass we don't notice anything we sit side by side in every class teacher thinks that I sound funny but she likes the way you sing

Tonight I'll dream while I'm in bed when silly thoughts go through my head about the bugs and alphabet and when I wake tomorrow I'll bet that you and I will walk together again cause I can tell that we're going to be friends

We are Going to be Friends – The White Stripes

iv

ACKNOWLEDGMENTS

Graduate school is a strange place. I owe many people my gratitude for their patience and support during this time: mentors, colleagues, friends, family, and students.

Without their help and encouragement, I would not be the scholar that I am today.

I would like to thank my advisor, Dr. John W. Wenzel, for his investment in me.

His commitment to my dissertation work and belief in me as a scientist have given me ability to continue my work. I am grateful for the many hours of discussion, writing, proofreading, and editing he has provided. There has not been one day in the last seven years that Dr. Wenzel has been in the museum and has not been available for his graduate students.

Dr. Christopher Randle, Husband and Botanist, has been with me since the first day of my post-high school education. With him I have grown as a person and a scientist.

Above all, his intellect and admiration inspire me to persevere. Just listen to your heart, that's what I do.

I would like to thank my committee members, Dr. Hans Klompen, Dr. Daniel

Herms, and Dr. Steven Passoa, who have provided me with vital evaluation of this document. In particular, Dr. Passoa has been critical to the completion of this project. A mentor in my studies of Lepidoptera, he has aided in identification, supplied and assisted

v

in the collection of specimens, and provided discussion. He has also been a magnificent supplier of items useful to my dissertation, many of which I had no idea I needed.

I would like to thank Dr. Jean-François Landry for his attention: it was he who revealed to me the subtle allure of . My graduate career began (with a study of ) and ended (with a study of Scythris) under his tutelage.

Many other scientists have shaped this work, as well. Jerry Powell, Daniel

Rubinoff, David Wahl, Hugo Kons, Lauri Kaila, and David Horn have provided specimens. Ronald Hodges, Richard Brown, Jean-François Landry, David Wagner, John

Rawlins, Joël Minet, Patrice Leraut, Lauri Kaila, Jerry Powell, David Adamski, Paul

Goldstein, and Fred Stehr have provided comments through the years regarding this work. David Adamski and John Brown were always helpful during museum study at the

NMNH. Brian Scholtens provided data from the GSMNP ATBI, and David Wagner provided unpublished data for Lepidoptera of Connecticut. John Herbert, Foster

Purrington, Chad Schone, and Eric Dotseth provided helpful comments on the LAWCO manuscript. John Herbert and Foster Purrington helped with the initial moth sorting.

Christopher Randle was immensely helpful in DNA alignment, phylogenetic analysis, and discussion. John Wenzel translated text written in French, and Christopher Randle translated text written in German. Mary Daniels provided technical assistance in the initial stages of the molecular projects.

My friends have had an equally significant influence. Conversations with my lab mates of the past and present have hugely benefited my work as a graduate student. In particular, Todd Blackledge, Kurt Picket, Hojun Song, Joe Raczkowski, and Ryan Caesar have played a central role in entomological and phylogenetic discussions. Hojun Song,

vi

with whom I have shared an office for several years, has not only provided stimulating discussion regarding the evolution of genitalia but also helped to keep me sane (or at least minimized a drastic increase in madness). I thank all of my friends at the Museum of Biological Diversity, especially members of the Phylogenetic Discussion Group, who helped to polish my phylogenetic skills. I also thank all of my friends in the departments of Entomology and EEOB who have helped me through the years. Mark Mort and

Christopher Randle shared their limited space with me while I was in Kansas.

I thank my family, the Randles (Biz, Noël, Jinx (my own private editor!),

Shonkie, William, Alex, Lizzie, Jackson, Dan-O, Fiona, Bridget, Jack, and Violet) and the Buchelis (Momma, Dad, Grammy, Pat, Brad, Patrice, Baby Joey, and Emma), for their love and support.

Finally, I thank my mother, Christina Arlia, who I believe is responsible for my love of . When I was a small child, I would bring to her tokens of my love: hideous six-legged tokens that bit and stung and made terrible smells. Instead of freaking out and killing them, she would tell me that they were "mommy-bugs" that had to go home to their babies. I grew up thinking that all insects were wonderful and caring creatures, just like my mother.

vii

VITA

May 22, 1973……………………………..………..……… Born – Tampa, Florida 1996………………………………………………..………. B.A. Biology, Hiram College 1999………………………………………………………... M.S. Entomology, The Ohio State University

PUBLICATIONS

Bucheli, S. R., J.-F. Landry, and J. W. Wenzel. 2002. "Cladistic Analysis of Larval Case Architecture and Implications of Host-Plant Associations for North American Coleophora (LEPIDOPTERA: )", 18, 71-93.

Bucheli, S. R. and J. W. Wenzel. 2005. Gelechioidea (Insecta: Lepidoptera) systematics: A reexamination using combined morphology and mitochondrial DNA data. Molecular Phylogenetics and Evolution 35, 380-394.

Bucheli, S. R., D. Horn, and J. W. Wenzel. In Press. Biodiversity of Gelechioidea (microlepidoptera): An assessment of a re-established Appalachian forest in southern Ohio. Biodiversity and Conservation.

FIELDS OF STUDY

Major Field: Entomology

Specialization: Systematics of Lepidoptera, especially Gelechioidea and other microlepidoptera, phylogenetics, morphological evolution, and host-plant evolution.

viii

TABLE OF CONTENTS

Abstract………………………………………………………………………………ii

Dedication…………………………………………………………………………...iv

Acknowledgements…………………………………………………………………..v

Vita………………………………………………………………………………….viii

List of Tables………………………………………………………………………..xiv

List of Figures……………………………………………………………………….xv

Chapters:

1. Introduction……………………………………………………………………….1

2. Taxonomic and Systematic History of Families and of Gelechioidea

Fracker 1915 (Lepidoptera)…………………....…………………………………….12

2.1. Introduction……………………………………………………………..12

2.1.1. Taxonomic and Systematic Treatment of Gelechioidea……...14

2.2. Discussion………………………………………………………………29

2.2.1. Significant trends in Gelechioidea …………………………...29

3. Annotation and Analysis of Phylogenetically Important Characters for Families and

Subfamilies of Gelechioidea Fracker 1915 (Lepidoptera..………………………….46

3.1. Introduction……………………………………………………………..46

ix

3.2. Discussion………………………………………………………………52

3.2.1. Monophyly of Gelechioidea………………………………….52

3.2.2. Adults…………………………………………………………53

3.2.3. Larvae…………………………………………………………68

3.2.4. Pupae………………………………………………………….75

4. Gelechioidea systematics: A reexamination using combined morphology and mitochondrial DNA data…………………………………………………………….83

4.1. Introduction……………………………………………………………..83

4.2. Materials and Methods………………………………………………….89

4.2.1. Taxon Sampling and Morphological Analysis……………….89

4.2.2. DNA Amplification, Sequencing and Alignment…………….93

4.3. Analyses and Discussions………………………………………………96

4.3.1. General perspective…………………………………………..96

4.3.2. Morphology…………………………………………………..96

4.3.3. MtDNA……………………………………………………….99

4.3.4. Combined analysis…………………………………………….101

4.3.5. Comparison with Kaila (2004)………………………………..105

4.3.6. Character Evolution…………………………………………..107

4.4. Conclusions……………………………………………………………..101

5. North American Flat-Body (: Depressariinae: Depressaria

Haworth): Morphological evolution, host-plant selection, and geographic distribution………………………………………………………….……………….113

5.1. Introduction……………………………………………………………..113

x

5.1.1. Host-Plant Use………………………………………………..114

5.1.2. Species Group Definitions……………………………………119

5.2. Materials and Methods………………………………………………….122

5.2.1. Taxon Sampling……………………………………………….122

5.2.2. Character Sampling……………………………………………126

5.2.3. Morphological characters……………………………………...126

5.2.4. Phylogenetic Analysis…………………………………………153

5.3. Results…………………………………………………………………...154

5.3.1. Taxon Descriptions……………………………………………159

5.4. Discussion……………………………………………………………….209

5.4.1. Generic Relationships…………………………………………213

5.4.2. Monophyly of Depressaria, the Ingroup………………………215

5.4.3. Evolution of Species Groups of Depressaria………………….215

5.4.4. Major Morphological Trends…………………………………221

5.4.5. Evolution of Host-Plant Associations………………………...230

5.4.7. General Remarks………………………………………………231

5.4.8. The use of Genital Characters in Phylogenetics………………234

5.5. Conclusions………………………………………………………….234

6. New species of Scythris Hübner 1825 (Gelechioidea: Xyloryctidae: Scythridinae) from the Galápagos Islands………………………………………………………….236

6.1. Introduction……………………………………………………………..236

6.2. Materials and Methods………………………………………………….238

6.3. Taxonomy……………………………………………………………….240

xi

6.3.1. Key to species of Galápagos Island Scythris…..……………...241

6.4. Discussion………………………………………………………………302

7. Sampling to assess a re-established Appalachian forest in Ohio based on gelechioid moths (Lepidoptera: Gelechioidea)………………………………………… ………304

7.1. Introduction……………………………………………………………..304

7.1.1. Lepidoptera as Survey Specimens…………………………….307

7.2. Materials and Methods………………………………………………….308

7.2.1. Study Sites…………………………………………………….308

7.2.2. Collection……………………………………………………..312

7.2.3. Identification………………………………………………….314

7.2.4. Cumulative Totals for Lawrence County Gelechioidea………314

7.3. Results…………………………………………………………………..315

7.3.1. Gelechioidea Diversity of Lawrence County…………………315

7.4. Discussion………………………………………………………………319

7.4.1. Extrapolation to total species…………………………………323

7.4.2. Assessment of Site Quality…………………………………...325

7.5. Conclusions……………………………………………………………..329

Bibliography…………………………………………………………………………332

Appendix A ………………………………………………………………………….346

Appendix B…………………………………………………………………………..353

Appendix C…………………………………………………………………………..356

Appendix D………………………………………………………………………….379

Appendix E…………………………………………………………………………..378 xii

Appendix F…………………………………………………………………………..386

xiii

LIST OF TABLES

Table Page

2.1 Taxonomy of Gelechioidea ………………………………………………... 32

xiv

LIST OF FIGURES

Figure Page

1.1. Relationships of Ditrysian Lepidoptera………………………………………..3

1.2. Adults of Gelechioidea………………………………………………………...5

1.3. enormis (: Batrachedrinae) showing synapomorphies

of Gelechioidea as defined by traditional characters…………………………..9

3.1. Characters of special interest used by various authors to define families and

subfamilies of Gelechioidea.………………………………...... ……....48

3.2. Coleophora trifolli showing split valves, presence of juxta, presence of

gnathos as fused, spined knob, absence of uncus……………………………77

3.3. showing free aedeagus, presence of socii and

uncus (reduced), presence of membranous transtilla, entire valves…………78

3.4. Scythris showing ankylosation of aedeagus to valvae and vinculum……….79

3.5. showing patches of spiniform setae…………………….80

3.6. Coleophora trifolii showing pupal antennae not meeting at the meson…….81

3.7. Lateral condyles present or absent ………………………………………....82

4.1. Phylogenic tree showing sister-group relationships within

Gelechioidea redrawn from Passoa 1995……………………………………85

xv

4.2. Phylogenic tree showing sister-group relationships within

Gelechioidea redrawn from Hodges 1998…………………………………..87

4.3. Names and collection localities for taxa used in total evidence

analysis..………………………………………..……………………………92

4.4. One of 18 most parsimonious trees from the combined morphology matrix,

showing one of two main topologies………………………………………..98

4.5. The single most parsimonious tree (L=2147, CI=0.29, RI=0.35) produced from

molecular matrix (CO-I+CO-II) showing branch lengths…………………..100

4.6. Consensus phylogeny (L= 2331) of two most parsimonious trees

(L= 2321, CI = 0.29, RI = 0.38) produced from the combined analysis

(recoded morphology+CO-I+CO-II)...... 104

4.7. produced by combining Kaila’s morphological data

with the molecular data of Figure 4.5……………………………………….107

5.1 Hypothetical phylogeny of Depressariinae based on the published phylogeny by

Berenbaum and Passoa (1999) and cladistic analysis of Depressaria species

groups (unpublished work by Passoa reanalyzed by Bucheli (See Appendix D

for the data matrix))………………………………………………………..118

5.2. Species of Depressaria used in the analysis……………………………….124

5.3. Genital features of male Depressaria douglasella (Gelechioidea: Elachistidae:

Depressariinae)………………………………………………………………132

5.4. Genital features of Depressaria absynthiella………………………………..134

5.5. Genital features of Depressaria albipunctella...... 135

5.6. Genital features of Depressaria angustati…………………………………..136

xvi

5.7. Genital features of Depressaria atrostrigella……………………………….137

5.8. Genital features of ……………………………………138

5.9. Genital features of Depressaria cinereocostella…………………………….139

5.10. Genital features of Depressaria constancei…………………………………140

5.11. Genital features of ……………………………………141

5.12. Genital features of Depressaria discipunctella………………………………142

5.13. Genital features of Depressaria eleanorae…………………………………..143

5.14. Genital features of Depressaria haydenii……………………………………144

5.15. Genital features of Depressaria multifidae…………………………………..145

5.16. Genital features of Depressaria petronoma…………………………………146

5.17. Genital features of ………………………………..147

5.18. Genital features of Depressaria silesiaca……………………………………148

5.19. Genital features of Depressaria ultimella……………………………………149

5.20. Genital features of Depressaria weirella…………………………………….150

5.21. Genital features of Depressaria yakinae…… ……………………………….151

5.22. Genital features of Depressaria libanoditella………………………………..152

5.23. Variation of vinculi within Depressaria……………………………………..153

5.24. Consensus phylogeny (L = 167, C.I. = 0.35, R.I. = 0.76) of 24 most parsimonious

trees (L = 162, CI = 0.37, RI = 0.77) produced from morphological matrix in

Appendix E…………………………………………………………………..155

5.25. Consensus phylogeny showing evolution of Depressaria species and species

groups...... 211

xvii

5.26. Consensus phylogeny of Depressaria showing evolution of key morphological

features………………………………………………………………………219

5.27. Evolution of host-plant associations for Depressariinae…………………….223

5.28. Consensus phylogeny showing distribution of Depressaria species………..228

6.1. Generalized male genitalic features of Galápagos Island Scythris

(Gelechioidea: Xyloryctidae: Scythridinae) in lateral view…………………243

6.2. Details of Galápagos Island male genitalia in posterior view……….………245

6.3. Abdominal features of Scythris cuneata…………………………………….250

6.4. Genitalic features of Scythris cuneata………………………………………251

6.5. Distribution of Scythris cuneata…………………………………………….252

6.6. Abdominal features of Scythris galapagosensis…………………………….257

6.7. Genitalic features of Scythris galapagosensis………………………………258

6.8. Distribution of Scythris galapagosensis…………………………………….259

6.9. Abdominal features of Scythris falcata……………………………………..263

6.10. Genitalic features of Scythris falcata…………………………………….....264

6.11. Distribution of Scythris falcata……………………………………………..265

6.12. Abdominal features of Scythris pistillata……………………………….….269

6.13. Genitalic features of Scythris pistillata…………………………………….270

6.14. Distribution of Scythris pistillata………………………………………..…271

6.15. Abdominal features of Scythris ancystra………………………………….275

6.16. Genitalic features of Scythris ancystra……………………………………276

6.17. Distribution of Scythris ancystra………………………………………….277

6.18. Abdominal features of Scythris sinuosa…………………………………..281

xviii

6.19. Genitalic features of Scythris sinuosa…………………………………….282

6.20. Distribution of Scythris sinuosa…………………………………………..283

6.21. Abdominal features of Scythris furculata…………………………………288

6.22. Genitalic features of Scythris furculata……………………………………289

6.23. Distribution of Scythris furculata……………………………………….…290

6.24. Abdominal features of Scythris darwini…………………………………...294

6.25. Genitalic features of Scythris darwini…………………………………..….295

6.26. Distribution of Scythris darwini………………………………………...….296

6.27. Abdominal features of Scythris bernardlandryi…………………………….299

6.28. Genitalic features of Scythris bernardlandryi………………………………300

6.29. Distribution of Scythris bernardlandryi…………………………………….301

7.1. Map of Lawrence County, Ohio, showing location of

historical furnaces………………………………………………………….310

7.2. Photograph of Ridge Furnace…………………………………………311

7.3. Comparison of collection methods between Lawrence County, Ohio (LAWCO),

and the Great Smoky Mountains National Park Lepidopteran All Taxon Bio

Inventory sponsored by Discover Life In America (GSMNP ATBI, data

extrapolated from Wagner and Scholtens 2002)…………………………..313

7.4. Comparison of Gelechioidea species collected during the LAWCO study and

GSMNP Lepidoptera ATBI (data provided by Brian Scholtens)………….316

7.5. List of species recovered from Lawrence County in June…………………317

7.6. Number of species for LAWCO study per township and site……………...319

xix

7.7. Comparison of the LAWCO study to the GSMNP ATBI (Wagner and Scholtens

2002), “total Ohio” study (Summerville and Crist 2003) and Connecticut (D. L.

Wagner, unpublished) for location, sampling methods, time of study and area of

study; total and projected number of Gelechioidea and percent LAWCO diversity

versus the other areas………………………………………………………..323

7.8. Actual and projected cumulative totals for Lawrence County Gelechioidea

recovered using passive, plot-based blacklight collection methods...... 325

7.9. Species accumulations as a proportion of total in regenerated Lawrence County,

Ohio sites……………………………………………………………………327

xx

CHAPTER 1

INTRODUCTION

Knowledge of Lepidoptera comes mainly from the study of charismatic or

economically significant groups of moths and . Much of the information that

remains to be discovered regarding behavior, ecology, systematics, and host-plant choice

of Lepidoptera will be from less notorious groups, in particular the microlepidoptera.

Until we understand their biology fully, we cannot claim to understand the order as a whole.

The cosmopolitan superfamily Gelechioidea is one of these little-known groups.

Gelechioidea are one of the great radiations toward the crown of the evolutionary tree of

Lepidoptera, consisting of approximately 1,425 genera and 16,250 described species

(Hodges, 1998). Hodges (1998) estimates that only 25% of the species diversity of

Gelechioidea has been described. If this estimate is accurate, diversity of Gelechioidea

would rival that of the , with total species reaching counts of almost 65,000

species (Kristensen and Skalski 1998). The radiation of Gelechioidea is believed to be

1

rather recent, although the origin of the lineage may be at least 120 million years old

(Labandeira et al, 1994).

The position of Gelechioidea within the order Lepidoptera remains uncertain. To date, no comprehensive cladistic treatment of the order is available, although modern treatments based on Hennigian argumentation (where derived character states define relationship but numerical analysis based on a matrix is not preformed) or phylogenies loosely pieced together are available. In a influential work by Minet in 1991,

Gelechioidea are considered to be a lineage within , an hypothesis that is based on groundplan coding. Most notable of these characteristics that unite Gelechioidea with the Ditrysia are females having a copulatory opening separate from the ovipore with an internal duct system connecting the sperm-receiving organ to the oviduct. According to

Minet, there are four Ditrysian superfamilies: ( (

(Yponomeutidae + Gelechioidea))) (Figure 1.1, A). Kristensen and Skalski (1998), based on Minet’s 1991 study, placed Gelechioidea as part of a derived unresolved polytomy:

((Simaethistoidea, Tineoidea)(Gracillarioidea, Gelechioidea, )) (Figure

1.1, B). In either scenario, Gelechioidea are possibly a sistergroup of the Apoditrysia; other possibilities are Yponomeutoidea and Gracillarioidea. Of course, sister-groups are necessary for character polarization, and position of the Gelechioidea within the order in general and relative to Apoditrysia in particular has serious implications not only for morphological character evolution but behavior and ecological statements, as well.

2

Figure 1.1. Relationships of Ditrysian Lepidoptera, A. Redrawn from Minet (1991); B.

Redrawn from Kristensen and Skalski (1998).

It is difficult to characterize Gelechioidea cohesively in terms of morphology, behavior and ecology due to its species richness. Gelechioidea moths range in size from very small to medium and are generally cryptically colored, nocturnal . There are, of course, exceptions, and one finds large, brightly colored diurnal animals (Figure 1.2).

Even within families, it is difficult to make species diagnoses and the main working taxonomic unit for Gelechioidea seems to be the subfamily level, although many researchers use the taxonomy of family and subfamily interchangeably. Some researchers believe the superfamily should be split into several superfamilies to better diagnose 3

lineages. Sinev (1993) and Kuznetsoz and Stekol’nikov (1979, 1984, 2001) promote the idea of a supergroup or infraorder with four to six superfamilies within it. However, they disagree with eachother about nomenclature and placement of smaller lineages within more inclusive units. The problem with splitting Gelechioidea is that, until recently, no cladistic analyses existed for the superfamily and delineation of supertaxa would have been a priori, certainly making taxonomy and systematics even more of a nightmare than it already is. Gelechioidea systematists have two primary fears: the creation of monotypic lineages or the creation of paraphyletic groups. I base this last statement on conversations I have had with many experienced taxonomists and systematists. While all agree that Gelechioidea is in dire need of attention, all warn against hasty family and subfamily treatments. After years of study, still superficial in comparison to so many experts past and present, I understand what was meant. A common attitude among

Gelechioidea researchers is that smaller taxonomic units may not be better taxonomic units. In the future, we may be faced with the challenge of deciding between several monotypic taxa or a large, heterogeneous superfamily, but such decisions should be based on very careful cladistic analyses. Several recent studies agree that a likely topology for

Gelechioidea is a Hennigian comb and a monophyletic crown, making delineation of superfamilies within an infraorder unadvisable as it would create what we seem to fear most: more monotypic taxa and paraphyletic groups, undoing what little progress has been made. In opposition to the above topological findings, Kaila’s recent treatment recognizes two main monophyletic groups but he wisely decides not to make any taxonomic revisions.

4

Figure 1.2. Adults of Gelechioidea. All specimens shown in plate are on loan from

Museum National d'Histoire Naturelle, Paris, France. A. Coleophoridae (Coleophorinae).

Coleophora species, Adult (top) Larval case on Rosa species (bottom); B.

(Lecithocerinae). C. (Stathmopodinae). Tortillia fllavella Chrét; D.

Stathmopodidae (Stathmopodinae).Stathmopoda jidella Chrét; E.

(Holcopogoninae). Holcopogon bubulcellus Stau.; F.

(). salicella (Hb.) Male (top), Female (bottom); G.

Xylorictidae (Xylorictinae). Stilbaromacha ratella H.-S.; H. Xyloryctidae

(Cryptophasidae). Cryptophasa sp. Meyr; I. Xyloryctidae (Cryptophasidae). Enolmis

kollarella Costa; J. Hypertrophinae. Thudaca sp.; K. Coleophoridae (Coleophorinae)

Pterolonche albescens Zell.; L. Schistonoeidae oecophila Stgr.; M. Lecithoceridae

(Lecithocerinae) Lecithocera orsoviella Hein.; O. (Symmocinae)

nigromaculla Rag.

5

6

Fracker first proposed Gelechioidea in 1915 using primarily larval characters. In

1916, Mosher redefined Gelechioidea using pupal characters and stated that Gelechioidea

was closely related to Yponomeutoidea. According to Mosher, two different developmental forms suggested that Gelechioidea should be divided into two

superfamilies, those where the fronto-clypeal suture is retained and those where it is

reduced or absent, but she did not have enough evidence to “warrant such a conclusion.”

Forbes (1923), utilizing characters from all life stages, was the first to recognize the

superfamily Gelechioidea as having “palpi almost always upturned beyond middle of

front, the third segment long and pointed, regularly tapering for most of its length”,

“tongue usually moderate, scaled at base”, and “maxillary palpi, when present,

characteristic, minute, but of the folded type, and curving over base of tongue.” He also

treated larval characters of the superfamily, stating that the larvae always have three setae

on the prespiracular wart, and a single subventral setae on the meso- and metathorax with

the prothoracic spiracle normal. He noted that abdominal setae iv and v are closely

approximated, i and ii are separated and usually at nearly the same level, and that ii of the

ninth abdominal segment is not much nearer the mid-dorsal line than the other subdorsals

and is usually nearer i than to “its mate” (See Stehr and Martinat, pg 324, couplet 108,

part b for discussion of this character). In his treatment, Forbes stated, “The Gelechioidea

form the most homogeneous of the subordinate groups of microlepidoptera, and the

largest of those groups.” Taxonomy of Gelechioidea remained relatively unstable after

Forbes’ definition and many authors made revisions (Comstock, 1920; Meyrick, 1928;

McDunnough, 1938; Common, 1970; Bradley, 1972). Hodges (1978) who investigated

7

world taxa with a primary focus on North America presented the first modern concept of the group and asserted his classification scheme was tentative because characters, such as wing venation and genitalia, were often in disagreement with each other. He based family names on distinctive groups of genera, stating that when genital characters are considered in conjunction with other characters, the relationships and relative rank of many names must be reconsidered. Hodges recognized Gelechioidea as having the head smooth scaled, the labial palpus upturned and sickle-shaped, the tongue scaled on the anterior surface basally, and the maxillary palpus one to four segmented and folded over the base of the tongue. Hodges characterization has remained the working definition. Kaila

(2004) challenges Hodges superfamily definition based on a parsimony analysis that does not support any of these characters but he does not provide an alternative definition. As of this date, Gelechioidea are still defined by tradition characters (Figure 1.3).

8

Figure 1.3. Batrachedra enormis (Batrachedridae: Batrachedrinae) showing synapomorphies of Gelechioidea as defined by traditional characters. A. Upturned labial palpus; B. Scaled haustellum; C. Folded maxillary palpus of Batrachedra enormis (Batrachedrinae).

9

Gelechioidea are ideal for questions regarding the evolution of host-plant choice

because of their wide array of larval life histories, including scavenging, gall forming,

-mining, seed-mining, leaf-tying, leaf-rolling, stem-boring, -boring and case-

making, mostly on gymnosperms and angiosperms (Powell, 1980; Powell et al., 1998).

Intimate connection to the host plant (such as in gall-makers, miners, or borers) produces

obligate specialization in some lineages, whereas others seem to colonize new hosts

freely despite similar habits. Phylogenetic study is only now producing successful synthetic theories of the evolution of host preference. In Bucheli et al. (2002) I used phylogenetic analysis of Coleophora (Gelechioidea: Coleophoridae: Coleophorinae),

integrating and extending earlier ecological models (Denno, 1995; Feeny, 1975; Janz and

Nylin, 1998), to demonstrate that these moths prefer certain plant tissues (seeds versus

) and growth forms (herbaceous versus woody) with exploitation of different plant

taxa, rather than preferring certain plant taxa with exploitation of different plant tissues or

forms. Mechanistic explanations are available from the earlier models, but none covered

the spectrum of larval feeding habits that I did. I rejected coevolution as an explanatory

process in the diversity of feeding syndromes in Coleophora. It seems likely that similar,

great advances in our understanding of gelechioid ecology will follow, as the general

phylogenetic structure of Gelechioidea is refined.

My research focuses on the phylogenetics, systematics, taxonomy, and biology of

Gelechioidea. In my dissertation, I focus on five main topics: 1) history of taxonomy and

character use for Gelechioidea, 2) molecular and morphological evolution of

10

Gelechioidea, 3) diversity of Gelechioidea in eastern North America, 4) in-depth study of

morphological evolution, host-plant selection, and geographical distribution of a

medium-sized genus of Gelechioidea (Elachistidae: Depressariinae: Depressaria), and 5) description of new species from a poorly known genus of Gelechioidea (Xyloryctidae:

Scythridinae: Scythris).

11

CHAPTER 2

TAXONOMIC AND SYSTEMATIC HISTORY OF FAMILIES AND SUBFAMILIES

OF GELECHIOIDEA FRACKER 1915 (LEPIDOPTERA)

INTRODUCTION

Gelechioidea is a large, heterogeneous superfamily of Ditrysian microlepidoptera

that is difficult to characterize due to its diversity of adult and larval morphologies, as

well as a variety of disparate ethologies and ecologies. If one thing must be said to

describe this lineage to an inexperienced entomologist, I would say that they are probably

the most commonly encountered moths at light traps at night, as common as Noctuoidea

and , but probably the most commonly overlooked, as well, due to their

small size and cryptic brownish-gray coloration. The average entomology student feels the same about Gelechioidea as they feel about microhymenoptera, acalypterate muscoid

flies, and small, black beetles: dread and anguish at the task of preparation and

identification. Perhaps the reason such a feeling persists may be lack of easily accessible

reference material; information is rarely considered to be high-priority and is frequently

12

published in obscure journals and is hardly ever found in one comprehensive, easily

accessible volume.

However; Gelechioidea are considered by many lepidopterists to be the one of the

most beautiful and intriguing animals, largely due to their small size, delicate bodies,

intricately colored wings, and ephemeral and mysterious lifestyles. They have received

much attention by passionate entomologists. Taxonomic and systematic progress has

been slow overall since the creation of the superfamily with many scholars undoing or

completely changing the previous system. Only recently have there been attempts at

comprehensive treatments that aim to address world fauna of the entire superfamily rather

than focusing on a few local families. One reason, of course, is the accumulation of data.

It is easy to treat a fauna where only a handful of species within six or so families exist,

but as technology advances so must we. The concept of Gelechioidea has gone from

being part of the catch-all lineage Tineoidea to having its own questionable superfamily status, to being considered a robust lineage, back to having a questionable status as a

monophyletic superfamily.

This chapter traces the higher level taxonomy of Gelechioidea in terms of family

and subfamily starting with its description in 1915 by Fracker and concluding in 2004

with Kaila’s most recent phylogenetic investigation. While many authors have treated

Gelechioidea, I include here only those who have made an impact on concepts of families

and subfamilies that later influenced more contemporary works on the superfamily.

13

Taxonomic and Systematic Treatment of Gelechioidea

Before their official definition in 1915, most species of Gelechioidea were considered to be part of Tineoidea, as were most microlepidoptera. Gelechioidea was first proposed by Fracker in 1915 using primarily larval characters. He included in the superfamily: Ethmiidae, Stenomidae, , Oecophoridae, and

Cosmopterygidae (sic). He considered Scythridinae a subfamily of Yponomeutidae

(Yponomeutoidea). He did not place Elachistidae or Coleophoridae in any superfamily.

His classification does not include a treatment of characters for the superfamily. He gives a brief treatment of characters at the family level, stating that families are difficult to separate at any stage , “…the interrelations of the families cannot be worked out from them altho (sic) there are a significant number of differences to separate them more or less completely from each other.”

In 1916, Mosher used pupal characters to define lepidopteran relationships. She stated that Gelechioidea were closely related to Yponomeutoidea, and placed

Coleophoridae in Yponomeutoidea, while including in Gelechioidea the following families: Lavernidae (, in part), Scythridae (sic), Gelechiidae,

Chrysopeleiidae, Oecophoridae, Stenomidae, Cosmopterygidae (sic), and Elachistidae.

According to Moser, there were two forms of developments which divided

Gelechioidea into two groups: the group retaining the fronto-clypeal suture (Lavernidae,

Scythridae (sic), Gelechiidae, and Chrysopeleiidae), and those in which it is not distinct

14

or absent (Oecophoridae, Stenomidae, Cosmopterygidae (sic), and Elachistidae). Mosher

believed that the latter group may represent a distinct superfamily, but did not have the

evidence needed to “warrant such a conclusion.”

Forbes (1923) utilized characters from all life stages. He placed Coleophoridae back in Tineoidea, and placed in part back in Yponomeutidae

(Yponomeutoidea). He proposed the superfamily Cycnodioidea and included

Cycnodiidae (Elachistidae) and (Incurvarioidea; Elachistidae, in part). He included in Gelechioidea the following families: Oecophoridae, Xyloryctidae,

Gelechiidae, Blastobasidae, and Lavernidae (Momphidae; Cosmopterygidae (sic);

Elachistidae, in part).

Forbes was the first to recognize the superfamily Gelechioidea as having “palpi almost always upturned beyond middle of front, the third segment long and pointed, regularly tapering for most of it’s length”, “tongue usually moderate, scaled at base”, and

“maxillary palpi, when present, characteristic, minute, but of the folded type, and curving over base of tongue.” He also treated larval characters of the superfamily, stating that the larvae always have three setae on the prespiracular wart, and a single subventral setae on the meso- and metathorax with the prothoracic spiracle normal. He noted that abdominal setae iv and v are closely approximated, i and ii are separated and usually at nearly the same level, and that ii of the ninth abdominal segment is not much nearer the mid-dorsal line than the other subdorsals and is usually nearer i than to “its mate.”

15

In his treatment, Forbes stated, “The Gelechioidea form the most homogeneous of the subordinate groups of microlepidoptera, and the largest of those groups”, but as noted above, his definition of Gelechioidea was less inclusive than the modern treatments and excluded Coleophoridae and Scythrididae.

In the family , Handlirsch (1924) placed the following subfamilies:

Tineinae (Augasmini; Coleophoridae in part), Hyponomeutinae (Sythridini)

(Depressariini, Chimabachini, Gelechiini, Oechophorini, Amphisbatini, Coleophorinae

(Momphini, Coleophorini, Cosmopterigini), Elachistinae (Elachistini, Cemiostomini) and incertae sedis in the family he placed: Xyloryctinae, , Metachandinae

(), Physoptilinae, Epimarptinae, , and Pterolonchinae.

Tillyard 1926 placed Elachistidae, Oecophoridae, Xyloryctidae, and Gelechiidae in Tineoidea saying that most species are small with hindwings that are narrow, usually have a haustellum and labial palpi, and have reduced venation (M usually absent) with long fringe.

In his Checklist of Lepidoptera of Canada and the United States, McDunnough

(1939) kept Coleophoridae in Tineoidea, and elevated Scythridae (sic) to a family within

Yponomeutoidea, along with Elachistidae. He included in Gelechioidea the following families: Cosmopterygidae (sic), (Copromorphoidea), Gelechiidae,

16

Oecophoridae, Blastobasidae, Stenomidae and Ethmiidae. McDunnough did not include a

treatment of characters used to define his categorization.

In Comstock’s (1940) “An Introduction to Entomology”, he classifies the

families Coleophoridae, Elachistidae, Oecophoridae, Ethmiidae, Stenomatidae,

Gelechiidae, Blastobasidae, and Scythrididae with in the AA. The

Frenatæ Lepidoptera (where the two wings of each side are united by and frenulum or its

substitute and the hind wing has a large humeral area), subcategory BB. The Specialized

Frenatæ (where the base of vein M has been lost and the branches of this vein are joined

to veins R and Cu), subcategory C. The Specialized Microfrenatæ (where frenulum

bearing moths are usually small and have an anal area with 3 anal veins or an area of

fringe acts as a substitute when the hind wings are very narrow). The families are not

united together in a superfamily, however, Comstock notes that while other authors may

group The Specialized Microfrenatæ into various superfamilies, he did not believe these groups were well established enough to be used in his book. Comstock comments that

Scythrididae are closely allied with Yponomeutidae, and writes, “I do not find that any tangible characters of the adults insects separating the two families have been pointed out; but there appear to be differences in the setal characters of the larvae (see Fracker

15).”

Turner (1947) included in Tineoidea (Subdivision Microptila, Division

Asthenochorda) the families Elachistidae (Coleophorinae, Scythrinae (sic), Elachistinae,

17

Douglasianae (sic), and Cosmopteryginae (sic)), and Gelechiidae (Thalmarchellinae (sic)

(Thalamarchellinae, Depressariinae), Oecophorinae, Blastobasinae, Gelechiinae and

Xyloryctinae). Other than stating that Meyrick 1928 was wrong about character interpretation (i.e. convergence through asthenogenesis1) and therefore, wrong about the

classification of Tineoidea, Turner gives no reason other than “neuration and other

characters” for his classification scheme.

Common (1970) was the first to treat world taxa using larval, pupal and adult

characters. All prior treatments were based on taxa of America, north of Mexico.

Common was the first to include Coleophoridae in the superfamily Gelechioidea. It also

included: Agonoxenidae, Elachistidae, Scythridae (sic), Stathmopodidae, Oecophoridae,

Ethmiidae, Timyridae (Lecithoceridae), Cosmopterigidae (Cosmopteriginae, Momphinae,

Walshiinae), Metachandidae (Oecophoridae), Anomologidae, ,

Blastobasidae, Xyloryctidae, Stenomidae, Gelechiidae, Physoptilidae, and

Strepsimanidae.

Bradley (1972) investigated the British fauna and in his checklist proposed the

following classification for Gelechioidea: Coleophoridae, Elachistidae, Oecophoridae

(Oecophorinae, Depressariinae), Ethmiidae, Gelechiidae (Gelechiinae, Symmocinae),

Blastobasidae, Stathmopodidae, Momphidae (Batrachedrinae, Momphinae,

Cosmopteriginae, Blastodacninae, and Chrysopeleiinae) and Scythrididae.

1 Turner’s use of the term asthenogenesis is referring to convergence through the loss of characters or neotinization, I think.

18

Hodges (1978) investigated world taxa with a focus on primarily North American

species. He asserted his classification scheme was tentative stating that characters, such

as wing venation and genitalia, were often in disagreement with each other. He based

family groups names on distinctive groups of genera, stating that when genital characters are considered in conjunction with other characters, the relationships and relative rank of

many names must be reconsidered. Hodges recognized Gelechioidea as having the head

smooth scaled, the labial palpus upturned and sickle-shaped, the tongue scaled on the

anterior surface basally, and the maxillary palpus one to four segmented and folded over

the base of the tongue.

Hodges treated world taxa proposing both a classification and suggested a

systematic scheme for some families in the superfamily. He recognized the following

families, subfamilies and tribes in Gelechioidea: Oecophoridae (Depressariinae

(Amphisbatini, Depressariini), , Peleopodinae, , Xyloryctinae,

Stenomatinae, Oecophorinae (Oecophorini, Stathmopodini, Pleurotini), Chimabachinae,

Hypertrophinae), Elachistidae (Coelopoetini, Elachistinae), Pterolonchidae,

Coleophoridae (Coleophorinae, Batrachedrinae), Agonoxenidae (Agonoxeninae,

Blastodacninae (Blastodacnini, Parametriotini), Blastobasidae (Blastobasinae

(Blastobasini, Pigritiini), Symmocinae), Momphidae, Scythridae (sic), Cosmopterigidae

(Cosmopteriginae, , Chrysopeleiinae), and Gelechiidae (,

Gelechiinae, , , Chelariinae, Lecithocerinae,

Physoptilinae). He transferred Strepsimanidae to Noctuoidea.

19

Kuznetsov and Stekol’nikov (1979) investigated the systematic position and

phylogenetic relationships of the superfamily Coleophoroidea based on functional

morphology of the male genitalia. They created the superfamily Coleophoroidea and

placed in it the families Coleophoridae, Ethmiidae and Oecophoridae (Pleurotinae,

Symmocinae, Deuterogoniinae, Oecophorinae, Depressariinae, Chimabachinae). They

stated that affinity between Gelechioidea and Coleophoroidea was not confirmed, and

that Coleophoroidea was most closely related to Tortricoidea based on the insertion of the

flexors of the gnathos (M2) and the tergal flexors of the valves (M4) muscles.

In 1984, Kuznetsov and Stekol’nikov investigated Gelechioidea and

Coleophoroidea more wholly based on functional morphology of the male genitalia. They

created two more superfamilies, Elachistoidea and Copromorphoidea. They removed

Ethmiidae from Coleophoroidea and placed it in Elachistoidea along with Elachistidae

(Elachistinae, Agonoxeninae), and Blastodacnidae (Blastodacninae, Parametriotinae).

They kept Coleophoridae and Oecophoridae in Coleophoroidea, adding the families

Batrachedridae, Blastobasidae, and Momphidae. Subfamilies of Oecophoridae remained unchanged. The Gelechioidea were then composed of the following families:

Cosmopterigidae (Cosmopteriginae, Antequerinae, and Chrysopeleiinae),

Stathmopodidae, Scythridae (sic), Pterolonchinae, and Gelechiidae (Gelechiinae,

Teleiodinae, , Helariinae, Dichomeridinae, Metzneriinae). In

Copromorphoidea, they placed Xyloryctinae (Xyloryctinae, Acriinae), Aeolanthidae

20

(fam. n.), and Lecithoceridae (Ceuthomadarinae, ). Aeolanthidae was created

on the basis of an extreme reduction of the tergal appendages and the origin of flexors of

the gnathos (M2) from a special sclerotized strand.

The Coleophoroidea are closely related to the Elachistoidea by the presence of the

M2 and the tergal flexors of the valves (M4) muscles. The Elachistoidea are united by the primitive presence of the juxto-sacculalis (M13), the vinculo-valvalis ventralis (M14), and tergo-sternalis dorsoventralis (M22) muscles, not present in Coleophoroidea.

Based on larval characters, Stehr (1987) placed in Gelechioidea: Oecophoridae,

Depressariinae (Ethmiinae, Stenomatinae, Batrachedrinae (inc. Batrachedra and

Homaledra noting that “they don’t fit well here, either”), Elachistinae, Oecophorinae,

Symmocinae, Elachistinae, Coelopoetinae), Blastobasidae, Coleophoridae, Momphidae,

Agonoxenidae (inc. Bastodacna), Cosmopterigidae, Scythrididae, and Gelechiidae

(Anomologinae, Gelechiinae, Chelariinae, Dichomeridinae). Stehr did not give a diagnosis of Gelechioidea, but did include a family key to larvae of Lepidoptera. He stated that “the delineation of families within Gelechioidea is difficult at best, and next to impossible at worst, even when a relatively restricted geographical region is considered.”

Minet (1990) changed concepts of Gelechioidea by revising the Elachistidae. He made these changes in taxonomy based on Henigian argumentation and using characters of the adults, pupae and larvae. His classification is as follows: Pterolonchinae,

Coleophoridae (Coleophorinae, Amphisbatinae), Elachistidae (Agonoxeninae,

21

Elachistinae, Stenomatinae, Cryptolechiinae, Hypertrophinae, Ethmiinae,

Depressariinae), Peleopodidae, Chimabachidae, Xyloryctidae, Batrachedridae,

Oecophoridae, Symmocidae, Lecithoceridae, Epimarptidae, Blastobasidae,

Stathmopodidae, Momphidae, Cosmopterigidae, and Gelechiidae.

In the Insects of , Nielson and Common (1991) recognize the following

families of Gelechioidea: Oecophoridae, Batrachedridae, Hypertrophidae, ,

Coleophoridae, Elachistidae, Blastodacnidae, Agonoxenidae, Ethmiidae, Blastobasidae,

Momphidae, Cosmopterigidae, Gelechiidae, Symmocidae, Holcopogonidae,

Lecithoceridae, and Scythrididae.

Leraut (1992) expanded Minet’s 1990 concept of Gelechioidea using characters

of the apodemes, spiniform setae, genitalia and associated structures, palpi, and antennae in addition to Minet’s pupal characters. Leraut defined Gelechioidea as: Pterolonchidae,

Coleophoridae, (Amphisbatinae, Coleophorinae), Elachistidae (Agonoxeninae,

Elachistinae, Stenomatinae, Aeolanthinae, Cryptolechiinae, Hypertrophinae (Tonicini,

Hypertrophini, Hypercalliini) Ethmiinae, Depressariinae (Depressariini, Epigraphiini,

Fushiini, Telechrysidini)), Peleopodidae, Chimabachidae, Carcinidae, Xyloryctidae,

Batrachedridae, Oecophoridae (Oecophorinae, Philobotinae), Symmocidae

(Symmocinae, Oegoconiinae), Lecithoceridae, Scythrididae, Epimarptidae,

Blastobasidae, Stathmopodidae, Momphidae, Cosmopterigidae, and Gelechiidae

(Aristoteliinae, Holcopogoninae, Apatetrinae, Gelechiinae, Dichomeridinae).

22

Fetz (1994) investigated larvae and pupae of selected gelechioid taxa,

Oecophoridae, Gelechiidae, Symmocidae, Scythrididae and Blastobasidae, in terms of

phylogenetic systematics. He believed all species examined belong to a “gelechiod super-

taxon” („gelechioiden Groβtaxons”), however, he could not demonstrate the delimitation.

He used the following characters as ground-plan synapomorphies for his “gelechioid

super-taxon”: SD 1 of A9 hairlike or secondarily setiform, and additional secondary setae

in ventral and dorsoventral area of anal prolegs.

He designated two partial groups within “gelechiod super-taxon.” Oecophorid

Group I is composed of “Depressariinae”, Chimabachinae, Ethmiinae, Anchiinae,

Carcina quercana, , , Cacophyia permixtella, and

“Gelechiidae” (where quotation marks indicate probable paraphyletic groups). This group

is united by the characters of having the adfrontal suture reaching the cranial suture,

cranial setae P1 and P2 in a horizontal line. Oecophorid Group II is composed of

Oecophorinae, united by the character of having the SV1 setae on abdominal segments 3

through 6 distinctly separate from SV2 setae. His phylogenetic relationship of

Gelechioidea can be summarized as follows: Oecophorid groups I + II + Symmocinae,

Pleurota + Topeutis +Scythrididae + Blastobasidae, with the remaining unexamined taxa

as a basal . Fetz unites these taxa by 6 apomorphies: the adfrontal suture not

reaching cranial incision, postmentum with pit-shaped depression, setae VI on the

prothoracic segment half as long as on the meso- and metathoracic segments,

development of chordotonal receptors on SD1 setae of abdominal segments 1 through 8

23

(with the following characters associated with this character: microscopic SD2 near

pinaculum of SD1, development of an apodeme at the base of SD1 on A 1-8 and fusion

with chordotonal organ, and pinaculum of SD1 on A 1-8 appearing as uninterrupted

ring), AV2 setae 1 through 4 on the anal prolegs well isolated, L setae 1 and 2 on

abdominal segments 1 through 8 arranged perpendicular to one another L1 more than twice as long as L2.

Using Hennigian argumentation and studying “all taxa of the family rank of the world fauna of gelechioid moths” (250 total genera), Sinev (1993) designated the following relationship for six superfamilies (also designated Groups A-F) in the infraorder Coleophoromorpha: ((Coleophoroidea and Oecophoroidea) ((Elachistoidea and

Gelechioidea) (Chrysopeleioidea and Cosmopterigoidea))). Sinev used genera from the

Yponomeutoidea and Copromorphoidea as “comparative material.” He justified this classification scheme on the idea of these six main morphological groups (his new superfamilies) that were delineated on the basis of eight main characters and other descriptive features. (Coleophoroidea + Oecophoroidea) is maintained by the gnathos being sclerotized and unpaired, the uncus fused, and the terga with spines. The relationship of ((Elachistoidea + Gelechioidea) (Chrysopeleioidea + Cosmopterigoidea)) is supported by the femur of the first pair of legs of the being concealed. The relationship of Chrysopeleioidea + Cosmopterigoidea is supported by having the gnathos

2 This is Fetz’s nomenclature. The AV setae of the anal prolegs are equivalent to SV setae of the abdominal prolegs.

24

and uncus absent, gnathos sclerotized and paired, and the aedeagus ankylosed. The

relationship of Elachistoidea + Gelechioidea is not well supported.

Sinev’s classification creates 6 new superfamilies under the purported monophyletic infraorder Coleophoromorpha, and elevates Chrysopeleioidea to a

monotypic superfamily.

Sinev suggests that the ancestral condition for Gelechioidea was to be rather

large-bodied with wide, lanceolate, wings that had a rounded apex and complete,

homoneurous venation with a trend Lepidoptera towards a decline in body size and a

resultant reduction in wing size and venation as well as an increase in the relative amount of fringe on the hind-wing. He considers feather-winged forms to be a parallelism and rather derived within families. He also notes a few instances of increased body size and a trend towards a morphology similar to that of the (a condition I like to refer to as pyraloidism) within these lineages.

The ancestral genital apparatus was “characterized by exceptional complexity” and making its characters “indispensable” in the search for reliable apomorphies. The terminal lobes were well-developed with a trend towards an unpaired, fused uncus; the gnathos, unlike the structures which become the uncus, was not part of the original groundplan, and appears from the tegumen and diaphragma. Sinev considers sclerotization of the gnathos to be more derived. The ancestral phallus was freely moving

and consisted of three layers: a sclerotized aedeagus and a sclerotized phallotheca within

the folds of the intersegmental membrane. The trend was for a decrease in sclerotization

towards more membranous structures that are either free or ankylosed (fixed). Sinev

25

suggests that many taxa have secondarily sclerotized phallic structures with a submergence of the aedeagus within the body wall, and a further sclerotization of the anellus leading to a reduction of the aedeagus in these lineages.

Passoa (1995), working almost independently from Fetz, studied Gelechioidea in a cladistic framework using mainly larval and pupal characters, many similar to those used by Fetz (1994). His results were similar in some regards. He also saw two main divisions within Gelechioidea as well as two groups of oecophorid taxa. His first main group contains ((Oecophoridae I (Stenomatinae, Amphisbatinae, Ethmiinae,

Depressariinae)) + (Oecophoridae II (Blastobasinae, Oecophorinae, Autostichinae)) +

Deocloninae)) sister to Scythrididae. This group (perhaps another case pf pyraloidism?

Passoa, pers. comm.) is sister to a clade which he called “Gelechiiformes”, containing

((Cosmopterigidae + Gelechiidae) + Batrachedridae + Momphidae + Coleophoridae +

Pterolonchidae). Passoa united Scythrididae + Deocloninae + Oecophoridae (s. l.) based

on the autapomorphic character of the larvae having sclerotized rings around SD1 on A

1-7. He believed Oecophoridae (s. l.) to be united by two synapomorphies, both

homoplastic: the M7 muscle (of genitalia) present and pupal labial palpi hidden.

Oecophoridae I is united by pupae having lateral condyles while Oecophoridae II is

characterized by having larvae that are mainly scavengers. The Gelechiiformes are united

by the character of the larvae having A9 with D2, D1, and SD1 (or at least D1 and SD1)

in a vertical line.

26

Hodges (1998) designated 15 families with 32 subfamilies from 37 family- and

subfamily-level terminals. Hodges revised, extended and synonymized many family and

subfamily relationships, and created three new families and four new subfamilies (see

Appendix I). Hodges’ efforts towards creating a comprehensive understanding of world

fauna of Gelechioidea have been greatest contribution to Gelechioidea systematics so far.

He was the first to investigate the whole superfamily in a cladistic framework.

Kuznetsov and Stekol’nikov (2001), again working on functional morphology of

male genitalia, established the monophyletic super taxon Gelechiiformes consisting of

four superfamilies. Elachistoidea, which contains (((((Elachistidae + Agonoxenidae) +

Ethmiidae) + Hypertrophidae) + Aeolanthidae) +Stenomidae). Coleophoroidea, which

contains (((((((((((Batrachedridae + Blastobasidae) + Peleopodidae) + Coleophoridae) +

Momphidae) + Deoclonidae) + Oecophoridae) + Schistonoeidae) + Glyphidoceridae) +

Stathmopodidae) + Lecithoceridae) + Xyloryctidae). Pterolonchoidea which contains

only Pterolonchidae, and finally, Gelechioidea, which contains (Cosmopterigidae +

Chrysopeleiidae) + ((((Gelechiidae (Metzneriinae, Anacampsinae, Teleiodinae,

Chelariinae, Dichomeridinae)) + Scythridae (sic)).

Kaila (2004) investigated the megadiverse lineage Gelechioidea in a cladistic

frame work using species as terminals rather than groundplan coding for lineages, providing the most comprehensive phylogeny to date for 156 taxa with 193 characters.

He is the first to treat Gelechioidea using species as terminals and presents the most

27

detailed morphological study to date. Kaila is also the first to thoroughly investigate

outgroup relationships. While Kaila’s treatment has many novel characters (such as those

of the thorax), it is based largely on Hodges’ and Passoa’s data matrices, expanding many

of Passoa’s and Hodges multi-state characters into separate characters (he states that, “No originality is claimed for the characters and their state definitions, although the author is unaware of literature references for many of them.”) Kaila designates a “gelechiid lineage” and an “oecophorid lineage”, each being monophyletic and weakly supported

(Bremer support of 2 (Bremer 1988)) in his analysis. His phylogeny is wisely not a revision and he does not make any changes to the superfamily. He does not include three genera in the final analysis due to their shifting of positions within Gelechioidea:

Epimarptis philocoma (Batrachedridae: Epimarptinae (Hodges, 1998)) shifted between

Coelopoeta, Stathmopoda and Batrachedra; Martyringa ussuriella (Oecophoridae:

Oecophorinae (Hodges, 1998)) shifted between Oecophorinae and the xyloryctid assemblage; and Letogenes festalis (Peleopodidae (Hodges 1998)), was associated with

Lecithoceridae and Symmocinae, or as the most basal lineage of Elachistidae.

The gelechiid lineage includes Deoclona (Deocloninae), Epimarptis

(Epimarptinae [not included in the final analysis]), (Syringopainae),

Coelopoeta (Oecophorinae), (Batrachedrinae), Stathmopoda

(Stathmopodinae), Idioglossa and Batrachedra (Batrachedrinae), and

Coleophora (Coleophorinae), Momphidae, Pterolonchidae, Scythrididae, Gelechiidae and

Cosmopterigidae. The oecophorid lineage includes Holcopogoninae, Symmocinae,

Glyphidocerinae, the , Autostichinae and Lecithocerinae; the xyloryctid

28

assemblage (with Hierodoris, Izatha and Phaeosaces) and with Deuterogoniinae and

Blastobasinae nested among ‘‘true’’ xyloryctids (Scieropepla, Nemotyla, Uzucha,

Tymbophora, Lichenaula and Xylorycta), a narrowly delimited ‘‘core’’ Oecophoridae;

Amphisbatidae s.s.; Carcinidae; Stenomatidae; Chimabachidae and Elachistidae (with

Hypertrophinae as delimited by Minet (1990)), Depressariinae, Ethmiinae (with

Orophia), Aeolanthinae, Parametriotinae, Agonoxeninae and Elachistinae. Some

interesting sister-group relationships include close association of Scythrididae to

Momphidae, Pterolonchidae, Gelechiidae and Cosmopterigidae; Batrachedridae and

Coleophoridae; and Blastobasidae and the xyloryctid lineage.

DISCUSSION

Significant trends in Gelechioidea

One re-occurring issue of systematics is whether Gelechioidea should be one or many superfamilies. Several authors have proposed a classification where Gelechioidea has been broken into two or more superfamilies (Costa Lima, 1939; Kuznetsov and

Stekol’nikov, 1979, 1984, 2001; Sinev, 1993). Support for the splitting is given by

Kuznetsov and Stekol’nikov (1979, 1984, and 2001) based on musculature of the male genitalia and Sinev (1993), who named several reasons including larval feeding strategy.

Most frequently, it is recommended by these authors that coleophorid taxa, pterolonchid

taxa, and elachistid taxa represent their own, independent lineages that are unique enough

to merit superfamily status.

29

More recent studies suggest that Gelechioidea is probably one superfamily united by several synapomorphies (see Chapter 3). This is strongly supported by Hodges’ research (1974, 1978, 1983, 1986, and 1998). Most recently, however, Kaila (2004) has recovered through phylogenetic analysis a monophyletic Gelechioidea (although not upheld by traditional, uncontroverted synapomorphies) with two main lineages: a gelechiid lineage and an oecophorid lineage. His findings are similar to those of Passoa’s

(1995). Kaila does not suggest that the superfamily be split as of yet.

Another trend, more common in distant Gelechioidea history, is the question of which families are part of Gelechioidea, as opposed to belonging to purportedly closely related and definitively non-Gelechioid lineages, such Yponomeutoidea. Coleophoridae is one such family; others include Scythrididae, Elachistidae, Xyloryctidae, and

Cosmopterigidae. Prior to its creation by Fracker, taxa of Gelechioidea belonged to

Tineoidea (as did most microlepidoptera) or Yponomeutoidea.

Another issue is the question of taxonomic status of lineages within Gelechioidea, particularly what status should be granted. It is frequent that an author elevates a lineage, such as Momphidae (Hodges, 1978) to a family level, and then almost immediately relegates it back to a subfamily (Momphinae, Hodges 1998).

A final and perhaps most significant problem is internal rearrangement of families and subfamilies by all authors, i.e. inclusion or exclusion of lineages within families, subfamilies, and tribes. It has historically been difficult when authors make changes to the taxonomy but do not include a justification (frequently lists are provided without explanation of rearrangement). It is only recently that this problem has become tractable

30

as taxonomy becomes more intertwined with phylogenetics and seldom are taxonomic revisions made without support from a phylogeny. Authors have characters and data matrices to back such critical decision.

31

Fracker 1915 Mosher 1916 ______YPONOMEUTOIDEA YPONOMEUTOIDEA Yponomeutidae Coleophoridae Scythridinae GELECHIOIDEA GELECHIOIDEA Lavernidae Ethmiidae Scythridae (sic) Stenomidae Gelechiidae Gelechiidae Chrysopeleiidae Oecophoridae Oecophoridae Blastobasidae Stenomidae Cosmopterygidae (sic) Cosmopterygidae (sic) Elachistidae Not placed Elachistidae Coleophoridae (continued on next page)

Table 2.1. Taxonomy of Gelechioidea.

32

Table 2.1 continued.

Forbes 1923 Handlirsch 1906 ______TINEOIDEA TINEOIDEA Coleophoridae Tineidae Tineinae YPONOMEUTOIDEA Augasmini (Coleophoridae in part) Yponomeutidae (Scythrididae, in Hyponomeutinae part) Scythridini Gelechiinae CYCNODIOIDEA Depressariini Cycnodiidae (Elachistidae) Chimabachini Heliozelidae (Elachistidae, in part) Gelechiini Oechophorini GELECHIOIDEA ? Amphisbatini Oecophoridae (Depressariidae; Coleophorinae Gelechiidae, in part) Momphini Xyloryctidae Coleophorini Gelechiidae Cosmopterigini Blastobasidae Elachistinae Lavernidae (Momphidae; Elachistini Cosmopterygidae (sic); Cemiostomini Elachistidae, in part) Tineidae incertae sedis Xyloryctinae Stenomatinae Metachandinae (Oecophorinae) Physoptilinae Epimarptinae Chrysopeleiinae Pterolonchinae

(continued on next page)

33

Table 2.1 continued.

Tillyard 1926 McDunnough 1938 ______TINEOIDEA GELECHIOIDEA Elachistidae Cosmopterygidae (sic) Oecophoridae Epermeniidae Depressariinae Gelechiidae Eulechriinae Oecophoridae Philobotinae Blastobasidae Oecopchoriinae Stenomidae Blastobasinae Ethmiidae Xyloryctidae Gelechiidae YPONOMEUTOIDEA Scythridae (sic) Elachistidae

TINEOIDEA Coleophoridae

(continued on next page)

34

Table 2.1 continued.

Costa Lima 1939 Comstock and Herrick 1940 ______TINEOIDEA AA. The Frenatæ Lepidoptera Coleophoridae BB. The Specialized Frenatæ C. The Specialized Microfrenatæ ELACHISTOIDEA Coleophoridae Elachistidae Elachistidae Oecophoridae GELECHIOIDEA Ethmiidae Agonoxenidae Stenomatidae Blastobasidae Gelechiidae Cryptophasidae (Xyloryctinae) Blastobasidae Epimarptidae Cosmopterigidae Ethmiidae Scythrididae Gelechiidae Lavernidae (Momphidae; Elachistidae, in part) Oecophoridae Stenomatidae Hyposmocomidae (Cosmopterigidae) (continued on next page)

YPONOMEUTOIDEA Scythrididae

35

Table 2.1 continued.

Turner 1947 Obenberger 1952 ______TINEOIDEA Coleophoridae Subdivision Microptila Cosmopterigidae Elachistidae Coleophorinae HYPONOMEUTOIDEA Scythrinae (sic) Xyloryctidae Elachistinae Scythrididae Cosmopteryginae (sic) Scythridinae Gelechiidae Amphisbatinae Thalmarchellinae (sic) (Thalamarchellinae, GELECHIOIDEA Depressariinae) Blastobasidae Oecophorinae Amphitheridae Blastobasinae Agonoxeniidae Gelechiinae Gelechiidae Xyloryctinae Gelechiinae Chimabacchinae (sic) Stenomatidae Ethmiidae Oecophoridae Depressariinae Oecophorinae Epermeniidae

ELACHISTOIDEA Elachistidae Heliozelidae

(continued on next page)

36

Table 2.1 continued.

Common 1970 Bradley 1972 ______GELECHIOIDEA GELECHIOIDEA Coleophoridae Coleophoridae Agonoxenidae Elachistidae Elachistidae Oecophoridae Scythridae (sic) Oecophorinae Stathmopodidae Depressariinae Oecophoridae Ethmiidae Ethmiidae Gelechiidae Timyridae (Lecithoceridae) Gelechiinae Cosmopterigidae Symmocinae Cosmopteriginae Blastobasidae Momphinae Stathmopodidae Walshiinae Momphidae Metachandidae (Oecophoridae) Batrachedrinae Anomologidae Momphinae Pterolonchidae Cosmopteriginae Blastobasidae Blastodacninae Xyloryctidae Chrysopeleiinae Stenomidae Scythrididae Gelechiidae Physoptilidae

(continued on next page)

37

Table 2.1 continued.

Hodges, 1978 ______GELECHIOIDEA Oecophoridae Gelechiidae Depressariinae Anomologinae Depressariini Gelechiinae Amphisbatini Anacampsinae Ethmiinae Dichomeridinae Peleopodinae Chelariinae Autostichinae Lecithocerinae Xyloryctinae Physoptilinae Stenomatinae Oecophorinae Oecophorini Stathmopodini Pleurotini (continued on next page) Chimabachinae Deuterogoniinae Hypertrophinae Elachistidae Coelopoetinae Elachistinae Pterolonchidae Coleophoridae Coleophorinae Batrachedrinae Agonoxenidae Agonoxeninae Blastodacninae Blastodacnini Parametriotini Blastobasidae Blastobasinae Blastobasini Pigritiini Symmocinae Momphidae Scythridae (sic) Cosmopterigidae Cosmopteriginae Antequerinae Chrysopeleiinae

38

Table 2.1 continued.

Zimmerman 1978 Kuznetsov and Stekol’nikov 1979 ______GELECHIOIDEA COLEOPHOROIDEA Scythrididae Coleophoridae Agonoxenidae Ethmiidae Cycnodiidae (Elachistidae) Oecophoridae Gelechiidae Pleurotinae Oecophoridae Symmocinae Ethmiidae Deuterogoniinae Xyloryctinae Oecophorinae Blastobasidae Depressariinae Chrysopeleiinae Chimabachinae Momphinae Cosmopteriginae GELECHIOIDEA Gelechiinae Not treated

(continued on next page)

39

Table 2.1 continued.

Kuznetsov and Stekol’nikov 1984 ______COPROMORPHOIDEA GELECHIOIDEA Xyloryctidae Stenomidae Xyloryctinae Xyloryctidae Acriinae, new subfamily Aeolanthidae Aeolanthidae, new family Peleopodidae Lecithocerinae Ethmiidae Ceuthomadarinae Oecophoridae Torodorinae Elachistidae Coleophoridae ELACHISTOIDEA Agonoxenidae Elachistidae Batrachedridae Elachistinae Momphidae Agonoxenidae Cosmopterigidae Blastodacnidae Scythrididae Blastodacninae Lecithoceridae Parametriotinae Epimarptidae Ethmiinae Blastobasidae Stathmopodidae COLEOPHOROIDEA Symmocidae Batrachedridae Gelechiidae Blastobasidae Blastobasinae Oecophoridae Symmocinae Pleurotinae (continued on next page) Oecophorinae Deuterogoniinae Chimabachinae Depressariinae Momphidae Coleophoridae

GELECHIOIDEA Cosmopterigidae Antequerinae Cosmopteriginae Chrysopeleiinae Stathmopodinae Scythridae (sic) Pterolonchinae Gelechiidae Gelechiinae Teleiodinae Apatetrinae Helariinae, new status Dichomeridinae Metzneriinae, new status

40

Table 2.1 continued.

Stehr 1987 Minet 1990 ______GELECHIOIDEA GELECHIOIDEA Oecophoridae Pterolonchinae Depressariinae Coleophoridae sensu nov. Ethmiinae Coleophorinae Stenomatinae Amphisbatinae Batrachedrinae (inc. Homaledra) Elachistidae sensu nov. Elachistinae Agonoxeninae Oecophorinae Elachistinae Symmocinae Stenomatinae Elachistinae Cryptolechiinae stat. nov. Coelopoetinae Hypertrophinae sensu nov. Blastobasidae Ethmiinae Coleophoridae Depressariinae Momphidae Peleopodidae Agonoxenidae (inc. Bastodacna) Chimabachidae Cosmopterigidae Xyloryctidae Scythrididae Batrachedridae Gelechiidae Oecophoridae Anomologinae Symmocidae Gelechiinae Lecithoceridae Chelariinae Epimarptidae Dichomeridinae Blastobasidae Stathmopodidae Momphidae Cosmopterigidae Gelechiidae

(continued on next page)

41

Table 2.1 continued.

Nielson and Common 1991 Leraut 1992 ______GELECHIOIDEA GELECHIOIDEA Oecophoridae Pterolonchidae Batrachedridae Coleophoridae Hypertrophidae Amphisbatinae Depressariidae Coleophorinae Coleophoridae Elachistidae Elachistidae Agonoxeninae Blastodacnidae Elachistinae Agonoxenidae Stenomatinae Ethmiidae Aeolanthinae Blastobasidae Cryptolechiinae Momphidae Hypertrophinae Cosmopterigidae Tonicini Gelechiidae Hypertrophini Symmocidae Hypercalliini Holcopogonidae Ethmiinae Lecithoceridae Depressariinae Scythrididae Depressariini Epigraphiini Fushiini Telechrysidini Peleopodidae Chimabachidae Carcinidae Xyloryctidae Batrachedridae Oecophoridae Oecophorinae Philobotinae Symmocidae Symmocinae Oegoconiinae Lecithoceridae Scythrididae Epimarptidae Blastobasidae Stathmopodidae Momphidae Cosmopterigidae Gelechiidae Aristoteliinae Holcopogoninae Apatetrinae Gelechiinae Dichomeridinae

(continued on next page)

42

Table 2.1 continued.

Sinev 1993 ______COLEOPHOROMORPHA Group A. OECOPHOROIDEA Group D. GELECHIOIDEA Oecophoridae s. str. Gelechiidae Amphisbatinae Gelechiinae Oecophorinae Anacampsinae Deuterogoniinae Aristoteliinae Pleurotinae Metzneriinae Xyloryctidae Teleiodinae Symmocidae Stomopteryginae Chimabachidae Anomologinae Autostichidae Brachmiinae Dichomeridinae Group B. COLEOPHOROIDEA Lecithoceridae Pterolonchidae Lecithocerinae Epimarptidae Torodinae Blastobasidae Ceuthomadarinae Pigritiinae Scythrididae Blastobasinae Metachandidae Ashinagidae Stathmopodidae Group E. CHRYSOPELEIOIDEA Stathmopodinae Chrysopeleidae (sic) Cuprininae Batrachedridae Group F. COSMOPTERIGOIDEA Momphidae Scaesophidae Coleophoridae Diplosaridae Cosmopterigidae Group C. ELACHISTOIDEA Stenomidae Stenominae Aeolanthinae Depressariidae (continued on next page) Depressariinae Cryptolechiinae Hypertrophinae Ethmiidae Peleopodidae Elachistidae Agonoxenidae Blastodacnidae Chrysoclistinae Blastodacninae Parametriotinae

43

Table 2.1 continued.

Fetz 1994 Passoa 1995 ______GELECHIOIDEA GELECHIOIDEA Oecophoridae I Scythrididae “Depressariinae” Oecophoridae I Chimabachinae Stenomatinae Ethmiinae Amphisbatinae Anchiinae Ethmiinae quercana Depressariinae Pseudatemelia Oecophoridae II Orophia ferrugella Blastobasinae Cacophyia permixtella “Gelechiinae” Oecophorinae Oecophoridae II Autostichinae Oecophorinae Cosmopterigidae Symmocinae Batrachedridae Pleurota Momphidae Topeutis Coleophoridae Scythrididae Pterolonchidae Blastobasidae Amphisbatidae Cosmopterigidae Remaining gelechioid taxa not treated. Chrysopeleiinae Cosmopteriginae “ ” designates probable paraphyletic groups Antequerinae Gelechiidae Physoptilinae Gelechiinae Dichomeridinae Pexicopiinae

(continued on next page)

44

Table 2.1 continued.

Hodges 1999 Kuznetsov and Stekol’nikov 2001 ______GELECHIOIDEA GELECHIIFORMES Elachistidae ELACHISTOIDEA Stenomatinae Elachistidae Ethmiinae Agonoxenidae Depressariinae Ethmiidae Elachistinae Hypertrophidae Agonoxeninae Aeolanthidae Hypertrophinae Stenomidae Deuterogoniinae Aeolanthinae COLEOPHOROIDEA Xyloryctidae Xyloryctidae Xyloryctinae Lecithoceridae Scythridinae Stathmopodidae Chimabachidae Glyphidoceridae Glyphidoceridae, new family Schistonoeidae Schistonoeidae, new family Oecophoridae Oeciinae, new subfamily Deoclonidae Schistonoeinae, new Momphidae subfamily Coleophoridae Oecophoridae Peleopodidae Oecophorinae Batrachedridae Stathmopodinae Blastobasidae Lecithoceridae Batrachedridae PTEROLONCHOIDEA Epimarptinae Pterolonchidae Batrachedrinae Deoclonidae, new family GELECHIOIDEA Deocloninae, new Scythridae (sic) subfamily Gelechiidae Syringopainae, new Metzneriinae subfamily Anacampsinae Coleophoridae Teleiodinae Coleophorinae Chelariinae Momphinae Dichomeridinae Blastobasinae Cosmopterigidae Pterolonchinae Chrysopeleiidae Autostichidae Autostichinae Symmocinae Holcopogoninae Peleopodidae (continued on next page)

45

CHAPTER 3

ANNOTATION AND ANALYSIS OF PHYLOGENETICALLY IMPORTANT

CHARACTERS FOR FAMILIES AND SUBFAMILIES OF GELECHIOIDEA

FRACKER 1915 (LEPIDOPTERA)

INTRODUCTION

Numerous authors made great advancements in Gelechioidea systematics in the past 20 years. However, character state delineation and terminology remain ambiguous for many terms and many taxa. There is little reason to rely on modern numerical methods if the biological features represented by numerals are not considered in detail.

Therefore, it is necessary to create a character analysis that discusses alternative interpretations or conflicting analyses. It is, after all, the variation of real morphology that we aspire to explain, not the numerical placeholders in our cladistic matrices.

The following treatment is a summary only of characters that have been used in a cladistic framework employing Hennigian argumentation or data matrix construction. I do not trace characters back to their original authors; rather, I investigate their utility as cladistic characters. Definitions of terms not available from the author in discussion are

46

from Tuxen (1970), Stehr (1987), and Triplehorn and Johnson (2005). Explanation of characters with appropriate details and images are provided when warranted. Some terms are interchangeable and their use relevant to this text is addressed in Figure 3.1. When authors have built off of their previous work with no substantial changes, only their most recent and comprehensive treatments are considered here.

47

Monophyly of Gelechioidea

Minet Leraut Fetz Kuznetsov & Passoa Sinev Hodges Kaila Stekol’nikov Haustellum √ √ √ √ with scales at base (Adult)

Labial palpi √ √ recurved (Adult)

Maxillary √ palpi folded (Adult)

Abdomen √ with modified scales (Adult)

Female anales √ papillae telescopic (Adult)

Sternal rod √ (Adult)

Antennae √ meeting mesially (Pupal)

Mesothoracic √ legs with invagination (Pupal)

Figure 3.1. Characters of special interest used by various authors to define families and subfamilies of Gelechioidea. See text for further details. Continued on next page.

48

Figure 3.1. Continued from previous page.

Monophyly of Gelechioidea, con’t.

Minet Leraut Fetz Kuznetsov & Passoa Sinev Hodges Kaila Stekol’nikov L1 and L2 √ √ approximate (Larval)

SD 1 of A9 √ hairlike (Larval)

Monophyly of Lineages within Gelechioidea

Minet Leraut Fetz Kuznetsov & Passoa Sinev Hodges Kaila Stekol’nikov M7 (Adult) √ √

Valves split √ √ √ √ (Adult)

Juxta (Adult)

Gnathos √ √ √ (Adult)

Uncus (Adult) √ √ √ √

Socius (Adult) √ √

Transtilla √ √ (Adult)

Figure 3.1. Continued from previous page.

49

Monophyly of Lineages within Gelechioidea, con’t.

Minet Leraut Fetz Kuznetsov & Passoa Sinev Hodges Kaila Stekol’nikov Aedeagus √ √ ankylosed (Adult)

Spiniform √ √ √ √ √ setae (Adult)

Apodemes & √ √ √ √ Venulae of A2 (Adult)

Antenna not √ √ √ meeting at the meson (Pupal)

D2, D1, and √ √ √ SD1 (Larval)

SD 1 of A9 √ √ √ √ hairlike (Larval)

SD1 w/ √ √ √ √ pinacular rings (Larval)

L Group √ (Larval)

P Group √ √ (Larval)

Stipular setae √ √ (Larval)

Figure 3.1. Continued from previous page.

50

Monophyly of Lineages within Gelechioidea, con’t.

Minet Leraut Fetz Kuznetsov & Passoa Sinev Hodges Kaila Stekol’nikov SD group √ √ with pore (Larval)

Adfrontal √ √ √ suture (Larval)

Postmentum √ √ √ √ with pit- shaped depression (Larval)

Lateral √ √ √ √ condyles (Larval)

A9 with √ √ paired ventromesial lobes (Larval)

Figure 3.1. Continued from previous page.

51

DISCUSSION

Monophyly of Gelechioidea

Minet (1990), using Hennigian argumentation, supported the monophyly of

Gelechioidea with the following groundplan synapomorphies: dense imbricated scales covering the base of the haustellum, strong recurved labial palpi in most taxa, larval abdominal segments 1-8 with L1 and L2 approximate or on same pinaculum, and pupae with apical or subapical invagination on the mesothoracic legs.

While scholars have used similar characters to diagnose the superfamily

Gelechioidea since Forbes first defined their utility in 1923, Passoa (1995) was the first to use the characters of adults with haustellum scaled at least halfway, adults with maxillary palpi folded, adults with labial palpi upturned, larvae with L1 and L2 setae approximate on abdominal segments, and larvae with SD1 setae on A9 hairlike in a cladistic analysis thereby establishing their importance as potential synapomorphies for Gelechioidea. As well, Passoa was the first to establish monophyly of Gelechioidea using outgroups

Yponomeutoidea and .

Hodges (1998) considered the scaled haustellum (at least halfway) and folded maxillary palpi to be synapomorphies for Gelechioidea, but did not use them for phylogenetic reconstruction. He considered the remaining of the above characters to be potential synapomorphies for the superfamily, but stated they would need further investigation in more taxa before their value were fully recognized.

52

Kaila’s (2004) work does not support the monophyly of Gelechioidea with any of

the above characters, and it is based only on homoplastic characters (abdomen with scales

modified as setae in tergum; sternal rod of the second sternum present as sharply

delimited narrow ridge; membrane between papillae anales and the eight segment in the

female telescopic; and pupal antennae meeting mesially. The first three of these

characters are found amongst the outgroup taxa, but reversed within Gelechioidea.

Adults

Intravalvalis muscle (M7) – In male Lepidoptera, the intravalvalis muscle (M7) of the

genitalia is responsible for movement of the valves3 and valvulae4 and is located in the

sacculus (Kuznetsov and Stekol’nikov 1979, 1984, 2001). Kuznetsov and Stekol’nikov

(1979, 1984, 2001) investigated functional morphology of Gelechioidea, focusing on the

musculature of genitalia. Although they did not conduct a cladistic analysis,

parsimonious reconstruction of the states present or absent on their evolutionary tree

suggests presence as a plesiomorphic condition of the super group Coleophoroidea

(Coleophoridae, Oecophoridae, Momphidae, Blastobasidae and Batrachedridae) with

losses in Blastodacninae, Amphisbatinae, Scythridae (sic), and Gelechiidae. The

3 The valves or valvae (p.) (valve or valva (s.)) are the paired, clasping organs caudal of all the genitalia that appear to be derived in part from the styli, coxites or parameres of the gonopods of the 9th abdominal segment (Klotts, 1970).

4 The valvulae (p.) (valvula (s.)) are the dorsal, lightly sclerotized lobe of the valves, perhaps part of the costa. See text “valves entire or divided” for discussion.

53

phylogenetic utility of this character needs to be reinvestigated in light of Kuznetsov and

Stekol’nikov’s (2001) recent functional morphology work.

Passoa (1995) employed characters from Kuznetsov and Stekol’nikov 1976 work in a cladistic framework, focusing only on the absence or presence of the intravalvalis muscle (M7). In Passoa’s (1995) phylogeny, the absence of the M7 muscle allies

Gelechioidea with Yponomeutoidea. The presence of the M7 muscle within Gelechioidea unites Momphidae, Coleophoridae (including Batrachedridae) and Pterolonchidae, although M7 is coded as missing data in Pterolonchidae in this treatment. In a later work,

Kuznetsov and Stekol’nikov (2001) demonstrated M7 to be present in Pterolonchidae.

Hodges (1998) did not include an analysis of M7 in his cladistic treatment.

Kaila (2004) does not use the M7 character explicitly, or any other musculature character for that matter, but rather codes “mobility of valvae” with states as “valvae mobile”, and “movement restricted.” In Kaila’s (2004) phylogeny, restricted mobility of valves is the ancestral condition for his “gelechiid lineage” and is reversed (mobility gained) in the ancestors to Momphidae (Mompha), some Coleophoridae (Stathmopoda,

Idioglossa, and B. pinicolellla but not B. eustola, Goniodoma,

Coleophora), and some Parametriotinae (Trachystola, Microcolona) in the oecophorid lineage.

Kaila’s use of the term “valvae” can be taken to mean reference to major regions of the valves: ampulla, clasper, costa, cucullus, sacculus and valvula (split regions of the valve or the hypertrophied sacculus). This character would seem to correlate with musculature of male genitalia, as defined by Kuznetsov and Stekol’nikov (1979, 1984,

54

2001); however it is difficult to interpret Kalia’s (2004) coding due to his vague character

definition. Movement of the valves and valvulae is controlled by the following six

muscles: the tegmino-valvalis superior (M2), which function as flexors of the valves; the

vinculo-juxtalis (M3), which function as extensors of the valves; the tegmino-valvalis

inferior (M4), which functions as flexors of the valves; the intravalvalis (M7), which

functions as flexors of the harpe or flexors of the cucullus and is located in the sacculus; the vinculo-valvalis (M14), which function as flexors of the valves; and the intervalvalis

(M26), which function as adductors of the valves and replaces M4 in Blastobasidae

(Kuznetsov and Stekol’nikov 1979). Movement of the valvulae is described as,

“Characteristic of all the families [of the Coleophoroidea], except the Batrachedridae, is the hypertrophy of the sacculi and the retention or proliferation of the intravalvar muscles

(M7) contained within the sacculi, as a result a result of the augmentation of the function of grasping the abdomen of the female by the sacculi of the male” (Kuznetsov and

Stekol’nikov 1979). Kaila’s (2004) coding does not correlate with the presence or absence of any particular muscle or group of muscles, although it can be very loosely associated with presence of the M7 (mobility seen in Mompha, Stathmopoda, Idioglossa,

Batrachedra praeangusta, B. pinicolellla, Goniodoma, Coleophora, Trachystola,

Microcolona). His coding is not clear and it is difficult to decipher what is meant by

“mobility” relative to the function of the M2, M3, M4, M7, M14, and M26 muscles.

Valves entire or split – (Figure 3.1, 3.2) In several Gelechioidea families, the males have

valves that are divided and are used as key taxonomic and phylogenetic features.

55

However; the state of being “divided” has been characterized differently by taxonomists making definitions of this condition less than precise. Hodges (1998) refers to the state of division as having, “the costa developed as a free lobe that often appears to arise from the sacculus”; while Razowski (1989) simply refers to the divided area (in Coleophoridae) as

“the sacculus”, an area that has also been referred to as the “hypertrophied sacculus” by

Kuznetsov and Stekol’nikov (1979). Toll (1939) defines the valvulae (of Coleophoridae) as the, “dorsal, lightly sclerotized lobe of the valva.” Klotts (1970) refers to the “valvula” as part of the valvae and delineates it as the ventro-distal portion of the valvae (in accordance with Sibatani et al 1954).

Minet (1990) discusses the presence of a small ventral sclerite independent of the sacculus, but it is unclear to me what the significance of this character is in his analysis.

Passoa (1995) and Hodges (1998) both used this character in their cladistic analyses, coding it as valva undivided or divided. In Passoa’s (1995) analysis, divided valves evolves twice, once in the ancestor to the clade ((Coleophoridae + Pterolonchidae)

+ Momphidae) with a reversal in Pterolonchidae, and once in the ancestor to

Blastobasidae. In Hodges (1998) analysis, split valves only evolve once, in the ancestor to (Momphidae + Coleophoridae) + Blastobasidae).

Kaila (2004) found the valva to be divided as either with costa separated from cucullus and sacculus, or as sacculus separated from cucullus and costa, and coded it as

two separate absence or presence characters: “costa not separate or separate” and the

“sacculus not separate or separate.” Kaila states that Hodges split valve character

emphasized the division of the valva but did not differentiate whether the costa or the

56

sacculus was divided from the remaining valva. In Kaila’s phylogeny, the “costa separate

/ not separate” character evolves seven times in his tree (in the ancestors of Goniodoma +

Coleophora (Coleophoridae); the ancestor of Mompha (Momphidae); the ancestor of

Paratheta (Scythrididae), the ancestor of Gelechiidae + Cosmopterigidae; the ancestor of

Symmocinae + Glyphidoceridae + Oditinae; the ancestor of Blastobasinae; and the

ancestor of Carcinae) and the “sacculus separate / not separate” evolves five times (in the

ancestors of Stathmopoda (Stathmopodinae); the ancestor of Gelechiidae, the ancestor of

Sorhagenia (Chrysopeleiinae); the ancestor of + Symmoca (Symmocinae);

and the ancestor of Scieropepla (not placed in family).

Kaila (2004) also noted another condition: valva divided into three lobes with the

costa and sacculus separate from a median lobe. In his analysis, he does not code this character explicitly but what he does instead is code janiszewskae

(Chrysopeleiinae), (Symmocinae), peliella,

juniperella and malvella (Gelechiidae) as having the costa separate and the

sacculus separate. He stated that Hodges (1998) interpreted the separated valval costa of

gelechiids as a distinctive structure but that Hodges’ interpretation requires, “the total

reduction of the costa, and the appearance of a new structure in its place. No

morphological support could be found for this view in the present material, and the

appendix appendicular was interpreted to be homologous to valval costa.”

While Kaila (2004) makes it a point to differentiate among the types of possible

divisions of valvae (arising from sacculus, arising from costa, or arising from cucullus)

he codes the mobility of the valvae as simply “mobile” or “restricted” for all his possible

57

valve types. It is interesting to note that Kaila’s characters of mobility and division almost no corroboration on his phylogeny.

Juxta – (Figure 3.1, 3.2) The juxta is a sclerotized plate located ventrad of the aedeagus, which helps to support it. It is often attached to the sacculi and ventral portion of the vinculum and sometimes connected to the anellus5. Both Passoa (1995) and Hodges

(1998) code this character as absent or present. Passoa (1995) coded the juxta as absent in

Yponomeutoidea and absent within Gelechioidea except in Cosmopterigidae and

Gelechiidae. Hodges (1998) the juxta as plesiomorphically present within Gelechioidea and polymorphic in the ancestor to Cosmopterigidae + Gelechiidae. It is absent in all subfamilies of Cosmopterigidae (Antequerinae, Chrysopeleiinae, and Cosmopteriginae) and absent in Gelechiinae and Pexicopiinae, coded as ambiguous in Dichomeridinae, and present in Physoptilinae.

Kaila (2004) codes the juxta as three separate characters addressing the ventral shield of the juxta, the dorsal shield of the juxta, and connection between the juxta and the valva. Kaila coded the ventral shield of juxta as absent, present as a free sclerotization ventrad of aedeagus, present and surrounding the aedeagus, present and connected to vinculum. The juxta was considered to be present in most Cosmopterigidae and

Hypatima rhomboidella (Gelechiinae). He also coded a differentiated dorsal shield of

5 The anellus (s.) (anelli (p.)) is the funnel-like cone that surrounds the vesica where the vesica emerges from the body. The manica, or innermost layer, fastens around the aedeagus at that point. The anellus is formed from the central part of the diaphragma, which has been evaginated and invaginated around the vesica. The vesica (s.) (also penis (s.)) is the flexible, eversible, and caudad tube lying within the aedeagus; used for insemination.

58

juxta as absent, present as a lobe or pair of lobes connected to ventral shield of juxta, or present and surrounding aedeagus. He codes the connection between juxta and valva as juxta not connected to valva, a narrow sclerotized valval process connecting valval costa and juxta, or valva broadly connected to juxta.

Gnathos – The gnathos is defined as a pair of arms derived from the tenth sternite and

articulated with the tegumen ventro-laterally of the articulation of the uncus and socii (if

present). In Gelechioidea, the gnathos may be highly variable and very difficult to

interpret.

Sinev (1993) hypothesizes that having the gnathos not sclerotized and spiny is plesiomorphic for Gelechioidea and the derived condition is sclerotized and unpaired or paired. In his scenario, Elachistoidea retains the ancestral condition, while

Oecophoroidea, Coleophoroidea and Gelechioidea s. s. have a sclerotized, unpaired

gnathos and Cosmopterigoidea and Chrysopeleoidea have a sclerotized, paired gnathos.

Passoa (1995) coded this character for taxa only within the oecophorid clade. In

his analysis, the presence of a spined gnathos unites taxa in the oecophorid group 1:

Peleopodinae, Xyloryctinae, Hypertrophinae, Agonoxeninae, Amphisbatinae,

Depressariinae, Ethmiinae, and Elachistinae. The absence of spines and the presence of a

gnathos in Xyloryctinae and Agonoxeninae respectively are considered reversals.

Ethmiinae is coded as polymorphic due to what Passoa thought was a plesiomorphic

absence of a gnathos in basal ethmiines. Passoa (1995) believed the presence of a spined

gnathos in both Coleophoridae and Deoclonidae to be the result of parallel evolution.

59

Hodges (1998) coded this character with eight states, and the ninth as absent in

Aeolanthinae, Deuterogoniinae, Antequerinae, Chrysopeleiinae, Cosmopteriginae,

Hypertrophinae, Syringopaidae, and Momphinae. The remaining states are coded with

attention paid to presence of an articulated or sclerotized band, articulated rami or

articulated sclerites, with finer variations within each state. Hodges did not treat the presence or absence of spines on the gnathos. This character evolves several time in his phylogeny.

Kaila codes the gnathos as seven separate characters with a total of 20 states. In his analysis, he codes the following seven characters: absence/presence of gnathos, the articulation with tegumen, the articulation with uncus, shape of mesial part of gnathos, scobination of mesial part of gnathos, symmetry of gnathos, and division of gnathos.

These characters are highly homoplastic, except for a few. In his analysis, the character

“mesial part of gnathos articulated from basal arms, with sickle-shaped, ascending hook” is a synapomorphy for Gelechiidae, “mesial part of gnathos scobinate with large thorns” is a synapomorphy for , “mesial part of gnathos laterally compressed, downwards directed and fused to basal arms, without distinct limit” is a synapomorphy for oecophorine taxa: fuscescens, Hofmannophila pseudospretella, bractella, Harpella forficella, similella, Bisigna procerella, Promalactis venustella, Polix coloradella, Tingena hemimochla, Tingena armigerella, and “basal arms of gnathos basally fused to uncus” is a synapomorphy for Xyloryctinae,

Deuterogoniinae, and Blastobasinae.

60

Uncus – (Figure 3.1, 3.2) The uncus is derived from the tenth tergite and is an extension

of the caudal, mid-dorsal margin of the tegumen. It may be membranous to heavily

sclerotized and range in complexity of form from simple to bifid or trifid.

Sinev (1993) considers the presence of unpaired, posterior appendages of

tegumen as the plesiomorphic condition and having posterior appendages fused into the

uncus or posterior appendages of tegumen lost (or the uncus absent) as derived.

Passoa (1995) does not code this character.

Hodges (1999) coded the uncus as simply absent or present. In his analysis, the

uncus is primitively present for the superfamily; lost in the ancestors of Syringopainae,

Coleophorinae; and polymorphic in the ancestor of Gelechiinae.

Kaila (2004) focused on the presence and absence of the uncus, the fusion of the

uncus and the symmetry of the uncus. These characters are homoplastic and none serve

as synapomorphies in his analysis.

Socius – (Figure 3.1, 3.2) Defined as paired, weakly sclerotized pads on caudal margin of

tegumen ventrad of base of uncus. The structure is perhaps derived from the

intersegmental membranes of segments IX – X. It is paired structure (Klotts 1970).

Hodges (1999) coded the socius as absent or present. It is primitively absent for the superfamily, polymorphic in Gelechiinae, and present in Dichomeridinae.

Kaila (2004) codes the socius as two characters: present (as tongue-shaped or

variably-shaped setose free lobe) or reduced (present at most as small group of setae in

ventral margin of tegumen or base of uncus); or symmetrical or asymmetrical. The socius

61

is vestigial in his analysis but reversed in and in Scythrididae, Pexicopia

(Gelechiidae), Hierodoris (xyloryctid assemblage), and in Depressariinae,

Hypertrophinae and Donacostola in Elachistidae within Gelechioidea.

Transtilla – (Figure 3.1, 3.2) The transtilla is a sclerotized structure in the dorsal part of the diaphragma, sometimes a dorsal bar connecting the dorso-proximal angles of the valvae.

Hodges (1999) codes the transtilla as absent or present. In his analysis, it is plesiomorphically absent for the superfamily; present in the ancestors of Coleophorinae +

Momphinae, Chimabachidae, and Schistonoeinae; and polymorphic for Depressariinae +

Elachistinae.

Kaila (2004) codes the transtilla as absent or present as developed to some extent

(usually hook-shaped), present as a mesially differentiated band, or present as band with paired differentiated structures. Kaila states that the presence of a hook-shaped transtilla was sometimes ambiguous so it was not coded separately from the absence of the transtilla. It is a homoplastic character with each state evolving several times (uniting species of a single genus or serving as an autapomorphy for species).

Aedeagus ankylosed – (Figure 3.4) Hodges (1999) codes the aedeagus as free or ankylosed6, which he defines as “fused with surrounding diaphragma.” An aedeagus free of the surrounding diaphragma is the plesiomorphic condition within Gelechioidea. It is

6 Ankylosation is union of separate sclerites to form a single sclerite.

62

ankylosed in Aeolanthinae and Coleophorinae. It is polymorphic within Blastobasinae.

Having the aedeagus ankylosed is a synapomorphy for Cosmopteriginae. Within the

Gelechiidae, it is ankylosed in Physoptilinae, polymorphic in Gelechiinae and

Dichomeridinae, and free in Pexicopiinae.

Kaila (2004) codes the ankylosation of the aedeagus as four separate characters each with the two states of “not” and “yes”: ankylosation by the median plate of juxta, ankylosation by the juxtal lobes, ankylosation by the dorsal shield of juxta or ankylosation by the anellus. In his analysis, a free aedeagus is the plesiomorphic condition for the superfamily and ankylosation is derived. Ankylosation by the median plate of juxta is derived eight times all as autapomorphies for terminal species (

(Cosmopterigidae), Prionocris (Oecophoridae), Phaeosaces (Oecophoridae),

(Amphisbatidae), and Thudaca (Elachistidae)) except for that it unites Adelpha +

Orthotanea (Tortricidae) and species in Scythrididae (later lost in Paratheta calyptra).

Ankylosation by the juxtal lobes is derived one time in the ancestor of

(Glyphidoceridae). Ankylosation by the dorsal shield of juxta is derived 3 times, once in the ancestor of + Agriophara (Stenomatinae), once in the ancestor of

Aeolanthes (Aeolanthinae), and once in the ancestor of Goniodoma + Coleophora

(Coleophorinae). Ankylosation by the anellus is derived 3 times, Once in the ancestor of

Antequerinae + Cosmopteriginae, once in the ancestor of Pseudatemelia

(Amphisbatinae), and once in the ancestor of Trachystola (Parametriotinae). Kaila’s method of character coding distinguishes separate conditions of ankylosation, but does not actually yield a deeper level of clarity lacking in Hodges analysis.

63

Adults with patches or bands of spiniform setae on abdominal terga – (Figure 3.5)

Spiniform setae are present or absent on the abdominal terga within the Gelechioidea and are used by many authors as an important taxonomic character. They are present in

various states: in two parallel rows as in Coleophorinae, covering the entire terga as in

Pterolonchinae, centered in a mesial patch on the terga as in Holcopogoninae, on the

anterior edge of the terga as in Antequerinae, or on the posterior edge of the terga as in

Stathmopodinae.

Minet (1990) stated that spiniform setae are present in many families and uses the absence of spiniform setae to unite the Elachistidae.

Leraut (1992) stated that he used the presence and the arrangement of the tergal

spines to define and place taxa of Gelechioidea; however, his treatment is not detailed

with respect to synapomorphies of clades and polarization of states.

Passoa (1995) coded this character as present or absent within Gelechioidea, and

then expanded this character further to be more descriptive within the oecophorid taxa, so

that the two matrices differ. Batrachedridae, Momphidae, and Coleophoridae were coded as having stout setae present in parallel patches, while Pterolonchidae was coded as having a band of setae present which was considered to be a fusion of the patches. In this matrix, the oecophorid group 2 taxa were coded as having a band of setae present but treated as a parallelism. Within the oecophorid group 2 taxa, Blastobasinae,

Oecophorinae, Lecithocerinae, Symmocinae and Autostichinae were coded as having a stout band of spiniform setae, and treated as distinct from the band, which

64

have, while this character is coded as absent in all other oecophorid taxa. In his analysis, the absence of spiniform is plesiomorphic for the superfamily.

Hodges (1998) coded this character as multistate, nonadditive with four distinct states including absent. Coleophoridae, Batrachedridae, Epimarptinae and Momphinae are coded as having patches of spiniform setae, Autostichinae, Blastobasinae, Oeciinae, and Pterolonchinae were coded as having a band of spiniform setae, and Stathmopodinae was coded as having a band of spiniform setae on posterior margin of most segments.

The remaining taxa were coded as having only regular scales in this location except

Oecophoridae and Deoclonidae, which were coded as being polymorphic for this character. In Hodges’ analysis, spiniform setae are plesiomorphically absent for

Gelechioidea. Spiniform setae present as a band across the tergite is derived in the ancestor of Xyloryctinae and the ancestor of Schistonoeidae plus all remaining taxa. The ancestor of Oecophoridae was polymorphic for having spiniform setae absent or present as a band at the posterior margin of the tergite. This condition is different for the ancestor of Oecophorinae, which had spiniform setae either absent or present in a band across the tergite. Having spiniform setae present in two patches is a synapomorphy for

Batrachedridae and for the ancestor of Coleophorinae + Momphinae. Spiniform setae are lost in the ancestor of Peleopodidae + Amphisbatidae + Cosmopterigidae + Gelechiidae.

Kaila (2004) codes spiniform setae as absent, present in single area in each tergum or present as divided areas in each tergum. He states that the distinction of whether the setose area was located at posterior margin of terga or throughout the terga was unclear in his sample of taxa. He also states that distinction among the type of

65

spiniform setae was not made because the delineation of states was not clear and that

there was considerable variation between taxa. In his analysis, spiniform setae present in

a single area is the ancestral condition for Gelechioidea. There are two derivations of

having divided areas of spiniform setae (in the ancestor of Idioglossa + Batrachedra

(reversed back to a single area) + Goniodoma + Coleophora (Coleophoridae)) and the

ancestor of Mompha (Momphidae). Spiniform setae are lost in the ancestor of Scythris

(Scythrididae) + Gelechiidae + Cosmopterigidae; Phaeosaces (Oecophoridae),

Lecithocera (Lecithoceridae), (Autostichidae), Oditinae + Glyphidocerinae

and the ancestor of the clade that contains through the remaining oecophorid assemblage taxa included in his phylogeny (the ancestor of Dastystoma +

(Oecophoridae) regains a single area of spiniform setae).

Second sternum with apodemes and venulae – Within the Lepidoptera, the presence and

absence and overall morphology of apodemes and venulae has been used to define major

clades. Minet (1983) used the presence or absence of venulae (a pair of sclerotized

thickenings on the second abdominal sternite) within Ditrysia to ally superfamilies.

Gelechioidea, which are believed to have primitively both apodemes and venulae, were classified as having a “Tineoid type” arrangement. Those lacking venulae are said to have a “Tortricoid type” arrangement. This approach was criticized by Kyrki (1983) who believed that simple presence and absence of apodemes and venulae within Ditrysia was not a reliable enough character to infer relationships. Kyrki suggested that the presence of the anterolateral processes on S2 to delimitate apoditrysian Lepidoptera from ditrysian

66

Lepidoptera. Researchers of Gelechioidea have used characters of the apodemes and venulae to study the systematics of families and subfamilies, but this character seems to be rather variable.

Leraut (1992) studied the absence and presence of apodemes and venulae in male and female Gelechioidea in detail. Using this suite of characters, along with other traits, he classified lineages of Gelechioidea.

Hodges (1998) coded this character with four states: venula only; both venula and apodeme present; apodeme only; and both venula and apodeme absent. In his phylogeny, the character is homoplastic and does not serve as an uncontroverted synapomorphy to unite any groups but does provide some hierarchy.

Kaila (2004) codes this character as absent or present for males and females. In his matrix, he codes the apodemes and venulae (sternal rods) for: absence or presence of a sternal rod in S2 (and if present as broad indistinct delimited ridge or as a sharply delimited narrow long ridge); absence or presence of a triangular keel in anterior end of sternal rod of male; sternal rod anteriorly modified as a sharp-tipped sclerotization fused to integument or not; absence or presence of a T-shaped sclerotization mesially in S2; absence or presence of lateroposteriorly directed extension from sternal rod of S2; and absence or presence of obliquely directed lateroanterior sclerotization in S2. He finds the evolution of this character to be quite complex and not easily assessed as merely present of absent, and in some cases, he notes that species of the outgroup Tortricidae seem to him to have the “Tineoid type” arrangement of venulae of S2 and species of Gelechioidea seem to have Apoditrysian anterolateral processes. He also suggested that there was a

67

“tendency towards stouter apodemes with increasing body size”, stating that generally,

females seemed to have more substantial apodemes than males, elaborating that, “the

heavier abdomen of females could be thought of as requiring a more developed

supporting structure at the abdominal base than in males.” Kaila does not acknowledge

Leraut’s 1992 treatment in his 2004 phylogeny.

Larvae

D2, D1, and SD1 – The dorsal group is located along the midline of the larval body and

the subdorsal group is below the dorsal group (Stehr, 1987). The arrangement of D1, D2

and SD2 on the ninth abdominal segment (A9) is commonly used as taxonomic features

in larval keys (Stehr, 1987).

Minet (1990) considered the vertical arrangement of D1, D2, and SD1 to be an

autapomorphy for Coleophoridae, thus allying Coleophorinae and Amphisbatinae. He notes their vertical arrangement to be present in Hypertrophinae but not in

Cryptolechiinae and Elachistidae.

Passoa (1995) used the vertical arrangement of D1, D2, and SD1 to define his group “Gelechiiformes” ((Batrachedridae + Momphidae + Coleophoridae +

Pterolonchinae) + (Cosmopterigidae + Gelechiidae)) stating that the plesiomorphic condition for Ditrysia is not to have D2 and SD1 in a vertical line. Within

Gelechiiformes, Passoa codes the character as polymorphic for Batrachedridae

(Homaledra), Pterolonchinae, some Cosmopterigidae and some Gelechiidae stating the

observed variations are due to specializations. He considers the presence of the condition

68

in Setiosoma (Stenomatinae), a few tineids and yponomeutids as independent evolutionary events.

Kaila (2004) codes D1, D2 and SD1 to be in a vertical row for Coelopoeta and

Coleophora, but not in ‘‘batrachedrids’’ or Pterolonchidae, all of Gelechiidae, most

Cosmopterigidae species included in his analysis, Amphisbatidae and some Elachistidae.

He notes that his character interpretation is different than that of Minet’s (1990) or

Passoa’s (1995).

SD 1 of A9 – On A9, the subdorsal group differs slightly form other abdominal segments.

On A9, SD2 is usually absent and SD1 is below D1. Sometimes SD1 is hairlike. In some

Gelechioidea taxa, SD1 is secondarily setiform (not hairlike) (Stehr, 1987).

Minet (1990) noted that this condition was homoplasious in Gelechioidea: present

Carcina, Peleopodidae, and Chimabachidae but not in the basic scheme of Elachistidae or

Coleophoridae. He records its presence in five subfamilies of Elachistidae: Stenomatinae,

Cryptolechiinae, Hypertrophinae, Ethmiinae, and Depressariinae.

Fetz (1994) indicated SD1 was not hairlike in his oecophorid groups I and II, and suggested that reversal of this character to its plesiomorphic condition is a synapomorphy to unite these groups.

Passoa (1995) considered this character to be a potential synapomorphy for the superfamily (supported by his cladistic analysis).

Kaila (2004) codes SD1 of A9 as other setae or hairlike. The monophyly of the oecophorid lineage is supported by the hairlike nature of seta SD1 of segment A9

69

(evolving eight times), reversed four times, and also found in Scythrididae, Gelechiidae and Homaledra (Coleophoridae).

SD1 with pinacular rings – In some species of Gelechioidea, the first subdorsal seta has a sclerotized ring at its base (Stehr, 1987).

Minet (1990) used the association of SD1 and SD2 being partially or entirely on a pinaculum to unite the families Xyloryctidae, Batrachedridae, Oecophoridae s.s.,

Symmocidae, Lecithoceridae, Scythrididae, Epimarptidae, Blastobasidae and

Stathmopodidae.

Passoa (1995) codes this character as present within the Scythrididae,

Blastobasinae, Xyloryctinae, Autostichinae, and Stenomatinae. In his analysis, the absence of the ring is plesiomorphic for the Gelechioidea and presence of the ring is a synapomorphy for the Scythrididae + Oecophoridae.

Hodges (1998) also codes this character as present or absent. In his phylogeny, the ring is plesiomorphically absent and its presence is derived in the ancestor of

Xyloryctinae + Scythridinae, in the ancestor of Lecithocerinae and in the ancestor of

Autostichinae.

Kaila (2004) codes the presence or absence of a ring-shaped pinaculum around

SD1 leaving a non-sclerotized area around the seta at A1-8. The ring is plesiomorphically absent. The presence of a pinacular ring in abdominal SD1 in larvae (also present in some

Scythris, some Stathmopoda and Lecithoceridae) supports the monophyly of his xyloryctid assemblage.

70

L-Group – The position of the lateral group varies within Lepidoptera on various segments of the larval body (Stehr, 1987). There are usually three L setae present on all body segments (Stehr, 1987). Stehr (1987) states that in most Gelechioidea on A1-8, L1 and 2 are close together and below the spiracle. Passoa (1995) records three character states for the L setae within Gelechioidea: L group anteroventrad, posteroventrad or ventrad of the spiracle. Larvae with L group on A1-8 posteroventrad of spiracle is a synapomorphy for Momphidae + Coleophoridae + Pterolonchidae and a derived condition within Gelechioidea. Larvae with L setae anteroventrad of spiracle is a synapomorphy for Oecophoridae + Scythrididae.

P-Group – The posteriodorsal group of setae are located on the area of the upper face of the head.

Passoa (1995) codes the position of the P group and the distance between setae in his analysis. In his analysis, it is the plesiomorphic condition to have the P setae arranged horizontally. The vertical arrangement of the P group unites the Oecophorid I + II taxa.

Passoa also uses the distance between P2 and P1 in his analysis, but this character proves to be rather homoplastic.

Kaila (2004) codes the number of P setae as two or one (not distinguishing between which seta was absent (1 0r 2)). He states that he found Passoa’s use of the relative positions of the P setae difficult to define and not useful in characterizing groups.

In his paper, this character is not plotted on the phylogeny.

71

Stipular setae – The stipular setae are located on the spinnerets of larval Lepidoptera

(Stehr, 1987).

Passoa (1995) coded this character with two states: short or long and stout. He found long and stout stipular setae to be a synapomorphy for Coleophoridae +

Pterolonchidae.

Kaila (2004) coded this character with three states: minute, long and thin, or long and stout. His analysis supports Passoa’s findings, but also demonstrates the presence on long, thin stipular setae in Scythris spp.

SD group with pore – In Lepidoptera, the SD group on abdominal segments 1-8 is sometimes associated with a pore (Stehr, 1987).

Passoa (1995) finds the absence of a pore adjacent to SD 1 on A-8 to be plesiomorphic for the oecophorid lineages. Its presence unites the subfamilies

Autostichinae and Symmocinae.

In Hodges (1999) analysis, the presence of a pore unites Xyloryctinae and

Scythridinae, Glyphidoceridae (not Blastobasinae), Lecithoceridae, and polymorphic for

Autostichidae (Autostichinae and Symmocinae but not Holcopogoninae).

Kaila (2004) excludes this character from his analysis due to technical difficulties.

Adfrontal suture – In larvae, the adfrontal suture is an inverted Y-shaped suture with internal ridges for muscle attachment (Stehr, 1987). It is composed of a medial epicranial

72

suture or adfrontal suture (the medial arm) and the lateral adfrontal sutures (the Λ–shaped arms) which surround the frontoclypeal area. The length of the medial adfrontal suture depends on the position of the mouthparts and the angle at which the head is held relative to the long axis of the body of the . In hypognathous larvae, the adfrontal suture is long relative to the frontoclypeal area. In prognathous larvae, the adfrontal suture is greatly reduced or completely lost while the lateral adfrontal sutures and the frontoclypeal area are elongated. Not surprisingly, semiprognathous larvae have an adfrontal suture intermediate in length. The length of the adfrontal suture and associated characters such as the epicranial notch and frontocypeal region has been used as phylogenetic characters for Gelechioidea.

Fetz (1994) treated the adfrontal suture not reaching the cranial incision as a synapomorphy to unite Oecophorinae I, Oecophorinae II, Symmocinae, Pleurota,

Topeutis, Scythrididae, and Blastobasidae. He later uses the reversal of this character to unite the Oecophorinae I taxa, and the secondary elongation of this suture to establish monophyly of Blastobasidae.

Passoa (1995) coded a short adfrontal suture as a synapomorphy for

Peleopodinae, Agonoxeninae, Xyloryctinae, and Hypertrophinae within his oecophorid 1 taxa.

Kaila (2004) codes the ecdysial line of adfrontalia as reaching cranial incision or not reaching cranial incision. Monophyly of the oecophorid lineage is also supported ecdysial line of adfrontalia not reaching cranial incision (however highly a homoplastic characters with 16 steps in his phylogeny).

73

Postmentum with pit-shaped depression –

Minet (1990) states this character is an apomorphy for his group XS

(Xyloryctidae, Batrachedridae, Oecophoridae (sensu stricto) Symmocidae,

Lecithoceridae, Scythrididae, Epimarptidae, Blastobasidae and Stathmopodidae), lost in

Symmocidae, and rare in some Oecophoridae (Pleurota and Hofmannophila), and certain

Lecithoceridae and Scythrididae. While Stehr (1987:385) records Gelechiinae as having a submental pit, Minet notes that he has not seen in it those studied.

Fetz (1994) treated the presence of a pit-shaped depression on the postmentum as a synapomorphy to unite Oecophorinae I, Oecophorinae II, Symmocidae, Pleurota,

Topeutis, Scythrididae, and Blastobasidae, and its reduction as a synapomorphy of

Symmocidae.

Hodges (1998) coded this character as present only in Blastobasinae and

Epimarptinae. These taxa are not closely related in his phylogeny.

Kaila (2004) coded submental pit at ventral side of larval head as absent, present as rounded or oval pit, or present as sclerotized pair of grooves. In his analysis, the xyloryctid assemblage is supported by larva having a submental pit (both as a rounded/oval pit or as grooves; later lost in Uzucha (Xyloryctinae) and yuccaecolella (Blastobasidae) and present in many Coleophoridae, Oecophoridae, and

Lecithoceridae)).

74

Pupae

Antenna not meeting at the meson – (Figure 3.6) Minet (1988) first used this character in

a phylogenetic context to unite Pterolonchidae with Coleophoridae.

Passoa (1995) found a similar result: Gelechioidea plesiomorphically have pupae

whose antennae meet at the meson. Pterolonchidae and Coleophoridae are united by having pupae whose antennae do not meet at the meson.

Kaila (2004) says that pupal antennae touching mesially is unique to and

plesiomorphic for Gelechioidea but it is reversed three or four times within the

Gelechioidea (depending on the optimization) in Holcopogon, Pterolonche, Scythris and

some Batrachedra and Coleophora species.

Lateral condyles – (Figure 3.7) Minet (1990) discusses the lateral mobility of gelechioid

pupae and says that the movement is restricted by the presence of lateral condyles, and is

related to orientation of the pupa to the sun. The presence of lateral condyles and

subsequent restricted movement was used to unite his expanded concept of Elachistidae.

Passoa (1995) codes this character as related to restricted pupal movement. In his

analysis, this character evolves several times.

Hodges (1998) codes this character as pupae without or with lateral condyles on

segments 5/6, 6/7. It is plesiomorphically absent for Gelechioidea. It is a synapomorphy

for his Elachistidae clade and polymorphic for Xyloryctinae.

Kaila (2004) codes this character as absent or present. Its absence is

plesiomorphic for Gelechioidea and it is present in Enteremna, Hypertrophia + Eupselia

75

(Hypertrophinae), and the clade that contains (“unplaced Elachistidae” + (Depressariinae

+ (Ethmiinae + (Aeolanthinae + (Parametriotinae + (Agonoxeninae + Elachistinae)))))).

A9 with paired ventromesial lobes

Hodges (1998) codes absence or presence of paired ventrolateral projections

(“pupal legs”) on segment 8/9. In his analysis, the absence of pupal legs is plesiomorphic

for Gelechioidea and the presence is derived in the ancestor to the clade that contains

(Agonoxeninae + (Hypertrophinae + (Deuterogoniinae + Aeolanthinae))).

Kaila (2004) codes this character as absent or present for “pupal legs ventrally at abdominal segment 9; sometimes only present as a pair of swellings.” Pupal legs are plesiomorphically absent for Gelechioidea. Within Gelechioidea, the character is homoplastic and its absence and presence unites several clades. The presence of pupal legs unites Hypertrophinae and the clade that contains (Ethmiinae + (Aeolanthinae +

(Parametriotinae + (Agonoxeninae + (Elachistinae))))). It is lost in the ancestor of

Elachistidae and regained in the ancestor of adscitella (Elachistidae).

76

Figure 3.2. Coleophora trifolli showing split valves, presence of juxta, presence of gnathos as fused, spined knob, absence of uncus.

77

Figure 3.3. Depressaria douglasella showing free aedeagus, presence of socii and uncus

(reduced), presence of membranous transtilla, entire valves.

78

Figure 3.4. Scythris showing ankylosation of aedeagus to valvae and vinculum.

79

Figure 3.5. Coleophora trifolii showing patches of spiniform setae.

80

Figure 3.6. Coleophora trifolii showing pupal antennae not meeting at the meson.

81

A.

B.

Figure 3.7. Lateral condyles present or absent. A. Lateral condyles present (redrawn from

Hodges 1998); B. Coleophora trifolii showing pupal lateral condyles absent.

82

CHAPTER 4

GELECHIOIDEA SYSTEMATICS: A REEXAMINATION USING COMBINED

MORPHOLOGY AND MITOCHONDRIAL DNA DATA

INTRODUCTION

Classification of Gelechioidea has been based on phylogenetic principles only recently. To date, specialists have necessarily focused on few taxa globally or many locally. Minet (1990) studied Gelechioidea using larval, pupal and adult characters. Fetz

(1994) then studied Gelechioidea using mainly larval characters. Both used Hennigian argumentation to designate characters and justify sister group relationships, but did not present a matrix or numerical analysis and showed little resolution.

Passoa (1995) relied mainly on larvae and pupae, with some adult characters. He

represented subfamilies by a single character vector for each, intending to represent the

groundplan for the group, as opposed to using several actual species as terminals for

each. Passoa's was the first treatment to include an outgroup (Yponomeutoidea) and

establish monophyly of Gelechioidea through the use of characters concerning the

83

haustellum, maxillary palpi, labial palpi, L1 and L2 setae of larvae, SD1 setae of larvae,

pupal antennae (Figure 4.1). He defined ((((Coleophoridae + Pterolonchinae) +

Momphidae) + Batrachedridae) + (Cosmopterigidae + Gelechiinae)) as a clade, sister to

the clade containing (Scythrididae + (Oecophoridae I + Oecophoridae II)) based on

morphological and ecological traits.

Hodges' (1998) landmark study represents the first effort to include many taxa globally with 37 family-level groups with extensive taxon sampling and a matrix of 38 characters with 100 states. Hodges used a variety of characters from all lifestages. From a constrained analysis, he presented one of 2,456 most parsimonious trees, chosen in part because it preserved homology among characters he favored. He designated 15 families with 32 subfamilies from 37 family- and subfamily-level terminals that he coded according to groundplan. Hodges redefined many family and subfamily relationships

(Figure 4.2).

84

Figure 4.1. Phylogenic tree showing sister-group relationships within Gelechioidea redrawn from Passoa 1995.

85

Kaila (2004) very recently presented a cladistic analysis of Gelechioidea for 156

taxa with 193 characters. He is the first to treat Gelechioidea using species as terminals

and presents the most detailed morphological study. Kaila is also the first to thoroughly investigate outgroup relationships. While Kaila’s treatment has many novel characters

(such as those of the thorax), it is based largely on Hodges’ and Passoa’s data matrices;

Kaila expanded many of Passoa’s and Hodges multi-state characters to separate

characters stating that, “The work of Hodges (1998) may also suffer from clustering

independent characters as artificial character complexes that do not always reflect true

observations.” Kaila criticizes Hodges for assuming subfamilies are monophyletic and

not comprehensively testing Gelechioidea family and subfamily identities. I do not agree

entirely with this sentiment. Hodges elevates monotypic genera to subfamily level which

means that he was treating generic identity. Kaila finds two main monophyletic lineages,

an Oecophorid Lineage and a Gelechiid Lineage. Monophyly of each of these lineages is

weakly supported by a Bremer value of 2. Internally, many deeper nodes have low

Bremer support values of 1 and 2 while generic lineages that are not in dispute have

higher values. This indicates that the addition of new taxa or new characters could cause

rearrangement of these deeper nodes.

86

Figure. 4.2. Phylogenic tree showing sister-group relationships within Gelechioidea redrawn from Hodges 1998. O1; Oecophoridae 1, O2; Oecophoridae 2, as designated by

Passoa 1995.

87

While each of these studies represents an advance, there are shortcomings to be

addressed, whether they be through taxon sampling, character coding, or the strategy of

analysis. Both Passoa and Hodges heavily rely on ground-plan coding, and Hodges uses a

hypothetical ancestor (both were standard practices at the time of the publications).

Hodges tree is the product of a constrained analysis and therefore, is not actually a most parsimonious solution to the matrix. Kaila’s large analysis suffers gravely from

ambiguous character coding, making assessment of evolution of those characters very

difficult. In many instances, despite the provided illustration or brief explanation, it is

unclear what he meant in his character definition and why taxa were coded as they were.

It is possible, of course, to decipher state assignment to taxa by reading the matrix, it is

not always clear what is meant by characters and states. Kaila excludes three taxa from the analysis that cause severe loss of resolution. While he does report this in the methods section of his manuscript, which is more than perhaps most researchers are willing to admit to, it would have been interesting to have a discussion of character optimization for those taxa when included in the matrix.

Relationships proposed by the authors vary greatly and have different implications for character evolution. Classification has been historically unstable within in the superfamily; very few entities have remained unchanged by authors working on the group. Nonetheless, several independent analyses agree with Passoa’s 1995 and Hodges’

1998 phylogenies in certain regards, (Lee and Brown unpubl., Landry unpubl). In particular, Kaila (2004) concluded that his analysis is similar in many ways to previous hypotheses including that of Hodges (1998) and Passoa (995), although he presented

88

several new groupings as well. Kaila’s matrix incorporated several of Hodges’ (1998)

and Passoa’s (1995) characters, if not explicitly stated as such, thus it is not surprising

that results are similar (below). To date, there has been no molecular approach to

compare with these morphological treatments. Building on the advancements of Passoa

and Hodges morphological studies, I base this analysis on their data matrices, coding

species as terminals, and adding the novel contribution of molecular data in the form of

Coenzyme I and II sequence data.

This study is a reexamination of terminals presented in Hodges 1998 phylogeny for which I have morphological and molecular data and primarily focuses on the subfamily Coleophorinae and historical putative sister groups. It is the first molecular analysis of the superfamily and it illustrates the reality that current phylogenetic treatments remain unstable; the addition of new taxa or new characters changes relationships, sometimes drastically.

MATERIALS AND METHODS

Taxon Sampling and Morphological Analysis.

Based on studies by Landry (1991), Minet (1991), and Passoa (1995),

Yponomeuta (Yponomeutoidea; Yponomeutidae) was chosen as the outgroup for this

study simply to root the tree. Definitive sister-group relationships to the Gelechioidea are

beyond the scope of this paper. Kaila’s (2004) analysis demonstrates monophyly of

Gelechioidea with respect to his chosen outgroups (including Yponomeutoidea).

89

Taxon sampling varies with each partition of data. Because of groundplan coding

(above), actual species were not terminals of earlier analyses (except for Kaila, 2004), so

sampling per se is not evaluated. The combined analysis of the Passoa’s and Hodges’

data matrices includes all taxa common to both analyses. Restricting the analysis to those

species for which I have DNA data (below) results in 42 vectors of morphology, CO-I,

and CO-II data. Because many genera are ephemeral as adults, they are difficult to collect

and generally not available except as pinned specimens. Ability to amplify good quality

DNA from such historical specimens determined to some extent which ones of many

possible species serve as the molecular terminals.

Fresh and dried specimens were used for this analysis (Figure 4.3). Fresh

specimens were collected and killed by freezing. Dried specimens were available from

museum and private collections. In most cases, the morphology and DNA vouchers are

the same individual.

States of morphological characters published Hodges (1998) and Passoa (1995)

were verified for available specimens and developmental stages. Morphology was studied

by making wing, whole body (adults and larvae), and genitalic preparations. Specimens

were prepared for identification according to Clarke (1941) using a standard 10%

potassium hydroxide solution. In some cases, structures were stained with

Mercurochrome. Preparations were mounted in euparol on slides following Robinson

(1976) and vouchered. Male structures were prepared variously using techniques

specifically aimed to preserve particular taxonomic features; females were left undissected and mounted whole. Larval skins were prepared similarly by clearing away

90

soft tissues and mounting whole skins on slides. Wings were prepared according to

Borror, Triplehorn and Johnson (1989). Identifications were made using the following:

Adamski and Brown, 1989; Adamski and Hodges, 1996; Covell, 1984; Forbes, 1923;

Hodges, 1974; Hodges, 1978; Hodges, 1983; Hodges, 1985; Hodges, 1986; Hodges,

1998, as well as museum study and unpublished work by Jean-François Landry on

Coleophora.

I compiled a morphological matrix consisting of 46 informative characters and

117 states from published data matrices by Hodges (1998) and Passoa (1995) for 23 ingroup taxa. Repeated characters were synonymized. Several of the published characters were recoded in a manner that makes better use of cladistic methods for this analysis: spiniform setae, gnathos, and the apodemes and venulae (See Appendix A, Appendix B,

Appendix C and Discussion). Taxa ambiguous for states were coded as missing or polymorphic in some cases.

91

Coleophoridae: Momphinae Mompha circumscriptella (Zell., 1873) Columbus, OH Mompha eloisella (Clem., 1860) Columbus, OH Mompha new species Kill Deer Plains, OH Elachistidae: Ethmiinae Ethmia trifurcella (Cham., 1873) Laurelville, OH (Wlsm., 1880) Tucson, AZ Elachistidae: Depressariinae robinella (Pack., 1869) Laurelville, OH Depressaria pastinacella (Dup., 1838) Columbus, OH sp 1 Lancaster, OH Elachistidae: Stenomatinae (Zell., 1854) Lancaster, OH Oecophoridae: Oecophorinae Epicallima aregenticinctella Clem., 1860 Laurelville, OH Mathildana newmanella (Clem., 1864) Laurelville, OH Fabiola shaleriella (Cham., 1875) Laurelville, OH Decantha boraesella (Cham., 1873) Laurelville, OH Autostichidae: Symmocinae Oegoconia qradripuncta (Haw., 1828) Columbus, OH Amphisbatidae: Amphisbatinae Clem., 1860 Lancaster, OH Cosmopterigidae: Cosmopteriginae sp 1 Laurelville, OH pergandeella Bsk., 1901 Columbus, OH serratella Treitschke., 1833 Cosmopterigidae: Chrysopeleiinae (Cham., 1875) Laurelville, OH juniperae Hodges, 1978 Redmond, OR Xyloryctidae: Scythridinae Asymmetura sp 1 Luskville, Quebec, Canada Scythris limbella (Fab., 1775) Urbana, Il

Figure 4.3. Names and collection localities for taxa used in total evidence analysis.

Taxonomy according to Hodges 1998. Continued on next page.

92

Figure 4.3. Continued from previous page.

Gelechiidae: Gelechiinae Gelechiinae 2 Portal, AZ sp 1 Portal, AZ Gelechia sp 4 Portal, AZ sp 1 Portal, AZ sp 1 Portal, AZ Gelechia sp 3 Lancaster, OH Teleiodini sp 1 Laurelville, OH Gelechiidae: Dichomeridinae Dichomeris ligullela Hbn., 1818 Lancaster, OH Gelechiidae: Pexicopiinae cerealella (Olivier, 1789) Columbus, OH

DNA Amplification, Sequencing and Alignment.

DNA samples were extracted from fresh specimens following the protocol for

Qiagen’s DNeasy Tissue Kit (Qiagen Inc. USA). A modified CTAB protocol (Phillips and Simon, 1995) was used for both fresh and pinned specimens. DNA from dried material more than 15 years old was successfully extracted and amplified using the modified CTAB protocol. A single back leg is sufficient for most extractions and is preferred for dried material. Sequences from all, but in particular, dried specimens were checked in Gen-Bank with the blast function to ensure that there was no contamination in our samples from bacteria or fungus.

PCR protocols were modified from Saiki (1990) (below) and carried out in 50 μl volumes; with 0.7 μl of Taq polymerase (added after 2 min at 80o in the PCR reaction,

93

see below), 5 μl of 10 x PCR buffer, 1.5 μl 1.5 mM MgCl2, 1.5 μl 20 mM dNTP, 1 μl of

each of amplification primer, and 0.5 – 1.5 μl of template. Sterile water was added to

make a total solution of 50 μl.

Choice of genes follows known utility in Lepidoptera systematics (see Appendix

D) and the ability to amplify high quality DNA from dried specimens (see Table 4.1). I

screened species to test the signal of 16S, 18S, CO-I, and CO-II. Mitochondrial protein

coding genes CO-I and CO-II provide resolution at the generic level (Brower, 1994);

mitochondrial ribosomal 16S is informative of recent divergences (Pashley and Ke, 1992;

Wahlberg and Zimmermann, 2000) nuclear ribosomal 18S provides resolution of basal

lineages (Wiegmann et al., 2000). While all these genes have potential for elucidating

phylogenetic patterns of Gelechioidea, CO-I and CO-II are most consistently amplified

from the dried specimens that I must rely upon.

Amplification primers for CO-I and CO-II were selected from Simon et al.

(Simon et al., 1994) and Brower (1994), respectively. CO-I primers used were as follows:

CO1P1 5’-TTG ATT TTT TGG TCA YCC WGA AGT-3’ and CO1R4 5’-CCW VYT

ARD CCT ARR AAR TGT TG -3’. Additional, internal primers for CO-I were designed

using existing sequences: CO1sibF 5’-TTC HCA AGA AAG AGG A-3’ and CO1sibR

5’-CCT AGG AAG TGT TGA GG-3’. CO-II primers used were as follows: CO2-S 5’-

TAA TTT GAA CTA TYT TAC CIG C-3’ and CO2-A 5’-GAG ACC ATT ACT TGC

TTT CAG TCA TCT-3’. Additional, internal primers for CO-II were designed using existing sequences: CO2sibF 5’-TTT ACC GGC WWT TAC WTT -3’ and CO2-Q 5’-

CCA CAA ATT TCT GAA CAT TGA CCA-3’. The amplification profile was 2 min at

94

94º, 15 min at 80º (Taq polymerase added at this stage), and 2 min at 94º for denaturing,

40 cycles of 1 min at 94º, 1 min at 45º and 1 min at 65º, and last 6 min at 65º for final extension.

PCR products were purified using Wizard PCR Preps (Promega Corp. Madison,

WI) and then loaded into 96-well plates for external sequencing using BigDye

Terminator Cycle Sequencing chemistry. Products were run on an automated ABI Prism

3700 DNA analyzer (Applied Biosystems, Inc.) and output proof-read using Sequencher

4.0.5 (Gene Codes Corp., Ann Arbor, MI). Nucleotide data were aligned using Clustal X

(Thompson et al., 1994) aligned sequences were proofread using Se-al (Rambaut, 1996).

Because the primary goal of this paper is to investigate the utility of adding molecular data to existing morphological data sets instead of completely restructurings existing data sets, parsimony was chosen as the only method of analysis for this study.

Phylogenetic analysis was performed using maximum parsimony searches in WinClada

(Nixon, 1999) implementing NONA (Goloboff, 1994) and the parsimony ratchet (Nixon,

1999). Bremer support was calculated with in Nona for trees up to 5 steps longer than the optimal trees (Bremer, 1998; 1994). Because there is evidence that a Bremer value of 3 or better corresponds to high bootstrap values (Davis 1995; but see DeBry 2001), bootstrap values were not calculated and Bremer values higher than 5 were not calculated (Wenzel,

2001).

95

ANALYSES AND DISCUSSIONS

General perspective

While Bayesian analyses may be capable of analyzing both morphological and

molecular data, I choose not to use this method of analysis because an increasing body of

evidence suggests that it is deeply flawed as implemented today. The posterior

probabilities of Bayesian analysis are excessively high (Suzuki et al. 2002, Cummings et

al. 2003, Simmons et. al. 2004; Pickett et. al. 2004), and flat prior probabilities do not model accurately a bias-free calculation (Pickett and Randle 2004). It is also shown that under circumstances of site-specific rate heterogeneity (a condition that is both common and difficult to demonstrate), both maximum likelihood and Bayesian analysis become strongly biased and statistically inconsistent, whereas parsimony performs consistently better under a wide range of conditions (Kolaczkowski and Thornton, 2004).

Clade support is represented by consensus cladograms and Bremer support

(Bremer 1994) because the goal of the paper is to analyze the combined, morpho- molecular data matrix rather than the molecular data alone.

Morphology

I was not able to reproduce Hodges' result in an unconstrained analysis of his matrix, whereas Passoa's matrix and result are rather straightforward. I used the 23 taxa common to both studies and recoded the combined matrix to account for redundancy between partitions or imprecise coding. Running the parsimony ratchet in WinClada followed by max* in NONA, or a separate analysis using mult*100 in NONA, produces

96

only 18 most parsimonious trees, with good resolution (length 135 steps, CI = 0.54, RI =

0.59). Although the strict consensus of these topologies is a basal 16-tomy (and a consensus length of 178 indicates a costly step away from optimality), diagnosis shows that the 18 solution trees fall into only two main topologies (see Figure 4.4). The distinction between the main topologies is based on trading components across three localities. These two topologies form the basis for all 18 with local rearrangements of

Pterolonchinae and Batrachedrinae; Oecophorinae is most unstable, appearing in five

locations including most basal.

97

Figure 4.4. One of 18 most parsimonious trees from the combined morphology matrix, showing one of two main topologies. Problematic taxa in boxes. Arrows indicate simultaneous rearrangement that generates the alternative main topology: hence, replace (Hypertrophinae + Agonoxeninae) with Chimabachidae; replace (Peleopodinae + Amphisbatinae) with (Hypertrophinae + Agonoxeninae); and replace Chimabachinae with (Peleopodinae + Amphisbatinae). Oecophoridae, Batrachedrinae, and Pterolonchinae take differing positions to generate nine equally parsimonious trees in each of the main two topologies. Other topological features are identical in all 18 trees

98

MtDNA

Our taxonomic knowledge of the Gelechioidea is largely based on historical

specimens of adults that are dried and pinned. Mitochondrial protein coding genes CO-I

and CO-II from such material can be of high quality, and may provide good characters

for the molecular analysis at the generic level (Brower, 1994). For 34 taxa spread across

Hodge's tree, I have 565 bases of CO-I, 218 of these positions parsimony informative. I

also have 508 bases of CO-II for 19 of these 34 taxa (plus 15 more that test the

monophyly of certain terminals). This 19 terminal matrix gives 1073 sites for CO-I+CO-

II, 396 are informative, and the matrix produces a single most parsimonious tree of 2147

steps with the ratchet and max* (CI = 0.29, RI = 0.35). From this phylogeny (Figure 4.5),

there are only a few expected relationships. Some results reinforce the preliminary

morphological analysis (Figure 4.4) and stand in contrast to Hodges, such as the disparate

positions of Coleophorinae versus either Blastobasinae or Batrachedrinae, but the tree in

general contrasts with most expectations, including loss of monophyly of many groups.

99

Figure 4.5. The single most parsimonious tree (L=2147, CI=0.29, RI=0.35) produced from molecular matrix (CO-I+CO-II) showing branch lengths. Black circles indicate uncontroverted synapomorphies, white circles indicate parallelisms.

100

Combined analysis

Combining the recoded morphology matrix that produced Figure 4.4 with the CO-

I+CO-II matrix gives 453 parsimony informative characters for the 42 taxa for which I

have sequence. Running with ratchet and max* (as above) produces two trees of 2321

steps (CI = 0.29, RI = 0.38). The consensus dissolves one node and is 10 steps longer,

shown in Figure 4.6. In our combined analysis, the monophyly of several families and

subfamilies of Gelechioidea is challenged, as well as the global topology of this

superfamily. It is very interesting that the combined matrix gives some results that differ

from the solutions to either the morphology or CO-I+CO-II partitions discussed above.

Coleophoridae are not monophyletic, but the component subfamilies Blastobasinae,

Momphinae and Coleophorinae are monophyletic (each has high Bremer support values

of 5 or greater). Batrachedrinae, also monophyletic (support of 4), allies with

Coleophorinae in clade G with a low Bremer support value of 2. This relationship is

consistent with Hodges 1978 and similar to Passoa 1995 but unlike Hodges 1998 or the

DNA data alone (Figure 4.5). This novel result indicates that the molecular data and

Hodges 1998 morphological data contain coherent secondary signals that did not emerge

from the preliminary separate analyses, but emerged from the combined matrix (Barrett

et al., 1991; Baker et al 1998; Sober and Steel, 2002). Blastobasinae is sister to clade b,

clade Z, and has very low support (1). This relationship is similar to relationships in

Passoa 1995, but not Hodges 1998. Similarly, the relatively basal placement of clade B

that includes Pterolonchinae and Momphinae is unexpected (Bremer support of 5).

Pterolonchinae are sister with (Teleiodini + Scythris limbella clade E; Bremer 5), to make

101

clade C (Bremer support of 5). Scythridinae are not monophyletic. Scythris limbella allies with Pterolonchinae and Teleiodini in clade C and Asymmetura allies with Antaeotricha schlaegeri in clade K with high support of 5. The relationship of clade K to

Coleophoridae is not highly supported (2). Cosmopterigidae are monophyletic (clade N;

Bremer support 2), but has a novel position in our phylogeny, sister to clade T. This position is not highly supported (2) and is not consistent with either Passoa 1995 or

Hodges 1998, where Cosmopterigidae are closely related to the Gelechiidae.

Chrysopeleiinae are monophyletic, but Cosmopteriginae are not, Eteobalea serratella is sister to Chrysopeleiinae, clade P (Bremer support 1). Ethmiinae and Depressariinae, both monophyletic (each with low support of 1 and 3 respectively), are sisters (clade V,

Bremer support 2) and that clade is sister to clade Z (Blastobasinae + clade b). The relationship of clade V is similar to both Passoa 1995 and Hodges 1998. Clade V + clade

Z, or clade U, is supported by a Bremer value of 4 and is similar to relationships in

Passoa 1995, but drastically different to Hodges 1998 (see below). Of particular concern are the Gelechiidae and Oecophoridae, neither of which is monophyletic in our phylogeny, and which are separated by the components that includes Ethmiinae,

Depressariinae, and Blastobasinae. The majority of taxa from the Gelechiidae are monophyletic, clade S, but Teleiodini allies with Pterolonche sp 1 and Scythris limbella

(clade C), also rendering Gelechiinae polyphyletic. The previous relationship is unexpected and may change with the addition of more taxa to the analysis. Dichomeris ligullela (Dichomeridinae) falls within the Gelechiinae (Bremer value of 3). The

Oecophorinae are polyphyletic within clade b (support 2), and are allied with Psilocorsis

102

reflexella (Amphisbatinae) and (Symmocinae). This arrangement is novel and different from both partitions. Support for internal nodes in

Figure 4.6 is variable, indicating that there is room for improvement (Figure 4.6).

In our analysis, the male split valve character is homoplastic, evolving at least four times (Figure 4.6 and below), suggesting that this character is in need of further investigation to assess morphological homology of associated features. The male gnathos, recoded from Hodges analysis as five characters with 14 states, provides synapomorphy for several lineages (Figure 4.6 and below), suggesting that this character is useful in detecting hierarchical patterns if not used as a multi-state character. In this analysis, the adult spiniform setae character is recoded as three characters. Spiniform setae are plesiomorphic for Gelechioidea. In this analysis, this character is lost three times and regained twice. The adult apodemes and venulae of the second sternum is uninformative at this level.

103

Figure 4.6. Consensus phylogeny (L= 2331) of two most parsimonious trees (L= 2321, CI = 0.29, RI = 0.38) produced from the combined analysis (recoded morphology+CO- I+CO-II). Numbers above nodes indicate Bremer support values.

104

Comparison with Kaila (2004)

After this paper was submitted to this journal, Kaila’s (2004) analysis was

published and available to us for the first time. Kaila’s paper is the most thorough

morphological paper to date. Nonetheless, many of his characters are substantially

synonymous with those we attribute to Hodges or Passoa, reinforcing the idea that there

is apparently limited additional morphological variation to harvest even by cautious and

thorough new investigators. Cursory comparison shows that the adjacency of taxa in the morphological tree of Figure 4.4 is rather similar to that of Kaila’s tree, but Kaila’s tree is rooted in a very different place. Of special interest to the present study is that the DNA data were able to produce certain results in our combined analysis (Figure 4.6) that are consistent with Kaila’s independent analysis of morphology, and that differ from the morphological tree of Figure7. Both Kaila and Figure 4.6 have the “Oecophorid lineage” monophyletic and apical in the tree (Figure 4.6, Clade U), whereas in Figure 4.4 it is paraphyletic with Amphisbatinae always rather basal in the tree and sometimes

Oecophorinae most basal (see discussion of Figure 4.4). Both Kaila and Figure 4.2 have

Blastobasine in the Oecophorid lineage, in contrast to Figure 4.4 (or Hodges 1998). Both

Figure 4.6 and Kaila have a close association of Pterolonche, Scythris and Mompha; both

Figure 4.6 and Kaila, show a close association of Batrachedra and Coleophora. Each of

these combinations differs from the morphological solution of Figure 4.4. Thus, it is

encouraging to see that several of the results of Figure 4.6 that appear to be due to DNA

105

data are corroborated by an independent worker with an alternative morphological data

set. There are differences between Figure 4.6 and Kaila, for example Figure 4.6 has a

paraphyletic “Gelechiid lineage”, sensu Kaila (Figure 4.6, everything but Yponomeuta and Clade U) whereas Kaila has it monophyletic and the sister to the Oecophorid lineage.

Combining our DNA data with the corresponding taxa of Kaila’s matrix (giving a

reduced matrix of the 19 terminals common to both) produces a monophyletic

Oecophorid lineage, and a basal paraphyly of the Gelechiid lineages (not shown). This matches what we find with our own combined analysis, indicating that the DNA signal is strong enough that it can shape Kaila’s morphological topology just as it shaped the

Hodges-Passoa data of Figure 4.4. We expect to continue this work in closer cooperation with Kaila.

106

Figure 4.7. Phylogenetic tree produced by combining Kaila’s morphological data with the

molecular data of Figure 4.5.

Character Evolution

Male valves divided or entire – Passoa (1995) and Hodges (1998) both used this character in their cladistic analyses. In Passoa’s (1995) analysis, split valves evolves twice, once in the ancestor to the clade ((Coleophoridae + Pterolonchidae) + Momphidae) with a reversal in Pterolonchidae, and once in the ancestor to Blastobasidae. In Hodges’ (1998)

analysis, split valves only evolve once, in the ancestor to (Momphidae + Coleophoridae)

107

+ Blastobasidae). In the combined evidence phylogeny (Figure 4.6), this character evolves at least four separate times suggesting the possibility of a homoplastic character in need of further investigation.

Male with gnathos, and shape and position of gnathos – Passoa (1995) coded this

character for taxa only within the oecophorid clade. In his analysis, the presence of a

spined gnathos unites taxa in the oecophorid group 1: Peleopodinae, Xyloryctinae,

Hypertrophinae, Agonoxeninae, Amphisbatinae, Depressariinae, Ethmiinae, and

Elachistinae. The absence of spines and the absence of a gnathos in Xyloryctinae and

Agonoxeninae respectively are considered reversals. Ethmiinae is coded as polymorphic

due to what Passoa thought was a plesiomorphic absence of a gnathos in basal ethmiines.

Passoa (1995) believed the presence of a spined gnathos in both Coleophoridae and

Deoclonidae to be the result of parallel evolution. Hodges (1998) coded this character

with eight states, and the ninth as absent in Aeolanthinae, Deuterogoniinae,

Antequerinae, Chrysopeleiinae, Cosmopteriginae, Hypertrophinae, Syringopaidae, and

Momphinae. The remaining states are coded with attention paid to presence of an

articulated or sclerotized band, articulated rami or articulated sclerites, with finer

variations within each state. Hodges did not treat the presence/absence of spines on the

gnathos. This character evolves multiple times in his phylogeny. For our analysis, I

recoded it as five characters with 14 states resulting in less homoplasy overall. In Figure

4.6, the gnathos is plesiomorphically present (68,1) and lost twice (0) in the ancestors of

Momphinae and (Cosmopteriginae+ Chrysopeleiinae). If the gnathos is present (69), it is

plesiomorphically a band (0), and having articulated symmetric sclerites (2) is derived

108

twice in the ancestor of Teleiodini sp 1 and the ancestor of (Sitotroga cerealella

(Gelechia sp 3 (Gelechia sp 1 (Dichomeris ligullela (Prolita group (Gelechiinae 2

(Gelechia sp 4 (Filatima sp 1)))))))). A sclerotized band (70,0) is basal, while an articulated (1) is derived and evolves 5 times in the ancestors of Pterolonchinae,

Batrachedrinae, Depressariinae, Psilocorsis reflexella (Amphisbatinae), and Oegoconia

quadripuncta (Symmocinae). If the gnathos is a sclerotized band fused to tegumen (71),

then having it be informally wide (0) is basal; having it be mesially turned down and

laterally compressed (1) evolves once in the ancestor of Coleophorinae; and having a

mesial bulb with a parallel row of spines (3) evolves once in the ancestors of Epicallima

aregenticinctella (Oecophorinae), Fabiola shaleriella (Oecophorinae), and (Mathildana newmanella + Decantha boraesella (Oecophorinae)). If the gnathos is an articulated band

(72), an unarticulated mesial hook (1) is basal; having the mesial region slightly expanded (0) is derived the ancestor of Batrachedrinae; and having a mesial bulb bearing row of short spines (2) is derived in the ancestor of Depressariinae.

Adults with patches or bands of spiniform setae on abdominal terga – Passoa

(1995) coded this character as present or absent within Gelechioidea, and then expanded

this character further within the oecophorid taxa, so that there is no continuity between

the two matrices. Batrachedridae, Momphidae, and Coleophoridae were coded as having

patches of stout setae, while Pterolonchidae was coded as having a band of setae present

but treated as a fusion of the patches. In this matrix, the oecophorid group 2 taxa were

coded as having a band of setae present but treated as a parallelism. Within the

oecophorid group 2 taxa, Blastobasinae, Oecophorinae, Lecithocerinae, Symmocinae and

109

Autostichinae were coded as having a stout band of spiniform setae, and treated as distinct from the band, which Pterolonche have, while this character is coded as absent in all other oecophorid taxa. Hodges (1998) coded this character as multistate, nonadditive with four distinct states including absent. Coleophoridae, Batrachedridae, Epimarptinae and Momphinae are coded as having patches of spiniform setae, Autostichinae,

Blastobasinae, Oeciinae, and Pterolonchinae were coded as having a band of spiniform

setae, and Stathmopodinae were coded as having a band of spiniform setae on posterior

margin of most segments. The remaining taxa were coded as having only regular scales

in this location except Oecophoridae and Deoclonidae, which were coded as being polymorphic for this character. In this analysis, the character is coded as three characters.

In Figure 4.6, adult abdominal segments with spiniform setae (63) present (1) is plesiomorphic in this analysis. It is lost 4 times and regained twice, once in the ancestor of Blastobasinae and once in the ancestor of Oegoconia quadripuncta (Symmocinae).

Spiniform setae (65) in patches (0) evolves twice and spiniform setae in band (1) evolves twice. The character of spiniform setae, when present, (64) in patches or band medially

(0) or in band along posterior margin (1) is uninformative in this analysis but could be so with the addition of more taxa.

Second sternum with apodemes and venulae – Hodges (1998) coded this character

with four states: venula only; both venula and apodeme present; apodeme only; and both venula and apodeme absent. In his phylogeny, the character is homoplastic and does not appear as a synapomorphy to unite any groups as sisters. Passoa did not include this character. In this analysis, Hodges’ character was recoded as 2 two characters, each with

110

two states (absent or present). The venulae character is uninformative in this analysis while the apodemes evolve twice and are lost four times (Figure 4.6).

CONCLUSIONS

This is the first cladistic analysis of Gelechioidea to include molecular data for a total evidence analysis. The addition of Cytochrome oxidase I and II to revised published morphological matrices gives 453 parsimony informative characters for the 42 taxa for which I have sequence data. The analysis resulted in two trees with mostly novel sister- group relationships based on this relatively small sample of taxa. These results challenge current concepts of Gelechioidea, suggesting that traditional morphological characters that have united taxa may not be homologous and are in need of further investigation. It also indicates that a total evidence approach may be the most robust method of study for

Gelechioidea because morphological characters are limited in number and utility due the difficulty of making appropriate homology statements across all terminals.

This study originally focused on the definition of Coleophoridae. It became clear during the analysis that most gelechioid taxa would need to be included in the analysis to make statements regarding character evolution and to define Coleophoridae confidently.

All matrices agree that Coleophoridae is not a monophyletic taxon. There is disagreement on the placement of Blastobasinae in relation to coleophorid taxa, and total evidence and combined morphology matrices suggest that they are not closely related. In our total evidence analysis, while the subfamilies of Coleophoridae (Momphinae, Coleophorinae,

111

and Blastobasinae) are monophyletic, the family is not. Batrachedrinae allies with

Coleophorinae, Momphinae allies with Pterolonchinae and Blastobasinae allies with the

oecophorine taxa. There is a novel placement of Cosmopteriginae. Our analysis also

indicates that Scythridinae, Elachistidae, Oecophoridae sensu latu, and Gelechiidae are

not monophyletic, either. Scythris limbata allies Pterolonche while Asymmetura allies

with Coleophorinae and Stenomatinae. Within Elachistidae, the Ethmiinae and

Depressariinae ally with the oecophorines and Stenomatinae allies with Coleophorinae.

Within Oecophoridae, Oecophorinae are polyphyletic and ally with Amphisbatinae,

Symmocinae, and Blastobasinae. Within Gelechiidae, the Gelechiinae are not

monophyletic – Dichomeridinae are within the Gelechiinae and Teleiodini ally with

Scythris limbata and Pterolonche. In our total evidence analysis, the male split valve

character is homoplastic, evolving at least four times, the male gnathos is a synapomorphy for several lineages and spiniform setae are plesiomorphic for

Gelechioidea. While we regard the present work as important progress, it is clear we have a long way to go before Gelechioidea is understood.

112

CHAPTER 5

NORTH AMERICAN FLAT-BODY MOTHS (ELACHISTIDAE: DEPRESSARIINAE:

DEPRESSARIA HAWORTH): MORPHOLOGICAL EVOLUTION, HOST-PLANT

SELECTION, AND GEOGRAPHIC DISTRIBUTION

INTRODUCTION

Depressaria Haworth 1812 (Gelechioidea: Elachistidae: Depressariinae) is a

Holarctic genus of microlepidoptera with nearly 100 described species. Seventy-six species are from the Palearctic region, 21 species are endemic to North America and 3 species are Holarctic (Hodges, 1974). Adults are morphologically very similar and rather unremarkable animals. They are drably colored with mottled brown wings, sometimes with an overall orangeish tinge. They are difficult to differentiate based on wing maculation alone because there is slight variation within species as well as between species. Taxonomists have relied on characteristics of male and female genitalia as well as host-plant data to designate species; rearing records and dissection of adults are necessary for identification of Depressaria (a statement that also applicable to species

113

within the closely related genera, for that matter). Not only are species of Depressaria

morphologically similar to eachother, but they are morphologically similar to species in the closely related genera of Agonopterix, , Nites and Apachea. Depressaria can

be distinguished from these genera based on characters of wing venation (Cu1 and Cu2

stalked and curved posterad in Agonopterix and Exaeretia versus separate basally in

Depressaria), the labial palpus (with anterior directed tuft in Apachea versus tuft lacking

in Depressaria), and the ocelli (lacking in Nites and present in Depressaria) (Clarke,

1941b; Hodges, 1974). Clarke (1941b) describes the adults as having the abdomen dorsoventrally flattened. In several species, especially Depressaria, the abdominal tergites appear to be smaller in length than the abdominal sternites causing them to roll up laterally giving them their flattened appearance (Bucheli observation). The etymology of the genus name Depressaria may refer to this character, but the concept of

Depressaria has changed rather drastically since its establishment and not all species that were once included in this genus are still included, making this character an unreliable feature for identification purpose but perhaps a phylogenetically informative one at the higher level.

Host-Plant Use

Depressaria, as well as the other members of the Depressariinae, are active mainly at night and are attracted to lights. They are well represented in areas where their host-plants occur; most of the diversity of Nearctic Depressaria occurs in the west

(purportedly new species are still being collected (Mckenna and Berenbaum, 2003)). The

114

larvae are leaf-tiers and feed in the umbels or meristems of their host tissue. First instar

larvae construct a small shelter of silk in the flower heads or between leaves of their host

(Hodges, 1974). Depressaria may be best known for their intimate association with their

larval host-plants; all described Nearctic Depressaria species whose larval biology is

known feed on plants in the families Apiaceae or Asteraceae (Hannemannn, 1995;

Hodges, 1974; Mckenna and Berenbaum, 2003). The majority of Nearctic species feed on plants in the family Apiaceae (17 species) (Hodges, 1974). Apiaceous, rutaceous

(Berenbaum and Zangerl, 1991), moraceous (Abegaz et al., 2004) and fabiaceous

(Innocenti et al., 1997) plants are known to contain a class of secondary plant chemicals called furanocoumarins, some of which are phytotoxins and react through an increase in toxicity when exposed to ultraviolet light. Phytotoxic furanocoumarins have been demonstrated to react to proteins, lipids, and DNA in a range of organisms including bacteria, insects, and mammals, causing in humans a reaction ranging in expression from a mild, itching skin irritation to blisters similar to those caused by third degree burns.

Many of those who have attempted to collect for their dissertation work Depressaria pastinacella from their larval shelters constructed in the umbels of Heracleum maximum can attest to phytotoxicity of furanocoumarins.

The ability to feed on and metabolize toxic host-plants in Depressariinae lead

Berenbaum to speculate that populations of Depressaria pastinacella could have coevolved with populations of its toxic host-plant sativa (Berenbaum, 1983) based on Ehrlich and Raven’s classic mechanism of coevolution through escape and radiation (1964). She has shown that chemically complex, toxic angular furanocoumarins

115

have evolved from less complex, less toxic coumarins, with two intermediate steps of intermediate toxicity, hydroxycoumarins and linear furanocoumarins. Plants containing angular furanocoumarins support fewer herbivores than plants with hydroxycoumarins.

These plants, in turn, support fewer herbivores than plants with linear furanocoumarins.

Plants with coumarins support the most herbivores (Berenbaum, 1983). Berenbaum and her colleagues have shown that Depressaria pastinacella larvae ingest large amounts of furanocoumarins daily, the majority of which are metabolized and excreted. They are able to metabolize linear furanocoumarins rapidly but angular furanocoumarins more slowly (Berenbaum and Zangerl, 1994). There seem to be interesting behavioral responses by Depressaria pastinacella to avoiding ultraviolet light, as well. Larvae that feed on their host plant are more orange in color than those that feed on a diet low in carotenoids. Larvae that have a diet low in carotenoids avoid sunlight while those that have a diet high in carotenoids do not (Carroll, 1997). This data suggests that the carotenoids potentially confer protection from ultraviolet light to larvae of Depressaria pastinacella. It has also been noted that the silken shelters of Depressaria pastinacella are thought to be an adaptation to avoiding ultraviolet light, as well. By constructing these shelters, it is theorized that the larvae are shaded from sunlight and the furanocoumarins are not activated. These hypotheses have not been discussed in a phylogenetic context.

Berenbaum and Passoa (1999) investigated in a cladistic framework the possibility that genera within Depressariinae have coevolved with their host-plants. They investigated the number of independent colonization events of Depressariinae on

116

Apiaceae and Asteraceae. They found no over-all pattern suggesting a potential case of

coevolution where basal lineages of Depressariinae would feed on less toxic plants

containing coumarins. According to their results, there have been two independent

colonization events of Apiaceae by Depressariinae genera (by species within Depressaria

and Agonopterix), potentially more if monophyly of purported species groups is not supported. Figure 5.1 illustrates host-plant use by Depressariinae. This phylogeny is a hypothetical synthesis of Berenbaum and Passoa’s (1999) phylogeny and a cladistic analysis of Depressaria species groups. According to this hypothetical phylogeny, the ancestral condition for host-use is feeding on the leaves of tree species. Species of

Apachea feed on species of Ptelea, a genus within the furanocoumarin-producing family

Rutaceae. Species of Nites feed on many different genera of trees, but are commonly recorded feeding on species of Betula, Corylus, and Ostrya. Within Depressaria, there

are two main species group lineages, each containing asteraceous-feeders and Apiaceous-

feeders (these data are based on an unpublished cladistic analysis of species groups by

Passoa and reanalyzed by Bucheli (See appendix E for the data matrix)). Apically in the

generic level phylogeny, both species of Agonopterix and Exaeretia feed on plant families containing both growth types of woody and herbaceous. Both have been recorded from species of Asteraceae, but only Agonopterix has been recorded from species of Apiaceae. It is equivocal as to the ancestral condition for host-plant use within

Depressaria. According to this phylogeny, Nites, the sister lineage of Depressaria, does not utilize plants that produce furanocoumarins; however the most basal lineage of the analysis, Apachea, does. Within Depressaria, ancestral host-plant use for Depressaria is

117

ambiguous according to this analysis because neither of the outgroups use Apiaceae or

Asteraceae as larval host-plants, making polarization of characters difficult. Optimization

of larval host-plants onto the species group phylogeny results in an equally most

parsimonious choice for either host-plant family being ancestral and then lost twice. In no

scenario are all of the asteraceous- or Apiaceous-feeder clades monophyletic. This

phylogeny suggests that a more detailed analysis of Depressaria and sister groups is

required to make any statement regarding host-plant selection.

Figure 5.1. Hypothetical phylogeny of Depressariinae based on the published phylogeny by Berenbaum and Passoa (1999) and cladistic analysis of Depressaria species groups (unpublished work by Passoa and Bucheli (See Appendix D for the data matrix)). A pink “” indicates a derivation of asteraceous-feeding while a blue “x” indicates loss of asteraceous-feeding; a blue “”indicates a derivation of apiaceous-feeding while a pink “x” indicates loss of apiaceous-feeding. Orange indicates a genus with species that feed on plants that produce furanocoumarins. 118

Species Group Definitions

Nearctic species of Depressaria were arranged into five groups by Clarke mainly

based on genitalic characters (1941b):

1. atrostrigella, artemisiae, palousella - process of the costa of the valve (in males)

and the broad somewhat dilated sclerotized ductus bursae (in females).

2. juliella, eleanorae, heracliana (now pastinacella in part), cinereocostella- strong

basal process from the sacculus and no “clasper” (or distal process) (in males) and

an elongated sclerotized section of the ductus bursae posteriorly (in females).

3. artimisiella, alienella - distal process present and basal process absent (in males)

and ductus bursae is entirely membranous (in females).

4. togata, angustati, multifidae - spined basal process (in males).

5. maculatella, betullela, grotella - divided distal process (in males) and spiraled

ductus bursae (in females).

Hannemann delineated and formally named species groups in 1953 for the European taxa (and later revised them, Hannemannn, 1995).

1. The douglasella-Group – basal process thorny, hairy, shrub-like or whip-like

process (clavus), distal process cone-shaped, or hook-like. Anellus flattened, often

heart-shaped. Vinculum pointed or rounded. Transtilla membranous or a finely

thorny band. Socii are band- or lobe shaped. Aedeagus curved with a wing-like

119

process on many species. Ductus bursae strongly sclerotized and finely structured

in front of the ostium. Signum flattened, cord-shaped or rhomboid, and densely

thorny.

2. The artemisiae-Group - basal process absent, distal process is absent or reduced.

Cucullus with a pointed or rounded process. Anellus varying. Gnathos is spindle-

shaped and furrowed at the median. The socii are band-like. Ductus bursae and

signum variable.

3. The pastinacella-Group – basal process curved and more or less densely thorny

and distal process absent or strongly reduced. Vinculum pointed or rounded.

Gnathos is spindle-shaped and rarely divided in two. Aedeagus is either straight

or curved, and the cornuti are always substantial. Ductus bursae sack-like and

widened before the ostium and finely thorny. Signum is variously toothed and

irregular in outline.

4. The discipunctella-Group – Distal process cone-like. Valves sickle-like.

Vinculum more or less long and pointed. Anellus strongly sclerotized plate.

Transtilla membranous. Gnathos oval. Socii more or less band-like. Aedeagus

long and narrow, most species with a long cornutus. Signum plate-like, broader

than tall, and is strongly thorny. Hannemann places D. leucocephala in this group

saying, “is only treated here for purely practical reasons. In the structure of the

male copulation apparatus, this species differs from all others.”

Hodges (1974) later segregated described Nearctic into species groups using Clarke’s and Hannemann's system. He named Clarke’s groups based on Hannemann’s taxonomy

120

and expanded Clarke’s groups to include all known Nearctic species. He created a new

group, the Betina Group, which includes only two endemic North American species

distinct morphologically from the European taxa. He removed five species from

Depressaria (Clarke’s fifth group that included the species maculatella, betullela, grotella) to form the genus Nites. Following Hodges, species groups are capitalized, although earlier authors did not do so.

1. Artemisiae Group – Males without basal or distal process, but often with small

lobe on valva. Costal margin with of valva often with broad projection at apex.

Aedeagus stout, vesica with long and stout cornuti. Females with anterior

margin of A8 sternite convex or slightly produced medially. Ostium bursae

near anterior margin surrounded by a membranous region. Ductus bursae short

to long and varyingly sclerotized.

2. Pastinacella Group – Males with basal process but lacking distal process and

lacking lobes on valva. Vesica with stout cornuti. Vinculum somewhat pointed.

Females with ostium bursae on anterior half of A8 sternum, anterior margin of

A8 sternum heavily sclerotized. Ductus bursae varyingly sclerotized,

sometimes bulbous near base.

3. Thomaniella Group – Males with distal process but lacking basal process.

Aedeagus long, slightly curved, with small flange anteriorly. Vesica with

numerous fine cornuti. Females with base of ductus bursae shaped like an ogee

arch7. Ostium bursae at middle of A8, behind it membranous.

7 This is Hodges’ terminology. ‘Ogee arch’ is defined by the Oxford English Dictionary as: In architecture, an arch whose curve is formed by two S-shaped or double curves meeting at its apex. 121

4. Betina Group – Males with free distal process but lacking basal process.

Aedeagus straight or twisted, base with flange. Vesica with stout cornuti.

Females with ostium bursae membranous and in a broad w-shaped outline.

Ductus bursae with base rigidly walled.

5. Douglasella Group – Males with basal and distal process. Basal process usually

with scale-like projections. Aedeagus long or short, with anterior and posterior

flange. Vesica without cornuti. Females with ostium bursae at middle of A8

sternum, sternum membranous posterad of ostium. Base of ductus bursae

rigidly walled.

Despite their limited fame, no modern species level phylogeny exists for Depressaria.

Here I present a morphological analysis using parsimony to investigate species level relationships for Nearctic Depressaria and make conclusions regarding monophyly of the

genus, host-plant use within the plant families of Asteraceae and Apiaceae, and

distribution of species in the worlds of old and new.

MATERIALS AND METHODS

Taxon Sampling

A generic level analysis for Depressariinae by Passoa (1999) and Berenbaum and

Passoa (1999) delineates the Depressariinae as including Nites, Apachea, Depressaria,

Agonopterix, and Exaeretia. This differs from Hodges (1974) treatment

122

which included all the above genera, but also and Himmacea. I have chosen to include species representatives from all genera included in Hodges (1974) treatment.

This is because of two recent treatments of Gelechioidea by Kaila (2004) and Bucheli and

Wenzel (2005) demonstrating that Gelechioidea systematics is highly unstable. Kaila’s

analysis places Semioscopis as more closely related to the Depressariinae than it is to the

Amphisbatinae. Because the analysis is concerned with establishing the monophyly of the

genus Depressaria and its species groups, as well as investigating evolution of host-plant

selection within the Depressariinae, representatives from all genera in Hodges 1974

definition of Depressariinae were included. I am treating Psilocorsis reflexella

(Amphisbatinae) as the distant outgroup. I am also treating species of Palearctic

Depressaria as outgroups, but have included as many of these species as possible in the

analysis (Figure 5.2).

123

TAXA ARTEMISIAE GROUP 1 • artemisiae NICKERL, 1864 (s.n. dracunculi CLARKE, 1933) 2 • atrostrigella CLARKE, 1941 2 • palousella CLARKE, 1941 • absynthiella HERRICH-SCHÄFFER, 1865 (s.n. tenerifae WALSHINGHAM, 1907; absynthiella 3 HEINEMANN, 1870; absynthiella STAINTON, 1870) absinthivora Frey. 3 • haydenii ZELLER, 1854 (s.n. haydenii FREY, 1856; haydenii STAINTON, 1861) • depressana (FABRICIUS, 1775) (s.n. Tinea depressella HÜBNER, 1813; Tinea depressana 3 FABRICIUS, 1798; depressella ZELLER, 1854; depressana HERRICH-SCÄFFER, 1854) • chaerophylli Zeller, 1839 (s.n. chaerophyllinella HERRICH-SCÄFFER, 1851; chaerophylli 3 HERRICH-SCÄFFER, 1854)

PASTINACELLA GROUP • pastinacella (DUPONCHEL, 1839) (s.n. Phalaena Tortrix heracliana LINNAEUS sensu auct.; heraclei HAWORTH,1811; heracleana ZELLAR, 1854; heracleana STAITON, 1861; ontariella BETHUNE, 1870) • daucella (DENIS & SCHIFFERMÜLLER, 1775) (s.n. Tinea daucella [DENIS & SCHIFFERMÜLLER, 1 1775]; Tinea rubricella [DENIS & SCHIFFERMÜLLER, 1775]; Tinea apiella HÜBNER, 1976) 2 • cinereocostella CLEMENS, 1864 (s.n. clausella WALKER, 1864) 2 • juliella BUSCK, 1908 2 • eleanorae CLARKE, 1941 3 • ultimella STAINTON, 1849 • rubricella (DENIS & SCHIFFERMÜLLER, 1775) (s.n. Tinea daucella DENIS & SCHIFFERMÜLLER, 3 1775; Tinea apiella HÜBNER, 1796; nervosa STEPHANS, 1834; nervosa ZELLER, 1854) 3 • bupleurella HEINEMANN, 1870 • pimpinellae Zeller, 1839 (s.n. Haemylis pulverella EVERSMANN, 1844; pimpinellae ZELLER, 3 1846; reichlini HEINEMANN, 1870) 3 • libanotidella SCHLÄGER, 1849 (s.n. libanotidella ZELLER, 1854; libanotidella STAINTON, 1861) 3 • silesiaca HEINEMANN, 1870 (s.n. Schistodepressaria freyi HERING, 1924) • badiella (HÜBNER, 1796) (s.n. Tinea badiella HÜBNER, 1796; frigidella TURATI, 191; brunneella 3 RAGONOT, 1874; : uhrykella FUCHS, 1903; frustratella REBEL, 1936) 3 • velox STAUDINGER, 1859 (s.n. tortuosella CHRÉTIEN,1908)

Figure 5.2. Species of Depressaria used in the analysis.1 = Nearctic; 2 = Introduced to North America; 3 = Palearctic. Specimens were borrowed from the Muséum national d'Histoire naturelle, in Paris, France, and the National Museum of Natural History, Washington, D.C. Continued on next page.

124

Figure 5.2. Continued from previous page.

THOMANIELLA GROUP 1 • alienella BUSCK, 1904 (s.n. nymphidia BUSCK, 1904; corystopa, MEYRICK, 1927) 1 • artimisiella MCDUNNOUGH, 1927

BETINA GROUP 1 • betina CLARKE, 1947 1 • constancei CLARKE, 1947

DOUGLASELLA GROUP 1 • whitmani CLARKE, 1941 1 • schellbachi CLARKE, 1947 1 • angelicivora CLARKE, 1952 1 • leptotaeniae CLARKE, 1933 1 • yakinae CLARKE, 1941 1 • multifidae CLARKE, 1933 1 • moya CLARKE, 1947 1 • besma CLARKE, 1947 1 • pteryxiphaga CLARKE, 1957 1 • togata WALSHINGHAM, 1889 (s.n. trusta CLARKE, 1947) 1 • armata CLARKE, 1952 • angustati Clarke, 19411 3 • douglasella STAINTON, 1849 (s.n. miserella HERRICH-SCHÄFFER, 1854) 3 • weirella STAINTON, 1849 (s.n. gudmanni REBEL, 1927) 3 • nemolella SVENSSON, 1982 3 • beckmanni HEINEMANN, 1870 • pulcherrimella STAINTON, 1949 (s.n. pulcherrimella ZELLAR, 1854; • emeritella STAINTON, 1849 (s.n. emeritella HERRICH-SCHÄFFER, 1854; emeritella, 3 • ZELLER, 1854; emeritella, STAINTON, 1861) 3 • hofmanni STAINTON, 1861 • albipunta HAWORTH, 1811 (s.n. Tinea albipunctella, HÜBNER, 1796; T. albipunctella DENIS & SCHIFFERMÜLLER, 1775; Agonopterix aegopodiella HÜBNER, 1825; albipunctella STEPHANS, 3 1834; albipunctella FREY, 1856; albipunctella STAINTON, 1861) 3 • olerella ZELLER, 1854 3 • ululana RÖSSLER, 1866

DISCIPUNCTELLA GROUP 3 • leucocephala SNELLEN, 1884 3 • cervicella HERRICH-SCHÄFFER, 1851 3 • gallicella CHRÉTIEN, 1908 3 • discipunctella HERRICH-SCHÄFFER, 1851

125

Character Sampling

For the ingroup, twenty-three Nearctic species were included in the analysis (20

endemic, 3 Holarctic), as well as 25 species from the Palearctic region. Specimens used

for this analysis were borrowed from the Muséum national d'Histoire naturelle, in Paris,

France, and the National Museum of Natural History, Washington, D.C. Morphology was

studied by genitalic preparations when necessary. Specimens were prepared for study

according to Clarke (1941a) using a standard 10% potassium hydroxide solution. In some

cases, structures were stained with Mercurochrome. Preparations were mounted in

euparol on slides following Robinson (1976). Male structures were prepared variously

using techniques specifically aimed to preserve particular taxonomic features; females

were left undissected and mounted whole. Slides were viewed using compound and

stereoscopic microscopes. Illustration of genitalia in this manuscript were drawn first as

pencil sketches with the aid of a drawing tube mounted on a compound microscope and

magnified at 100x, digitalized by scanning, and then finalized using Adobe® Illustrator

CS to create scalable vector files (Figures 5.3 – 5.23).

Morphological characters

A list of morphological characters used in parsimony analysis to generate the data matrix in Appendix F follows. Characters 0 - 4 of this analysis were originally coded in the analysis by Passoa (1995) and Berenbaum and Passoa (1999) and applied here.

Although they used groundplan coding, it is still useful to include these characters. The

126

remaining list of characters is my original interpretation of many key taxonomic features

used by Clarke, Hodges, and Hannemann to identify species and species groups plus my

original observations. See figures 5.3 – 5.23 for illustrations of characters and states.

0. D1 and D2 on T2 and T3 of larva: united = 0; separate = 1. (From Passoa

(1995)/Berenbaum and Passoa (1999)).

1. Number of teeth on mandibles of larva: greater than 4 = 0; 4 = 1. (From Passoa

(1995)/Berenbaum and Passoa (1999)).

2. Pinaculum of larva: pale and not well developed = 0; pigmented and well developed =

1. (From Passoa (1995) and Berenbaum and Passoa (1999)).

3. Intersegmental membrane of pupa: without sclerotized toothed ridges laterally = 0;

with sclerotized toothed ridges laterally = 1. (From Passoa (1995) and Berenbaum

and Passoa (1999)).

4. Prothoracic femur of pupa: hidden = 0; exposed = 1. (From Passoa (1995)/Berenbaum

and Passoa (1999)).

5. Second segment of labial palpus: smooth scaled = 0; with tuft = 1.

6. Ocelli: absent = 0; present = 1.

7. Antennal pectin: absent = 0; present = 1.

8. Cu1 and Cu2 of forewing: not stalked = 0; stalked = 1.

9. Abdomen: not flattened dorsoventrally = 0; flattened dorsoventrally = 1.

10. Uncus: absent = 0; reduced = 1 (Figure 5.3); present = 2. [additive].

11. Uncus: short rounded = 0 (Figure 5.3); well developed triangular = 1.

12. Socii: absent = 0; reduced = 1 (Figure 5.3); present = 2. [additive].

127

13. Uncus and socii: separate = 0; fused = 1.

14. Gnathos: narrow ovoid = 0 (Figure 5.3); broad triangular = 1.

15. Ductus bursa meets corpus bursa: at right angle = 0; straight or curved = 1.

16. Aedeagus: long = 0 (Figure 5.13, C); short = 1 (Figure 5.20, C).

17. Aedeagus: strongly curved = 0 (Figure 5.4, B); c-shaped = 1 (Figure 5.5, B); nearly

straight = 2 (Figure 5.12, B); twisted = 3; s-shaped = 4 (Figure 5.10, C).

[nonadditive].

18. Aedeagus flange: absent = 0 (Figure 5.7, B); single = 1 (Figure 5.10, C); double = 2

(Figure 5.6, C). [nonadditive].

19. Vesica of aedeagus: bare = 0 (Figure 5.16, C); with cornuti = 1 (Figure 5.14, B).

20. Cornuti: scale-like numerous in patch = 0 (Figure 5.7, B); stout and finger-like and

either single or few in row = 1 (Figure 5.9, C); saw-tooth like around base of

vesica = 2. [nonadditive].

21. Cornuti fingerlike: limited to middle of vesica = 0 (Figure 5.13, C); extending near tip

of vesica = 1 (Figure 5.14, B).

Hodges used presence and type of cornuti to decribed species groups of

Depressaria. In this study, I found presence or absence, type of cornuti and location of cornuti to be informative.

22. Dorsal margin of sacculus: with lobe or process basally (short or long) = 0; without

lobe or process basally = 1 (Figure 5.11, A).

23. Basal process: small hardly distinct = 0 (Figure 5.4, A); elongate = 1 (Figure 5.13, A);

short = (Figure 5.18, A) 2. [nonadditive].

128

24. Basal process: without scalelike projections = 0 (Figure 5.4, A); with scalelike

projections = 1 (Figure 5.20, A).

25. If basal process with scalelike projections: at tip = 0 (Figure 5.9, A); covering entire

process = 1 (Figure 5.21).

26. Scale-like projections of basal process: stout = 0 (Figure 5.8, B); long = 1 (Figure

5.19, B).

27. Basal process of sacculus: straight or nearly so or with inner margin slightly concave

= 0 (Figure 5.8, A); highly curved its inner margin convex = 1 (Figure 5.9, A).

28. Basal process of sacculus: tapering to acute apex = 0 (Figure 5.9, A); parallel

margined nearly to blunt apex = 1 (Figure 5.13, A).

29. Valve: without lobe = 0 ; with lobe = 1 (Figure 5.14, A).

30. Valve with lobe: clearly distinct from sacculus = 0 (Figure 5.18, A); not clearly

distinct from sacculus = 1 (Figure 5.14, A).

31. Distal process of sacculus (claspers of Clarke): absent = 0; present = 1 (Figure 5.15,

A).

Clarke called the most distal saccular process “claspers” while Hodges called this process simply the “distal process.” I adopt Hodges terminology because the use of the term clasper may suggest that it is potentially homologous with the claspers of other

Lepidoptera.

32. Distal process: linear to curved = 0 (Figure 5.15, A); quadrad or branched = 1.

33. Distal process: more or less straight = 0 (Figure 5.10, A); s-shaped = 1 (Figure 5.5,

A); c-shaped = 2 (Figure 5.15, A). [nonadditive].

129

34. If s-shaped or c-shaped apical region of distal process: strongly curved toward

cucullus = 0 (Figure 5.9, B); strongly curved toward juxta = 1 (Figure 5.22, A).

35. Apex of distal process: rounded or pointed = 0 (Figure 5.6, A); branched like antlers

= 1 (Figure 5.20, A).

36. Costal margin of valva: straight or nearly so = 0 (Figure 5.20, A); noticeably concave

= 1 (Figure 5.7, A).

37. Shape of valve: valva broad at base tapering at 1/2 to 1/3 length = 0 (Figure 5.20, A);

broad at base to 5/6 its length (nearly straight) = 1 (Figure 5.10, A); broadest at

middle = 2 (Figure 5.4, A). [nonadditive].

38. Cucullus: rounded = 0 (Figure 5.21); pointed = 1 (Figure 5.13, A).

39. Costal margin of valve: without process = 0 (Figure 5.10, A); with process = 1

(Figure 5.18, A).

40. Costal process of valve: located at least 3/4 of middle of costa to tip of costa = 0

(Figure 5.4, A); located basally or at least to 3/4 of middle of costa = 1 (Figure

5.18, A).

41. Valve hairs: long and slender, only = 0 (Figure 5.20, A); short and stout, only= 1

(Figure 5.4, A); has an area of both long and slender setae and short and stout

setae = 2 . [nonadditive].

42. Valve hairs: diffuse covering valve = 0 (Figure 5.16, A); localized = 1 (Figure 5.7,

A).

43. If valve hairs localized: primarily at cucullus = 0 (Figure 5.7, A); evenly at costa

130

sacculus and/or cucullus = 1 (Figure 5.10, A); centrally located ampulla = 2

(Figure 5.15, A). [nonadditive].

44. Transtilla: membranous = 0; sclerotized = 1 (Figure 5.3).

45. Lateral lobes of transtilla: well developed = 0 (Figure 5.3); weakly developed diffuse

= 1.

46. Vinculum: rounded = 0 (Figure 5.23, C); pointed = 1 (Figure 5. 23, A and B).

47. Vinculum anterodorsal process: membranous not well developed absent = 0; well

developed = 1.

131

Figure 5.3. Genital features of male Depressaria douglasella (Gelechioidea: Elachistidae:

Depressariinae). Regions and structures that labeled are used in this chapter after my interpretation of Hodges (1974) and Clarke (1941b). Illustration is provided to show symmetry and position of features relative to others.

132

133

Figure 5.4. Genital features of Depressaria absynthiella. A. Right Valve; B. Aedeagus.

134

Figure 5.5. Genital features of Depressaria albipunctella. A. Right Valve; B. Aedeagus.

135

Figure 5.6. Genital features of Depressaria angustati. A. Right Valve; B. Distal process enlarged; C. Aedeagus

.

136

Figure 5.7. Genital features of Depressaria atrostrigella. A. Right Valve; B. Aedeagus.

137

Figure 5.8. Genital features of Depressaria badiella. A. Right Valve; B. Distal process enlarged; C. Aedeagus

.

138

Figure 5.9. Genital features of Depressaria cinereocostella. A. Right Valve; B. Distal process enlarged; C. Aedeagus.

139

Figure 5.10. Genital features of Depressaria constancei. A. Right Valve; B. Distal process enlarged; C. Aedeagus.

140

Figure 5.11. Genital features of Depressaria daucella. A. Right Valve; B. Aedeagus.

141

Figure 5.12. Genital features of Depressaria discipunctella. A. Right Valve; B. Aedeagus

142

Figure 5.13. Genital features of Depressaria eleanorae. A. Right Valve; B. Basal process; C. Aedeagus.

143

Figure 5.14. Genital features of Depressaria haydenii. A. Right Valve; B. Aedeagus.

144

Figure 5.15. Genital features of Depressaria multifidae. A. Right Valve; B. Basal process.

145

Figure 5.16. Genital features of Depressaria petronoma. A. Right Valve; B. Basal process; C. Aedeagus.

146

Figure 5.17. Genital features of Depressaria pimpinellae. A. Right Valve; B. Aedeagus.

147

Figure 5.18. Genital features of Depressaria silesiaca. A. Right Valve; B. Aedeagus.

148

Figure 5.19. Genital features of Depressaria ultimella. A. Right Valve; B. Basal process;

C. Aedeagus.

149

Figure 5.20. Genital features of Depressaria weirella. A. Right Valve; B. Basal process;

C. Aedeagus.

150

Figure 5.21. Genital features of Depressaria yakinae. Right valve.

151

Figure 5.22. Genital features of Depressaria libanoditella. A. Right Valve; B. Basal process; C. Aedeagus.

152

Figure 5.23. Variation of vinculi within Depressaria. A. depressana; B. silesiaca; C. weirella.

Phylogenetic Analysis.

I compiled a morphological matrix consisting of 58 terminals (10 outgroup taxa, three Holarctic species, 20 Nearctic species, and 25 Palearctic species) and 47 characters with 108 states based on personal observation, four published characters by Berenbaum and Passoa (1999), and published descriptions by Clarke (1941b), Hannemann (1995), and Hodges (1974) (See Figure 5.2 for a complete list of species included in the analysis) using WinClada. Parsimony searches were implemented in Ratchet to find the shortest trees using the following commands: 100 iterations/rep; 1 tree/iteration; 8 characters to sample. Shortest trees were thoroughly searched in NONA using the commands: max*; ksv*; best. Bremer Support values were calculated in NONA using the command: Sub 5. 153

WinClada was used to generate the strict consensus phylogeny and view character transformations.

RESULTS

The analysis resulted in 24 most parsimonious trees (l= 162, C.I. = 0.37, R.I. =

0.77; Figure 5.24). A strict consensus collapsed four nodes (L= 167, C.I. = 0.35, R.I. =

0.76; Figure 5.24) as a result of rearrangement of species with two clades: the clade that contains the 6 species haydenii, depressana, atrostrigella, chaerophylli, artemisiae, and absynthiella and the clade that contains the 4 species albipunta, olerella, hofmanni, and ululana. Several analyses were preformed as the characters and matrixes were revised when questions of homology assessment arose. Larval host-plant data and distribution of species (Palearctic versus Nearctic) were mapped onto the consensus tree to study the relationship between host-plant families used by species groups of Depressaria and monophyly of species groups relative to location respectively. Figure 5.24 shows the consensus phylogeny and characters evolution for species of Depressaria. Bremer support values were calculated up to five and are given in Figure 5.25 below the nodes.

Overall, Bremer support values were low for all clades of the analysis.

154

Figure 5.24. Consensus phylogeny (L = 167, C.I. = 0.35, R.I. = 0.76) of 24 most parsimonious trees (L = 162, CI = 0.37, RI = 0.77) produced from morphological matrix in Appendix E. Solid black circles represent uncontroverted synapomorphies, white circles represent homoplasies. Number above circles represent characters, number below circles represent character states. Clades with alternative topologies are indicated with brackets and elaborated; A. Six most parsimonious solutions for Clade 1; B. Two most parsimonious solutions for Clade 2; C. Two most parsimonious solutions for Clade 3.

155

156

Figure 5.24. Continued from previous page.

157

Figure 5.24. Continued from previous page.

158

Taxon Descriptions

Abbreviations of Institutions and Private Collections: USNM National Museum of Natural History, Washington, D.C. GSMNP Great Smokey Mountains National Park voucher collection; in part, Columbus, OH. SPIC Steven Passoa Insect Collection, Columbus, OH. SRB Sibyl Rae Bucheli Collection, Columbus, OH.

Psilocorsis reflexella

GSMNP, ♂, TN: Sevier Co., Indian Gap, N35.604, W83.26.298, 9/10 JUN 2002,

UV trap. (OSUC 0165198)

Second segment of labial palpus ambiguous; ocelli ambiguous; antennal pectin

ambiguous; Cu1 and Cu2 of forewing ambiguous; abdomen not flattened dorsoventrally;

uncus present; uncus well developed triangular; socii absent; uncus and socii fused; gnathos broad triangular; ductus bursa meets corpus bursa ambiguous; aedeagus short, c- shaped, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row; dorsal margin of sacculus without lobe or process basally; valve without lobe; distal process of sacculus absent; costal margin of valva noticeably concave; valve broad at base tapering at 1/2 to 1/3 length; cucullus rounded; costal margin of valve without process; valve hairs long, slender; valve hairs diffuse and covering valve; transtilla sclerotized; lateral lobes of transtilla weakly developed diffuse; vinculum rounded; vinculum anterodorsal process well developed.

Himmacea huachucella

USNM, ♂, Madera Canyon, Santa Rita Mts. 5600’, Santa Cruz Co., Ariz., 20

JUN 1963, G. J. Franclemont. (OSUC 0165138)

159

USNM, ♂, Madera Canyon, 4880’, Santa Rita Mts., ariz., August 20, 1959, R.

W. Hodges. (OSUC 0165117)

Second segment of labial palpus smooth scaled; ocelli absent; antennal pectin absent; Cu1 and Cu2 of forewing not stalked; abdomen not flattened dorsoventrally; uncus present;

uncus well developed triangular; socii absent; uncus and socii separate; gnathos broad

triangular; ductus bursa meets corpus bursa ambiguous; aedeagus long, nearly straight,

lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row; dorsal margin of sacculus without lobe or process basally; valve without lobe; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender; valve hairs diffuse and covering valve; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Semioscopis megamicrella

SPIC, ♂, NH: Grafton Co., Hanover, April 15.1977, Coll. J.C. Schultz. (OSUC

0165196)

Second segment of labial palpus with tuft; ocelli ambiguous; antennal pectin absent; Cu1

and Cu2 of forewing not stalked; abdomen not flattened dorsoventrally; uncus absent;

socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa meets corpus

bursa ambiguous; aedeagus short, c-shaped, lacking flange; vesica of aedeagus with

160

cornuti; cornuti scale-like and numerous in patch; dorsal margin of sacculus without lobe

or process basally; valve without lobe; distal process of sacculus present, more or less

straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at middle; cucullus rounded; costal margin of valve without process; valve hairs long, slender; valve hairs diffuse and covering valve; transtilla membranous; lateral lobes of transtilla weakly developed diffuse; vinculum rounded; vinculum anterodorsal process well developed.

Nites maculatella

GSMNP, ♂, TN: Sevier Co., Tremont, West Prong trailhead, N35.48, W83.41.38,

June 9/10 2002, UV light, Brown, Pogue, Garret. (OSUC 0165139)

Second segment of labial palpus with tuft; ocelli absent; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus present; uncus

short, rounded; socii absent; uncus and socii fused; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, twisted, lacking flange; vesica of

aedeagus with cornuti; cornuti scale-like and numerous in patch; dorsal margin of

sacculus without lobe or process basally; distal process of sacculus present, quadrad or

branched; apex of distal process rounded or pointed; costal margin of valva straight or

nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal

margin of valve without process; valve hairs long, slender, localized primarily at cucullus; transtilla sclerotized; lateral lobes of transtilla weakly developed diffuse;

161

vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Apachea barberella

USNM, ♂, Madera Canyon, Santa Rita Mts. 4880’, Santa Cruz Co., Ariz., 2

JUN 1963, G. J. Franclemont. Reared from Ptelea angustifolia (v.)

cognate (Greene). (OSUC 0165137)

Second segment of labial palpus with tuft; ocelli absent; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced; socii absent; uncus and socii separate; gnathos narrow ovoid; ductus bursa meets corpus bursa straight or curved; aedeagus short, twisted, lacking flange; vesica of aedeagus with cornuti; cornuti saw tooth-like around base of vesica; dorsal margin of sacculus without lobe or process basally; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, diffuse and covering valve; transtilla sclerotized; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Bibarrambla allenella

USNM, ♂, Newfield, N.Y., 27.v.1959, R. W. Hodges. (OSUC 0165140)

162

Second segment of labial palpus smooth scaled; ocelli present; antennal pectin present;

Cu1 and Cu2 of forewing stalked; abdomen not flattened dorsoventrally; uncus reduced; socii reduced; uncus and socii fused; gnathos narrow ovoid; ductus bursa meets corpus bursa straight or curved; aedeagus long, s-shaped, lacking flange; vesica of aedeagus with cornuti; cornuti scale-like and numerous in patch; dorsal margin of sacculus without lobe or process basally; valve without lobe; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broad at base tapering at 1/2 to 1/3 length; cucullus rounded; costal margin of valve without process; valve hairs long, slender; valve hairs diffuse and covering valve; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Exaeretia fulvus

Coded from image in Clarke 1941b.

Second segment of labial palpus smooth scaled; ocelli present; antennal pectin present;

Cu1 and Cu2 of forewing stalked; abdomen not flattened dorsoventrally; uncus reduced, short, rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa meets corpus bursa straight or curved; aedeagus long, strongly curved, lacking flange; vesica of aedeagus with cornuti; cornuti scale-like and numerous in patch; dorsal margin of sacculus without lobe or process basally; distal process of sacculus present, quadrad or branched; costal margin of valva straight or nearly so; valve broadest at base

163

to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without

process; valve hairs long, slender, localized primarily at cucullus; transtilla sclerotized;

lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal

process membranous, not well developed or absent.

Exaeretia lutosella

USNM, ♂, Liguria, Capo mele, el lume UV, macchia, 13.vi.61, E. Jäckh.

(OSUC 0165141)

USNM, ♂, Liguria, Capo mele, el lume UV, macchia, 13.vi.61, E. Jäckh.

Second segment of labial palpus smooth scaled; ocelli present; antennal pectin present;

Cu1 and Cu2 of forewing stalked; abdomen not flattened dorsoventrally; uncus reduced, short, rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti scale-like and numerous in patch; dorsal margin of sacculus without lobe or process basally; valve without lobe; distal process of

sacculus present, quadrad or branched; costal margin of valva straight or nearly so; valve

broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve

without process; valve hairs short, stout, localized evenly at costa, sacculus, and/or

cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum

rounded; vinculum anterodorsal process membranous, not well developed or absent.

164

Agonopterix flavicomella

GSMNP, ♂, TN: Sevier Co., Indian Gap, N35.604, W83.26.298, 9/10 JUN 2002,

UV trap. (OSUC 0165142)

GSMNP, ♂, TN: Sevier Co., Indian Gap, N35.604, W83.26.298, 9/10 JUN 2002,

UV trap. (OSUC 0165143)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing stalked; abdomen flattened dorsoventrally; uncus absent; socii present;

uncus and socii separate; gnathos narrow ovoid; ductus bursa meets corpus bursa straight

or curved; aedeagus short, strongly curved, lacking flange; vesica of aedeagus with

cornuti; cornuti scale-like and numerous in patch; dorsal margin of sacculus without lobe

or process basally; distal process of sacculus present, more or less straight; apex of distal

process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs short, stout, localized primarily at cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Agonopterix gelidella

Coded from images in Clarke 1941b.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing stalked; abdomen flattened dorsoventrally; uncus absent; socii present;

uncus and socii separate; gnathos narrow ovoid; ductus bursa meets corpus bursa straight

165

or curved; aedeagus short, strongly curved, lacking flange; vesica of aedeagus with

cornuti; cornuti scale-like and numerous in patch; dorsal margin of sacculus without lobe

or process basally; valve without lobe; distal process of sacculus present, more or less

straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal

margin of valve without process; valve hairs long, slender, diffuse and covering valve;

transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded;

vinculum anterodorsal process membranous, not well developed or absent.

Depressaria atrostrigella

USNM, ♂, 25 mi. S. of Bitter Creek, Sweetwater Co., Wyoming, C.M. Acc.

13019, July 22-31.19??. (OSUC 1065152)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of vesica; dorsal margin of sacculus without lobe or process basally; valve with lobe not clearly distinct from sacculus; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve with process located at least 3/4 of middle of costa to tip of costa; valve hairs long, slender, localized primarily at

166

cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum

rounded; vinculum anterodorsal process well developed.

Depressaria artemisiae

USNM, ♀, Truax, Whitman Co., 5.v.35, J.F.G. Clarke, Reared from Artemisia

dracunculoides. (OSUC 0165149)

USNM, ♂, Hart’s Pass, Washington, Ocanogan Co., Aug.2.1940, J.F.G. Clarke.

(OSUC 0165150)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal process small, hardly distinct, without scalelike projections; valve with lobe not clearly distinct from sacculus; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve with process; costal process of valve located at least 3/4 of middle of costa to tip of costa; valve hairs short, stout, localized primarily at cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process well developed.

167

Depressaria absynthiella

USNM, ♀, Z Vedlis, Hofmann, 17.1.89., BHamfelt Collection. (OUSC

0165101).

USNM, ♂, Banaria, Shaud, Collection Snallen, Museum Leiden, Deprassaria

absynthiella, det. H.S. (OSUC 0165102)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row extending near tip of vesica; dorsal margin of sacculus with lobe or process basally; basal process small, hardly distinct, without scalelike projections; valve with lobe clearly distinct from sacculus; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve with process; costal process of valve located at least 3/4 of middle of costa to tip of costa; valve hairs long, slender, localized primarily at cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process well developed.

Depressaria haydenii

USNM, ♂, Collection OHofmann, D. haydenii, Z., St. Mositz/{illegible}.

(OSUC 0165133)

168

USNM, ♀, Collection OHofmann. (OSUC 0165134)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of vesica; dorsal margin of sacculus without lobe or process basally; valve with lobe clearly distinct from sacculus; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus ambiguous; costal margin of valve with process located at least

3/4 of middle of costa to tip of costa; valve hairs long, slender, localized primarily at cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria chaerophylli

USNM, ♂, Mittelrhein, Umgebung, d. Lorely, Z. Anthr.cerefollum, 15.vii.42, E,

Jäckh. (OSUC 0165108)

USNM, ♀, 2.4.93, Cliocer: tien Golia, Leicthe, Gothea Himch, 23.11.99,

BHamfelt Collection. (OSUC 0165109)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

169

meets corpus bursa straight or curved; aedeagus short, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row

extending near tip of vesica; dorsal margin of sacculus without lobe or process basally;

valve with lobe not clearly distinct from sacculus; distal process of sacculus absent;

costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus ambiguous; costal margin of valve with process; costal process

of valve located at least 3/4 of middle of costa to tip of costa; valve hairs long, slender

localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral lobes of

transtilla well developed; vinculum rounded; vinculum anterodorsal process well

developed.

Depressaria depressana

USNM, ♂, Basses Alpes, Digne, Mt. Courbons a.l., 16.vii.69, E. Jäckh (OSUC

0165110)

USNM, ♀, Piemonte Val Susa, Meana, Collina Mesa, e.l. Las. Gallicum,

22.vi.61, E. Jäckh (OSUC 1165111)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row extending near tip of vesica; dorsal margin of sacculus without lobe or process

170

basally; valve with lobe not clearly distinct from sacculus; distal process of sacculus

absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus rounded; costal margin of valve with process; costal

process of valve located at least 3/4 of middle of costa to tip of costa; valve hairs long,

slender localized primarily at cucullus; transtilla sclerotized; lateral lobes of transtilla

well developed; vinculum pointed; vinculum anterodorsal process membranous, not well

developed or absent.

Depressaria pastinacella

SRB, ♂, OH: Madison Co., Cedar Bog, 12 Jun 2001, Rf. Parsnip. (OSUC

0165197\)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus absent; socii

present; uncus and socii separate; gnathos narrow ovoid; ductus bursa meets corpus bursa

straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with

cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of

vesica; dorsal margin of sacculus with lobe or process basally; basal process elongate,

with scalelike projections at tip; scale-like projections of basal process long; basal

process of sacculus straight or nearly so or with inner margin slightly concave, tapering to acute apex; valve without lobe; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at middle; cucullus rounded; costal margin of valve without process; valve hairs short, stout, localized primarily at cucullus; transtilla

171

sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum

anterodorsal process membranous, not well developed or absent.

Depressaria bupleurella

USNM, ♂, Grumsdould, Eppelsh, 26.12.83, BHamfelt Collection. (OSUC

0165107)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal

process short; basal process with scalelike projections at tip; scale-like projections of

basal process long; basal process of sacculus straight or nearly so or with inner margin

slightly concave, tapering to acute apex; valve without lobe; distal process of sacculus

absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus rounded; costal margin of valve with process; costal

process of valve located at least 3/4 of middle of costa to tip of costa; valve hairs long,

slender, localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral

lobes of transtilla weakly developed diffuse; vinculum rounded; vinculum anterodorsal

process membranous, not well developed or absent.

172

Depressaria pimpinellae

USNM, ♂, Collection OHofmann. (OSUC 01651184)

USNM, ♀, Weser-berge, Bad Eisen, Z. magna, 16.viii.42, E. Jäckh.

(OSUC 0165185)

USNM, ♀, Mittelrhein, Umbebung, d.Loreley, Z. Bupl. falcatum, 16.viii.44, E.

Jäckh. (OSUC 0165186)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal process short, with scalelike projections at tip; scale-like projections of basal process long; basal process of sacculus straight or nearly so or with inner margin slightly concave, tapering to acute apex; valve without lobe; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus rounded; costal margin of valve with process located at least 3/4 of middle of costa to tip of costa; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral lobes of transtilla weakly developed diffuse; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

173

Depressaria libanotidella

USNM, ♂, BHamfelt Collection. (OSUC 0165128)

USNM, ♀, 18.6.1923, Regensburg, Libanotis montana. (OSUC 0165129)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal process short, with scalelike projections at tip; scale-like projections of basal process long; basal process of sacculus straight or nearly so or with inner margin slightly concave, tapering to acute apex; valve without lobe; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus rounded; costal margin of valve with process; costal process of valve located at least 3/4 of middle of costa to tip of costa; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral lobes of transtilla weakly developed diffuse; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria silesiaca

USNM, ♂, Suecia, Norrbolten, Boden, {illegible}, 30.7.65, R. Johansson.

(OSUC 0165189)

174

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal process small, hardly distinct with scalelike projections at tip; scale-like projections of basal process long; basal process of sacculus straight or nearly so or with inner margin slightly concave; basal process of sacculus parallel margined nearly to blunt apex; valve with lobe; valve with lobe not clearly distinct from sacculus; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus pointed; costal margin of valve with process located basally or at least to 3/4 of middle of costa; valve hairs long, slender, localized evenly at

costa, sacculus, and/or cucullus; transtilla sclerotized; lateral lobes of transtilla well

developed; vinculum rounded; vinculum anterodorsal process membranous, not well

developed or absent.

Depressaria badiella

USNM, ♂, Livon Tuch, 11.90, BHamfelt Collection. (OSUC 0165105)

USNM, ♀, Collection OHofmann. (OUSC 0165106)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

175

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal

process elongate; basal process with scalelike projections at tip; scale-like projections of

basal process long; basal process of sacculus straight or nearly so or with inner margin

slightly concave, parallel margined nearly to blunt apex; valve without lobe; distal

process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at

base to 5/6 its length (parallel sided); cucullus ambiguous; costal margin of valve with

process located basally or at least to 3/4 of middle of costa; valve hairs long, slender,

localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes

of transtilla weakly developed diffuse; vinculum pointed; vinculum anterodorsal process

membranous, not well developed or absent.

Depressaria velox

Coded from images in Hannemann 1955.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, lacking flange; vesica of aedeagus with cornuti; cornuti stout and finger-like and either single or few in row, limited to middle of vesica; dorsal margin of sacculus with lobe or process basally;

176

basal process elongate, with scalelike projections at tip; scale-like projections of basal

process long; basal process of sacculus straight or nearly so or with inner margin slightly

concave, parallel margined nearly to blunt apex; valve without lobe; costal margin of

valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided);

cucullus rounded; costal margin of valve with process located basally or at least to 3/4 of

middle of costa; valve hairs long, slender, localized evenly at costa, sacculus, and/or

cucullus; transtilla membranous; lateral lobes of transtilla weakly developed diffuse;

vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria cinereocostella

USNM, ♂, 13 mi. E. of NoPlatte Nebr., Em 14.viii.50, J.F.G. Clarke, Rf Cicuta

maculata. (OSUC 0165157)

USNM, ♀, Lucas, Iowa, Em. 14.viii.50, J.F.G. Clarke, Rf Cicuta maculate

(OSUC 0165158)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal

process elongate, with scalelike projections at tip; scale-like projections of basal process

177

long; basal process of sacculus highly curved with its inner margin convex, parallel

margined nearly to blunt apex; valve without lobe; distal process of sacculus absent;

costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria ultimella

USNM, ♂, Basilicata-PZ, dit. Di Monticchio, 14.vi.70, E. Jäckh. (OSUC

0165190)

USNM, ♀, Dania, Rihhali, Maribo, Straus, 25.8.1920. (OSUC 0165192)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

extending near tip of vesica; dorsal margin of sacculus with lobe or process basally; basal process short, with scalelike projections at tip; scale-like projections of basal process long; basal process of sacculus highly curved with its inner margin convex, tapering to acute apex; valve without lobe; distal process of sacculus absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus

178

rounded; costal margin of valve without process; valve hairs long, slender, localized

primarily at cucullus; transtilla membranous; lateral lobes of transtilla well developed;

vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria rubricella

Coded from images in Hannemann 1955.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal

process elongate, with scalelike projections at tip; scale-like projections of basal process

long; basal process of sacculus highly curved with its inner margin convex. parallel

margined nearly to blunt apex; valve without lobe; distal process of sacculus absent;

costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

179

Depressaria juliella

USNM, ♀, ex Cicuta sp., Gunnison Colo., 2.July.1994, J.F. Gates Clarke.

(OSUC 0165164)

USNM, ♂, ex Cicuta sp., Gunnison Colo., 2.July.1994, J.F. Gates Clarke.

(OSUC 0165165)

USNM, ♀, Pullman, Wn., J.F. Clarke, 21.vii.33, Rf Cicuta occidentalis. (OSUC

0165166)

USNM, ♀, Rock Creek, Ore., E 10.viii (OSUC 0165167

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal

process elongate, with scalelike projections at tip; scale-like projections of basal process

long; basal process of sacculus highly curved with its inner margin convex, parallel

margined nearly to blunt apex; valve without lobe; distal process of sacculus absent;

costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, diffuse and covering valve; transtilla membranous; lateral lobes of transtilla

well developed; vinculum pointed; vinculum anterodorsal process membranous, not well

developed or absent.

180

Depressaria daucella

USNM, ♂, Lawrence Wash., Whatcom Co., Em. 22.vii.50, J.F.G. Clarke, Rf

Oenanthe sarmentosa. (OSUC 0165161)

USNM, ♀, Toad Lake, Wash., Whatcom County, Em. 24.vii.50, J.F.G. Clarke,

Rf Oenanthe sarmentosa. (OSUC 0165162)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal

process elongate, with scalelike projections at tip; scale-like projections of basal process

long; basal process of sacculus highly curved with its inner margin convex, parallel

margined nearly to blunt apex; valve without lobe; distal process of sacculus absent;

costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender; valve hairs diffuse and covering valve; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process

membranous, not well developed or absent.

181

Depressaria eleanorae

USNM, ♂, Ithaca, N.Y. (OSUC 0165163)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti stout and finger-like and either single or few in row,

limited to middle of vesica; dorsal margin of sacculus with lobe or process basally; basal

process elongate; basal process with scalelike projections at tip; scale-like projections of

basal process long; basal process of sacculus highly curved with its inner margin convex,

parallel margined nearly to blunt apex; valve without lobe; distal process of sacculus

absent; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus rounded; costal margin of valve without process; valve

hairs long, slender; valve hairs diffuse and covering valve; transtilla membranous; lateral

lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process

membranous, not well developed or absent.

Depressaria alienella

USNM, ♀, near Baddeck Bridge, Baddeck River, Victoria Co., N. S., August

17, 1967, D.C. Ferguson. (OSUC 0165145)

182

USNM, ♂, Washington:hr, Mt. Stuart, T22N, R14E, ms?c, Kittatas Co., 27

JULY 1981, reared on flower buds of Angelica, larva 11JULY1981, pupa

16JULY1981, along boulder Creek Trail, coll. Bookman. (OSUC USNM,

♂, Madera Canyon, Santa Rita Mts. 4880’, Santa Cruz Co., Ariz., 2 JUN

1963,0165146)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa at right angle; aedeagus long, nearly straight, with single flange;

vesica of aedeagus with cornuti; cornuti scale-like and numerous in patch, near tip of

vesica; dorsal margin of sacculus without lobe or process basally; distal process of

sacculus present; distal process more or less straight; apex of distal process rounded or

pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus rounded; costal margin of valve without process; valve

hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla

sclerotized; lateral lobes of transtilla well developed; vinculum pointed; vinculum

anterodorsal process membranous, not well developed or absent.

Depressaria artimisiella

USNM, ♂, Deer Creek, Provo Canyon, Ut., IX.8.18, Tom Spaulding. (OSUC

0165151)

183

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa at right angle; aedeagus long, nearly straight, with single flange;

vesica of aedeagus with cornuti; cornuti scale-like and numerous in patch, extending near

tip of vesica; dorsal margin of sacculus without lobe or process basally; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel

sided); cucullus rounded; costal margin of valve without process; valve hairs long,

slender localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral

lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process

membranous, not well developed or absent.

Depressaria discipunctella

USNM, ♂, Basilicata-PZ, dit. Di Monticcio, a.l., 16.vi.70, E. Jäckh. (OSUC

0165112)

USNM, ♀, Basilicata-PZ, dit. Di Monticcio, Val Ofamto a.l., 15.vi.70, E. Jäckh.

(OSUC 0165113)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

184

aedeagus with cornuti; cornuti saw tooth like around base of vesica, limited to middle of

vesica; dorsal margin of sacculus without lobe or process basally; valve without lobe;

distal process of sacculus present, linear to curved; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus pointed; costal margin of valve without process; valve

hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla

membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria leucocephala

USNM, ♂, Schuls – Tarasjo, 10.vii.16, Thomanni Rbl., Paratype. (OSUC

0165130)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti saw tooth like around base of vesica, limited to middle of

vesica; dorsal margin of sacculus without lobe or process basally; valve without lobe;

distal process of sacculus present, linear to curved; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus pointed; costal margin of valve without process; valve

hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla

185

membranous; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process well developed.

Depressaria cervicella

Coded from images in Hannemann 1955.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti saw tooth like around base of vesica, limited to middle of

vesica; dorsal margin of sacculus without lobe or process basally; valve without lobe;

distal process of sacculus present, linear to curved; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus pointed; costal margin of valve without process; valve

hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla

membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process well developed.

Depressaria gallicella

Coded from images in Hannemann 1955.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

186

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, lacking flange; vesica of

aedeagus with cornuti; cornuti saw tooth like around base of vesica, limited to middle of

vesica; dorsal margin of sacculus without lobe or process basally; valve without lobe;

distal process of sacculus present, linear to curved; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broad at base tapering at 1/2 to

1/3 length; cucullus pointed; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process well developed.

Depressaria emeritella

USNM, ♀, Nieder-Weser, Oldenbüttel, Z. Tanacetum, 22.vii.45, E. Jäckh,

emeritella Stt., det. E. Jäckh, 1955. (OSUC 0165135)

USNM, ♂, Nieder-Weser, Neubrunchhauser, z., 31.vii.45, 12.vi, 45, E. Jäckh,

emeritella, Stt., det. E. Jäckh.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, c-shaped, with double flange;

vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections covering entire process; scale-

187

like projections of basal process stout; basal process of sacculus straight or nearly so or

with inner margin slightly concave, tapering to acute apex; valve without lobe; distal

process of sacculus present, more or less straight; apex of distal process rounded or

pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its

length (parallel sided); cucullus pointed; costal margin of valve without process; valve

hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla

membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria betina

USNM, ♂, Lyle, Klickitat Co., Wash., 5.vi.46, J.F.G. Clarke. Paratype. (OSUC

0165154)

USNM, ♂, Lyle, Klickitat Co., Wash., 7.vi.46, J.F.G. Clarke. Paratype. (OSUC

0165155)

USNM, ♀, Sawmill Flat, Kittitas Co., Wn., Em. 10.vii.50 J.F.G. Clarke (OSUC

0165156)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with single flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus without lobe or process basally; distal process of sacculus present, more or less straight; apex of distal process

188

rounded or pointed; costal margin of valva noticeably concave; valve broad at base

tapering at 1/2 to 1/3 length; cucullus rounded; costal margin of valve without process;

valve hairs long, slender, localized primarily at cucullus; transtilla membranous; lateral

lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria constancei

USNM, ♂, Yreka, Siskiyou Co., Calif., Em. 28.v.46, J.F.G. Clarke, Rf

Lumatium californicum. Paratype. (OSUC 0165159)

USNM, ♀, California, Hilt, Em. 31.v.70, J.F.G. Clarke, Lumatium californicum.

(OSUC 0165160)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with single flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus without lobe or process basally; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva noticeably concave; valve broad at base tapering at 1/2 to 1/3 length; cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized primarily at cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

189

Depressaria beckmanni

Coded from images in Hannemannn, 1995.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections covering entire process; scale-

like projections of basal process stout; basal process of sacculus straight or nearly so or

with inner margin slightly concave, tapering to acute apex; valve without lobe; distal

process of sacculus present, more or less straight; apex of distal process branched like

antlers; costal margin of valva straight or nearly so; valve broad at base tapering at 1/2 to

1/3 length; cucullus pointed; costal margin of valve without process; valve hairs long, slender, localized centrally located ampulla; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria nemolella

Coded from images in Hannemannn, 1995.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

190

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, nearly straight, with double flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process basally; basal process short, with scalelike projections covering entire process; scale-like projections of basal process stout; basal process of sacculus straight or nearly so or with inner margin slightly concave, tapering to acute apex; valve without lobe; distal process of sacculus present, more or less straight; apex of distal process branched like antlers; costal margin of valva straight or nearly so; valve broad at base tapering at 1/2 to 1/3 length; cucullus pointed; costal margin of valve without process; valve hairs long, slender, localized centrally located ampulla; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria pulcherrimella

USNM, ♂, {illegible} 7.93, BHamfelt Collection. (OSUC 0165186)

USNM, ♀, Odershausen, Wiidungen, Juli 1925. (OSUC 0165187)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, nearly straight, with double flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process basally; basal process short, with scalelike projections, covering entire process; scale-like

191

projections of basal process stout; basal process of sacculus straight or nearly so or with

inner margin slightly concave, tapering to acute apex; valve without lobe; distal process

of sacculus present, more or less straight; apex of distal process branched like antlers; costal margin of valva straight or nearly so; valve broad at base tapering at 1/2 to 1/3 length; cucullus pointed; costal margin of valve without process; valve hairs long, slender, localized centrally located ampulla; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria douglasella

USNM, ♂, Friuli, Italia, Lago di Cavazzo, Mt. Festa, m700 a.l., 27.vi.68, E.

Jäckh. (OSUC 0165114)

USNM, ♀, Weser-Bergland, Bad Eilsen, Z. carota, 3.vii.44, E. Jäckh.

(OSUC 0165115)

USNM, ♀, Basilicata-PZ, dit. Di Monticcio, a.l., 16.vi.70, E. Jäckh. (OSUC

0165116

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections covering entire process; scale-

192

like projections of basal process stout; basal process of sacculus straight or nearly so or

with inner margin slightly concave, tapering to acute apex; valve without lobe; distal

process of sacculus present, more or less straight; apex of distal process branched like

antlers; costal margin of valva straight or nearly so; valve broad at base tapering at 1/2 to

1/3 length; cucullus pointed; costal margin of valve without process; valve hairs long, slender, localized centrally located ampulla; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria weirella

USNM, ♂, Dania, Vang, 21.7.57. (OSUC 0165194)

USNM, ♂, Jungshoved, 1.8.61 NLW. (OSUC 0165195)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections, covering entire process; scale-

like projections of basal process stout; basal process of sacculus straight or nearly so or

with inner margin slightly concave, tapering to acute apex; valve without lobe; distal

process of sacculus present, more or less straight; apex of distal process branched like

antlers; costal margin of valva straight or nearly so; valve broad at base tapering at 1/2 to

193

1/3 length; cucullus pointed; costal margin of valve without process; valve hairs long,

slender, localized centrally located ampulla; transtilla sclerotized; lateral lobes of

transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous,

not well developed or absent.

Depressaria hofmanni

USNM, ♂, BHamfelt Collection. (OSUC 0165131)

USNM, ♀, exol. 14.6.98, Rebemk, Flank, 4.12.99, BHamfelt Collection.

(OSUC 0165132)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, c-shaped, with double flange;

vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections covering entire process; scale-

like projections of basal process stout; basal process of sacculus straight or nearly so or

with inner margin slightly concave, tapering to acute apex; valve without lobe; distal

process of sacculus present, s-shaped with apical region of distal process strongly curved

toward cucullus; apex of distal process rounded or pointed; costal margin of valva

straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus

pointed; costal margin of valve without process; valve hairs both stout and long, localized

evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of

194

transtilla well developed; vinculum rounded; vinculum anterodorsal process

membranous, not well developed or absent.

Depressaria ululana

USNM, ♂, Trentino, Pieve di Ledro, A.ix.1981, E. Jäckh. (OSUC 0165192)

USNM, ♂, Trentino, Pieve di Ledro, A.ix.1981, E. Jäckh. (OSUC 0165193)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections covering entire process; scale-

like projections of basal process stout; basal process of sacculus straight or nearly so or

with inner margin slightly concave, tapering to acute apex; valve without lobe; distal

process of sacculus present, c-shaped with apical region of distal process strongly curved

toward cucullus; apex of distal process rounded or pointed; costal margin of valva

straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus

pointed; costal margin of valve without process; valve hairs both stout and long, localized

evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

195

Depressaria albipunta

USNM, ♂, Liguria, Andora, Val Merula, al lume uv, 19.iv.63, E. Jäckh, (OSUC

0165103)

USNM, ♀, Umgeb.v.Kassel, Kaufunger Wald, Hg-Damphmischlicht, 12.iv.52,

E. Jäckh (OSU 0165104)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections covering entire process; scale-

like projections of basal process stout; basal process of sacculus highly curved with its

inner margin convex, tapering to acute apex; valve without lobe; distal process of

sacculus present, s-shaped with apical region of distal process strongly curved toward juxta; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus pointed; costal margin of valve without process; valve hairs both stout and long, localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

196

Depressaria olerella

USNM, ♂, Depr. olerella, Z. (OSUC 0165124)

USNM, 19/286, {illegible}, potsdm, himuli, 2.2.92, BHamfelt Collection.

(OSUC 0165125)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process elongate, with scalelike projections covering entire process; scale-

like projections of basal process stout; basal process of sacculus highly curved with its

inner margin convex, tapering to acute apex; valve without lobe; distal process of

sacculus present, s-shaped with apical region of distal process strongly curved toward juxta; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus pointed; costal margin of valve without process; valve hairs both stout and long, localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum pointed; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria armata

Coded from images in Hannemann, 1955.

197

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus straight or nearly so or with

inner margin slightly concave, tapering to acute apex; valve without lobe; distal process

of sacculus present, s-shaped with apical region of distal process strongly curved toward cucullus; apex of distal process rounded or pointed; costal margin of valva straight or

nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal

margin of valve without process; valve hairs long, slender, evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria angelicivora

USNM, ♂, Washington, Hart’s Pass, 4300’, Okanogan Co., Em 30.vii.1962,

J.F.G. Clarke. (OSUC 0165147)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

198

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped with apical region of distal process strongly curved toward cucullus; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla weakly developed diffuse; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria angustati

USNM, ♂, Washington, Hurricane Ridge, Olympie Mnts., 5500’, Em.

11.vii.1955, J.F.G. Clarke. Rf. Lomatium angustatum flavum (OSUC

0165148)

USNM. 2♀, no label data

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

199

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped with apical region of distal process strongly curved toward cucullus; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria togata

USNM, ♂, 15 mi. SE. Snowville, Utah, Hway 305, Em. 18.vi.55. (OSUC

0165177)

USNM, ♀, 2 mi. So. Midvale, Ida, Hway 95, Em. 21.vi.55. (OSUC 0165178)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, scalelike projections covering entire process; scale-like

200

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped, apical region of distal process strongly curved toward cucullus; apex of

distal process rounded or pointed; costal margin of valva straight or nearly so; valve

broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve

without process; valve hairs long, slender, localized evenly at costa, sacculus, and/or

cucullus; transtilla membranous; lateral lobes of transtilla well developed; vinculum

rounded; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria whitmani

USNM, ♀, 8.25.04, Oslar, Platte Canyon, Col. (OSUC 0165179)

USNM, ♂, Dabis Creek, Modoc Co., Calif., 16 – 24 JUN 1922, A. W. Lindsey,

Coll. (OSUC 0165180)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped, with apical region of distal process strongly curved toward cucullus;

201

apex of distal process rounded or pointed; costal margin of valva straight or nearly so;

valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of

valve without process; valve hairs long, slender, hairs localized evenly at costa, sacculus,

and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed;

vinculum rounded; vinculum anterodorsal process well developed.

Depressaria schellbachi

USNM, ♂, Washington, 6 mi. NW New Castle, 23 JUN 1965, R. W. Hodges.

(OSUC 0165176)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, strongly curved, with double

flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped with apical region of distal process strongly curved toward cucullus; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus,

202

and/or cucullus; transtilla membranous; lateral lobes of transtilla well developed;

vinculum rounded; vinculum anterodorsal process well developed.

Depressaria multifidae

USNM, ♂, {no collection data}. (OSUC 0165171)

USNM, ♀, Goodman Springs, Blue Mt., 23.vii.35, J.F.G. Clarke, Rf. Lomatiam

grayi, Collection J.F.G. Clarke. (OSUC 0165172)

USNM, ♀, USNM, ♀, Goodman Springs, Blue Mt., Em. 3.viiii.55, #3.

(OSUC 0165173)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, nearly straight, with double flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process basally; basal process short, with scalelike projections covering entire process; scale-like projections of basal process stout; basal process of sacculus highly curved with its inner margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped with apical region of distal process strongly curved toward cucullus; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus,

203

and/or cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process well developed.

Depressaria leptotaeniae

USNM, ♂, Pullman, Wn., J.F. Clarke, x.23.25. (OSUC 0165168)

USNM, ♀, Park Lake, Grant Co., 11.v.35, J.F.G. Clarke, Rf. Leptottaenia

salmoniflora. (OSUC 0165169)

USNM, ♀, Kamiack Butte, 10.vi.35, J.F. Clarke, Rf. Leptotaenia multiflora

Collection J.F.G. Clarke. (OSUC 0165170)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, with double flange;

vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped with apical region of distal process strongly curved toward cucullus; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus,

204

and/or cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

Depressaria yakimae

USNM, ♂, Walla Walla, Wn., 8.vi.31, D.R. Brannon, Collection J. F. G. Clarke.

(OSUC 0165181)

USNM, ♀, Moses lake, Wn., 8 mi. W. on Hiway 10, Em. 21.vi.50, Rf. Pteryxia

terebinthina. (OSUC 0165182)

USNM, ♀, Truax, 4-v, Rf. Lomatium grayi. (OSUC 0165183)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, with double flange;

vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, s-shaped with apical region of distal process strongly curved toward cucullus; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided); cucullus rounded; costal margin of valve without process; valve hairs long, slender, localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral lobes of transtilla weakly developed diffuse;

205

vinculum rounded; vinculum anterodorsal process membranous, not well developed or

absent.

Depressaria moya

Coded from images in Clarke 1941b.

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, with double flange;

vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short, with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided);

cucullus pointed; costal margin of valve without process; valve hairs long, slender,

localized primarily at cucullus; transtilla sclerotized; lateral lobes of transtilla well

developed; vinculum rounded; vinculum anterodorsal process membranous, not well

developed or absent.

206

Depressaria besma

USNM, ♀, Port Lewis Wash., 6 mi. E. Main Gate, 29.v.44, reared from

Hamacium utricula, Paratype. (OSUC 0165153)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus short, nearly straight, with double flange;

vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process

basally; basal process short with scalelike projections covering entire process; scale-like

projections of basal process stout; basal process of sacculus highly curved with its inner

margin convex, tapering to acute apex; valve without lobe; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length (parallel sided);

cucullus pointed; costal margin of valve without process; valve hairs long, slender,

localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral lobes of

transtilla well developed; vinculum rounded; vinculum anterodorsal process

membranous, not well developed or absent.

Depressaria pteryxiphaga

USNM, ♂, Ten Sleep, Wyo., 9.vii.50, J.F.G. Clarke, Rf. Pterixio terebinthin

calcarea. Paratype. (OSUC 0165174)

207

USNM, ♀, 15 mi. SE. Snowville, Utah, Hway 305, Em. 18.vi.55. (OSUC

0165175)

Second segment of labial palpus with tuft; ocelli present; antennal pectin present; Cu1 and

Cu2 of forewing not stalked; abdomen flattened dorsoventrally; uncus reduced, short,

rounded; socii present; uncus and socii separate; gnathos narrow ovoid; ductus bursa

meets corpus bursa straight or curved; aedeagus long, nearly straight, with double flange; vesica of aedeagus lacking cornuti; dorsal margin of sacculus with lobe or process basally; basal process short, with scalelike projections covering entire process; scale-like projections of basal process stout; basal process of sacculus straight or nearly so or with inner margin slightly concave, tapering to acute apex; valve without lobe; distal process of sacculus present, more or less straight; apex of distal process rounded or pointed; costal margin of valva straight or nearly so; valve broadest at base to 5/6 its length

(parallel sided); cucullus pointed; costal margin of valve without process; valve hairs long, slender; valve hairs localized; if valve hairs localized evenly at costa, sacculus, and/or cucullus; transtilla sclerotized; lateral lobes of transtilla well developed; vinculum rounded; vinculum anterodorsal process membranous, not well developed or absent.

208

DISCUSSION

This phylogeny relies heavily on genital characters, especially those of the male for practical reasons. First, species delineations rely heavily on characteristics of the male genitalia, and to date, few other character systems have been generally useful. Second, larvae (which are useful) are less frequently collected than adults and are not well represented in many collections. I tried to include as many Nearctic and Palearctic taxa as possible, limiting the degree to which I could rely on different life stages. Third, I have relied heavily on museum specimens that are sometime unique and it is not desirable to sacrifice that specimen for whole body mounts that require removal of scales, maceration of tissues and mounting of material on slides. Because species descriptions rely on genitalic characters, the genitalia of many specimens are already dissected and mounted.

Psilocorsis and Himmacea are the most basal lineages in the analysis. Depressariinae form a monophyletic clade. Depressaria is not monophyletic with respect to the genera

Agonopterix and Exaeretia. Each forms a monophyletic genus in this analysis and together they form a monophyletic clade (clade c) within Depressaria that is sister to the

Artemisiae Group and the Pastinacella Group (Figure 5.25). This relationship is weakly supported (Bremer value of 1) and may change with the addition of larval characters.

All species groups of Depressaria are monophyletic except for one, the Douglasella

Group (Figure 5.25). There are two main divisions of species groups within Depressaria: the clade which contains Agonopterix + Exaeretia, the Artemisiae Group, and the

Pastinacella Group (clade b), and the clade that contains the Thomaniella Group, the

209

Discipunctella Group, the Betina Group, and the Douglasella Group (clade g). Both are supported by a single character and have a Bremer support value of 1. Clade b is supported by the single character of a rounded vinculum (46, 1) and clade g is supported by the single polymorphic character of having the valve hairs located evenly at costa sacculus and/or cucullus (43, 1). It is possible that the addition of more taxa or more characters would result in a different topology.

210

Figure 5.25. Consensus phylogeny showing evolution of Depressaria species and species groups. Bremer support values for clades below node and clade designations above node.

211

212

Generic Relationships

Hodges (1974) defined Depressariinae as containing Himmacea, Semioscopis,

Nites, Apachea, Bibarrambla, Agonopterix, Exaeretia (Martyrhilda), and Depressaria.

Passoa (1995) and Berenbaum and Passoa (1999) suggested that Himmacea and

Semioscopis were more closely related to genera within the Amphisbatinae than they are to genera within the Depressariinae. Passoa (1995) and Berenbaum and Passoa (1999) find that genera of Depressariinae form a comb with Depressaria falling out in the middle: (Nites (Apachea (Depressaria (Bibarrambla (Exaeretia (Agonopterix))))))

(Figure 5.2). Kaila’s (2004) analysis suggests Semioscopis to be more closely allied with other Depressariinae taxa (Exaeretia, Levipalpus, Depressaria, and Agonopterix) than it is to the Amphisbatinae. This analysis, like Kaila’s (2004), also suggests that Semioscopis may be a basal lineage of Depressariinae, but the results are not conclusive. More thorough taxon sampling of Amphisbatinae would be necessary to make final conclusions. For the sake of this analysis and discussion, Semioscopis will be considered to be within the Depressariinae.

The current analysis and Passoa (1995) and Berenbaum and Passoa (1999) find

Agonopterix to be sister Exaeretia, unlike in Kaila’s (2004) analysis, where Agonopterix

is sister to Depressaria and Exaeretia is basal to that clade, part of an unresolved polytomy with Levipalpus. The results of this analysis are different in nearly all other regards to the findings of Passoa (1995) and Berenbaum and Passoa (1999). In this analysis, Semioscopis, Nites, Apachea, and Bibarrambla form a comb basal to the

Depressaria-Agonopterix-Exaeretia complex (Figure 5.25; discuss below, also).

213

Depressaria is not monophyletic with respect to Agonopterix and Exaeretia in this

analysis and may not represent a natural taxonomic unit. Agonopterix is a medium-large

genus (125 species) with a rather disjunct distribution, with 42 species occur in America

(north of Mexico), one occurs in Central America, 80 occur in the Palearctic Region, and

one occurs in South Africa (Hodges, 1974). There are only 28 described species of

Exaeretia, 11 occur in the Palearctic region, 10 occur in the Nearctic region, one is

Holarctic and six are Neotropical (Hodges, 1974). Historically, there has been a lot of

shuffling of species between Agonopterix, Exaeretia and Depressaria by authors working

on these genera (Clarke, 1941b; Hannemannn, 1995; Hodges, 1974), suggesting a degree

of historical uncertainty about the identity of species. In his 1941 treatment, Clarke

creates the genus Bibarrambla from Semioscopis, Martyrhilda (Exaeretia) from

Depressaria and Agonopterix, Apachea from Depressaria, and Himmacea from

Cryptolechia (eventually elevated to the subfamily status, Cryptolechiinae of the family

Elachistidae (Minet, 1990)). In his 1974 treatment of the Oecophoridae, Hodges

transferred three species of Agonopterix to Exaeretia, as well as creating the new genus

Nites from Depressaria.

In general, it is surprising to find such discrepancies between the generic

phylogeny of Passoa (1995) and Berenbaum and Passoa (1999) and the generic

relationships presented in this phylogeny because several of Passoa’s 1995) characters

were included here. Below, I discuss several discrepancies in turn.

214

Monophyly of Depressaria, the Ingroup

Clade a: Agonopterix-Exaeretia-Depressaria - In this analysis, unlike Passoa (1995) and

Berenbaum and Passoa (1999) and Kaila (2004), Depressaria is not monophyletic with

respect to Agonopterix and Exaeretia. This clade is supported by uncontroverted

synapomorphy of having, in the pupal stage, the prothoracic femur exposed (4, 1), and

the parallelisms of having the socii present (12, 2) and having short, stout hairs on the valve (42, 1).

Evolution of Species Groups of Depressaria

This analysis is very similar in topology to the species group phylogeny presented in Figure 5.1, with one major difference, Agonopterix and Exaeretia in the ingroup, and one minor difference, the paraphyletic Douglasella Group.

I. Clade b: Exaeretia-Agonopterix, the Artemisiae Group, and the Pastinacella Group –

This clade is supported by the single character of a rounded vinculum (46, 1).

II. Clade d: The Artemisiae Group and the Pastinacella Group – This clade is supported by two polymorphic characters: stout, finger-like cornuti present on the vesica of the aedeagus (20, 1) and the valve has a lobe that is not clearly distinct from the sacculus (31,

0).

1. Clade e: The Artemisiae Group – (artemisiae, palousella, absynthiella,

haydenii, depressana, chaerophylli) This group is supported by a single

uncontroverted synapomorphy of having a basal process that lacks scale-like

projections (24, 0), and three parallelisms: the basal process is small and hardly

215

distinct (23, 0), the valve has a small lobe (29, 1), and the vinculum has a well

developed anterodorsal process (47, 1).

2. Clade f: The Pastinacella Group – (pastinacella, daucella, cinereocostella,

juliella, eleanorae, ultimella, rubricella, bupleurella, pimpinellae, libanotidella,

silesiaca, badiella, velox) This group is supported by two characters: the larvae

have pinacula that are pigmented and well developed (2, 1 uncontroverted

synapomorphy) and the dorsal margin of the sacculus is lacking a lobe or process

basally (22, 0).

III. Clade g: The Thomaniella Group, the Discipunctella Group, the Betina Group, and the Douglasella Group – This clade is supported by the single polymorphic character of having the valve hairs located evenly at costa sacculus and/or cucullus (43, 1).

3. Clade h: The Thomaniella Group – (alienella, artimisiella) This species group

is supported by the uncontroverted synapomorphy of the ductus bursa meeting

corpus bursa at right angle (15, 0) and two parallelisms: the aedeagus has a single

flange (18, 1) and the cornuti are stout and finger-like and either single or few in

row (21, 1).

IV. Clade i: The Discipunctella Group, the Douglasella Group, and the Betina Group –

This clade is supported by the character of having the aedeagus strongly curved (17, 0).

4. Clade j: The Discipunctella Group – (discipunctella, leucocephala, cervicella,

gallicella) – This species group is supported by the parallelism of having the

vinculum anterodorsal process well developed (47, 1).

216

V. Clade k: emeritella, The Betina Group + douglasella, weirella, nemolella, beckmanni, pulcherrimella and the Douglasella Group – This clade is supported by the two

uncontroverted synapomorphies of having the aedeagus with a double flange (18, 2) and

the vesica of the aedeagus bare (19, 0), and one parallelism of having the dorsal margin of sacculus with a lobe or process basally (short or long) (22, 0).

VI. Clade l: The Betina Group + douglasella, weirella, nemolella, beckmanni, pulcherrimella and the Douglasella Group – This clade is supported by the parallelism of having the aedeagus strongly curved (17, 0).

VII. Clade m: Betina Group + douglasella, weirella, nemolella, beckmanni, pulcherrimella – This clade is supported by the parallelism of having the valva broadest at the base and tapering at 1/2 to 1/3 length (37, 0).

5. Clade n: The Betina Group – (betina, constancei) – This group is supported by

four parallelisms: the aedeagus has a single flange (18, 1), the dorsal margin of

the sacculus is lacking a lobe or process basally (22, 1), the costal margin of valva

is noticeably concave (36, 1), and the cucullus is rounded (38, 0).

6. Clade o: douglasella, weirella, nemolella, beckmanni, pulcherrimella – The

monophyly of this clade of species is supported by the uncontroverted

synapomorphy of having the apex of distal process branched like antlers (35, 1).

7. Clade p: hofmanni, ululana, albipunta, olerella, armata, angelicivora,

angustati, togata, whitmani, schellbachi, multifidae, leptotaeniae, yakinae, moya,

pteryxiphaga, besma, – The monophyly of this clade of species is supported by

the two uncontroverted synapomorphies of having the distal process s-shaped (33,

217

1) and having valve hairs that are both long and slender and short stout (41, 2) and the parallelisms of having the vinculum rounded (46, 0).

218

Figure 5.26. Consensus phylogeny of Depressaria showing evolution of key morphological features. A. Vesica with cornuti; B. Flange absent; C. Distal process present; D. Uncus reduced (lost in Agonopterix, Exaeretia and D. pastinacella); E.

Gnathos narrow, ovoid; F. Socii present (absent in Nites and Apachea, reduced in

Bibarrambla); G. Pupal coxae exposed (reversed in Agonopterix; image redrawn from

Mosher 1916); H. Distal process lost; I. Basal process with scale-like projections at tip; J.

Single flange; K. Vesica bare; L. Double flange; M. Basal process with scale-like projections covering entire process (except betina-constancei); N. Single flange; O.

Distal process branched.

219

220

Major Morphological Trends

It is plesiomorphic for all the taxa included in this analysis to have the vesica with

cornuti (Figure 5.26, A). A bare vesica evolves only once, in the ancestor to the clade that

contains emeritella + the Betina Group + the remaining Douglasella Group (Figure 5.26,

K). An aedeagus without a flange is also plesiomorphic for all the taxa included in this analysis (Figure 5.26, B). Having an aedeagus with a double flange evolves only once, in the ancestor to the clade that contains emeritella + the Betina Group + the remaining

Douglasella Group (Figure 5.26, L). Having the aedeagus with a single flange evolves

twice within the Depressaria, once in the ancestor of the Thomaniella Group (Figure 5.26,

J) and once in the ancestor of the Betina Group (Figure 5.26, N). The presence of a distal

process (or the “claspers” of Clarke) is also plesiomorphic for the family (Figure 5.26,

C). The loss of the distal process unites the clade that contains the Artemisiae Group and

the Pastinacella Group (Figure 5.26, H). Having the uncus reduced unites Bibarrambla

and the Depressaria - Agonopterix - Exaeretia Complex (Figure 5.26, D). It is lost again

in Agonopterix, Exaeretia and D. pastinacella. The presence of a narrow, ovoid gnathos unites this clade as well (Figure 5.26, E). Having socii present unites the Agonopterix-

Exaeretia-Depressaria Complex (Figure 5.26, F). It is absent in Nites and Apachea,

reduced in Bibarrambla. Exposed pupal coxae (from Passoa 1995 and Berenbaum and

Passoa 1999) unites the Agonopterix-Exaeretia-Depressaria Complex (Figure 5.26, G). It

is reversed in Agonopterix. The presence of a basal process with scale-like projections at

the tip only is a synapomorphy for the Pastinacella Group. The basal process with scale-

like projections covering entire process unites emeritella, the Betina Group and the

221

remaining species for the Douglasella Group (Figure 5.26, M). It basal process is lost in

the Betina Group. The presence a distal process with a branched apex unties beckmanni – weirella (Figure 5.26, O).

222

Figure 5.27. Evolution of host-plant associations for Depressariinae. Orange indicates feeding on woody or herbaceous plants within the Apiaceae, pink indicates feeding on

herbaceous plants within the Apiaceae, blue indicates feeding on herbaceous plants

within the Asteraceae, and grey indicates ambiguity or unknown records.

223

224

Evolution of Host-Plant Associations

There is no clear and obvious pattern for host-plant selection within the

Depressariinae; however, some general comments can be made. Figure 5.27 show the

host-plant association mapped onto the phylogeny through color-coding of branches and

taxon names. Bucheli et al. (2002) demonstrated that for the genus Coleophora, there

was a trend to specialize in habitats and tissue types. Coleophora was faithful to plant

growth forms of particular habitats and tissue types rather than host-plant clades. The

ancestral feeding condition for Coleophora was to feed on seeds and leaves of

herbaceous plant families and the more derived trend was to feed on leaves of woody

plant families. Host fidelity was conserved more in the herbaceous-feeders than in the

woody-feeders. Within the herbaceous-feeders, host fidelity seems to be conserved more

in seed miners than in leaf miners. Once leaf-feeding evolved, there were no derivations

of seed-feeding, and once woody-feeding evolved, there was one to zero derivations of

herbaceous-feeding (depending the topology of the equally most parsimonious trees).

There seems to be no similar trend within the genus Depressaria and closely related

genera.

Woody- versus Herbaceous-Feeding

There are two main divisions within the Agonopterix-Exaeretia-Depressaria

Complex. Clade b, which contains Agonopterix + Exaeretia, the Artemisiae Group, and

the Pastinacella Group, and clade g, which contains the Thomaniella Group, the

Discipunctella Group, the Betina Group, and the Douglasella Group. Many of the

225

outgroup taxa are woody-feeders. Bibarrambla alienella, the most closely related of the outgroups that are not in the ingroup, feeds on trees in the plant family . Species within both Agonopterix and Exaeretia feed on a wide variety of woody and herbaceous plants families. Species of Depressaria are herbaceous-feeders (Figure 5.27). This trend would indicate that tree-feeding is the ancestral condition for the Agonopterix-Exaeretia-

Depressaria Complex and clade b, but not necessarily for clade g.

Furanocoumarin-Feeding

Species of Apachea feed on species of Ptelea, a genus within the furanocoumarin-

producing family . Species of Agonopterix feed on many different plant

families, including the Apiaceae, Rutaceae, and , all of which produce

furanocoumarins as secondary metabolites. Species of Exaeretia for which larval data

exists are not known to feed on plants that produce furanocoumarins. This overall trend

suggests that the Agonopterix-Exaeretia-Depressaria Complex and species of

Depressaria are ancestrally furanocoumarin-feeding, and this association is a rather old

one.

Apiaceae versus Asteraceae Feeding

Species of Depressaria primarily feed on plants in the family Apiaceae, and it is possible that feeding on this plant family is the ancestral condition (Figure 5.27). Within the Artemisiae Group, haydenii, chaerophylli and depressana sometimes form a monophyletic group and are Apiaceae-feeders. Species of the Pastinacella group, except

226

for the unknown or ambiguous records, are all Apiaceae-feeders. The clade that contains

emeritella, The Betina Group + douglasella, weirella, nemolella, beckmanni,

pulcherrimella and the Douglasella Group are also all Apiaceae-feeders, except for

emeritella and olerella. According to these results, there are potentially as many as five

independent derivations to asteraceous-feeding, and as little as four. In only one species

group, the Thomaniella Group, does asteraceous-feeding form a monophyletic clade in all most parsimonious topologies. In some most parsimonious trees, artemisiae and

absynthiella, both asteraceous-feeders, are sister species. Within the non-monophyletic

Douglasella Group, two species, emeritella and olerella, feed on Asteraceae, but in no

topologies are they a monophyletic group, and, according to this matrix, the Douglasella

Group may be polyphyletic, with emeritella potentially representing a separate lineage.

In summary, the Depressariinae were probably tree-feeders and have more

recently switched to feeding on herbaceous plants. It also suggests the possibility that

feeding on plants that produce toxic furanocoumarins was ancestral for the subfamily, but

it is not conclusive. This pattern of host-plant choice also suggests that Depressaria were

ancestrally Apiaceae-feeders and that Asteraceae-feeding is more derived.

227

Figure 5.28. Consensus phylogeny showing distribution of Depressaria species. Pink indicates Nearctic species, blue indicates Palearctic species, and orange indicates species common to both areas.

228

229

Geographic Associations

The majority of Depressaria are Palearctic in distribution (76 species).Twenty-

one species are Nearctic and three species are Holarctic. This analysis included 10

outgroup taxa (both Palearctic and Nearctic species were used when possible), three

Holarctic species, 20 Nearctic species, and 25 Palearctic species. Figure 5.28 show the geographic distribution of Depressaria mapped onto the phylogeny through color-coding of branches and taxon names. There are three monophyletic clades of Depressaria that are Nearctic: The Thomaniella Group, the Betina Group, and the clade of the Douglasella

Group that contains the species of armata through besma. In some equally most parsimonious solutions, atrostrigella is sister the Holarctic species artemisiae.

Depressaria juliella and eleanorae are part of a polytomy with the Holarctic species daucella. In no topologies is cinereocostella part of the juliella-daucella-eleanorae polytomy. Depressaria pastinacella, a Holarctic species, is the basal lineage of that

Pastinacella Group. Palearctic species are, generally speaking, the basal most lineages in the majority of the monophyletic clades of Depressaria. Within the clade that contains the Artemisiae and Pastinacella groups, it is equivocal as to the plesiomorphic distribution, but in some alternate topologies of the equally most parsimonious trees, the

Palearctic species are the basal most lineages, and in some, the Nearctic species are the

basal most lineages. Within the clade that contains the Thomaniella, Discipunctella,

Betina and Douglasella groups, the Nearctic Betina species group is basal. This overall trend suggests Depressaria is plesiomorphically Palearctic in distribution with multiple

230

colonization events into the Nearctic region (resulting in species which are strictly

Nearctic in distribution and species which are Holarctic).

General Remarks

Clarke begins his 1941 Revision of North American Moths of the Family

Oecophoridae… by saying, “In the beginning I had intended only to do a specific revision of the genera Agonopterix and Depressaria. It soon became apparent, however, that it would be necessary to study carefully all the species known from North America together with many from other parts of the world.” Even after Clarke creates four new genera with in the Depressariinae (two from Depressaria), he goes on to conclude:

In a few genera, it is not entirely clear what we are dealing with. For example, in Depressaria there are five distinct species groups, which may actually represent different genera. The leptotaeniae-multifidae group is especially interesting and represents a series of “species” that may represent only simple Mendelian variants of one species.

In fact, as previously stated, Hodges (1974) later creates Nites from Clarke’s fifth species group, and I have demonstrated a very close association of the leptotaeniae-multifidae group.

As Clarke, I had intended only to study the systematics of just the North

American species of Depressaria. Even before reading the introduction to his revision where he states his purpose (which, of course, I had done midway through this analysis and only out of the frustration I felt towards my “ingroup”), I had abandoned the idea and announced that I was driving to the National Museum of Natural History to, “get the rest.”

231

According to this analysis, Depressaria is not monophyletic with respect to

Agonopterix and Exaeretia. It suggests the possibility that there are at least two main lineages within Depressaria, and that not only might Depressaria be polyphyletic, but potentially, Agonopterix and Exaeretia may represent paraphyletic lineages, as well. As

Clarke suggests, this analysis demonstrates that Depressaria may represent possibly two distinct genera, perhaps more, and is therefore, in need of further revision.

Within Depressaria, all species groups form monophyletic lineages except for the Douglasella Group. The Douglasella Group is the most specious group of

Depressaria. It is polyphyletic with respect to the Betina Group, which is nested within it. The clade that is composed of the taxa beckmanni through weirella is more closely related to species of the Betina group than it is to the other species of the Douglasella

Group. Depressaria emeritella is the most basal lineage of this assemblage. Distribution data and morphological evidence supports the idea that the Douglasella Group is potentially four distinct lineages that have been united by authors by plesiomorphic characters. One lineage is represented by a single species, emeritella. The second lineage is the one that is composed of taxa beckmanni through weirella, which are the only taxa to have a distal process that is branched apically. The third lineage is the polytomy that contains species hofmanni through olerella. The last lineage is the monophyletic clade that is composed of armata through besma, which is Nearctic in distribution.

This analysis presents a generic phylogeny of the Depressariinae that is very different in topology than the one presented by Passoa (1995) and Berenbaum and Passoa

(1999). Although the species group topology is similar to the one in Figure 5.1, inclusion

232

of Palearctic species and species groups results in a novel hypothesis for the evolution of

species groups, specifically the Douglasella Group and the Betina Group. I make no

formal changes to these families without further study. A more comprehensive phylogenetic analysis must be conducted with more characters and more thorough taxon sampling of Agonopterix and Exaeretia, including, as Clarke says, those species known from North America together with many from other parts of the world, and a data set incorporating more larval and pupal characters.

These findings, in general, support the findings of Passoa (1995), who suggested

that the Depressariinae were ancestrally associated with coumarin-producing host-plants,

and that the switch to (primarily) Apiaceae-feeding in the Depressaria lead to an adaptive radiation, as outlined by Erlich and Raven’s classic work (1964). This analysis extends

Passoa’s study by constructing a species level phylogeny focusing on North American species of Depressaria, and sheds further light on the most interesting topic, demonstrating that, within the Depressaria, there is a plesiomorphic association with

Apiaceae, but that Asteraceae-feeding is a more labile association.

This analysis represents the most comprehensive phylogeny of Depressaria to date.

These results clarify our understanding of the morphological evolution of Depressaria, host-plant evolution for the Depressariinae and distribution of species groups in the

Nearctic and Palearctic regions.

233

The use of Genital Characters in Phylogenetics

In entomology, genitalia are commonly used by taxonomists to define species due to high intraspecific variation, usually greater than seen in other characters (Eberhard

1985). Combined with support from empirical studies that genitalia are under sexual selection (Arnqvist and Danielsson, 1999; Eberhard. 1993; Eberhard, 2001; Huber, 1999;

Rowe and Arnqvist, 2002) and theoretical studies that characters under sexual selection evolve rapidly Kirkpatrick, 1982; Lande, 1981; West-Eberhard, 1983), it is believed that genitalia evolve too rapidly and divergently to be useful in a phylogenetic analysis

(Eberhard, 2004; Lowe and Arnqvist, 2002). To demonstrate that genitalia are informative in morphological phylogenetics, Song and Bucheli (in preparation) survey only two primary journals, Systematic Entomology and Annals of the Entomological

Society, for morphological analyses that include genitalia in the matrix. In the 89 papers published in these two journals between the years 2000 and 2004, 74 papers (80.9%) found genitalia to be useful in phylogenetic analyses. This data suggests that genitalia are hierarchical in nature and provide useful characters for cladistic analysis.

CONCLUSION

Though not complete, this phylogenetic study is the most comprehensive to date of the Depressaria, a relatively well-known Gelechioid genus. This analysis resulted in

24 equally most parsimonious phylogenies, and the consensus tree is the result of four nodes being collapsed. The results of this analysis suggest that Semioscopis may be a

234

very basal lineage of the Depressariinae. It also shows that Depressaria is not

monophyletic with respect to the genera Agonopterix and Exaeretia (which are each monophyletic and together form a monophyletic clade within Depressaria). The historical species groups of Depressaria are monophyletic except for the Douglasella

Group, which the Betina Group is nested within. The Douglasella Group may represent four distinct lineages that have been united by authors by plesiomorphic characters: the

emeritella clade (which is monotypic in this analysis), the clade that contains the taxa

beckmanni through weirella, the polytomy that contains the species hofmanni through

olerella, and clade that is composed of armata through besma. Investigation of host-plant

preference suggests that woody-feeding and furanocoumarin-feeding may ancestral

conditions for genera in the subfamily Depressariinae. It also suggests that Apiaceous-

feeding may be the ancestral condition for Depressaria and that there have been as many

as five or as little as four derivations of asteraceous-feeding. Investigation of distribution

patterns suggests multiple colonization events from the Palearctic region to the Nearctic

region, or perhaps range compression or extinction of a genus that was primarily

Holarctic in distribution. No formal changes are made to Depressaria without further

investigation of Agonopterix and Exaeretia.

235

CHAPTER 6

NEW SPECIES OF SCYTHRIS HÜBNER 1825 (GELECHIOIDEA: XYLORYCTIDAE:

SCYTHRIDINAE) FROM THE GALÁPAGOS ISLANDS

INTRODUCTION

The Galápagos Islands present a fascinating arena for the observation of evolution, as is well known from the countless examples of microevolution and speciation that have been offered up by biologist since the time of Darwin. It is comforting to me to be able to add the microlepidopteran genus Scythris Hübner 1825

(Gelechioidea: Xyloryctidae: Scythridinae) to this list. While the Galápagos Island

Scythris may never be elevated to the sacred status of the finch, or even the Galápagos

Island Galagete (Gelechioidea: Autostichidae) (Landry, 2002), the present work potentially represents the beginning of a series of treatments for the Galápagos Island

Scythris.

The significance of insects on islands in studies of evolutionary biology is easily seen in the Hawaiian and Caribbean Island Drosophila, the poster-child of insect

236

radiation on island systems. Other less well-known, yet equally titillating, examples of

insect radiations can be seen in the superfamily Gelechioidea. On the Hawaiian Islands,

an endemic genus Hyposmocoma (Cosmopterigidae), with approximately 350 identified

species, has semi-aquatic larvae that construct cases. This case-building behavior is not

documented within the family outside of the Hawaiian archipelago (Rubinoff, in

preparation). On the Galápagos Islands, eight species of Cosmopterigidae (Landry, 2001)

and two species of Blastobasidae have been recorded (one new) (Adamski and Landry,

1997). Galagete, a new genus of Autostichidae, has also been described and represents

the most extensive radiation of Galápagos Island Gelechioidea to date with 11 new species (Landry, 2002).

The genus Scythris belong to the subfamily Scythridinae of the family

Xyloryctidae. Worldwide, this subfamily is composed of 26 genera with over 700 described species (Hodges, 1998), representing potentially only 10% of the world scythridine fauna (Landry, 1991), with most of the species diversity located in arid regions.

The material used in this study was collected Bernard Landry and collaborators from the Galápagos Islands from 1989 until 2004 and is part of an ongoing study with the long term goal of documenting endemic microlepidopteran fauna of the Galápagos

Islands. Over 150 specimens of Scythris have been collected from the Galápagos Islands in that period of time. This project began with an invitation extended to me from Dr.

Jean-François Landry to describe in collaboration with him the slowly accumulating mass of Galápagos Islands Scythris and a visit to the Canadian Nation Collection in Ottawa,

237

Ontario in the dead of winter to sort and identify available material. After a week of

work, we had dissected the bulk of the material and had sorted it into 10 questionably

distinct morphospecies and I was sent home to prepare this manuscript.

MATERIALS AND METHODS

This current work represents a collaborative effort with Dr. Jean-François Landry

of the Canadian National Collection to describe new species of Scythris from the

Galápagos Islands. My portion of the manuscript focuses only on the males of the above mentioned material. Dr. Landry portion focuses on the females. For this document, I have

studied a total of 42 Galápagos Islands Scythris specimens on loan from institutions and private collections (listed below). The majority of the specimens were collected from

1989 until 2004 by B. Landry, L. Roque, and P. Schmitz with the use of a Mercury Vapor

Light (MVL) at night. A variety of habitats on majority of the islands belonging to the

Galápagos Islands were sampled.

Types of Ecuador and were examined by Jean-François Landry.

Species of Scythris from the Galápagos Islands did not match closely any of the type material.

Abbreviations of Institutions and Private Collections: Blan Bernard Landry Collection; Genève, Suisse. CDRS Charles Darwin Research Station; Santa Cruz Island, Galápagos Islands, Ecuador. CNC Canadian National Collection; Ottawa, Ontario, Canada. MHNG Muséum d’Histoire Naturelle; Genève, Suisse.

238

Specimens were sorted into morphospecies based on collection location and wing coloration. Ultimately, genitalic and wing morphology were used to identify species.

Specimens were prepared following the standard methods of (Clarke, 1941) and

(Robinson, 1976). Dissections proceeded and followed closely the methods used by

Landry (1991) developed specifically for study of Scythridinae with some exceptions.

Adult abdomens were wetted with 70% ethanol and then macerated in a solution of 10%

Potassium Hydroxide (KOH). The solution was gently warmed to just under boiling and the material was allowed to clear for 5 to 6 minutes. After maceration, the abdomens were transferred to 70% ethanol to be cleaned and dissected.

The male genitalia were separated from the abdomen but the modified eighth tergites and sternites were left attached to the abdomen. Aedeagi were either left attached to the vinculum/valvae or carefully removed. Prepared abdomen and genitalia were stained with orange G and then transferred to lactic acid (in microcentrifuge tubes) for temporary storage. The female genitalia were prepared following closely Landry (1991), but were stained with orange G and chlorazol black and then transferred to lactic acid (in microcentrifuge tubes) for temporary storage.

Because the genitalia of Scythris are highly dimensional and not easily flattened without distortion of key features, (Landry 1991) the genitalia were not mounted on slides but examined by placing them in ring slides; submerged in lactic acid (Landry

1991) and held in place with vinyl slide props.

239

Adult moths were examined through a stereomicroscope at magnifications of 10x

– 20x. Abdominal and genitalic morphology was examined through a compound

microscope at magnifications of 100x – 450x. Illustration of the abdomen and genitalia in

this manuscript were drawn first as pencil sketches with the aid of a drawing tube

mounted on a compound microscope and magnified at 100x, digitalized by scanning, and

then finalized using Adobe® Illustrator to create scalable vector files and Adobe®

Photoshop CS (Figures 6.1 - 6.20).

TAXONOMY

The Galápagos Island Scythris treated here are small, tan, light brown, or brown

moths frequently with white, creamy white, or tan scales on the body, and legs. The

wings are plain, flecked, streaked or mottled with white, creamy white, tan, light brown,

or brown scales.

The males have the abdominal tergites and sternites sclerotized with the terga

larger than sterna. The eighth tergite and sternite are modified, with T8 usually strongly concave and S8 strongly convex with an elongate process (Fig. 6.1; A, B.).

The genitalia are symmetric. The uncus is reduced relative to gnathos, sclerotized, and V-shaped when seen in dorsal view. The gnathos is wedge-shaped. The uncus and gnathos are strongly associated. The socii are small and hairy to very hairy lobes. The tegumen is narrowly U-shaped when seen in dorsal view. The valvae are ankylosed and to the vinculum. The vinculum has a sclerotized V-shaped region when seen in posterior

240

view. The aedeagus is ankylosed basally to base of vinculum-valvae and curves to the right and then back toward median when seen in dorsal view (Fig. 6.2; A, B).

Key to species of Galápagos Island Scythris

1a. Gnathos with small projection ventrally ...... 2

1b. Gnathos lacking small projection ventrally ……………………………………...…...3

2a. Small projection of gnathos with spines or stout setae ……….Scythris bernardlandryi

2b. Small projection of gnathos lacking spines or stout setae …………..Scythris furculata

3a. Valvae with ventrally recurved process that originates internally from ventral surface

……...……………………………………………………………………………………...4

3b. Valvae without recurved process that originates internally from ventral surface

……………………………………………………………………………………………..5

4a. Valvae transversely expanded ………………………………………Scythris cuneata

4b. Valvae not transversely expanded …………………………………Scythris pistillata

241

5a. Valvae with acutely pointed process just beyond 1/2 its length ……………………...6

5b. Valvae with a rounded process or lobe just beyond 1/2 its length …..………………..7

6a. Apex of aedeagus with anterior region not twisted …………………Scythris darwini

6b. Apex of aedeagus with anterior region twisted ………………………Scythris falcata

7a. Distal region of valvae squarely spatulate ……………………………………………8

7b. Distal region of valve narrowly spatulate ……………………………Scythris sinuosa

8a. Aedeagus L-shaped, arched only once just beyond 1/2 its length ……………………..

…….………………………………………………………………Scythris galapagosensis

8b. Aedeagus J-shaped, arched twice: once just beyond 1/2 its length and again at 5/6 its length …………………………………………………………………….Scythris ancystra

242

Figure 6.1. Generalized male genitalic features of Galápagos Island Scythris

(Gelechioidea: Xyloryctidae: Scythridinae) in lateral view. A. Abdomen and genitalia showing position of modified eighth tergite and sternite and genitalia; B. Structures of male genitalia. T = tergite; S = sternite.

243

244

Figure 6.2. Details of Galápagos Island male genitalia in posterior view. A.

Representation of articulation of major portions of the male genitalia showing

ankylosation of valvae + vinculum + aedeagus; B. Major portions of the male genitalia disarticulated.

245

246

Scythris cuneata, new species

Figures 6.3, 6.4, 6.5

Etymology.

This species is named from the Latin cuneus, which means wedge, referring to the shape

of the apex of the valva.

Specimens examined.

Holotype.

CNC (CNC LEP 00004171), ♂, Ecuador, Galápagos Islands, Isabella, 8.5 km N. Pto

Villamil; 11.III.1989, MVL, leg. B. Landry.

Paratytpe.

CNC (CNC LEP 00004079), ♂, Ecuador, Galápagos Islands, Isabella, 8.5 km N. Pto

Villamil; 11.III.1989, MVL, leg. B. Landry.

Diagnosis. Scythris cuneata can be easily identified by the valvae, which are distinct

from all other species described here and are broadened anteriorly and transversely

broadened posteriorly.

Description. Forewing length 4 mm to 5.5 mm, wing span 1 cm to 1.1 cm. Moth ground

color brown with creamy white and dark-brown scales.

247

Males with head with brown scales extending to approximately 1/4 length of eye,

white medially, brown at base of frons near fronto-clypeal ridge. Neck tufts creamy

white. Pilifers brown. Haustellum base with white scales. Labial palpi brown; first segment predominantly with white, second and third segments with white basally and apically. Antennae brown, extending beyond 1/2 length of forewing.

Dorsum of thorax brown with creamy white scales throughout. Male forewing upper surface ground color brown, mottled with large areas of creamy white and dark brown, flecked with beige scales. Male hindwings upper surface beige basally darkening to brown apically. Fringe of both wings light brown. Legs light brown to brown with creamy white to white scales throughout.

Male abdomen with T8 trapezoid, slightly longer than wide, wider anteriorly and narrowing posteriorly. S8 greatly enlarged, 1/3 as long as abdomen, heavily sclerotized, entirely surrounding genitalia ventrally, with elongate distal process strongly upcurved extended upward over genitalia; anterior region transverse, anterior margin with deep and broad U-shaped cleft to 1/3 of length and forming two broad lobes; distal process constricted mesially, its dorsal surface keel-like when seen in lateral view, apex bulbous.

Male genitalia valvae with anterior half vertical and broad, posterior half narrowed, transversely expanded with ventrally directed recurved process that originates internally from ventral surface and with setose apex. Vinculum almost as wide as it is long, extended posteriorly to about 1/2 length of valvae. Aedeagus c-shaped, arched

downward, dilated at 5/6 apex tapering to very fine, recurved acute point.

248

Similar Species. The forewing pattern of Scythris cuneata superficially resembles that of the following new species: Scythris galapagosensis, Scythris pistillata, Scythris furculata,

Scythris darwini and Scythris bernardlandryi. The aedeagus and sternite 8 of New

Species 1 are most similar to that of species 4.

249

Figure 6.3. Abdominal features of Scythris cuneata. A. Tergite 8 in dorsal view; B.

Sternite 8 in ventral view; C. Sternite 8 in lateral view.

250

Figure 6.4. Genitalic features of Scythris cuneata. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in dorsal view.

251

Figure 6.5. Distribution of Scythris cuneata.

252

Scythris galapagosensis, new species

Figures 6.6, 6.7, 6.8

Etymology.

This species is named after the Islands because it is most abundant of the material and widespread. It is found on eight of the islands.

Specimens examined.

Holotype.

Blan (CNC LEP 00004117), ♂, Ecuador, Galápagos Islands, Santiaga, Cerro Inn,

28.III.1992, leg. B. Landry.

Paratypes.

Blan (CNC LEP 00004100), ♂, Ecuador, Galápagos Islands, Seymour Norte, 29.III.1992,

MVL, leg. B. Landry.

Blan (CNC LEP 00004134), ♂, Ecuador, Galápagos Islands, Española, Punta Suarez,

2.V.1992, MVL, leg. B. Landry.

Blan (CNC LEP 00004130), ♂, Ecuador, Galápagos Islands, Española, Punta Suarez,

2.V.1992, MVL, leg. B. Landry.

Blan (CNC LEP 00004128), ♀, Ecuador, Galápagos Islands, Española, Punta Suarez,

2.V.1992, MVL, leg. B. Landry.

Blan (CNC LEP 00004131), ♂, Ecuador, Galápagos Islands, Española, Punta Suarez,

2.V.1992, MVL, leg. B. Landry.

253

Blan (CNC LEP 00004160), ♂, Ecuador, Galápagos Islands, Santa Fe Tourist Trail,

28.V.1992, MVL, leg. B. Landry.

CDRS (CNC LEP 00004175), ♂, Ecuador, Galápagos Islands, Santa Cruz, C.D.R.S.,

Barranco, MVL, 11.XI.1999, leg. R. Roque.

CNC (CNC LEP 00004068), ♀, Ecuador, Galápagos Islands, Isabella, 8.5 km N. Pto

Villamil; 9.III.1989, MVL, leg. B. Landry.

CNC (CNC LEP 00004069), ♀, Ecuador, Galápagos Islands, Isabella, 8.5 km N. Pto

Villamil; 9.III.1989, MVL, leg. B. Landry.

Diagnosis. Scythris galapagosensis can be identified by the valvae, which are squarely spatulate and the L-shaped aedeagus which is arched only once just beyond 1/2 its length.

Description. Forewing length 3 to 4 mm, wing span 9 mm to 1 cm. Moth ground color pale beige to dark brown with creamy white to white and light brown to dark-brown scales.

Males with head pale beige to light brown. Pilifers pale beige to light brown with white scales throughout. Haustellum base with creamy white to white scales. Labial palpi creamy white to white; second and third segments with pale beige to light brown scales ventrally. Antennae with scape pale beige with white scales ventrally, pectin filaments beige; pedicle beige with white scales ventrally, flagellum beige throughout, extending beyond 3/4 length of forewing.

254

Dorsum of thorax pale beige to light brown. Male Forewing upper surface ground color pale beige to light brown, mottled with large areas of creamy white to white and pale beige to light brown, some flecked with creamy white to tan scales. Male

Hindwing creamy white basally darkening to brown apically. Fringe of both wings pale beige to light brown. Legs pale beige to brown with creamy white to white scales.

Male abdomen with T8 rectangular, slightly longer than wide, broadly cleft along posterior margin. S8 greatly enlarged, 1/3 as long as abdomen with elongate process extending upward over genitalia, anterior region transverse, anterior margin with U- shaped cleft to 1/2 its length and forming two lobes; distal process constricted mesially, its dorsal surface keel-like when seen in lateral view.

Male genitalia valvae with basal half narrow and distal half squarely spatulate, spatulate region densely setose along apico-ventral margin. Vinculum narrower than it is long, extended posteriorly to about 1/4 length of valvae. Aedeagus L-shaped, strongly arched just beyond 1/2 its length, apex two distinct regions: anterior region tapered to an acutely pointed, twisted anteriorly directed apex; caudal region bulbous.

Variation. Adults of Scythris galapagosensis vary between locations from a pale beige form to a light brown form with wing maculation varying from pale beige ground color with mottled with creamy white and light brown areas to light brown ground color mottled with creamy white and dark brown areas with beige scales throughout. The genitalia vary little.

255

Similar Species. The forewing pattern of Scythris galapagosensis superficially resembles that of the following new species: Scythris cuneata, Scythris pistillata, Scythris furculata,

Scythris darwini, and Scythris bernardlandryi. The aedeagus of Scythris galapagosensis is most similar to that of Scythris falcata, Scythris ancystra, and Scythris darwini.

Sternite 8 of Scythris galapagosensis is most similar to that of Scythris ancystra.

256

Figure 6.6. Abdominal features of Scythris galapagosensis. A. Tergite 8 in dorsal view;

B. Sternite 8 in ventral view; C. Sternite 8 in lateral view.

257

Figure 6.7. Genitalic features of Scythris galapagosensis. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in lateral view.

258

Figure 6.8. Distribution of Scythris galapagosensis.

259

Scythris falcata, new species

Figures 6.9, 6.10, 6.11

Etymology.

This species is named from the Latin falcate, which means sickle-shaped, referring to the

sickle-like process on the dorsal edge of the valva.

Specimens examined.

Holotype.

MHNG (CNC LEP 0005935), ♂, Ecuador, Galápagos Islands, Isabella, Alcedo, Iado NE

700m, camp guayabillos, uvl, 16.IV.2002, leg. B. Landry and L. Roque.

Diagnosis. New species 3 can be identified by the valvae, which have an acutely pointed

process just beyond 1/2 their length, and the apex of the aedeagus, which has the anterior

region twisted.

Description. Forewing length 4 mm, wing span 8.5 mm. Moth ground color golden

brown with creamy white to white and tan scales.

Males with head golden brown with tan scales at vertex. Neck tufts tan. Pilifers

golden brown with creamy white scales throughout. Haustellum base with white scales.

Labial palpi creamy white; second and third segments with golden brown scales ventrally. Antennae with scape golden brown with creamy white scales ventrally, pectin

260

filaments tan; pedicle golden brown with creamy white scales ventrally, flagellum golden

brown throughout, extending beyond 3/4 length of forewing.

Dorsum of thorax golden brown flecked with a few tan scales. Male Forewing upper surface ground color golden brown flecked with tan scales. Male Hindwing creamy white basally darkening to golden brown apically. Fringe of both wings golden brown. Legs creamy white with brown scales.

Male abdomen with T8 rectangular, slightly longer than wide, broadly cleft on posterior margin. S8 greatly enlarged, 1/4 as long as abdomen with elongate process slightly extending upward over genitalia, anterior region wider than it is long and strongly convex; anterior region transverse, anterior margin with deep and broad U- shaped cleft to 3/4 its length and forming two lobes; distal process evenly wide along its entire length with apex slightly notched.

Male genitalia with anterior half of valvae vertical and broad, narrowing at 1/2 its length, costal margin with acutely pointed process just beyond 1/2 its length; posterior half spatulate posteriorly, spatulate region densely and evenly setose. Vinculum narrower than it is long, extended posteriorly to about 1/6 length of valvae. Aedeagus L-shaped, weakly arched just beyond 1/2 its length, apex with two distinct regions: anterior region

tapering to an acutely pointed apex, the caudal region bulbous.

Similar Species. The forewing pattern of Scythris falcata superficially resembles that of

Scythris sinuosa. The aedeagus of Scythris falcata is most similar to that of Scythris

261

galapagosensis, Scythris ancystra, and Scythris darwini. Sternite 8 of Scythris falcata is most similar to that of Scythris darwini.

262

Figure 6.9. Abdominal features of Scythris falcata. A. Tergite 8 in dorsal view; B.

Sternite 8 in ventral view; C. Sternite 8 in lateral view.

263

Figure 6.10. Genitalic features of Scythris falcata. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in lateral view.

264

7.11. Distribution of Scythris falcata.

265

Scythris pistillata, new species

Figures 6.12, 6.13, 6.14

Etymology.

This species is named from the Latin pistillum, which is the pestle used with a mortar,

referring to the process on the ventral side of the valva.

Specimens examined.

Holotype.

Blan (CNC LEP 00004166), ♂, Ecuador, Galápagos Islands, Pinta, 16.III.1992, 200 m

elev., MLV, leg. B. Landry.

Paratype.

Blan (CNC LEP 00004165), ♂, Ecuador, Galápagos Islands, Pinta, 16.III.1992, 200 m

elev., MLV, leg. B. Landry.

Diagnosis. Scythris pistillata can be easily identified by the valvae, which are distinct

from all other species described here and are notched ventrally and broadened posteriorly; and by sternite 8, which, when seen in lateral view, has the tip of process

swollen and bulbous with a small, anteriorly directed region.

Description. Adults with forewing length 4 mm, wing span 1 cm. Moth ground color

brown with creamy white to white and dark-brown scales.

266

Males with head with brown scales extending to approximately 1/4 length of eye,

white medially, brown at base of frons near fronto-clypeal ridge. Neck tufts white.

Pilifers light brown. Haustellum base with white to light brown scales. Labial palpi light

brown ground color; first segment predominantly to all white, second and third segments

with white basally, apically and dorsally; second segment with some white scales

dorsally. Antennae with scape brown with white scales at base, pectin filaments light

brown, some tipped with black; pedicle and flagellum brown throughout, extending

beyond ¾ length of forewing.

Dorsum of thorax brown with creamy white scales throughout. Male forewing

upper surface ground color brown, mottled with large areas of white scales. Male

hindwings upper surface light brown basally darkening to brown apically. Fringe of both

wings brown. Legs creamy white to white with brown throughout.

Male abdomen with T8 trapezoid, slightly longer than wide, wider anteriorly and

narrowing posteriorly, very strongly concave. S8 greatly enlarged, 1/3 as long as abdomen, heavily sclerotized, entirely surrounding genitalia ventrally, with elongate distal strongly upcurved process extended upward over genitalia; anterior region transverse, anterior margin with deep and broad U-shaped cleft to 3/4 of length and

forming two broad lobes; distal process constricted mesially with posterior region

enlarged along the dorsal surface when seen in lateral view and tip of process notched,

swollen and bulbous; with a small, anteriorly directed region.

Male genitalia valvae with anterior half vertical, posterior half broadened, ventral margin notched and with ventrally directed recurved process that originates internally

267

from ventral surface and with setose apex. Vinculum almost as wide as it is long, extended posteriorly to about half length of valvae. Aedeagus c-shaped, arched downward, dilated slightly at 5/6 apex tapering to very fine, recurved acute point.

Similar Species. The forewing pattern of Scythris pistillata superficially resembles that of Scythris cuneata, Scythris galapagosensis, Scythris furculata, Scythris darwini and 10.

The aedeagus and sternite 8 of Scythris pistillata are most similar to that of Scythris cuneata.

268

Figure 6.12. Abdominal features of Scythris pistillata. A. Tergite 8 in dorsal view; B.

Sternite 8 in ventral view; C. Sternite 8 in lateral view.

269

Figure 6.13. Genitalic features of Scythris pistillata. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in lateral view.

270

Figure 6.14. Distribution of Scythris pistillata.

271

Scythris ancystra, new species

Figures 6.15, 6.16, 6.17

Etymology.

This species is named from the Greek ankistron, which means fish-hook, referring to the

apex of aedeagus.

Specimens examined.

Holotype.

CNC (CNC LEP 00004065), ♂, Ecuador, Galápagos Islands, Isabella, 2km W. Puerto

Villamil; 5.III.1989, MVL, leg. B. Landry.

Paratype.

MHNG (CNC LEP 00005972), ♂, Ecuador, Galápagos Islands, Isabella, NE slope

Alcendo, near shore, GPS: elev. 9m, S 00o 23.619’ W 90o 59.715’, 29.III.2004,

UVL, leg. B. Landry & P. Schmitz.

Diagnosis. Scythris ancystra can be identified by the unique forewing coloration, which have a golden brown ground color and are streaked medially with creamy white scales; and the valvae, which are squarely spatulate and the J-shaped aedeagus which is arched

twice, once just beyond 1/2 its length and again at 5/6 its length.

272

Description. Forewing length 3 to 4 mm, wing span 9 mm to 1 cm. Moth ground color

light golden brown to dark golden brown with creamy white and light brown to dark-

brown scales.

Males with head light golden brown. Pilifers light brown. Haustellum base with light golden brown scales flecked with a few white scales. Labial palpi light golden

brown. Antennae light golden brown.

Dorsum of thorax golden brown; collar and tegulae light golden brown. Male

Forewing upper surface golden brown ground color streaked medially with creamy white

scales and flecked with tan scales along the anterior margin of streak. Male Hindwing

creamy white basally darkening to golden brown apically. Fringe of both wings light

golden brown. Legs pale beige with brown.

Males abdomen with T8 rectangular, slightly longer than wide, broadly cleft on

posterior margin. S8 greatly enlarged, 1/3 as long as abdomen with elongate process

extending upward over genitalia, anterior region transverse, anterior margin with U-

shaped cleft to 1/2 its length and forming two lobes; distal process constricted mesially,

its dorsal surface keel-like when seen in lateral view.

Male genitalia valvae with distal half narrowly spatulate, spatulate region

densely setose along apico-ventral margin, costal margin with small, rounded process just

beyond 3/4 its length. Vinculum narrower than it is long, extended posteriorly to about

1/3 length of valvae. Aedeagus J-shaped, strongly arched just beyond 1/2 its length and

again at 5/6 its length ending anteriorly; apex with two distinct regions: anterior region

tapered to an acutely pointed, twisted, anteriorly directed apex; caudal region bulbous.

273

Similar Species. The aedeagus of Scythris ancystra is most similar to that of Scythris

galapagosensis, Scythris falcata, and Scythris darwini. Sternite 8 of Scythris ancystra is most similar to that of Scythris galapagosensis.

274

Figure 6.15. Abdominal features of Scythris ancystra. A. Tergite 8 in dorsal view; B.

Sternite 8 in ventral view; C. Sternite 8 in lateral view.

275

Figure 6.16. Genitalic features of Scythris ancystra. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in lateral view.

276

Figure 6.17. Distribution of Scythris ancystra.

277

Scythris sinuosa, new species

Figures 6.18, 6.19, 6.20

Etymology.

This species is named from Latin sinuosa, which means full of bends or winding,

referring to the shape of the aedeagus.

Specimens examined.

Holotype.

MHNG (CNC LEP 00005946), ♂, Ecuador, Galápagos Islands, Isabella,

Alcendo, lado NE 200m, camp arida alta, UVL, 14.IV.2002, leg. B. Landry & L.

Roque.

Paratypes.

MHNG (CNC LEP 00005951), ♂, Ecuador, Galápagos Islands, Isabella, NE slope

Alcendo, GPS: elev. 292m, S 00o 23.829’ W 91o 01.957’, 30.III.2004, UVL, leg.

B. Landry & P. Schmitz.

MHNG (CNC LEP 00005980), ♂, Ecuador, Galápagos Islands, Floreana Scalesias near

Cerro Pajas, GPS: S 01o 17.743’ W 90o 27.111’, 12.IV.2004, UVL, leg. P.

Schmitz.

278

Diagnosis. Scythris sinuosa can be identified by the valvae, which narrowly spatulate, and the apex of the aedeagus, which has the anterior region blunt and the caudal region bulbous.

Description. Forewing length 3.5 mm, wing span 9 mm. Moth ground color golden brown with creamy white and light brown to dark-brown scales.

Males with head light golden brown to golden brown, golden cream to light golden brown along lateral margins of frons from middle of eyes to clypeus. Neck tufts light golden brown to golden brown. Pilifers golden cream to golden brown, or creamy white scales throughout. Haustellum base with golden brown scales. Labial palpi third segment creamy white, second segment and third segment light brown flecked with white scales throughout or primarily located dorsally or dorsally and ventrally. Antennae with scape light golden brown to golden brown with creamy white to light golden brown scales ventrally, pectin filaments golden creamy white; pedicle light golden brown to golden brown with creamy white to light golden brown scales ventrally, flagellum golden brown throughout, extending beyond 3/4 length of forewing.

Dorsum of thorax golden brown. Male Forewing upper surface ground color golden brown flecked with tan scales. Male Hindwing creamy white basally darkening to golden brown apically. Fringe of both wings golden brown. Legs golden tan to light golden brown with brown scales.

279

Males abdomen with T8 rectangular, slightly longer than wide, slightly narrowed

anteriorly, broadly cleft on posterior margin. S8 greatly enlarged, 1/3 as long as abdomen

with elongate process extending upward over genitalia; anterior region transverse,

anterior margin with U-shaped cleft to 1/2 its length and forming two lobes; distal

process constricted mesially, its dorsal surface keel-like when seen in lateral view.

Male genitalia valvae with anterior half broadened and distal half narrowly spatulate, spatulate region densely setose along apico-ventral margin, costal margin with small, rounded process just beyond 3/4 its length. Vinculum narrower than it is long, extended posteriorly to about 1/4 length of valvae. Aedeagus L-shaped, strongly arched just beyond 1/2 its length, apex with two distinct regions: anterior region blunt, slightly twisted; caudal region bulbous.

Similar Species. The forewing pattern of Scythris sinuosa superficially resembles that of

Scythris falcata.

280

Figure 6.18. Abdominal features of Scythris sinuosa. A. Tergite 8 in dorsal view; B.

Sternite 8 in ventral view; C. Sternite 8 in lateral view.

281

Figure 6.19. Genitalic features of Scythris sinuosa. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in lateral view.

282

Figure 6.20. Distribution of Scythris sinuosa.

283

Scythris furculata, new species

Figures 6.21, 6.22, 6.23

Etymology.

This species is named from the Latin furcula, which means small fork, referring to the shape of the apex of the aedeagus.

Specimens examined.

Holotype.

CNC (CNC LEP 00004087), ♂, Ecuador, Galápagos Islands, Isabella, 3km N. Sto

Tómas, Agr. Zone, 8.III.1989, MLV, leg. B. Landry.

Paratypes.

Blan (CNC LEP 00004112), ♂, Ecuador, Galápagos Islands, Seymour Norte, 29.III.1992,

MLV, leg. B. Landry.

Blan (CNC LEP 00004107), ♂, Ecuador, Galápagos Islands, Seymour Norte, 29.III.1992,

MLV, leg. B. Landry.

MHNG (CNC LEP 00005968), ♂, Ecuador, Galápagos Islands, Santa Cruz, CDRS, wall

of Inverts. Lab., GPS: elev. 11m, S 00o 44.478’ W 90o 18.132’, 19.III.2004, UVL,

leg. B. Landry.

CDRS (CNC LEP 00004173), ♂, Ecuador, Galápagos Islands, Santa Cruz, C.D.R.S.,

Barranco, MVL, 7.X.1999, leg. L. Roque.

CDRS (CNC LEP 00004171), ♂, Ecuador, Galápagos Islands, Santa Cruz, C.D.R.S.,

284

Barranco, MVL, 7.X.1999, leg. L. Roque.

CDRS (CNC LEP 00004174), ♂, Ecuador, Galápagos Islands, Santa Cruz, C.D.R.S.,

Barranco, MVL, 11.X.1999, leg. L. Roque.

Blan (CNC LEP 00004144), ♂, Ecuador, Galápagos Islands, Pinta, 15.III.1992, arid

zone, MVL, leg. B. Landry.

Blan (CNC LEP 00004142), ♂, Ecuador, Galápagos Islands, Pinta, 14.III.1992, Plaja

Ibbeston, MVL, leg. B. Landry.

Blan (CNC LEP 00004141), ♂, Ecuador, Galápagos Islands, Pinta, 13.III.1992, Plaja

Ibbeston, MVL, leg. B. Landry.

Blan (CNC LEP 00004140), ♂, Ecuador, Galápagos Islands, Pinta, 13.III.1992, Plaja

Ibbeston, MVL, leg. B. Landry.

Blan (CNC LEP 00004153), ♂, Ecuador, Galápagos Islands, Pinta, 21.III.1992, ± 15m

elev., MVL, leg. B. Landry.

Diagnosis. Scythris furculata can be easily identified by the presence of a hook, bare of any spines, on ventral surface of the gnathos, the aedeagus, which has a claw-shaped apex, and sternite 8, which lacks a distal process.

Description. Forewing length 3 to 4 mm, wing span 7 to 9 mm. Moth ground color light brown to brown with creamy white to white scales.

Males with head light brown to brown with creamy white to white scales at base of frons. Pilifers light brown to brown, or with creamy white scales throughout.

285

Haustellum base with creamy white to white scales. Labial palpi third segment creamy

white, second segment with white scales basally and light brown to brown scales apically or with white scale dorsally and light brown to brown scales ventrally, third segment light brown to brown, or with creamy white to white scales. Antennae light brown to brown or with pedicle with white scales, extending beyond 3/4 length of forewing.

Dorsum of thorax light brown to brown. Male Forewing upper surface ground color light brown to brown flecked with creamy white to white scales, usually localized but rarely solid patches, or nearly to completely lacking any flecks of white scales. Male

Hindwing creamy white basally darkening to light brown to brown apically. Fringe of both wings light brown to brown. Legs creamy white to white with brown scales.

Males abdomen with T8 narrowly rectangular, longer than wide. S8 greatly enlarged, with narrow V-shaped cleft to 1/2 its length and forming two lobes; distal process lacking.

Gnathos of male genitalia with hook on ventral surface. Valvae narrow with distal half spatulate, spatulate region densely setose. Vinculum narrower than it is long, extended posteriorly to about 1/4 length of valvae. Aedeagus L-shaped, arched just beyond 1/2 its length, apex claw-like with two distinct regions, anterior region tapered to pointed apex, caudal region hooked.

Variation. Scythris furculata shows the most within-species variation. Adults vary between locations from a light brown form to a brown form with wing maculation varying from light brown ground color flecked with creamy white scales to brown ground

286

color flecked with white scales. Some forms have very little to no creamy white to white scales on wings and appear to have solid brown wings. The genitalia vary little.

Similar Species. The forewing pattern of Scythris furculata superficially resembles that of Scythris cuneata, Scythris galapagosensis, Scythris pistillata, Scythris darwini, and

Scythris bernardlandryi.

287

Figure 6.21. Abdominal features of Scythris furculata. A. Tergite 8 in dorsal view; B.

Sternite 8 in ventral view.

288

Figure 6.22. Genitalic features of Scythris furculata. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in lateral view.

289

Figure 6.23. Distribution of Scythris furculata.

290

Scythris darwini, new species

Figures 6.24, 6.25, 6.26

Etymology.

This species is named for Charles Darwin.

Specimens examined.

Holotype.

Blan (CNC LEP 00004136), ♂, Ecuador, Galápagos Islands, Española, Bahía

Manzanillo, 25.IV.1992, MVL, leg. B. Landry.

Paratypes.

MHNG (CNC LEP 00005958), ♂, Ecuador, Galápagos Islands, Santa Cruz, low

agriculture zone, GPS: elev. S 00o 42.132’ W 90o 19.156’, 13.III.2004, UVL,

leg. B. Landry & P. Schmitz.

MHNG (CNC LEP 00005976), ♂, Ecuador, Galápagos Islands, Isabella, NE slope

Alcendo, near pega-pega camp, GPS: elev. 483m, S 00 24.029’ W 91o 02.895’,

31.III.2004, UVL, leg. B. Landry & P. Schmitz.

Blan (CNC LEP 00004151), ♂, Ecuador, Galápagos Islands, Pinta, 20.III.1992, ± 15m

elev., MVL, leg. B. Landry.

Blan (CNC LEP 00004150), ♂, Ecuador, Galápagos Islands, Pinta, 20.III.1992, ± 15m

elev., MVL, leg. B. Landry.

Blan (CNC LEP 00004158), ♂, Ecuador, Galápagos Islands, Pinta, 21.III.1992, ± 15m

291

elev., MVL, leg. B. Landry.

CNC (CNC LEP 00004056), ♂, Ecuador, Galápagos Islands, Isabella, 1km W. Puerto

Villamil; 3.III.1989, MVL, leg. B. Landry.

MHNG (CNC LEP 00005959), ♂, Ecuador, Galápagos Islands, Floreana close to

Loberia, GPS: elev. 6m, S 01o 17.002’ W 90o 29.460’, 11.IV.2004, UVL, leg. P.

Schmitz.

Diagnosis. Scythris darwini can be identified by the valvae, which have an acutely pointed process just beyond 1/2 their length, and the apex of the aedeagus, which has the anterior region straight.

Description. Forewing length 4.5 mm, wing span 1 cm. Moth ground color light golden brown to golden brown with creamy white to white and tan scales.

Males with head light golden brown to golden brown at vertex and lightening to creamy white to white scales at clypeus. Neck tufts creamy white to white. Pilifers golden brown with creamy white scales throughout. Haustellum base with white scales. Labial palpi creamy white; second and third segments with golden brown scales ventrally.

Antennae with scape golden brown with creamy white scales ventrally, pectin filaments tan; pedicle and flagellum light golden brown to golden brown throughout, extending beyond 3/4 length of forewing.

Dorsum of thorax light golden brown to golden brown. Male Forewing upper surface ground color light golden brown to golden brown flecked with creamy white to

292

white scales, usually localized but rarely solid patches. Male Hindwing creamy white basally darkening to light golden brown to golden brown apically. Fringe of both wings light golden brown to golden brown. Legs creamy white with brown scales.

Male abdomen with T8 rectangular, slightly longer than wide, broadly cleft on posterior margin. S8 greatly enlarged, 1/4 as long as abdomen with elongate process slightly extending upward over genitalia, anterior region wider than it is long and strongly convex; anterior region transverse, anterior margin with deep and broad U- shaped cleft to 3/4 its length and forming two lobes; distal process evenly wide along its entire length.

Male genitalia with anterior half of valvae vertical and broad, narrowing at 1/2 its length, costal margin with acutely pointed process just beyond 1/2 its length; posterior half spatulate posteriorly, spatulate region densely and evenly setose. Vinculum narrower than it is long, extended posteriorly to about 1/4 length of valvae. Aedeagus L-shaped, arched just beyond 1/2 its length, apex with two distinct regions: anterior region tapering to an acutely pointed apex, the caudal region bulbous.

Similar Species. The forewing pattern of Scythris darwini superficially resembles that of

Scythris cuneata, Scythris galapagosensis, Scythris pistillata, Scythris furculata and

Scythris bernardlandryi. The genitalia and the eighth sternite are most similar to that of

Scythris falcata.

293

Figure 6.24. Abdominal features of Scythris darwini. A. Tergite 8 in dorsal view; B.

Sternite 8 in ventral view; C. Sternite 8 in lateral view.

294

Figure 6.25. Genitalic features of Scythris darwini. A. Aedeagus in lateral view; B.

Genitalia in lateral view; C. Right valve in lateral view.

295

Figure 6.26. Distribution of Scythris darwini.

.

296

Scythris bernardlandryi, new species

Figures 6.27, 6.28, 6.29

Etymology.

This species is named in honor of Bernard Landry, who collected the majority of the

specimens.

Specimens examined.

Holotype.

MHNG (CNC LEP 00005979), ♂, Ecuador, Galápagos Islands, Isabella, NE slope

Alcendo, near pega-pega camp, GPS: elev. 483m, S 00 24.029’ W 91o 02.895’,

31.III.2004, UVL, leg. B. Landry & P. Schmitz.

Diagnosis. Scythris bernardlandryi can be easily identified by the presence of a spined

hook on ventral surface of the gnathos, and the aedeagus, which is short and stout.

Description. Forewing length 3.5 mm, wing span 8 mm. Moth ground color brown with creamy white to white and dark-brown scales.

Males with head brown flecked with whites scales just below base of antennae.

Neck tufts white. Pilifers brown. Haustellum base with white scales. Labial palpi upcurved; first segment white, second segment brown with white basally, apically, and

297

dorsally, third segment brown with white dorsally. Antennae brown, extending beyond

1/2 length of forewing.

Dorsum of thorax brown, color flecked with creamy white scales. Male forewing

upper surface ground color brown, with areas of creamy white and white scales, flecked with white scales. Male hindwings upper surface beige basally darkening to brown apically. Fringe of both wings brown. Legs creamy white to light brown with brown scales throughout.

Male abdomen with T8 rectangular, slightly longer than wide, broadly cleft on posterior margin. S8 greatly enlarged, 1/3 as long as abdomen with elongate process, anterior region transverse, anterior margin with shallowly cleft to 1/4 its length and forming two lobes; distal process broadest at base and tapering to rounded apex.

Male genitalia gnathos with spinose ventral process. Valvae with anterior half of

valvae vertical and broad, narrowing at just beyond 1/2 its length, distal half constricted

and narrowly spatulate, spatulate region densely setose along apico-dorsal margin.

Vinculum narrower than it is long, extended posteriorly to about 1/4 length of valvae.

Aedeagus C-shaped, stout, tapered to an acutely pointed anteriorly directed apex.

Similar Species. The forewing pattern of Scythris bernardlandryi superficially resembles

that of species Scythris cuneata, Scythris galapagosensis, Scythris pistillata, Scythris

furculata, and Scythris darwini.

298

Figure 6.27. Abdominal features of Scythris bernardlandryi. A. Tergite 8 in dorsal view;

B. Sternite 8 in ventral view.

299

Figure 6.28. Genitalic features of Scythris bernardlandryi. A. Aedeagus in lateral view;

B. Genitalia in lateral view; C. Right valve in dorsal view.

300

Figure 6.29. Distribution of Scythris bernardlandryi.

301

DISCUSSION

The nine new species of described here are placed in Scythris (Gelechioidea:

Xyloryctidae: Scythridinae) based on several features of the male genitalia (Landry

1991). The males of the Galápagos Island new species posses the characteristic enlarged

S8, which extends beneath the male genitalia, bilobate uncus, and wedge-shaped gnathos.

The females have the ostium bursae on situated on S8.

Because the systematics of Scythridinae remains poorly understood and many new world taxa remain to be described, I do not attempt to construct a phylogeny for the nine new species. Finding a possible outgroup is a daunting task at this stage due to the number of undescribed species from South America. Also, at this stage, it is possible that several more new species of Scythris from the Galápagos Islands will surface.

However, based on similarity of the genitalia, tergite 8 and sternite 8, I recognize

overall similarities in following species:

Group 1. Scythris cuneata, Scythris pistillata. Aedeagus c-shaped and dilated at 5/6 length; Valvae with a ventrally directed recurved process that originates internally from the ventral surface of the valvae.

Group 2. Scythris galapagosensis, Scythris falcata, Scythris ancystra, Scythris sinuosa,

Scythris darwini. Valvae with a projection or lobe along costal margin; Aedeagus L- shaped of J-shaped with apex with two regions.

302

Group 3. Scythris furculata. Gnathos with hooked process; Aedeagus claw-shaped.

Group 4. Scythris bernardlandryi. Gnathos with spined process; Aedeagus short and stout.

303

CHAPTER 7

SAMPLING TO ASSESS A RE-ESTABLISHED APPALACHIAN FOREST IN OHIO

BASED ON GELECHIOID MOTHS (LEPIDOPTERA: GELECHIOIDEA).

INTRODUCTION

Biological surveys have become increasingly popular. Discover Life in America’s

All Taxon Bio-Inventory (ATBI) and similar efforts are popular with both the scientific community and the public for a number of reasons. They aim to identify all species of organisms inhabiting specific areas, document the locations of organisms, record life histories and critical habitats for recorded organisms and make available a database for professionals and the public (Sharkey, 2000). The GSMNP ATBI represents a grand mission: to document all organisms in the park and create a synoptic database for those species. This massive project requires a massive amount if resources including manpower, money, and time. Aside from merely documenting the organisms present in the park, the ATBI strives to name all undescribed species, as well. This creates another level of complexity: groups like fungi and are barely known; Sharkey

304

estimates that less than 6% of the total invertebrates are recorded. Sharkey project that it

would take a total of 2,225 years to describe the estimated 90,000 unnamed species in the

park. While this and similar efforts are well respected and have received very positive

feedback, such sampling methods are not the normal practice.

One universally understood problem with biological surveys, other than the

intensive and costly nature of projects likes the GSMNP ATBI, is that it is difficult to

know when, if ever, all potential diversity in an area has been sampled (Magurran, 1988;

Muona, 1999; Scharff et al., 2003; Sørenson et al., 2002). Preston’s (1948) classical

example showed for moths at light traps that although species with intermediate numbers of individuals are most prevalent in communities, a few abundant species account for

most of the records while many rare species account for few of the records. Thus, it is

difficult to know when to stop sampling because many rare species may remain

unmeasured. This problem can be severe, even under favorable circumstances, and an

asymptotic approach to total diversity appears to be the best we can expect. Muona

(1999) investigated whether sampling total beetle diversity was possible given various

trapping methods including but not limited to car-net, handpicking, pitfall traps, and

flight-intercept traps. He demonstrated that a variety of trapping methods was needed because the actual number of species in an area is dynamic and effectiveness of traps

depends on the species (biology) of the beetle being trapped. He showed that even

massive trapping schemes were not effective in recovering all species and in particular

rare species that live in isolated and patchy habitats. Muona’s (1999) results have the following implications: a) to trap rare species, optimal search strategies should be

305

devised, b) trapping results cannot be used as proof of absence of species, and c) because

of large numbers of rare species, it is unlikely that trapping saturation curves ever reach

the true maximum.

Sørenson et al. (2002) focused on the recovery of spider diversity using plotless and

plot-based sampling methods. Their study showed that even intensive sampling was

insufficient for recovery of the entire spider fauna in a given area and that, counter-

intuitively, enlarging the sample area decreased, rather than increased, the number of rare

species collected. In the case of plot-based sampling, Sørenson et al. (2002)

recommended that long-term monitoring should focus on a single or few species and use

standardized methods that are absolute and practical within standardized plots to provide

a baseline for surveys.

Considering the above studies, one may conclude that due to the presence of many

rare species in a given area at a given time, one can never anticipate recovering the total expected diversity using standard, cost efficient trapping efforts done by typical survey

teams (Muona, 1999; Preston, 1948; Sørenson et al., 2002). Such a conclusion makes

clear the importance of comparative survey data when we consider estimates of total

biodiversity. But, comparing across surveys introduces complications of its own, such as

unequal effort, unequal methods, and unequal landscape effects of community

composition. To date, one of the few practical recommendations addressing these

problems is to concentrate on target groups in focal localities rather than sampling

broadly and extensively (Sørenson et al., 2002) when exhaustive, ATBI-scale efforts are

not possible.

306

Lepidoptera as Survey Specimens

Lepidoptera have been demonstrated to be important indicators of habitat structure and community health for several diverse forest ecosystems, from Brazil (Brown and

Freitas, 2000) to Canada (Kerr et al., 2001), and Australia (Kitching et al., 2000). Eastern deciduous forests of North America support a high diversity of trees (Greller, 1988), but today are fragmented due to destructive historical land use. Remaining or regenerated forests with a higher diversity of host-plants, older plant communities, or those with fewer disturbances should be expected to support a more diverse community of

Lepidoptera (Summerville et al., 2003a, b; Summerville and Crist, 2003; Usher and

Keiller, 1998). Following the recommendation of Sørenson et al. (2002), perhaps labor- efficient sampling focused on target groups can be used as a means to survey potential diversity, and also to provide standards of comparison with other surveys. Good candidates would be moths of the superfamily Gelechioidea, many of which come readily to light traps and are often closely associated with individual species of plants or narrow forest niches.

For practical, political, and management reasons, researchers measure diversity in local parks, regenerated natural areas, or regions undergoing succession. As such, these areas themselves represent samples of a greater, historical diversity that may not have been measured on site. Thus, initial estimates of total expected diversity must be derived from surveys of different areas. In this study, we survey gelechioid moths in the Wayne

National Forest (Lawrence County, Ohio, USA), representing a large Appalachian area of 307

forest regenerated in less than 100 years from landscape completely denuded for industrial charcoal. We compare results from our traps to three different kinds of more exhaustive inventories. The Great Smoky Mountain National Park ATBI (Wagner and

Scholtens, 2002) is a labor intensive effort representing a snapshot of Appalachian diversity for a 24 hour period in June, as recorded by many expert lepidopterists.

Summerville and Crist’s (2003) study surveyed moths across different regions of Ohio throughout a summer season, and was based on passive traps only, but covered much more time than the GSMNP-ATBI. Finally, the most exhaustive inventory is that of state of Connecticut (DL Wagner, unpubl.). From comparison with these studies, we address two questions: 1) How well do passive surveys of Gelechioidea compare to more labor intensive surveys? 2) How does the regenerated Wayne National Forest compare to other well documented areas with respect to gelechioid diversity?

MATERIALS AND METHODS

Study Sites

Lawrence County, Ohio - Lawrence County (now referred to as LAWCO), the site of this research, is the southernmost county of Ohio, located at the nexus of Ohio, Kentucky and

West Virginia, and is Appalachian in flora and fauna. Central hardwood trees, primarily oak and hickory are dominant in southeastern Ohio. American , red maple and flowering dogwood are common midstory species, and greenbrier, viburnum and spicebush dominate the understory (Sutherland, 2003).

308

Lawrence County was once part of the extensive Eastern Deciduous forests which

were purported to have run unbroken from approximately east of the coastal piedmont to

approximately west of the Mississippi River.

Southern Ohio had no permanent settlements prior to European contact. Tribes such as the Shawnee, Wyandotte, and Delaware lived southeastern Ohio in far-ranging nomadic groups. The barrier of trees, mountains, and native Americans slowed colonists

but by the mid 1700's, traders and explorers frequented the area.

The Northwest Ordinance of 1787 provided for the survey, sale, and development

of the lands north and west of the Ohio River. These ordinances were a major step in

opening Ohio and other lands to these pioneers. Small towns and villages sprung up were

dependant on forests for their way of life. The virgin forests were logged for trees to

make to charcoal to run furnaces to smelt iron ore. Each company had about 3000 acres of land for the charcoal company (Figure 7.1). Forests were challenged by two primary adversaries: clear cutting of forests to harvest trees which were then burned to create charcoal, and forest fires that were a common event around smelting furnaces (Figure

7.2). Deforestation occurred rapidly and only the most resilient species of trees such as

oak and hickory could withstand the repeated fires associated with the coal and iron

industries. The iron and coal charcoal industry began to die down in Lawrence County by

1890, and by 1920, virtually no primeval forest remained uncut in Ohio.

309

Figure 7.1. Map of Lawrence County, Ohio, showing location of historical furnaces.

Redrawn from http://www.oldindustry.org.

In February 1935, the first tract of land in Lawrence County was acquired for the

National Forest System in Ohio. In September 1951, the Wayne Purchase Units officially became the Wayne National Forest, which today forms the core of the hill country of southeastern Ohio, the most heavily forested part of the state. Current ecosystem restoration goals for The Wayne National Forest include: Restore the mixed oak ecosystem to a sustainable level; Use vegetative management techniques to move toward 310

the desired future condition; Control non-native invasive species; Provide a range of ecological conditions to maintain diversity of native plants and animals (Department of

Agriculture, April 4, 2002).

Figure 7.2. Photograph of Oak Ridge Furnace. From http://www.oldindustry.org.

Great Smoky Mountains National Park – The Great Smoky Mountains National Park

(now referred to as GSMNP), on the border of Tennessee and North Carolina, serves as

311

the baseline against which to measure Lawrence County moth diversity. There are five

major forest habitats in the GSMNP. Nearly 80% of the park is comprised of deciduous

forest, which is the most diverse botanically. Above 1370m, Fraser fir and red spruce are

dominant. American beech, yellow and maple are indicators of the highest elevation deciduous forest of park. Eastern hemlock dominates streamsides and moist, shady slopes up to 1200m. Pine and oak dominate relatively dry, exposed slopes and ridges, especially on the west side of the park. Grassy and heath balds are present within all main forest habitats.

Other studies - Summerville and Crist (2003) studied nine locations in Ohio (unglaciated

and glaciated) from May to September, 1999 with passive blacklight traps. Sites were

chosen to span forest biomes and prairies. Wagner’s inventory (DL Wagner, unpubl.) for

Connecticut includes samples of all life stages in most available habitats over five years, plus extensive museum specimens. This represents a cumulative inventory of the entire state.

Collection

LAWCO - The diversity of nocturnal Gelechioidea was estimated by counting number of species present in the Wayne National Forest, Lawrence County, Ohio, as part of an ongoing study monitoring selected insect populations (Sutherland and Hutchinson 2003).

Six blacklight traps were placed in two locations for three years (1995, 1996, and 1997):

Aid Township 38o, 36' N, 82 o 31' W (Sharps Creek/Bluegrass A, B, and C), and Decatur

Township 38 o, 43' N, 82 o, 41' W (Young's Branch A, B, and C). To standardize

312

comparison, only trapped Gelechioidea data from June for the LAWCO study are

compared to the GSMNP Lepidopteran ATBI trap data.

GSMNP - Thirty lepidopterists, 24 volunteers and a team of llamas performed intense

and directed sampling, both plot-based and plotless, including approximately 30 trap

sites, for 24 hours starting at 15:00 hrs on June 9, 2002. Moths were collected and then

processed. A detailed comparison of trapping methods with respect to Gelechioidea

between the LAWCO study and the GSMNP Lepidopteran All Taxon Bio Inventory

(now referred to as ATBI, or jointly as the GSMNP ATBI), is presented in Figure 7.3.

The areas of deciduous hardwood forests were the most heavily sampled during the

ATBI.

LAWCO Study - GSMNP ATBI – Passive Trapping Scheme Labor Intensive Trapping Scheme June trap data for 1995, 1996, and 1997. 24 hour trapping period starting June 9 2002 at 3:00 pm, 24 hour identification session following. 6 trapping sites all within The Wayne National 30 trap sites placed throughout park. Forest. 1 to 2 collectors to retrieve traps. 30 lepidopterists, 24 volunteers and two llama teams. Plot-based sampling. Plot-less and plot-based sampling methods. Blacklight buckets. Blacklight buckets, blacklight sheets, mercury vapor traps, bait traps, pheromone traps, hand searching and picking, and net-collecting. 1 graduate student identifier. 1 professional identifier; 2 grad student assistants. Identified over 3 months. Identified for the majority on site within 24 hours and continued over a period of time.

Figure 7.3. Comparison of collection methods between Lawrence County, Ohio (LAWCO), and the Great Smoky Mountains National Park Lepidopteran All Taxon Bio Inventory sponsored by Discover Life In America (GSMNP ATBI, data extrapolated from Wagner and Scholtens 2002). The LAWCO study represents a passive trapping scheme while the GSMNP ATBI represents a more labor-intensive trapping scheme. 313

Both LAWCO and GSMNP are based on presence data, and frequency data were not collected.

Identification

Gelechioidea were identified using genitalic characters. Male and female abdomens were prepared for identification using a standard 10% potassium hydroxide solution. In some cases, structures were stained with Mercurochrome. Preparations were mounted in euparol on slides. Male structures were prepared using various techniques specifically aimed to preserve particular taxonomic features; females were left undissected and mounted whole. Moths were identified using the following: Adamski and

Brown, 1989; Adamski and Hodges, 1996; Covell, 1984; Forbes, 1923; Hodges, 1974,

1978, 1983, 1985, 1986, 1998, as well as museum study and unpublished work by Jean-

François Landry on Coleophora.

Cumulative Totals for Lawrence County Gelechioidea

Actual and projected species accumulation curves of Lawrence County Gelechioidea for June were generated. The actual accumulation was estimated by plotting total new species added per trap year and the potential diversity curve was produced by repeating the percent new species added until the asymptote. Frequency data were not collected.

314

RESULTS

Gelechioidea Diversity of Lawrence County

Fifty-five species of Gelechioidea were collected at the Lawrence County sites in

June of 1995, 1996, and 1997. Ten species, or 19%, appear to be undescribed. For a summary of genera and species recovered per family, see Figure 7.4. For Lawrence

County, most of the species diversity is within the families Coleophoridae and

Gelechiidae. For a full species list, see Figure 7.5.

Aid and Decatur townships showed no difference in number of species recovered

(Figure 7.6). There was a disproportionate number of species recovered from one trapping location (Figure 7.6).

315

Number of genera per family, number of species per family Gelechioidea LAWCO Gelechioidea GSMNP Study ATBI Species total 52 76 New county records all ca. 55 Undescribed species ca. 10 ca. 34 Amphisbatidae 1,1 0, 0 Coleophoridae 6, 19 2, 2 Cosmopterigidae 2,2 2, 2 Elachistidae 2,2 2, 2 Gelechiidae 11,19 8, 12 Oecophoridae 2,2 4, 4 Unplaced Gelechioidea 7 52

Figure 7.4. Comparison of Gelechioidea species collected during the LAWCO study and

GSMNP Lepidoptera ATBI (data provided by Brian Scholtens). Number of genera and species recovered per study.

316

FAMILY SUBFAMILY GENUS SPECIES Amphisbatidae Amphisbatinae Psilocorsis reflexella Clem. Coleophoridae Blastobasinae (Riley) Blastobasis 1 Blastobasis 2 Blastobasis 3 1 Holcocera 2 Holcocera 3 1 Hypatopa 2 1 Coleophorinae Coleophora tiliaefoliella Clem. Coleophora juglandella McD. Coleophora malivorella Riley Coleophora querciella Clem. Coleophora comptoniella (McD) Coleophora 1 Coleophora 2 Coleophora 3 Cosmopterigidae Cosmopteriginae Cosmopteryx 1 Stagmatomorpha 1 Elachistidae Ethmiinae Ethmia 1 Stenomatinae Antaeotricha schlaegeri (Zell.)

Figure 7.5. List of species recovered from Lawrence County in June. Continued on next page.

317

Figure 7.5. Continued from previous page.

Gelechiidae 1 2 3 4 5 6 7 Dichomeridinae 1 2 Dichomeris georgiella (Wlk.) Dichomeris liguella (Hub.) Dichomeris 1 Gelechiinae 1 2 4 5 6 7 1 obscurella (Cham.) Chionodes formosella-group Chionodes formosella (Murt.) Chionodes 1 Facista 1 Gelechia 1 Gelechia 2 Gelechia 3 Oecophoridae Oecophorinae Decantha 1 Martyringa latipennis (Wlsh.)

318

Aid Township Decatur Township Total = 30 Total = 22 Percent Total = 57% Percent Total = 42% Sharps Creek / Bluegrass A = 5 Young's Branch A = 3 Sharps Creek / Bluegrass B = 18 Young’s Branch B = 9 Sharps Creek / Bluegrass C = 7 Young’s Branch C = 10

Figure 7.6. Number of species for LAWCO study per township and site.

DISCUSSION

Gelechioid adults are generally short-lived and inconspicuous, but larvae are important in most terrestrial ecosystems (Stehr, 1987), sometimes abundant, and are a major source of food for small predators and many parasites. Larvae have a great diversity of feeding habits: scavenging, gall-forming, leaf-mining, seed-mining, leaf- tying, leaf-rolling, stem-boring, flower-boring and case-making, mostly on gymnosperms and angiosperms (Powell et al., 1998). Larvae of each species tend to be specialists or generalists on only a few plant species, making it simple to predict what species of moths should be present in a given locality at a given time of the year if host plants are known.

As a group, Gelechioidea represent a great diversity of forest niches.

Families of nocturnal, tree-feeding Gelechioidea are well represented in traps from Appalachian regions. Gelechioidea diversity of the Great Smoky Mountains 319

National Park is taken to represent the total diversity possible for Gelechioidea for June

in an undisturbed Appalachian forest with maximum collection effort. Because Lawrence

County, OH and particular sites in The Great Smoky Mountains National Park have similar habitats and host-plant species, we find similar species of Gelechioidea in both locations. In the GSMNP, 21% of the diversity comes from the families Gelechiidae and

Oecophoridae. In the LAWCO study, 73% of the total diversity comes from the families

Coleophoridae and Gelechiidae, which may serve as good indicators of recovery.

Coleophoridae (Coleophorinae and Blastobasinae) is a relatively large family with approximately 1418 described species (Hodges, 1998). Coleophorinae are crepuscular and diurnal moths that are collected at blacklight traps but are more commonly encountered flying nearer to host plants (Emmet, 1996; Forbes, 1923; Landry, 1998). In the Eastern deciduous forest, larvae of Coleophorinae feed on leaves of species of Salix,

Malus, Prunus, , Tilia, Quercus, and Juglans. Eastern deciduous forest larvae of

Blastobasinae feed on leaves of Malus, acorns of Quercus, cones Pinus, dead stems of

Salix, litter of Holodiscus (Adamski and Brown, 1989; Hodges, 1998), and unknown hosts potentially include dead and living plant matter (Adamski and Brown, 1989).

Gelechiidae (Gelechiinae and Dichomeridinae) is a large family with more than 4,000 described species (Hodges, 1998). Eastern deciduous forest larvae of Gelechiinae are leaf-tiers and -rollers of Quercus, Ostrya, Fagus, and Acer. Eastern deciduous forest larvae of Dichomeridinae are leaf-tiers of Populus, Tilia, Carya, Quercus, Acer, Betula,

Corylus, Juglans, and recorded from and Pinaceae (Hodges, 1978; Hodges,

1986; Hodges, 1998) (Figure 7.4).

320

Ecological or conservation studies aspire to measure diversity, or perhaps underlying biological factors that produce it, but these measures are necessarily influenced by sampling protocols. Sampling protocols vary in intensity and method of catch, and all those reported here are superficial compared to thorough, long term assessments. For example, DeVries et al. (1997) reports on 40 traps spread over 200 hectars in Amazonian

Ecuador, arranged as five replicate sampling sites in four habitat types, each site supporting an understory trap and a canopy trap, with traps baited for seven days and then left without bait for three weeks (baited traps were cleared daily, unbaited traps were cleared weekly), run from August of 1992 to August of 1993. The entire study was essentially repeated (DeVries and Walla, 1999) about 200km away from August 1993 to

August 1994. None of the sampling studies discussed here claim to represent this sort of effort.

For Gelechioidea, an inventory including all records spanning all seasons, several years, and many locations throughout Connecticut, (D. L. Wagner, personal communication) has recorded 205 species of Gelechioidea. Yet, no short term study can achieve such a total, so the relevant comparison becomes the proportional merits of differing, imperfect sampling schemes. Our results are evaluated with reference to two different measures of what might be recovered in a sampling program in region of

Eastern hardwood forest. First, we compare our results with those of intensive sampling from the Great Smoky Mountains National Park, estimated by a single “snapshot” in

June. We found similar species of Gelechioidea in both Lawrence County, OH and particular sites in The Great Smoky Mountains National Park that have similar overall

321

habitats and host-plant species. In the LAWCO study, we recovered 68% of the total

Gelechioidea diversity of the GSMNP ATBI (Figure 7.7). In the GSMNP, 21% of the

diversity came from Gelechiidae and Oecophoridae. In LAWCO study, 73% of the total

diversity came from Coleophoridae and Gelechiidae. Several families of Gelechioidea are

greatly underrepresented in Lawrence County, including Cosmopterigidae and

Oecophoridae. Nearly half of the total diversity was recovered from each Ohio township

(42% from Decatur and 57% from Aid, see Figure 7.6); however, considerably more

species were recovered from one site (34.6% from Sharps Creek, Bluegrass B in Aid

Township).

Second, a “total Ohio” estimate was produced through trapping methods more like those we used, but over a more extended period and across a greater area. Summerville

and Crist (2003) recorded 93 species of Gelechioidea, 86% of the diversity coming from the families Gelechiidae and Oecophoridae. The LAWCO study recovered 56% of this

diversity (see Figure 7.7). Thus, our sample of diversity, though very much more narrow in time and area, produced an assessment about half of what Summerville and Crist produced in their much more exhaustive coverage.

322

Location, Sampling Method, Time of Total Number Projected Percent LAWCO Study and Area of Study of Number of Diversity versus Gelechioidea Gelechioidea other Areas LAWCO – 52 60 for the month -- Passive, June for 3 yrs. in regenerated of June Appalachian forest.

GSMNP – 76 100 for 24 hour 68% Exhaustive, June for 24 hrs. in “snap period shop” Appalachian region (DLIA ATBI).

“total Ohio” – 93 --- 56% Passive, 1 year in all of Ohio (Summerville and Crist 2003).

Connecticut – 205 300 total 25% Exhaustive, most habitats of Connecticut for 5 years (D. L. Wagner, unpubl.)

Figure 7.7. Comparison of the LAWCO study to the GSMNP ATBI (Wagner and Scholtens 2002), “total Ohio” study (Summerville and Crist 2003) and Connecticut (D. L. Wagner, unpublished) for location, sampling methods, time of study and area of study; total and projected number of Gelechioidea and percent LAWCO diversity versus the other areas.

Extrapolation to total species.

Wagner and Scholtens (2002) estimate that the number of Gelechioidea recorded for the GSMNP ATBI is low, stating that “gelechioids are simply too small and too numerous to be thoroughly inventoried in 48 hours”, and that there were probably about

323

near 100 species of Gelechioidea in the park during the ATBI. Using the percentages of

other groups of Lepidoptera such as bombycids, notodontids and butterflies, Wagner and

Scholtens (2002) were able to extrapolate the total number of Lepidoptera present in the

park during the survey to be somewhere around 2000 species. Nonetheless, our much less intensive sampling compares favorably to the actual records recovered in the GSMNP

ATBI. When Ohio data are compared to each other, the LAWCO data represents 56% of

the total diversity of Summerville and Crist’s (2003) “total Ohio study” (Figure 7.7).

LAWCO samples come from June only and trapping sites within Lawrence County while

Summerville and Crist (2003) collected over 4 months from 9 sites in both glaciated and

unglaciated Ohio deciduous forests.

Assuming that our data represent the same progressive approximation of species

shown by Muona (1999) and others (above), we can estimate that the actual diversity is

higher than our measurements. Species accumulations show that for each year, roughly

half as many additional species were added to the LAWCO total as the year before (32,

12 and 8, respectively). The projected diversity for Lawrence County Gelechioidea for

June reaches 100% in 4 years with a total of 60 species (Figure 7.8). Measured June

diversity of Lawrence County Gelechioidea is at 93% of the projected June diversity

according to our simple estimates. We estimate that four more years of trapping would

recover only four more species of Gelechioidea (Figure 7.8). Even with this modest

correction, our results are surprisingly robust against the background of the enormously

more extensive (and presumably diverse baseline) of the GSMNP. This relationship

represents exactly what Sørenson et al. (2002) predict: that it may be a more efficient use

324

of time and resources to focus on target groups in focal localities rather than extensive sampling.

Figure 7.8. Actual and projected cumulative totals for Lawrence County Gelechioidea recovered using passive, plot-based blacklight collection methods. Actual data shown with a diamond and solid line, projected data shown with a triangle and dashed line.

Assessment of Site Quality

Just as biologists present samples from sites of interest, regenerated habitats present samples from an historical total biodiversity. As a result, the exact problem of asymptotic approach to “total” that we see in our site assessment protocols (above) must

325

obtain also for the regeneration of historical biodiversity itself. Thus, the problem of assessment is two-fold because we must establish the species diversity on site (both recorded and projected) and we must estimate how this relates to a separate estimate of a regional, historical baseline, perhaps one for which there is only an approximate measure.

We present here a provocative approach to this second approximation, based on our understanding of the first. Given that Lawrence County itself represents only a sample of

Appalachian diversity; we can assume that the regenerated forests can be plotted as points on as asymptotic approach to 100% of historical diversity. Assessment of what is

100% becomes a new challenge. Fortunately, we have several appropriate comparators.

If we take a generalized curve to represent an ideal, generated forest through time, we can imagine that Lawrence County is moving along this curve toward a hypothetical value of 100% regeneration (Figure 7.9). If we take the exhaustive historical sampling of

Connecticut as a measure of the total we could hope to find, then we see that Lawrence

County fares poorly at about one quarter regeneration of our index species (25%). It is important to note that Wagner was sampling many habitats at various times of the year while we are only interested in June samples. Of course, no one can actually trap total diversity in an assessment protocol (Muona, 1999); so that it would be more appropriate to compare our trap data to other trap data more similar in habitat. Judging the LAWCO study against the “total Ohio” survey (Summerville and Crist, 2003), we see about half of expected measurable diversity present (56%). Yet, it still might be more appropriate to limit the comparison to moths caught in June in an Appalachian forest, given that this best describes the LAWCO data. Comparing with the GSMNP ATBI data (a 24 hour

326

snapshot of intense sampling), our data fare better than before, at 68% diversity. If this

closet estimate of measurable diversity is used, then our low impact, low intensity samples appear to be very effective.

Figure 7.9. Species accumulations as a proportion of total in regenerated Lawrence

County, Ohio sites.

327

The areas of Lawrence County that were severely deforested for the production of

charcoal and allowed to re-establish under federal protection have regenerated just above

56% total Gelechioidea of the Summerville and Crist (2003) “total Ohio” fauna in less than 100 years (Figure 7.7). Little is known about the dispersal capabilities of most microlepidoptera except for species, which are usually introduced accidentally to an area and then spread from the point of introduction. Unlike many species of butterflies

and larger moths, gelechioid moths are not considered robust animals. Individuals are

short-lived, rather small and generally lack the capacity to fly great distances, as is the

case with many microlepidoptera. Larvae are internal feeders that are unlikely or unable

to leave the host-plant to travel. Potential for re-colonization is for Gelechioidea probably

not as great as it is for more mobile Lepidoptera species (Doak, 2000; Gutierrez et al.,

2001; Holl, 1996; Petit et al., 2001; Tscharntke et al., 2002; Wahlberg et al., 2002),

demonstrated to re-colonize even areas that do not possess all necessary habitat

requirements (Holl, 1996). A possible scenario of re-establishment of Gelechioidea of

Lawrence County is that small patches of forest remained during a period when

substantial industrial deforestation occurred (ETIB of MacArthur and Wilson, 1967).

Perhaps gelechioid moths persisted even in degraded forest patches and re-colonized

regenerated forest. Potential for re-colonization is therefore, dependent on distance between forest patches diversity and within each patch (Brotons et al., 2003; MacArthur and Wilson, 1967). Small patches of forest have been shown to act as stepping-stones for

species in colonization, or if incorporated into new forests, may serve as sources of

species for immediate colonization (Usher and Keiller, 1998).

328

CONCLUSIONS

Muona (1999) showed that massive trapping schemes were not effective in

recovering all species and in particular rare species that live in isolated and patchy

habitats. To trap rare species, optimal search strategies should be devised but because of

large numbers of rare species, it is unlikely that trapping saturation curves ever reach the

true maximum. Sørenson et al. (2002) showed that even intensive sampling was

insufficient to recovery of the entire spider fauna in a given area and recommended that

long-term monitoring should focus on a single or few species and use standardized

methods that are absolute and practical within standardized plots to provide a baseline for

surveys. Considering the above studies, one may conclude that due to the presence of

many rare species in a given area at a given time, one can never anticipate recovering the

total expected diversity using standard, cost efficient trapping efforts done by typical survey teams (Muona, 1999; Preston, 1948; Sørenson et al., 2002). Such a conclusion makes clear the importance of comparative survey data when we consider estimates of total biodiversity.

In this study, surveyed gelechioid moths in the Wayne National Forest are compared to results from different kinds of more exhaustive inventories. From comparison with these studies, we addressed two questions: 1) How well do passive surveys of Gelechioidea compare to more labor intensive surveys? 2) How does the

329

regenerated Wayne National Forest compare to other well documented areas with respect

to gelechioid diversity?

Our answer to the first question is that our passive surveys of Gelechioidea compare favorably to more labor intensive surveys. The Great Smoky Mountain National

Park ATBI (Wagner and Scholtens 2002) is a labor intensive effort representing a

snapshot of Appalachian diversity for a 24 hour period in June. In the LAWCO study, we

recovered 68% of the total Gelechioidea diversity of the GSMNP ATBI. Summerville

and Crist’s (2003) study surveyed of moths across different regions of Ohio throughout a

summer season, and was based on passive traps only, but covered much more time than

the GSMNP-ATBI. The LAWCO study recovered 56% of this diversity.

Our answer to the second question is that the regenerated Wayne National Forest compares favorably to other well documented areas with respect to gelechioid diversity.

Given that Lawrence County itself represents only a sample of Appalachian diversity; we can assume that the regenerated forests can be plotted as points on as asymptotic approach to 100% of historical diversity. If we take a generalized curve to represent an ideal, generated forest through time, we can imagine that Lawrence County is moving along this curve toward a hypothetical value of 100% regeneration. If we take the exhaustive historical sampling of Connecticut as a measure of the total we could hope to find, then we see that Lawrence County fares poorly at about one quarter regeneration of our index species (25%). Judging the LAWCO study against the “total Ohio” survey

(Summerville and Crist, 2003), we see about half of expected measurable diversity present (56%). Finally, comparing our data with the GSMNP ATBI data, we are at 68%

330

diversity. If this closet estimate of measurable diversity is used, then our low impact, low intensity samples appear to be very effective.

331

BIBLIOGRAPHY

Abegaz, B. M., B. T. Ngadjui, G. N. Folefoc, S. Fotso, P. Ambassa, M. Bezabih, E. Dongo, F. Rise and D. Petersen. 2004. Prenylated flavonoids, monoterpenoid furanocoumarins and other constituents from the twigs of Dorstenia elliptica (Moraceae). Phytochemistry 65: 221-226.

Adamski, D., Brown, R. L., 1989. Morphology and Systematics of North American Blastobasidae (Lepidoptera: Gelechioidea). Mississippi Agricultural and Forestry Experimental Station Technical Bulletin. Mississippi Entomological Museum No. 1, 65, 1-70.

Adamski, D., Hodges R. W., 1996. An annotated list of North American Blastobasinae (Lepidoptera: Gelechioidea: Coleophoridae). Proceedings of the Entomological Society of Washington 98, 708-740.

Adamski, D. and B. Landry. 1997. Review of the Blastobasinae (Lepidoptera: Gelechioidea: Coleophoridae) of the Galápagos Islands. Proceedings of the Entomological Society of Washington 99: 348-358.

Arnqvist, G. and I. Danielsson. 1999. Copulatory behavior, genital morphology, and male fertilization success in water striders. Evolution 53: 147-156.

Barrett, M., Donoghue, M. J., and Sober, E., 1991. Against consensus. Systematic Zoology 40, 486-493.

Berenbaum, M. 1983. Coumarins and : a case for coevolution. Evolution 37: 163-179.

Berenbaum, M. R. and S. Passoa. 1999. Generic phylogeny of North American

332

Depressariinae (Lepidoptera: Elachistidae) and hypotheses about coevolution. Annals of the Entomological Society of America 92: 971-986.

Berenbaum, M. R. and A. R. Zangerl. 1991. Acquisition of a Native Hostplants by an Introduced Oligophagous Herbivore. Oikos 62: 153-159.

Berenbaum, M. R. and A. R. Zangerl. 1994. Costs of inducible defense: protein limitation, and detoxification in parsnip webworms. Ecology 75: 2311-2317.

Bradley, J. D. 1972. A check list of British insects. Part 2, Lepidoptera. In G. S. Kloet and W. D. Hincks (eds), A check list of British insects. Part 2, Lepidoptera. Royal Entomological Society of London, London, pp. viii, 153 p.

Bremer, K., 1988. The limits of amino-acid sequence data in angiosperm phylogenetic Reconstruction. Evolution 42, 795-803.

Bremer, K., 1994. Branch support and tree stability. Cladistics 10, 295-304.

Brotons L., Monkkonen M., and Martin J.L. 2003. Are fragments islands? Landscape and density-area relationships in boreal forest birds. American Naturalist 162: 343-357.

Brower, A. V. Z., 1994. Phylogeny of Heliconius butterflies inferred from mitochondrial DNA sequences (Lepidoptera: ). Molecular Phylogenetics and Evolution 3, 159-174.

Brown K.S., and Freitas A.V. L. 2000. Atlantic Forest Butterflies: Indicators for Landscape Conservation. Biotropica 32: 34-956.

Bucheli, S., Landry, J. F., Wenzel, J., 2002. Larval case architecture and implications of host-plant associations for North American Coleophora (Lepidoptera; Coleophoridae). Cladistics 18, 71-93.

Bucheli, S. R. and J. W. Wenzel. 2005. Gelechioidea (Insecta: Lepidoptera)

333

systematics: A reexamination using combined morphology and mitochondrial DNA data. Molecular Phylogenetics and Evolution. 35, 380-394.

Carroll, M., A. Hanlon, T. Hanlon, A. R. Zangerl and M. R. Berenbaum. 1997. Behavioral effects of carotenoid sequestration by the parsnip webworm, Depressaria pastinacella. Journal of Chemical Ecology 23: 2707-2719.

Clarke, J.F.G., 1941. The preparation of slides of the genitalia of Lepidoptera. Bulletin of the Brooklyn Entomological Society 36, 149-161.

Clarke, J. F. G. 1941b. Revision of the North American moths of the family Oecophoridae, with descriptions of new genera and species. Proceedings of the United States National Museum 90: 33-286.

Common, I. F. B., 1970. Lepidoptera (Moths and Butterflies). Melborne University Press, Melborne, pp. 765-866.

Comstock, J. H., 1920. An introduction to entomology. N.Y. Comstock Publishing Co., Ithaca, New York.

Comstock, J. H. and G. W. Herrick. 1940. An introduction to entomology. Comstock Pub. Co., Ithaca, N.Y.

Costa Lima, A. M. d. 1939. Insectos do Brasil. Officinas de Obras do Jornal do Commercio, Rio de Janeiro.

Covell, C. V., 1984. A field guide to the moths of eastern North America. Houghton Mifflin, Boston.

Cummings, M. P., Handley, S. A., Myers, D. S., Reed, D. L., Rokas, A., Winka, K., 2003. Comparing bootstrap and posterior probability values in the four-taxon case. Systematic Biology 52, 477-487.

334

Davis, J. I., 1995. A phylogenetic structure for the monocotyledons, as inferred from chloroplast DNA restriction site variation, and a comparison of measures of clade support. Systematic Botany 20, 503-527.

DeBry, R. W., 2001. Improving interpretation of the decay index for DNA sequence data. Systematic Biology 50, 742-752.

Denno, R. F., McClure, M. S., Ott, J. R., 1995. Interspecific interactions in phytophagous insects: Competition reexamined and resurrected. Annual Review of Entomology 40, 297-331.

Department of Agriculture Forest Service. 2002. Notice of intent to prepare an environmental impact statement. Meyer D.L. (ed), Federal Register. Office of the Federal Register.

Doak P. 2000. Population consequences of restricted dispersal for an insect herbivore in a subdivided habitat. Ecology 81: 1828-1841.

Eberhard, W. G. 1985. Sexual selection and genitalia. Harvard University Press, Massachusetts.

Eberhard, W. G. 1993. Evaluating models of sexual selection: Genitalia as a test case. Am. Nat. 142: 564-571.

Eberhard, W. G. 2001. Species-specific genitalic copulatory courtship in sepsid flies (Diptera, Sepsidae, Microsepsis) and theories of genitalic evolution. Evolution 55: 93-102.

Eberhard, W. G. 2004. Male–female conflict and genitalia: failure to confirm predictions in insects and spiders. Biol. Rev. 79: 121-186.

Emmet A.M. 1996. Coleophoridae. Harley Books, Colchester, Essex, England.

Ehrlich, P. R. and P. H. Raven. 1964. Butterflies and Plants: a study in coevolution. Evolution 18: 586-608.

335

Experimental Station Technical Bulletin (Mississippi Entomological Museum No. 1) 165: 1-70.

Fracker, S. B., 1915. The classification of lepidopterous larvae. University of Illinois, Urbana.

Feeny, P., 1975. Biochemical coevolution between plants and their insect herbivores. University of Texas Press, Austin.

Fetz, R., 1994. Larvalmorphologische Beitrage zumphylogenetischen System der ehemaligen Oecophoridae (Lepidoptera, Gelechioidea). Neue ent. Nachr 33, 1- 273. Translation.

Forbes W.T.M. 1923. Superfamily Gelechioidea. The Lepidoptera of New York and Neighboring States: Primitive Forms, Microlepidoptera, Pyraloids Bombycoids. Cornell University, Agricultural Experiment Station, Ithaca.

Fracker, S. B. 1915. The classification of lepidopterous larvae. University of Illinois, Urbana.

Gaston, K. J., Reavy, D., and Valladares, G. R. (1992). “Intimacy and fidelity: internal and external feeding by the British microlepidoptera.” Ecological Entomology(17), 86-88.

Goloboff, P. A., 1994. NONA reference: documentation for version 1.15.

Greller A.M. 1988. Deciduous Forests. In Barbou M.G. and Billings W.D. (eds), North American Terrestrial Vegetation, pp 287-316. Cambridge University Press, New York, New York; Cambridge, UK.

Gutierrez D., Leon-Cortes J.L., Menendez R., Wilson R.J., Cowley M.J.R., and Thomas C.D. 2001. Metapopulations of four lepidopteran herbivores on a single host plant, Lotus corniculatus. Ecology 82: 1371-1386.

336

Handlirsch, A., and L. P. Bush. 1906. Revision of American Paleozoic insects. Gov't print. off., Washington.

Hannemannn, H.-J. 1995. Small butterflies or Microlepidoptera: IV. Broad-winged moths (Depressariidae).

Hawkins B.A. and Porter E.E. 2003. Does herbivore diversity depend on plant diversity? The case of California butterflies. The American Naturalist 161: 40-49.

Hodges, R. W., 1974. Gelechioidea: Oecophoridae. In: Dominick, R.B., Dominick, T., Ferguson, D.C., Franclemont, J.G., Hodges, R.W., Munroe, E.G. (Eds.), The Moths of America North of Mexico. E. W. Classey and The Wedge Entomological Research Foundation, London.

Hodges, R. W., 1978. Gelechioidea (in part): Cosmopterigidae. In: Dominick, R.B., Dominick, T., Ferguson, D.C., Franclemont, J.G., Hodges, R.W., Munroe, E.G. (Eds.), The Moths of America North of Mexico. E. W. Classey and The Wedge Entomological Research Foundation, London.

Hodges, R. W., 1983. Check list of the Lepidoptera of America north of Mexico including Greenland. E. W. Classey and The Wedge Entomological Research Foundation, London.

Hodges, R. W., 1985. A new species of Dichomeris from Costa-Rica (Lepidoptera, Gelechiidae). Proceedings of the Entomological Society of Washington 87, 456- 459.

Hodges, R. W., 1986. Gelechioidea: Gelechiidae (part), Dichomeridinae. In: Dominick, R.B., Dominick, T., Ferguson, D.C., Franclemont, J.G., Hodges, R.W., Munroe, E.G. (Eds.), The Moths of America North of Mexico. E. W. Classey and The Wedge Entomological Research Foundation, London.

337

Hodges, R. W., 1998. The Gelechioidea. In: Kristensen, N., (Ed), The Handbook of Zoology/Handbuch der Zoologie. Walter de Gruyter, Berlin & New York, pp. 132-158.

Holl, K.D. 1996. The effect of coal surface mine reclamation on diurnal lepidopteran conservation. Journal of Applied Ecology 33: 225-236.

Huber, B. A. 1999. Sexual selection in pholcid spiders (Araneae, Pholcidae): Artful chelicerae and forceful genitalia. J. Arachnology 27: 135-141.

Innocenti, G., A. Piovan, R. Filippini, R. Caniato and E. M. Cappelletti. 1997. Quantitative recovery of furanocoumarins from Psoralea bituminosa. Phytochemical Analysis 8: 84-86.

Janz, N., Nylin, S., 1998. Butterflies and plants: A phylogenetic study. Evolution 52, 486- 502.

Kaila, L., 2004. Phylogeny of the superfamily Gelechioidea (Lepidoptera: Ditrysia): an exemplar approach. Cladistics 20: 303-340.

Kallersjo, M., Albert, V. A., Farris, J. S., 1999. Homoplasy increases phylogenetic structure. Cladistics 15, 91-93.

Kerr J.T., Southwood T.R.E., and Cihlar J. 2001. Remotely sensed habitat diversity predicts species richness and community similarity in Canada. Proceedings of the Entomological Society of Washington 98: 11365-11370.

Kitching R.L., Orr A.G., Thalib L., Mitchell H., Hopkins M.S., and Graham A.W. 2000. Moth assemblages as indicators of environmental quality in remnants of upland Australian rain forest. Journal of Applied Ecology 37: 284-297.

Kirkpatrick, M. 1982. Sexual selection and the evolution of female choice. Evolution 36: 1-12.

Klotts, A. B. 1970. 20. Lepidoptera. In S. L. e. Tuxen (ed) 20. Lepidoptera. Munksgaard,

338

Copenhagen, pp. 359 p. illus. 26 cm.

Kolaczkowski, B., Thornton, J. W., 2004. Performance of maximum parsimony and likelihood phylogenetics when evolution is heterogeneous. Nature 431, 980-984.

Kristensen N.P. and Skalski A.W. 1998. Phylogeny and Paleontology. In: Kristensen N.P. (ed), The Handbook of Zoology/Handbuch der Zoologie. pp 7-25. Walter de Gruyter, Berlin and New York.

Kuznetsov, V.I., Stekol’nikov, A.A., 1979. The systematic position and phylogenetic relationships of the superfamily Coleophoroidea (Lepidoptera: Oecophoridae, Coleophoridae, Ethmiidae) as revealed by the functional morphology of the male genitalia. Entomol. Rev. 57, 91–103. Translation.

Kuznetsov, V.I., Stekol’nikov, A.A., 1984. System and phylogenetic associations of families and superfamilies of gelechioid Lepidoptera of the infraorder Papilionomorpha (Lepidoptera: Copromorphoidea, Elachistoidea, Coleophoroidea, and Gelechioidea) with use of functional morphology of genitalia of males. Trudy Zool. Instituta Akademija Nauk SSSR 112, 3–68.

Kuznetsov, V. I. and A. A. Stekol’nikov. 2001. New approaches to the system of Lepidoptera of World Fauna (on the basis of functional morphology of abdomen). Nauka, St Petersburg. Translation.

Labandeira, C. C., Dilcher, D. L., Davis, D. R., Wagner, D. L., 1994. Ninety-seven million years of angiosperm-insect association: Paleobiological insights into the meaning of coevolution. Proceedings of the National Academy of Sciences of the United States of America 91, 12278-12282.

Lande, R. 1981. Models of speciation by sexual selection on polygenic traits. Proc. Nat. Acad. Sci. USA 78: 3721-3725.

Landry, B. 2001. The Cosmopterigidae (Lepidoptera) of the Galápagos Islands, Ecuador. Revue Suisse De Zoologie 108: 513-539.

339

Landry, B. 2002. Galagete, a new genus of Autostichidae representing the first case of an extensive radiation of endemic Lepidoptera in the Galápagos Islands. Revue Suisse De Zoologie 109: 813-868.

Landry J.-F. 1998. Répartition géographique, plantes nourricières et notes taxonomiques sur 29 espèces de Coleophora (Lepidoptera: Coleophoridae) au Quebec. Fabreries 23: 25-104. Translation.

Landry, J. F. 1991. Systematics of Nearctic Scythrididae (Lepidoptera, Gelechioidea) - Phylogeny and Classification of Supraspecific Taxa, with a Review of Described Species. Memoirs of the Entomological Society of Canada: 3-338.

MacArthur R.H. and Wilson E.O. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, New Jersey; Kent, England, U.K.

Magurran A.E. 1988. Ecological diversity and its measurement. Princeton University Press, Princeton, New Jersey; Kent, England, U.K.

McDunnough, J. H., 1938. Check list of the Lepidoptera of Canada and the United States of America. California, Los Angeles.

Mckenna, D. D. and M. Berenbaum. 2003. A field investigation of Depressaria (Elachistidae) host-pants and ecology in the western United States. Journal of the Lepidopterists' Society 57.

Meyrick, E., 1928. A revised handbook of British Lepidoptera. Watkins and Doncaster, London.

Minet, J. 1990. Ramaniement partiel de la classification des Gelechioidea, essentiellement en fonction de caractères pre-imagginaux (Lepidoptera Ditrysia). Alexanor 16: 239-255. Translation.

Minet, J., 1991. Tentative reconstruction of the ditrysian phylogeny (Lepidoptera, ). Entomologica Scandinavica 22: 69-95.

340

Mosher, E., 1916. A classification of the Lepidoptera based on characters of the pupa. Bulletin of the Illinois State Museum of Natural History 12, 17-159.

Muona J. 1999. Trapping Beetles in boreal coniferous forest - how many species do we miss? Fennia 77: 11-16.

Nielson, E.S. and I.F.B. Common. 1991. Lepidoptera (moths and butterflies). In The Insects of Australia : a textbook for students and research workers Corp Author(s): Commonwealth Scientific and Industrial Research Organization (Australia).; Division of Entomology. Cornell University Press, Ithaca, N.Y.

Nixon, K. C., 1999. The Parsimony Ratchet, a new method for rapid parsimony analysis. Cladistics 15, 407-414.

Obenberger, J. 1952. Entomologie. Prírodovedecké vydavatelství, Praha.

Pashley, D. P., Ke, L. D., 1992. Sequence evolution in mitochondrial ribosomal and Nd-1 genes in Lepidoptera - Implications for phylogenetic analyses. Molecular Biology and Evolution 9,1061-1075.

Passoa, S. C., 1995. Larval and pupal systematics of Nearctic Amphisbatinae and Depressariinae (Lepidoptera; Oecophoridae). University of Illinois, Urbana, pp. 1- 300.

Petit S., Moilanen A., Hanski I., and Baguette M. 2001. Metapopulation dynamics of the bog fritillary butterfly: movements between habitat patches. Oikos 92: 491-500.

Phillips, A. J., Simon C., 1995. Simple, efficient and nondestructive DNA extraction protocol for arthropods. Annals of the Entomological Society of America 88, 281- 283.

Pickett, K. M., Simmons, M. P., Randle, C. P., 2004. Do Bayesian support values reflect probability of the truth? Cladistics 20, 92-93.

341

Pickett, K. M., Randle, C. P., In press. Strange bayes indeed: Uniform topological priors imply non-uniform clade priors. Molecular Biology and Evolution.

Powell, J., 1980. Evolution of larval food preferences in microlepidoptera. Annual Review of Entomology 25, 133-159.

Powell, J., Mitter, C., Farrell, B., 1998. Evolution of larval food preferences in Lepidoptera. In: Kristensen, N., (Ed), The Handbook of Zoology/Handbuch der Zoologie. Walter de Gruyter, Berlin & New York, 403-422 pp.

Preston F.W. 1948. The commonness, and rarity, of species. Ecology 29: 254-283.

Rambaut, A., 1996. Se-Al: Sequence Alignment Editor. Available at http://evolve.zoo.ox.ac.uk/.

Robinson, G.S., 1976. The preparation of slides of with special reference to the microlepidoptera. Entomologist’s Gazette 27, 127-132.

Rowe, L. and G. Arnqvist. 2002. Sexually antagonistic coevolution in a mating system: combining experimental and comparative approaches to address evolutionary

Saiki, R. K., 1990. Amplification of genomic DNA. In: Innis, M. A., Gelfand, D. H., Sninsky, J. J., White, T. J., (Eds.), PCR Protocols: A Guide to Methods and Applications. Academic Press, Inc, pp. 315-322.

Scharff N., Coddington J.A., Griswold C.E., Hormiga G., and Bjorn P.de P.. 2003. When to quit? Estimating spider species richness in a northern European deciduous forest. The Journal of the Arachnologists Society 31: 246-273.

Sharkey M.J. 2001. The All Taxon Biological Inventory of the Great Smoky Mountains National Park. Florida Entomologist 84: 556-564.

342

Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H., Flook P., 1994. Evolution, weighting, and phylogenetic utility of mitochondrial gene-sequences and a compilation of conserved polymerase chain-reaction primers. Annals of the Entomological Society of America 87, 651-701.

Simmons, M. P., Pickett, K. M., Miya, M., 2004. How meaningful are Bayesian support values? Molecular Biology and Evolution 21, 188-199.

Sinev, S. Y. 1993. On the systematics and phylogeny of the Gelechioidea s.l. (Lepidoptera). Entomological Review 72: 10-25.

Sober, E., Steel, M., 2002. Testing the hypothesis of common ancestry. Journal of Theoretical Biology 218, 395-408.

Sørenson L.L., Coddington J.A., and Scharff N. 2002. Inventorying and estimating subcanopy spider diversity using semiquantitative sampling methods in an afromontane forest. Environmental Entomology 31: 319-330.

Stanton R.C. 2000. Ground beetle abundance and diversity patterns within mixed-oak forests subjected to prescribed burning in southern Ohio. PhD Thesis, The Ohio State University, Columbus.

Stehr F.W. 1987. Superfamily Gelechioidea. In Stehr F.W. (ed), Immature Insects, pp 379-399. Kendall/Hunt, Dubuque, Iowa.

Summerville K.S., Boulware M.J., Veech J.A., and Crist T.O. 2003a. Spatial variation in species diversity and composition of forest Lepidoptera in eastern deciduous forests of North America. Conservation Biology 17: 1045-1057.

Summerville K.S. and Crist T.O. 2003. Determinants of lepidopteran community composition and species diversity in eastern deciduous forests: roles of season, eco- region and patch size. Oikos 100: 134-148.

Summerville K.S., Crist T.O., Kahn J.K., and Gerling J.C. 2003b. Community structure

343

of arboreal caterpillars within and among tree species of the eastern deciduous forest. Ecological Entomology 28: 747-757.

Sutherland E.K. and Hutchinson T.F. (eds). 2003. Characteristics of mixed-oak forest ecosystems in southern Ohio prior to the reintroduction of fire. USDA Forest Service General Technical Report NE-299.

Suzuki, Y., Glazko, G. V., Nei, M., 2002. Overcredibility of molecular phylogenies obtained by Bayesian phylogenetics. Proceedings of the National Academy of Sciences of the United States of America 99, 16138-16143.

Tillyard, R. J. 1926. The insects of Australia and . Angus & Robertson, Sydney.

Thompson, J. D., Higgins, D. G., Gibson T. J., 1994. Clustal-W - Improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position- specific gap penalties and weight matrix choice. Nucleic Acids Research 22, 4673-4680.

Triplehorn, C. A., and Johnson, N. F., 2004. Borror and DeLong's introduction to the study of insects. CBS College Publishing, Philadelphia. Seventh Edition.

Tscharntke T., Steffan-Dewenter I., Kruess A., and Thies C. 2002. Characteristics of insect populations on habitat fragments: A mini review. Ecological Research 17: 229-239.

Tuxen, S. L. 1970. Taxonomist's glossary of genitalia in insects. Munksgaard, Copenhagen.

Usher M.B. and Keiller S.W.J. 1998. The of farm woodlands: determinants of diversity and community structure. Biodiversity and Conservation 7: 725-748.

Wagner D.M. and Scholtens B. 2002. Lepidoptera Blitz nets 860 and counting. ATBI Quarterly 3: 6-8.

344

Wahlberg, N., Zimmermann, M., 2000. Pattern of phylogenetic relationships among members of the Melitaeini (Lepidoptera: Nymphalidae) inferred from mitochondrial DNA sequences. Cladistics 16, 347-363.

Wahlberg N., Klemetti T., Selonen V., and Hanski I. 2002. Metapopulation structure and movements in five species of checkerspot butterflies. Oecologia 130: 33-43.

Wiegmann, B. M., Mitter, C., Regier, J. C., Friedlander, T. P., Wagner, D. M., Nielsen, E. S., 2000. Nuclear genes resolve Mesozoic-aged divergences in the insect order Lepidoptera. Molecular Phylogenetics and Evolution 15, 242-259.

Wenzel, J. W., 2001. Phylogenetic analysis: The basic method. In: De Salle, R., Giribet, G., and Wheeler, W. (Eds) Phylogenetic analysis: The basic method. Birkhäuser Verlag Basel, Switzerland, pp. 6 - 30.

West-Eberhard, M. J. 1983. Sexual selection, social competition, and speciation. Q. Rev. Biol. 58: 155-183.

Zimmerman, E. C. 1978. Insects of ; a manual of the insects of the Hawaiian Islands, including an enumeration of the species and notes on their origin, distribution, hosts, parasites, etc. University of Hawaii Press, Honolulu.

345

APPENDIX A

MORPHOLOGICAL CHARACTERS USED IN CHAPTER 3: GELECHIOIDEA

SYSTEMATICS: A REEXAMINATION USING COMBINED MORPHOLOGY AND

MITOCHONDRIAL DNA DATA

Characters were coded by my interpretation, not as originally coded by original authors. Characters in matrix are left in the order presented by original authors as a matter of organization; rather than deleting repetitive characters, they were deactivated in

the Passoa matrix and left on in the Hodges matrix, or in some instances, deactivated in

both matrices and recoded in this analysis.

PASSOA (1995)

0. Setal arrangement of A9 of larva: D1 anteroventrad of D2 and anterodorsad of SD1 =

0; D2, D1 and SD1 in a vertical line = 1.

1. Spiniform Setae: none = 0; patches = 1; band = 2 [nonadditive].

2. M7 of genitalia: absent = 0; present = 1.

3. Valvae: undivided = 0; divided = 1 [deactivated].

346

4. Larvae with L group: ventrad = 0; posteroventrad = 1; anteroventrad = 2 [nonadditive].

5. Cremaster: absent = 0; present = 1.

6. Antennae in pupae: meeting = 0; not meeting = 1.

7. Stipular setae long and stout: normal = 0; long and stout = 1.

8. Juxta: present = 0; absent = 1 [deactivated].

9. Labial palpi: exposed = 0; hidden = 1.

10. Larvae with sclerotized rings: absent = 0; present = 1.

11. P setae arrangement: horizontal = 0; vertical = 1; polymorphic = 3 [nonadditive].

12. Larvae with distance between P2 subequal to distance between P1: longer = 0;

greater than or subequal = 1; [nonadditive].

13. Pupal movement restricted to vertical plane: full range of circular movement = 0;

movement restricted = 1; polymorphic = 3 [nonadditive].

14. Gnathos spined, oval: not spined = 0; spined = 1 [nonadditive; deactivated].

15. SV group of anal prolegs with secondary setae: 2nd setae covering whole body = 0;

2nd setae absent = 1; 2nd setae on SV group of anal proleg only = 2; 2nd setae

present on SV group of A3-6 and anal proleng only = 3; 2nd setae covering whole

body densely = 4 [nonadditive].

16. Larve with front short, not extending half distance to epicranial notch: extends more

than half the distance = 0; extends less than half the distance = 1.

17. Larvae with 2nd setae on A3-6: absent = 0; present = 1.

18. Male genitalia with strongly recurved aedeagus: straight = 0; curved = 1. .

19. Pubescent pupae: naked = 0; pubescent = 1.

347

20. Larval P setae with P2 posterolaterad to P1: P2 subequal to P1 = 0; P2 posterolaterad

to P1 = 1.

21. Adult abdomen with bands of stout setae: absent = 0; present = 1.

22. Pupae with elongate cremaster: absent = 0; present = 1.

23. Cremaster forked: not forked = 0; forked = 1.

24. Pupal mesothoracic spiracle ridgelike: slitlike, tubular projections = 0; ridgelike = 1.

25. Larvae with pits adjacent to SD1 setae on A1-8: lack pits = 0; have pits = 1.

HODGES (Hodges 1998)

26. Valva division: undivided = 0; divided = 1.

27. Juxta: present = 0; absent = 1.

28. Transtilla: absent = 0; present = 1.

29. Gnathos: sclerotized band that is generally uniformly wide, fused to tegumen,

sometimes incomplete mesially = 0; fused to tegumen, a sclerotized band that is

expanded mesially= 1; fused to tegumen, a sclerotized band with mesial region

down turned and laterally compressed = 2; fused to tegumen, a sclerotized band

with a mesial bulb bearing parallel rows of short spines = 3; an articulated band

with mesial region slightly expanded = 4; an articulated band with an

unarticulated mesial hook = 5; an articulated band with mesial bulb sometimes

bearing parallel rows of short spines = 6; a pair of articulated rami = 7; a pair of

lateral, articulated, symmetric sclerites = 8; absent = 9 [nonadditive; deactivated].

30. Uncus: present = 0; absent = 1.

348

31. Vinculum: generally and uniformly sclerotized, saccal region expanded (or not) = 0;

weakly sclerotized or open mesially = 1.

32. Sicae: absent = 0; present = 1.

33. Aedeagus: free = 0; ankylosed = 1.

34. Male genitalia: symmetric = 0; asymmetric = 1.

35. Abdominal second sternum: venula only = 0; venula + apodeme = 1; apodeme only =

2. [nonadditive; deactivated].

36. Female frenulum: three acanthi = 0; two acanthi = 1; one acanthus = 2 [nonadditive].

37. Female retinaculum consisting of: diffusely distributed, anteriorly directed, slightly

upturned scales = 0; anteriorly directed scales on CuA = 1; anteriorly directed scales

between CuA and R = 2; anteriorly directed scales starting in cell and extending to

area between Sc and R = 3; anteriorly directed scales between CuA and R and then on

R further out from base = 4; anteriorly directed scales on R = 5; anteriorly directed

scales between Sc and R = 6 [nonadditive].

38. Frenulum: absent = 0; present = 1.

39. Forewing with: R5 terminating on outer margin = 0; R5 terminating on costa = 1.

40. Forewing with CuA1/CuA2 : separate, directed towards outer margin = 0;

stalked/connate/approximate, directed towards posterior margin at origin from

end of cell= 1.

41. Forewing with CuP: Present = 0; absent = 1.

42. Forewing with pterostigma: absent = 0; present = 1.

43. Hindwing: Sc/R1 united from base of wing = 0; R1 running into Sc beyond base of

349

wing = 1.

44. Hindwing with RsM1: separate = 0; stalked = 1.

45. Hindwing with M1/M2: separate = 0; stalked = 1; fused = 2 [nonadditive].

46. Hindwing with M3CuA1: separate = 0; connate or stalked = 1; fused = 2

[nonadditive].

47. Hindwing angle of closer of cell by vein M2-M3 (M1-M2): perpendicular with long

o axis of wing = 0; directed at a 45 angle toward base of wing from M2= 1;

directed toward apex = 2; open= 3 [nonadditive].

48. Hindwing outer margin: rounded/straight = 0; slightly excavated just posterad of apex

= 1.

49. Hindwing: without scales or stout setae on costal margin = 0; with scales or stout

setae on costal margin = 1.

50. Ocellus: present = 0; absent = 1

51. Antennal pectin: absent = 0; present = 1.

52. Antennal notch: absent = 0; present = 1. .

53. Male genitalia with appendix appendicular: absent = 0; present = 1.

54. Haustellum: present = 0; absent = 1.

55. Pupa: without lateral condyles on segments 5/6, 6/7 = 0; with lateral condyles on

segments 5/6, 6/7 = 1.

56. Pupa: without ventrolateral projections on segment 8/9 = 0; with ventrolateral

projections on segment 9 or 8/9 = 1.

57. Pupal rows of spines on terga 4,5,6,7: absent = 0; present = 1.

350

58. Late instar larvae: without submental pit = 0; with submental pit = 1.

59. Larval secondary SV setae on A3-9: absent = 0; present = 1.

60. Male genitalia manica: membranous = 0; sclerotized = 1.

61. Larval abdominal segments A1-8: without a pore = 0; with a pore = 1.

62. Larval abdominal segments A1-8: without a pinaculum ring = 0; with a pinaculum

ring = 1.

RECODED FROM ABOVE CHARACTERS

63. Adult abdominal segments: without spiniform setae = 0; with spiniform setae = 1.

64. If spiniform setae present (38,0): spiniform setae in patches or band medially = 0;

spiniform setae in band along post. margin = 1.

65. If spiniform setae in patches or band medially (39,0): spiniform setae in patches = 0;

spiniform setae in band = 1.

66. Abdominal second sternum venula: absent = 0; present = 1.

67. Abdominal second sternum apodemes: absent = 0; present = 1.

68. Gnathos: Absent = 0; Present = 1.

69. If gnathos present (42,1): Band = 0; articulated rami = 1; articulated symmetric

sclerites = 2 [nonadditive].

70. If band (43,0): sclerotized = 0; articulated = 1.

71. If sclerotized band fused to tegumen (44,0): informally wide = 0; mesially turned

down, laterally compressed = 1; expanded mesially = 2; mesial bulb, parallel row

of spines = 3 [nonadditive].

351

72. If articulated band (44,0): mesial region slightly expanded = 0; unarticulated mesial

hook = 1; mesial bulb bearing row of short spines = 2 [nonadditive].

352

APPENDIX B.

COMPLETE MORPHOLOGICAL MATRIX FOR FAMILIES AND SUBFAMILIES

OF GELECHIOIDEA USED IN CHAPTER 4: GELECHIOIDEA SYSTEMATICS: A

REEXAMINATION USING COMBINED MORPHOLOGY AND MITOCHONDRIAL

DNA DATA

.

Actual specimens were coded as terminals and presented in Appendix A.

Subfamilies used for consistency with Hodges and Passoa’s matrices.

0 5 10 15 20 25 30 35 40 45 | | | | | | | | | | Yponomeutoidea 00000010000------Batrachedrinae *210200000000000000000000000040*000$$$*10*0001230 Coleophorinae 12111111000000000000000000101310010026011000*$000 Momphinae 121110000000000000000000001019000000110100000$000 Pterolonchinae 01?011110000000000000000000005000002$001000000000 Cosmopteriginae *0*000001100000000000000000109010111??0101001?0?0 Gelechiinae *00000001100000000000000000108?????1050101??????1 Scythridinae 000020000010000000000000000000??01?????00000000?0 Blastobasinae 011120000111200130010000001000000*01$$01*01000100 Oecophorinae 0110200001033001000310000000*1000*0*$$0*000000*$0 Autostichinae 0110200001111001100111111100?5000001130??000?010? Stenomatinae 00102000011111011000000000?0?0??0?0???00?0?0?0??0 Ethmiinae 0010200001012122011000000000?0??0000?20100?000??0 Depressariinae 0010200001012112011000000000?600000?020??00000??0 Amphisbatinae 00102000010111120100000000000600000?0?01000000100 Symmocinae 0010200001011001100311111100?500000?12???100?0?00 Stathmopodinae ------0001000000$101000000$30 Chrysopeleiinae ------0109??01?1??010100?0030 353

0 5 10 15 20 25 30 35 40 45 | | | | | | | | | | Dichomeridinae ------0?080?1??0?5011101?0??1 Pexicopiinae ------01080000???501010??0?01

354

50 55 60 64 69 | | | | | Yponomeutoidea ------Batrachedrinae *1*000000*0000100--101-0 Coleophorinae 01100000000000100101003- Momphinae 01100000000000100100---- Pterolonchinae 0110010000000010101101-1 Cosmopteriginae ??10000000?1000--110---- Gelechiinae ??00?0000000000--1112--- Scythridinae 0?1000000?11110----1000- Blastobasinae 011*00000100**101111000- Oecophorinae 01*000000*000**01--1002- Autostichinae 0100000000001110111101-1 Stenomatinae 0100001?0000000----1000- Ethmiinae 01?0001?0010000--101000- Depressariinae 0??000100000000----101-2 Amphisbatinae 010000000000000----101-- Agonoxeninae ?1?000110010000--101$--- Symmocinae 01?00000000010101--101-1 Stathmopodinae 0100000000*00011-101002- Chrysopeleiinae ???0?0000000000--110---- Dichomeridinae 0?0?10000000000--1012--- Pexicopeiinae ???0?0000000000----12---

355

APPENDIX C.

MOLECULAR MATRIX USED TO CONSTRUCT PHYLOGENIES PRESENTED IN

CHAPTER 4: GELECHIOIDEA SYSTEMATICS: A REEXAMINATION USING

COMBINED MORPHOLOGY AND MITOCHONDRIAL DNA DATA

Matrix aligned using CustalX and proof-read using Se-Al.

0 5 10 15 20 25 30 35 40 45 50 | | | | | | | | | | | Yponomeuta multipunctella gaagtatatattctaattttaccgggatttggaataatttctcatattattt Batrachedra enormis ------Batrachedra praeangusta ------ttttaccaggatttggaataatttctcatattattt Batrachedra calator ------Coleophora ericoides ------taatctctcatattattt Coleophora new species gaagtttatattttaattttaccaggatttggtataatttctcatattatcc Mompha circumscriptella ------tatattttaattttacctggatttggtataatttctcatattattt Mompha eloisella ----tttatattttaattttaccgggatttggaataatttctcatattattt Mompha new species gaagtttatattttaattttacctggatttggtataatttctcatattattt Pterolonche sp 1 ------Triclonella pergandeella ------ttttattttacctggatttggaataatctctcatattattt Eteobalea serratella gaagtttatattttaattttacctggatttggaataatttctcatattattt Cosmopterix sp 1 ------Periploca laetella ------Walshia miscecolorella -aagtatatattttaattttaccaggatttgggataatttcccatattattt Sitotroga cerealella ------Gelechiinae 2 ------cttccagggatttggaatnntctctcatattattt Gelechia sp 1 gaagtttccattttaattttacctggatttggtataatttctcatattattt Gelechia sp 4 gaagtttatattttaattttaccaggatttggtataatttctcatattattt Teleiodini sp 1 ------Filatima sp 1 gaagtttatattttaattttaccgggatttggaataatttctcacattattt Paralita group sp 1 gaagtttatattttaattttaccaggatttggaataatttctcatattattt Dichomeris ligulella gaagtttatattttaattttaccgggatttggtataatttctcatatcattt Gelechia sp 3 ------Asymmetura sp 1 gaagtttatattctaattctacctggatttggaataatttctcatattattt Scythris limbella ------Antaeotricha schlaegeri ------ttaccgggatttgggataatctcccatattattg Oegoconia quadripuncta gaagtttatattttaatcctaccaggatttggtataatttcccacattattt Decantha boraesella gaagtttatattttaattctcccaggatttggaataatttcccatattattt Psilocorsis reflexella ------attcttccaggatttggaataatttcacatattattt Mathildana newmanella gaagtttatattttaattttacctggatttggaataatttcacatattattt 356

0 5 10 15 20 25 30 35 40 45 50 | | | | | | | | | | | Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola ------ggaataatctctcatattattt Ethmia trifurcella ------Nites sp 1 gaagtttatattttaattttacctggatttggaataatttctcatattattt Depressaria pastinacella ------Agonpterix robinella ------Pigritia sp 1 ------Hypatopa sp 3 ------atatttttattttaccaggatttggaaaaatttctcacattatcc Blastobasis sp 3 ------Calosima sp 1 ------

357

52 57 62 67 72 77 82 87 92 97 102 | | | | | | | | | | | Yponomeuta multipunctella ctcaagaaagaggaaaaaaagaaacatttggtaatttaggaataatttacgc Batrachedra enormis ------tatgc Batrachedra praeangusta ctcaagaaagagggaaaaaagaaacttttggatgtttaggaataatttatgc Batrachedra calator ------Coleophora ericoides cacaggaaagtgggaaaaaagaaactttcgggtatttaggaataatttatgc Coleophora new species cccaagaaagaggtaaaaaagaaacttttgggtatttaggaataatttatgc Mompha circumscriptella accaagaaagaggaaaaaaagaaacatttggatctttaggaataatctatgc Mompha eloisella accaagaaagaggaaaaaaagaaacatttggatctttagggataatttatgc Mompha new species accaagaaagaggaaaaaaagaaacatttggatctttaggaataatctatgc Pterolonche sp 1 ------Triclonella pergandeella ctcaagaaagaggaaaaaaagaaacttttggttgtttaggaataatttatgc Eteobalea serratella cccaagaaagaggaaaaaaagaaacttttggttgtttaggaataatttatgc Cosmopterix sp 1 ------Periploca laetella -----aaagaggagaaaaggaaacatttggntgtcttaggaataatttatgc Walshia miscecolorella cccaagaaagaggaaaaaaagaaacttttggttgcttaggaataatttatgc Sitotroga cerealella ------ggaata-tttatgc Gelechiinae 2 cccaagaaagtggcaaaagggaaacttttggatcattaggaataatttatgc Gelechia sp 1 cccaagaaagaggtaaaaaagaaacatttggatctttagggataatttatgc Gelechia sp 4 ctcaagaaagtggaaaaaaggaaactttcggatctttaggaataatttatgc Teleiodini sp 1 ------aaaaattgaatcatttggaacactaggaataatttatgc Filatima sp 1 ctcaagaaagaggaaaaaaggaaacttttggatcactaggaataatttatgc Paralita group sp 1 cccaagaaagaggtaaaaaggaaacttttggggccttaggaataatttatgc Dichomeris ligulella cccaagaaagaggaaaaaaggaaacatttggatcattaggaataatttatgc Gelechia sp 3 ------Asymmetura sp 1 cccaagaaagaggaaaaaaagaaacttttggttgtttaggaataatttacgc Scythris limbella ------aaaaattgaatcatttggaacactaggtataatttatgc Antaeotricha schlaegeri cccaagaaagaggtaaaaaagaaacttttggttgtttaggaataatttatgc Oegoconia quadripuncta cccaagaaagaggaaaaaaagaaacttttggttgtttaggtataatttatgc Decantha boraesella cacaagaaagagggaaaaaagaaacttttgggtccttaggaataatctatgc Psilocorsis reflexella ctcaagaaagaggaaaaaaagaaacttttggttgtttaggaataatttatgc Mathildana newmanella cccaagaaagaggaaaaaaagaaacttttggctctttaggaataatttatgc Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola cccaagaaagaggaaaaaaagaaacttttgggtgtttaggtataatttatgc Ethmia trifurcella ------Nites sp 1 cccaagaaagaggaaaaaaagaaacttttggatgtttaggaataatttatgc Depressaria pastinacella ------aaaaagaaacatttggttgcttaggaataatttatgc Agonpterix robinella ------Pigritia sp 1 ------gaggctcttaggttgtttaggaataatttatgc Hypatopa sp 3 ctcaagaaagaggaaaaaaagaaacttttggttgtctaggaataatttacgc Blastobasis sp 3 ------Calosima sp 1 ------

358

104 109 114 119 124 129 134 139 144 149 154 | | | | | | | | | | | Yponomeuta multipunctella tataatagcaattggattattaggatttgttgtatgagctcatcatatattt Batrachedra enormis tataatagcaattggtcttcttgggtttattgtctgagctcatcatatattt Batrachedra praeangusta aataatagcaattggattattaggatttgttgtctgagctcatcatatattt Batrachedra calator ------Coleophora ericoides tataatagcaattggtttattaggatttgtagtttgagctcatcatatattt Coleophora new species tataatagcaattggattattaggatttgtagtttgagctcatcatatattt Mompha circumscriptella tataatggcaatcggtttattaggatttgtagtatgagctcatcatatattt Mompha eloisella tataatagcaattggattattaggatttgtagtttgagctcatcatatattt Mompha new species tataatggcaatcggtttattaggatttgtagtatgagctcatcatatattt Pterolonche sp 1 ------Triclonella pergandeella tataatagcaattggtttattaggatttgtagtttgagctcatcatatattt Eteobalea serratella tataatagcaattggtttattaggatttgtagtttgagctcatcatatattt Cosmopterix sp 1 ------Periploca laetella tataatagcaattggtttattaggatttgtagtttgagctcatcatatattt Walshia miscecolorella tataatagcaattggtcttttaggatttgtagtctgagctcatcatatattt Sitotroga cerealella tataatagcaattggacttttaggattcgttgtttgagctcatcatatattt Gelechiinae 2 tataatagcaattggattattaggatttgttgtttgagctcaccatatattt Gelechia sp 1 tataatagcaattggtttattaggatttgttgtttgagctcatcatatattt Gelechia sp 4 tataatagcaattggattattaggatttgttgtttgagcacatcatatattt Teleiodini sp 1 tatattatcaattggactaataggatttattgtatgagcacatcatatattc Filatima sp 1 tataatagcaattggattattaggatttgttgtatgagctcatcatatattt Paralita group sp 1 tataatagcaattggattattaggatttgtcgtatgagctcatcatatattt Dichomeris ligulella tataatagcaattggacttttaggatttgttgtttgagctcatcatatattt Gelechia sp 3 ------Asymmetura sp 1 tataatagcaattggattattaggatttattgtttgagcacatcatatattc Scythris limbella tatattatcaattggactaataggatttattgtatgagcacatcatatattc Antaeotricha schlaegeri aataatagcaattggtttgttaggatttgttgtttgagctcatcatatgttt Oegoconia quadripuncta aataatagcaattggtttacttggatttgttgtatgagctcatcatatattc Decantha boraesella tataatagctattggtttattgggatttgtagtatgagctcatcatatattt Psilocorsis reflexella tataatagcaattggattacttggatttgtagtttgagctcatcacatattt Mathildana newmanella tatattagcaattggattattaggatttgtagtatgagctcatcacatattt Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola tataatagcaattgggctattaggatttgtagtgtgagctcatcatatattt Ethmia trifurcella ------Nites sp 1 tataatagcaattggtttattaggatttgtagtgtgagcacatcatatattt Depressaria pastinacella tataatggcaattggattattaggatttgtagtatgagctcaccatatgttt Agonpterix robinella ------Pigritia sp 1 tataatagctattggtcttttagggtttgtagtttgagctcatcatatattt Hypatopa sp 3 tataatagcaattggattattaggatttgtagtttgagctcatcatatattt Blastobasis sp 3 ------Calosima sp 1 ------

359

156 161 166 171 176 181 186 191 196 201 206 | | | | | | | | | | | Yponomeuta multipunctella acagttggaatagatattgatacccgagcttattttacttctgcaactataa Batrachedra enormis acagtaggaatagatattgatacacgagcttactttacttcagcaactataa Batrachedra praeangusta actnttggaatagatattgatacacgagcttattttacctcagcaactataa Batrachedra calator ------Coleophora ericoides actgttggcatagatattgatactcgagcttactttacttctgctactataa Coleophora new species actgttggaatagatattgatactcgagcttattttacttcagctacaataa Mompha circumscriptella acggtaggaatagatattgacactcgagcttattttacctcagcaacaataa Mompha eloisella acagtaggaatagatattgatactcgagcttattttacttcagctaccataa Mompha new species acggtaggaatagatattgacactcgagcttattttacctcagcaacaataa Pterolonche sp 1 ------attgacacacgagcttattttacatctgcaacaataa Triclonella pergandeella acagtaggtatagatattgatacacgagcttattttacttcagcaactataa Eteobalea serratella actgttggaatagacattgatactcgagcttattttacttcagcaacaataa Cosmopterix sp 1 ------Periploca laetella acagtcggtatagatattgatacacgagcttattttacatcagctactataa Walshia miscecolorella actgttggaatagatattgatactcgagcttactttacttcagccaccataa Sitotroga cerealella acagttggtatagatatcgatactcgagcttatttcacttcagctactataa Gelechiinae 2 actgtaggaatagacattgatacccgagcttattttacctctgctactataa Gelechia sp 1 acagttggaatagatattgatactcgggcttattttacatcagctactataa Gelechia sp 4 actgtaggaatagatattgatactcgagcttattttacgtcagccacaataa Teleiodini sp 1 acagtaggaatagacgttgacacacgagcatactttacatcagcaacaataa Filatima sp 1 actgttggtatagatattgatactcgagcttattttacttcagctacaataa Paralita group sp 1 acagtagggatagatattgatactcgagcatactttacttctgctactataa Dichomeris ligulella acagttggtatagatattgatactcgagcttattttacttcagccactataa Gelechia sp 3 ------Asymmetura sp 1 actattggaatagatattgatactcgagcatatttcacttcagctaccataa Scythris limbella acagtaggaatagacgttgacacacgagcatactttacatcagcaacaataa Antaeotricha schlaegeri actgttggaatagatattgatactcgagcttattttacttcagctacaataa Oegoconia quadripuncta acagtaggtatagatattgatacacgagcttactttacctcagccacaataa Decantha boraesella actgttgggatagatattgatactcgagcatattttacctcagcgactataa Psilocorsis reflexella accgtaggaatagatattgatactcgagcttattttacatcagctacaataa Mathildana newmanella acagttggaatagatattgacacacgtgcttattttacttcagcaactataa Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola acagtaggtatagatattgatacacgagcttattttacatcagcaacaataa Ethmia trifurcella ------Nites sp 1 acagtaggaatagatattgatactcgagcttattttacttcagcaaccataa Depressaria pastinacella acagtaggaatagatattgatactcgagcttattttacttcagcaactataa Agonpterix robinella ------Pigritia sp 1 actgttggtatagatattgatacacgagcttactttacttcagcaactataa Hypatopa sp 3 actgtaggtatagatattgatactcgtgcttattttacttccgctacaataa Blastobasis sp 3 ------Calosima sp 1 ------

360

208 213 218 223 228 233 238 243 248 253 258 | | | | | | | | | | | Yponomeuta multipunctella ttattgctgtacctactgggattaaaatttttagttgattagcaactttaca Batrachedra enormis ttattgctgttcctacaggaattaaaatttttagttgattagctacattaca Batrachedra praeangusta ttattgcagttcctacaggaattaaaatttttagttgattagctactttaca Batrachedra calator ------Coleophora ericoides ttattgctgtacctactggtattaaaatttttagatgattagcaacattaca Coleophora new species ttattgctgtacctactggaattaaaatctttagatgattagcaacattaca Mompha circumscriptella ttattgctgttcctacaggaattaaaatttttagttgattagcaactcttca Mompha eloisella ttattgctgttcctacaggaattaaaatttttagttgattagccactttaca Mompha new species ttattgctgttcctacaggaattaaaatttttagttgattagcaactcttca Pterolonche sp 1 ttatggctattccaacaggaattaaagttttcagatgaatagcaactattta Triclonella pergandeella ttattgcagtaccaacaggtattaaaatttttagttgattagctactcttca Eteobalea serratella ttattgcagtacctacaggtattaaaatttttagttgattagcaactcttca Cosmopterix sp 1 ------Periploca laetella ttattgcagtacctacaggaattaaaatttttagatgattagctacattaca Walshia miscecolorella ttattgctgtaccaacaggaattaaaatttttaggtgattagctactcttca Sitotroga cerealella ttattgctgttccaaccggtattaaaatttttagttgacttgctactttaca Gelechiinae 2 ttattgctgtaccaacaggaattaaaatttttagttgattagcaacccttca Gelechia sp 1 ttattgctgttccaacaggaattaaaatttttagttgattagctactcttca Gelechia sp 4 ttattgctgtacctacaggaattaaaatttttagttgacttgcaactcttca Teleiodini sp 1 ttattgcagtacctacaggaatcaaggtattctgatgactagctacactata Filatima sp 1 ttattgcagttccaacaggaattaaaattttcagttgacttgcaactcttca Paralita group sp 1 ttattgcagtacctacgggaattaaaatttttagttgattagccactcttca Dichomeris ligulella ttattgctgtgcctacaggaattaaaatttttagttgattagctactcttca Gelechia sp 3 ------Asymmetura sp 1 ttattgcagtacctactggaattaaaatttttagatgattagctactttaca Scythris limbella ttattgctgtacctacaggaatcaaggtattcagatgactagctacactata Antaeotricha schlaegeri ttattgctgtaccaacaggtattaaaatttttagatgattggctactttaca Oegoconia quadripuncta ttattgcagtacctacaggaattaaaatttttagatgattagctactttcca Decantha boraesella ttattgcggttcctacaggaattaaaatttttagatgattagcaactcttca Psilocorsis reflexella ttattgctgtacctacaggtattaaaatttttagttgattagccactttaca Mathildana newmanella ttattgctgttccaacaggaattaaaatttttagatgattagcaacacttca Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola ttattgctgtgcctacaggaattaaaatttttagttgattagctactttaca Ethmia trifurcella ------Nites sp 1 ttattgccgttcctacaggaattaaaatttttagttgattagcaactcttca Depressaria pastinacella ttattgctgtgccaaccggaattaaaatctttagttgattagctacccttca Agonpterix robinella ------Pigritia sp 1 ttattgcagtaccaactggtattaaaattttcagttgacttgctactcttca Hypatopa sp 3 tcattgcagtacctacaggaatcaaaatttttagttgacttgctactcttca Blastobasis sp 3 ------Calosima sp 1 ------

361

260 265 270 275 280 285 290 295 300 305 310 | | | | | | | | | | | Yponomeuta multipunctella tggaactcaaattaattatagaccttcaattttatgaagattaggatttgta Batrachedra enormis tggaactcaaattaattatagtccttcaatattatgaagattaggatttgta Batrachedra praeangusta tggaactcaaattaattatagaccttctatactatgaagattaggatttgta Batrachedra calator ------Coleophora ericoides tggaactcaaatcaattatagaccttctatactttgaagattaggatttgta Coleophora new species cgggacacaaattaattatagtccatctatactttgaagattaggatttgta Mompha circumscriptella tggatcccaaattaactatagaccatcaattttatgaagactaggatttgtt Mompha eloisella tggctcacaaattaattatagtccatctattttatgaagattaggatttgtt Mompha new species tggatcccaaattaactatagaccatcaattttatgaagactaggatttgtt Pterolonche sp 1 tggatcaaaaaatattttatctttatcaataatttgaagtataggatttatt Triclonella pergandeella cggtacacaaattaattatagtccttctattttatgaagattaggatttgta Eteobalea serratella tggaactcaaattaattatagtccatcaattttatgaagattaggatttgta Cosmopterix sp 1 ------Periploca laetella tggaactcaaattaattatagtccttctattctatgaagattaggatttgtg Walshia miscecolorella tggaacacaaattaattatagaccttcaattctttgaagattaggattcgtt Sitotroga cerealella tggaactcaaattaattatagcccctctattctttgaagattaggatttgta Gelechiinae 2 tggaacacaaattaattatagtccttctattttatgaagcttaggatttgta Gelechia sp 1 tggaacacaaattaattatagtccatctatattatgaagtttaggattcgta Gelechia sp 4 tggaactcaaattaattatagtccttcaatcttatgaagattaggatttgtt Teleiodini sp 1 tggaactaaattcaaatttaatccaccgatattatgagctctaggatttatt Filatima sp 1 tggaacccaaattaattatagtccttcaattttatgaagtttaggatttgtt Paralita group sp 1 tggtacccaaattaattatagtccttcaattttatgaagattaggatttgta Dichomeris ligulella cggtactcaaattaactatagcccttcaattttatgaagcttaggatttgta Gelechia sp 3 ------Asymmetura sp 1 tggaacaaaaattaactatagtccttcaattctttgaagattaggatttgta Scythris limbella tggaactaaattcaaatttaatccaccgttattatgagctctaggatttatt Antaeotricha schlaegeri tggaactcaaattaactatagtccatctatattgtgaagattaggatttgtt Oegoconia quadripuncta tggaacacaaattaattttagaccgtctattctttgaagattaggatttgta Decantha boraesella tgggactcaaattaattatagcccttctatattatgaagattaggatttgtt Psilocorsis reflexella tggaactcaaattaattatagtccatctactttatgaagactaggatttgta Mathildana newmanella tggaactcaaattaactatagtccttctatattatgaagtcttggatttgta Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola tggaactcaaattaattatagaccttcaattctctgaagattaggatttgta Ethmia trifurcella ------Nites sp 1 tggaactcaaattaattatagaccttcaattttatgaagtttaggatttgta Depressaria pastinacella tggaactcaaattaattatagaccatcaatcttatgaagactagggtttgta Agonpterix robinella ------Pigritia sp 1 tggaactcaaattaattttagaccttcaactttatgaagattaggatttgtt Hypatopa sp 3 cggaactcaaattaattttagcccatccactttatgaagattaggatttgta Blastobasis sp 3 ------Calosima sp 1 ------

362

312 317 322 327 332 337 342 347 352 357 362 | | | | | | | | | | | Yponomeuta multipunctella tttttatttacagttggaggattaacaggtgtaattttagctaactcttcta Batrachedra enormis tttttatttacagtaggaggattaacaggagtaattttagctaattcatcta Batrachedra praeangusta tttttattcacagtaggaggattaacaggagtaattttagctaattcttcaa Batrachedra calator ------Coleophora ericoides tttttatttactgtagggggattaactggagtaattttagctaattcttcta Coleophora new species tttttatttactgtaggaggactaactggagtaattttagctaattcttcta Mompha circumscriptella tttttatttactgtaggaggattaacaggagtaattttagcaaattcatcta Mompha eloisella tttttattcacagttggtggattaacaggagtaattttagctaactcatcta Mompha new species tttttatttactgtaggaggattaacaggagtaattttagcaaattcatcta Pterolonche sp 1 tttttatttactttaggtggattaacaggaattattttatccaattcttcta Triclonella pergandeella tttttattcacagttggaggtttaactggagtaattttagctaattcttcta Eteobalea serratella tttttatttacagtaggaggtttaactggagtaattttagctaattcttcta Cosmopterix sp 1 ------Periploca laetella ttcttatttactgtaggaggattaacaggagtaattttagctaattcatcaa Walshia miscecolorella tttttatttacagtgggaggattaacaggagtaatcctagctaattcttcaa Sitotroga cerealella tttttatttactgtagggggattaacaggagtaattttagctaattcatcta Gelechiinae 2 tttctttttactgtaggaggattaacaggagtaattcttgctaattcctcaa Gelechia sp 1 tttttatttactgtaggaggattaacaggtgtaattttagctaattcttcta Gelechia sp 4 ttcttatttactgtagggggattaactggtgttattttagctaattcttcta Teleiodini sp 1 ttcctatttacagttgggggactaacaggattagtattagcaaattcatcac Filatima sp 1 tttttatttacagtaggtggattaactggtgtaattttagctaattcatcaa Paralita group sp 1 ttcttatttacagtaggaggattaactggagtaattttagccaattcttcaa Dichomeris ligulella tttttatttacagttgggggattaacaggagttattttagctaattcttcta Gelechia sp 3 ------Asymmetura sp 1 tttttattcactgtagggggattaactggagttattttagctaattcttcta Scythris limbella ttcctatttacaattggtggactaacaggattagtattagcaaattcatcac Antaeotricha schlaegeri ttcctttttactgtaggaggattaacaggagtaattttagctaactcttcta Oegoconia quadripuncta ttcttatttactgtcggaggtttaactggagtagttttagctaactcctcta Decantha boraesella tttttatttactgttggaggattaacaggtgtaattttagctaattcatcaa Psilocorsis reflexella tttttatttactgttggaggattaacaggagtaattttagctaattcttcta Mathildana newmanella tttttatttactgtaggaggattaacaggagtaattttagctaattcctcaa Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola tttctttttactgtaggtggactaacaggagtaattctagctaattcatcaa Ethmia trifurcella ------Nites sp 1 tttttattcacagtaggtggattaacaggagtaattttagcaaattcatcaa Depressaria pastinacella tttttatttacagtagggggattgacaggagttattttagctaactcttcaa Agonpterix robinella ------Pigritia sp 1 tttctatttactgttggaggattaacaggagttattttagctaattcttcta Hypatopa sp 3 tttttatttactgttggaggtttaactggagttattttagctaattcttcca Blastobasis sp 3 ------Calosima sp 1 ------

363

364 369 374 379 384 389 394 399 404 409 414 | | | | | | | | | | | Yponomeuta multipunctella ttgatgtatcattacatgatacttattatgttgttgctcatttccattatgt Batrachedra enormis ttgatgtatctcttcatgacacttactatgttgtagctcactttcattatgt Batrachedra praeangusta ttgatgtttctcttcacgacacatattatgttgtagctcattttcattatgt Batrachedra calator ------Coleophora ericoides ttgatgtaactttacatgatacatattatgttgtagcacattttcattatgt Coleophora new species ttgatgtaatattacatgatacatattatgttgtagctcattttcattatgt Mompha circumscriptella ttgatattacattacatgatacttattatgttgtagctcattttcactatgt Mompha eloisella ttgatattatattacatgatacctactatgtagttgcgcattttcattatgt Mompha new species ttgatattacattacatgatacttattatgttgtagctcattttcactatgt Pterolonche sp 1 ttgatattattcttcatgatacttattatgtaattggtca-ttccattatgt Triclonella pergandeella ttgatgtatctcttcatgatacttattatgttgttgctcattttcattatgt Eteobalea serratella ttgatgtatcattacatgatacatattatgttgtagctcattttcattacgt Cosmopterix sp 1 ------Periploca laetella ttgatgtttctcttcatgatacttattatgtagttgcccactttcattatgt Walshia miscecolorella ttgatgtatctcttcacgacacttattatgtagttgctcattttcattatgt Sitotroga cerealella ttgatattactttacatgatacttattacgtagttgctcattttcattatgt Gelechiinae 2 ttgatgttgcattacatgatacttattatgttgttgctcattttcattatgt Gelechia sp 1 ttgatgtagcactccatgatacttattatgtagttgcccattttcattatgt Gelechia sp 4 ttgatgttgctctccatgatacttattatgtagtagctcacttccattatgt Teleiodini sp 1 ttgacattgtattacatgacacatattatgtagatgccctcttccattatgt Filatima sp 1 ttgatgttgctctccatgatacttattatgtagtagcccactttcattacgt Paralita group sp 1 ttgatgttgctttacatgatacatactacgtagtcgctcattttcattatgt Dichomeris ligulella ttgatgtcgctcttcatgatacatattatgtagtagcacatttccattatgt Gelechia sp 3 ------Asymmetura sp 1 ttgatatcattttacatgatacttattatgtagtagcccactttcattatgt Scythris limbella ttgacattgtattacatgacacatattatgtagttgcccacttccattatgt Antaeotricha schlaegeri ttgatgtagcattacatgatacctattatgttgtagcacattttcattatgt Oegoconia quadripuncta ttgatattaccctccacgatacatattacgtagtagctcattttcactatgt Decantha boraesella ttgatgttactttacatgacacatattatgttgttgctcattttcattatgt Psilocorsis reflexella ttgatgttactttacatgatacttattatgtagttgctcattttcattatgt Mathildana newmanella ttgatgtcacattacatgacacttattatgttgtagctcattttcattacgt Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola ttgatattaccttacatgatacttattatgtagtagctcattttcattatgt Ethmia trifurcella ------Nites sp 1 ttgatgtaacattacatgatacttattatgtagtagcacactttcattatgt Depressaria pastinacella ttgatgtaactttacatgatacttattatgttgtagcacattttcattatgt Agonpterix robinella ------Pigritia sp 1 ttgatattactcttcatgatacttattatgtagttgctcattttcattatgt Hypatopa sp 3 tcgatattactcttcatgatacttattatgtagttgctcatttccattatgt Blastobasis sp 3 ------Calosima sp 1 ------

364

416 421 426 431 436 441 446 451 456 461 466 | | | | | | | | | | | Yponomeuta multipunctella tttatcaataggagcagtatttgctattatggcaggatttatccattgattt Batrachedra enormis cttatcaataggagccgtatttgctattataggaagatttattcattgatat Batrachedra praeangusta tctatctataggagctgtatttgctattataggaggatttattcattgatat Batrachedra calator ------Coleophora ericoides tctttctataggagctgtattcgctattataggaggatttattcactgatac Coleophora new species tctttctataggagctgtatttgctattataggaggatttattcattgatac Mompha circumscriptella attatctataggagctgtatttgctattataggaggatttattcattgatat Mompha eloisella attatctataggagcagtatttgctattataggaggatttattcactgatac Mompha new species attatctataggagctgtatttgctattataggaggatttattcattgatat Pterolonche sp 1 tctttcaataggagcagtatttgctatttttgctagattaattcaatgattt Triclonella pergandeella attatctatgggagccgtatttgctattataggaggatttattcattgatac Eteobalea serratella tctttcaataggagctgtatttgccattataggaggatttattcattgatat Cosmopterix sp 1 ------Periploca laetella tctttctataggagcagtatttgctattatagcaggatttattcattgatat Walshia miscecolorella tttatctataggagcagtatttgctattatagggggatttattcattgatat Sitotroga cerealella tttatctataggagcagtatttgcaatcttagggggattcattcattgatac Gelechiinae 2 tctatctataggagctgtattcgctattttaggaggatttattcattgatat Gelechia sp 1 attatctataggagcagtatttgctattttagggggatttattcattgatat Gelechia sp 4 tttatctatgggagctgtatttgccattttagggggatttattcacgggtac Teleiodini sp 1 attatctataggagcagatattgcaattataggaggagttattcaatgatac Filatima sp 1 tctttctataggagcagtatttgctattttagggggatttattcattgatat Paralita group sp 1 attatctataggagctgtatttgctattatgggaggttttattcattgatac Dichomeris ligulella tttatctataggagctgtatttgctattttaggaggatttattcactgatat Gelechia sp 3 ------Asymmetura sp 1 attatcaataggagcagtatttgctattataggaggatttattcattgatat Scythris limbella attatctataggagcagtatttgcaattataggaggtgttattcaatgatac Antaeotricha schlaegeri tttatcaataggggcagtatttgccattatagggggatttattcattgatac Oegoconia quadripuncta cctttctataggagctatctttgccattatagcaggatttgtacattgatat Decantha boraesella tctttccataggagctgtttttgctattataggagggtttattcattgatac Psilocorsis reflexella cttatctataggagctgtatttgcaattatagggggatttattcattgatat Mathildana newmanella attatcaataggggctgtatttgctattataggaggatttatccactgatat Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola tttatccataggagctgtatttgccattataggaggatttatccattgatac Ethmia trifurcella ------Nites sp 1 tttatctataggagctgtatttgcaattataggaggatttatccattgatac Depressaria pastinacella tttatctataggagccgtatttgctattataggaggatttattcactgatac Agonpterix robinella ------Pigritia sp 1 tttatcaataggagctgtatttgctattataggaggttttattcattgatat Hypatopa sp 3 tctttctataggagcagtttttgccattatagcaggatttattcactgatat Blastobasis sp 3 ------Calosima sp 1 ------

365

468 473 478 483 488 493 498 503 508 513 518 | | | | | | | | | | | Yponomeuta multipunctella cctttatttacaggattaacattaaattcttatatattaaaaattcaatttt Batrachedra enormis cctttatttactggattatcaataaatccttatttattaaaaattcaatttt Batrachedra praeangusta cccctatttacaggcttatttataaatccttatttattaaaaattcaatttt Batrachedra calator ------Coleophora ericoides ccattatttactggtctttctttaaatccttacttattaaaaattcaattta Coleophora new species ccattattcactggactttctttaaatccttatttgttaaaaattcaattta Mompha circumscriptella cctttatttacaggattaataataaatccgtatttattaaaaatccaattct Mompha eloisella cccctatttacaggaataataataaacccatatttattaaaaattcaatttt Mompha new species cctttatttacaggattaataataaatccgtatttattaaaaatccaattct Pterolonche sp 1 ccattattttttggtatatcattaaat------Triclonella pergandeella cctttattcacaggattatcaatgaattcctatttattaaaaattcaatttt Eteobalea serratella ccattatttacaggattatctttaaatccttatcttttaaaaattcaatttt Cosmopterix sp 1 ------Periploca laetella ccactttttactggaattactttaaataattatttattaaaaattcaatttg Walshia miscecolorella ccattatttacaggattaataataaataattacctattaaaaattcaatttt Sitotroga cerealella cccttatttacaggtttatctttaaatccttatcttttaaaaattcaatttt Gelechiinae 2 cctcttttcactggtttatctttaaacccttattttcttaaaatccaatttt Gelechia sp 1 cctctttttacaggactttctttaaatccttttttattaaaaattcatttta Gelechia sp 4 cccc------Teleiodini sp 1 ccattatttacaggattaactataaataatacatgattaaaaatccaattca Filatima sp 1 cctctttttactggtttatctttaaatccttatcttttaaaaattcaattct Paralita group sp 1 ccattatttactggattatctcttaatccttatatattaaaaattcaatttt Dichomeris ligulella cctttatttacaggactttctttaaacccatatttattaaaaattcaattta Gelechia sp 3 ------Asymmetura sp 1 ccattatttataggattaaatttaaattcatatttattaaaaattcaatttt Scythris limbella ccattatttacaggattaactataaataatacatgattaaaaatccaattca Antaeotricha schlaegeri cctttatttacaggattaaatataaatccttatttattaaaaattcaatttt Oegoconia quadripuncta cctttattttcaggcttaatattaaacccttacttattaaaaattcaatttt Decantha boraesella ccattatttacaggattaactttaaataattatcttctcaaaattcaattta Psilocorsis reflexella ccattatttaccggactttcaataaatccttatttattaaaaattcaatttt Mathildana newmanella ccattatttacaggacttttaataaataattacttattaaaaatccaattta Epicallima argenticinctella ------Fabiola shaleriella ------Ethmia monticola cctttatttacaggtttacttataaatccttacttattaaaaattcaatttt Ethmia trifurcella ------Nites sp 1 cctttatttacaggattaataataaatcaatatatattaaaaattcaatttt Depressaria pastinacella ccattatttacaggattaactataaatcaatatttattaaaaattcaatttt Agonpterix robinella ------Pigritia sp 1 cctttatttacaggattatctttaaattcttacttattaaaaattcaatttt Hypatopa sp 3 ccattatttactggtttatcactaaattcttacttattaaaaattcaatttt Blastobasis sp 3 ------Calosima sp 1 ------

366

520 525 530 535 540 545 550 555 560 565 570 | | | | | | | | | | | Yponomeuta multipunctella taattatatttattggagtaaatttaaca-ttttttccccaacac------Batrachedra enormis taattatattccttggagt------Batrachedra praeangusta taattatatttttaggagttaatttaaca-ttttttccccaacatcctgctt Batrachedra calator ------ccggcta Coleophora ericoides ttacaatattt-taggagt------Coleophora new species tttcaatatttattggtgtaaatttaact-ttttttcctcaacacccggcta Mompha circumscriptella ttacaatatttattggagtaaacttaac------Mompha eloisella taacaatatttattggggtaaatttaaca-tttt-ccctcaacac------Mompha new species ttacaatatttattggagtaaacttaact-tttttcccacaacac------Pterolonche sp 1 ------ccggctc Triclonella pergandeella ttattatatttattggagtaaatttaact-ttcttccctcaacacccggctt Eteobalea serratella ttacaatatttattggagttaatttaaca-ttttttccccaacatccagcaa Cosmopterix sp 1 ------ccggctt Periploca laetella ttactatatttattggggttaatttaact-ttttttcctcaacatccagcat Walshia miscecolorella ttattatatttattggggttaacttaaca-ttttttccacaacac------Sitotroga cerealella ttacaatatttattggagtaaacttaact-ttcttcccccaacatccagcaa Gelechiinae 2 ttacaatatttattggggttaatctaact-tttttcccccaacacccggcaa Gelechia sp 1 ttattatatttattggtgttaatttaaca-ttttttcccctac---cggcta Gelechia sp 4 ------ccggcta Teleiodini sp 1 caactatatttattggagtaaacttaaca-ttcttccctcaacac------Filatima sp 1 ttactatatttattggagttaatttaact-ttttttcctcaacac------Paralita group sp 1 ttacaatatttattggagttaaccttacg-ttctttcctcaacac------Dichomeris ligulella ttattatatttattggagttaatttaacc-ctttttccc------Gelechia sp 3 ------ccggcta Asymmetura sp 1 ttattatatttttaggagttaatttaacc-tttttccctttaattccggcta Scythris limbella caattatatttattggagtaaacttaaca-ttcttc------Antaeotricha schlaegeri ttactatatttattggggttaatttaact-tttttcccccaacatccagcaa Oegoconia quadripuncta tttcaatattcattggagttaacctaacc-tttttcccccaacatccagcta Decantha boraesella ttattatatttttaggggtaaattta------ccggcta Psilocorsis reflexella ttacaatatttattggagttaatttaacc-ttttttcct---tac--ggcta Mathildana newmanella ttattatatttttaggtgttaatttaaccatttttcccccaacac------Epicallima argenticinctella ------ccggcaa Fabiola shaleriella ------ccggcta Ethmia monticola ttacaatatttattggggtaaat------cta Ethmia trifurcella ------ccggcta Nites sp 1 ttttaatatttattggagtaaatttaact-tttttcccacaacacccggcaa Depressaria pastinacella tcactatatttattggagtaaatttaaca-ttcttccctcaacacccggcaa Agonpterix robinella ------ccggcaa Pigritia sp 1 tttctatattttttggagtaaatttaact-tttttccctcaacatcctgcaa Hypatopa sp 3 tttctatattcttaggagtaaatttaact-ttctttccccaacac------Blastobasis sp 3 ------ccggcta Calosima sp 1 ------ccggcta

367

572 577 582 587 592 597 602 607 612 617 622 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis ------cattgcccttccttctttacgattattatatttacttg- Batrachedra praeangusta taactttaatttttattgcaattccatctttacgattattatatttactcg- Batrachedra calator taactttaatttttattgctcttccttctttacgattattatatttattgg- Coleophora ericoides ------Coleophora new species ttacattaatttttattgctctcccttctcttcgtcttttatatttattag- Mompha circumscriptella ------catcattacgacttctttattgactag- Mompha eloisella ------attgcacttccttctttacgtttactttatttattag- Mompha new species ------Pterolonche sp 1 ttaccctaatttttattgctcttccatcattacgtcttctatatctactag- Triclonella pergandeella tcacattaatctttattgcacttccctctttacgattactttatttactag- Eteobalea serratella ttactttaatttttattgctcttccttctcttcgattactttatttattag- Cosmopterix sp 1 ttacattaatctttattgcattaccatcattacggttactttatttgttag- Periploca laetella ttactttaatttttattgctattccatcattacgattactatatttattag- Walshia miscecolorella ------Sitotroga cerealella tcacattaatttttattgctctaccatcattacgtctactctatcttttag- Gelechiinae 2 ttactttaatttttattgccttaccttctcttcgacttttatacttattag- Gelechia sp 1 ttacattaatttttattgctttaccttctctccgattactttatttattgg- Gelechia sp 4 ttactttaatttttattgctttaccctcactacgacttttatatttattag- Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 ttactttaatttttattgcactcccatctcttcgacttttatatttattag- Asymmetura sp 1 ttactttaatttttattgctttaccttctttacgattattatatttattag- Scythris limbella ------Antaeotricha schlaegeri ttactttaatttttattgcactaccttctttacgtttattatatcttttag- Oegoconia quadripuncta ttactttaattttcatcgctcttccttcccttcgtcttctctatcttcttg- Decantha boraesella tttctttaattttcatcgctcttccttcccttcgtcttctctatcttcttg- Psilocorsis reflexella ttactttaatttttattgctcttccctctcttcgattactttatcttttag- Mathildana newmanella ------Epicallima argenticinctella tcacattaatttttattgcattaccttctcttcgtcttctttatcttttag- Fabiola shaleriella ttactttaatttttattgctttaccttcacttcgccttctttatttattag- Ethmia monticola ttactttaatttttattgctttaccctctcttcgtttattatacctcttag- Ethmia trifurcella ttactctaatctttattgccctaccctctcttcgacttttatatcttttag- Nites sp 1 ttactctaatttttattgcattaccatctcttcgtcttctttaccttttag- Depressaria pastinacella ttactttaatttttattgctttaccatcactacgacttctttacctattag- Agonpterix robinella ttacattaatttttattgcattaccatctcttcgtctcctttatttattag- Pigritia sp 1 ttacattaatttttattgctcttccatctcttcgtttattatatttattagg Hypatopa sp 3 ------Blastobasis sp 3 ttacattaatttttattgctttaccttctcttcgtttattatatttattag- Calosima sp 1 ttacattaatttttattgctctcccttctcttcgtcttttatatttattag-

368

624 629 634 639 644 649 654 659 664 669 674 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis atgaacttaataatcctttaattactctaaaatctattggacatcaatgata Batrachedra praeangusta atgaacttaataatcctttaattactcttaaatccattggtcatcaatgata Batrachedra calator atgaattaaataatcctttaattaccttaaaatctattggtcatcaatgata Coleophora ericoides ------Coleophora new species atgaattaaataatcccttaattactttaaaatctattggtcatcaatgata Mompha circumscriptella atgaattaaataacccattaatcacattaaaaacaaaaggccatcaatgata Mompha eloisella atgatttaaataatcctttaattacattaaaaactatcggacaccaatgata Mompha new species ------Pterolonche sp 1 atgaaatcaataatccattaattactttaaaatcaattggacatcaatgata Triclonella pergandeella acgaactaaataatcctttaattacattaaaatctattggacatcaatgata Eteobalea serratella atgaattaaataatcctttaattactttaaaaacaattggacatcaatgata Cosmopterix sp 1 atgaattaaataatcctttaattactttaaaaacaattggacatcaatgata Periploca laetella atgaaattaataatcctttaattactttaaaatctattggtcatcaatgata Walshia miscecolorella ------Sitotroga cerealella aagaacttaataatcctttaattactttaaaatcaattggtcatcaatgata Gelechiinae 2 atgaacttaacaatcctttaattacccttaaatctattggccatcaatgata Gelechia sp 1 atgaattaaataatcctttaattaccttaaaatctattggtcatcaatgata Gelechia sp 4 atgaacttaataatcctttaattactttaaaatctattggccaccaatgata Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 atgaattaaataatccattaatcactttaaaatctattggacatcaatgata Asymmetura sp 1 atgaattaaataatcctattattacattaaaatctattgggcaccaatgata Scythris limbella ------Antaeotricha schlaegeri atgaattaaataatcctttaattaccttaaaatctattggacatcaatgata Oegoconia quadripuncta atgaattaaataatcctctaattactctaaaatctatcggacaccaatgata Decantha boraesella atgaattaaataatcctctaattactctaaaatctatcggacaccaatgata Psilocorsis reflexella atgaacttaataatcctttaatcacattaaaatcaattggtcatcaatgata Mathildana newmanella ------Epicallima argenticinctella atgaacttaataaacctttaattactttaaaatctattggacatcaatgata Fabiola shaleriella atgaattaaataaccccttaattacattaaaatcaattggtcatcaatgata Ethmia monticola atgaacttaataatcctcttattaccttaaaatcaattggtcatcaatgata Ethmia trifurcella atgaacttaataatcctttattaactttaaaatctattggtcatcagtgata Nites sp 1 atgaacttaataatccattaattacattaaaatcaattggtcatcaatgata Depressaria pastinacella atgaacttaataacccattaattactttaaaatctattggtcatcaatggta Agonpterix robinella atgaacttaataatcctttaattacattaaaatctattggccatcaatgata Pigritia sp 1 atgaattaaataatccattaattactttaaaatctattggtcatcaatgata Hypatopa sp 3 ------Blastobasis sp 3 atgaattaaataatcctttaattactctaaaatcaattggacatcaatgata Calosima sp 1 atgaattaaataatcccttaattactttaaaatctattggtcatcaatgata

369

676 681 686 691 696 701 706 711 716 721 726 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis ttgaagttatgaatattctgattttaaaaatattgaatttgactcttatata Batrachedra praeangusta ttgaagttatgaatattctgattttaaaaatattgaatttgattcttatata Batrachedra calator ttgaagatatgaatactctgattttaaaaacattgaatttgattcttatata Coleophora ericoides ------Coleophora new species ttgaagctatgaatattcagattttaataatattgaatttgattcttatata Mompha circumscriptella ttgaagatatgaatattctgattttaataatattgaatttgattcatacata Mompha eloisella ttgaagatatgaatactctgattttaataatattgaattcgattcttatata Mompha new species ------Pterolonche sp 1 ctgaagatatgaatattcagacttcaataatattgaatttgattcttatata Triclonella pergandeella ttgaagatatgaatattcagatttttacaatattgaatttgattcatatata Eteobalea serratella ttgaagatatgaatattctgattttaataatattgaatttgattcctatata Cosmopterix sp 1 ttgagcttatgaatattcagattttacaaatattcaatttgattcatatata Periploca laetella ttgaagatatgaatattcagattttaataatattgaatttgattcttatata Walshia miscecolorella ------Sitotroga cerealella ttgaagttatgaatactcagactttaaaaatattgaatttgattcatatata Gelechiinae 2 ctgaagttatgaatactcagatttcaaaaatattgaatttgactcatatata Gelechia sp 1 ttgaagttatgaatattcagattttaataatattgaatttgattcttatata Gelechia sp 4 ttgaagttatgaatactcagatttcaaaaatattgaatttgattcctatata Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 ttgaagttatgaatattctgattttaataatattgaatttgattcatatata Asymmetura sp 1 ttgaaattatgaatattctgattttaataatattgaatttgattcatacata Scythris limbella ------Antaeotricha schlaegeri ttgaagttatgaatattcagactttaaaaatattgaatttgattcttatata Oegoconia quadripuncta ttgaagctatgaatattcagattttcacaacattgaatttgattcttacata Decantha boraesella ttgaagctatgaatattcagattttcacaacattgaatttgattcttacata Psilocorsis reflexella ttgaagttatgaatattcagattttaataatattgaatttgactcatatata Mathildana newmanella ------Epicallima argenticinctella ttgaagttatgaatattcagattttaataatattgaatttgattcatatata Fabiola shaleriella ttgaagttatgaatattctgattttaataatattgaatttgactcttatata Ethmia monticola ttgaagttatgaatattcagattttaataatattgaatttgattcttatata Ethmia trifurcella ttgaagatatgaatattcagattttaacaatattgaatttgattcatatata Nites sp 1 ttgaagttatgaatactcagactttaataacattgaatttgattcatatata Depressaria pastinacella ttgaagttatgaatattcagatttcaataatatcgaatttgactcttatata Agonpterix robinella ttgaagttatgaatattcagattttaacaatattgaatttgattcttatata Pigritia sp 1 ttgaagctatgaatattctgattttaataatattgaatttgattcatatata Hypatopa sp 3 ------Blastobasis sp 3 ttgaagatatgaatattctgactttaataatattgaatttgactcatatata Calosima sp 1 ttgaagctatgaatattcagattttaataatattgaatttgattcttatata

370

728 733 738 743 748 753 758 763 768 773 778 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis atcccatcaaatgaattaaataaaaataataactttcgtcttcttgatgttg Batrachedra praeangusta atcccctctaatgaattaaataaaaataataacttccgtttacttgatgtag Batrachedra calator attccatcaaatgaattaaataaaaataataattttcgtttacttgatgtag Coleophora ericoides ------Coleophora new species atccctaataatcaaataaatcc---taataattttcgtttattagatgtag Mompha circumscriptella atttcatctaatgatttaaattc---ttctagatttcgtttattagatgtag Mompha eloisella atttcttcaaatgaattattacc---ttctagatttcgattattagatgttg Mompha new species ------Pterolonche sp 1 atcccctcaaatgacttatctta---taatgaatttcgtttattagatgttg Triclonella pergandeella attccctccaatgaattaaactc---taataattttcgattattagatgttg Eteobalea serratella atcccatataat-aattcttc-----aaataattttcgtttattagatgtag Cosmopterix sp 1 attaactttgataatttaaatca---aaatagattccgattattagatgtag Periploca laetella attccttcaaatgaattacctcaaaataat---tttcgattattagacgtag Walshia miscecolorella ------Sitotroga cerealella attccatctaatgaactttctaa---taataattttcgactattagaagttg Gelechiinae 2 attccctcaactgatttaaattc---aaataattttcgactattagatgttg Gelechia sp 1 atcccctcaaatgatttatcatt---aaataattttcgacttttagatgttg Gelechia sp 4 atccccacaaatgatttatcccc---taataattttcgactattagatgtgg Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 attccaactaacgatttatcaaa---taataatttccgattattagatgtag Asymmetura sp 1 attcctcaaaatgaaataaatat---taataattttcgattattagatgttg Scythris limbella ------Antaeotricha schlaegeri atccctcaaaatgaaataaataa---aaatagatttcgattattagatgttg Oegoconia quadripuncta attccaattaaaaatatatcccccaataat---tttcgtttattagatgtag Decantha boraesella attccaattaaaaatatatcccc---caataattttcgtttattagatgtag Psilocorsis reflexella attccatctaatgaaataatccc---aaataattttcgattattagatgttg Mathildana newmanella ------Epicallima argenticinctella atcccatctaatgaaatatcccc---taataattttcgattattagatgtcg Fabiola shaleriella atcccatcaaatgaaataatacc---taacaattttcgcttattagatgtag Ethmia monticola atcccctctaatgaattatctaa---taataatttccgattattagaagtag Ethmia trifurcella atcccatctaatgatttacttaa---taatagatttcgtttattagatgttg Nites sp 1 attccatcaaatgatttaacacc---taataattttcgtttattagatgtag Depressaria pastinacella atcccctcaactgatttatcaat---taatgggtttcgactattagatgtag Agonpterix robinella attccttctaatgaactttctcc---taataatttccgattactagatgttg Pigritia sp 1 atcccctcaaatgaaataaatat---taataactttcgtttattagaagttg Hypatopa sp 3 ------Blastobasis sp 3 attccttctaatgaaattaatat---taataattttcgtcttttagaggtag Calosima sp 1 atccctaataatcaaataaatcc---taataattttcgtttattagatgtag

371

780 785 790 795 800 805 810 815 820 825 830 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis ataatcgaattattttaccaataaataatcaaattcgaattttaattactgc Batrachedra praeangusta ataatcgaattgttcttcctataaataatcaaattcgtattttaattactgc Batrachedra calator ataatcgaattattttacctataaataatcaaattcgaattttaattacttc Coleophora ericoides ------Coleophora new species ataatcgaattattttaccaataaataaccaaattcgaattttagtaacagc Mompha circumscriptella ataatcgaattattctaccttacaataatcaaattcgaattcttgtaactgc Mompha eloisella ataatcgagttattttaccatttaataatcaaattcgaattttagtcactgc Mompha new species ------Pterolonche sp 1 ataatcgaattattttaccaataaataaccaaattcgaattttagttacagc Triclonella pergandeella ataatcgaattattctacctataaataatcaaattcgaattttaattacagc Eteobalea serratella ataatcgaattattttacctataaataatcaaattcgtatcttaataacatc Cosmopterix sp 1 ataatcgtgtagtattaccaataaataaccaaattcgaattttaattacagc Periploca laetella ataatcgaattattttacctataaataatcaaattcgaattttagttactgc Walshia miscecolorella ------Sitotroga cerealella ataatcgaattattttaccaataaataatcaaattcgaattatagtaactgc Gelechiinae 2 ataatcgaattattctccctataaataatcaaattcgaattttagttactgc Gelechia sp 1 ataatcgaattgttttacctataaataatcaaattcgtattttagtgacagc Gelechia sp 4 ataatcgtattattcttcccataaataaccaaattcgaattttagttacagc Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 ataatcgaattgtattacctataaaaaatcaaattcgtattttagttactgc Asymmetura sp 1 ataaccgaattactttacctataaaaaatcaaattcgaattatagttaccgc Scythris limbella ------Antaeotricha schlaegeri ataatcgaattattttaccaataaataaccaaattcgaattatagtaacagc Oegoconia quadripuncta ataatcgaattgttttacctataaataatcaaattcgaattataatcacagc Decantha boraesella ataatcgaattgttttacctataaataatcaaattcgaattataatcacagc Psilocorsis reflexella ataatcgtattattttacctataaataatcaaattcgaattttagttactgc Mathildana newmanella ------Epicallima argenticinctella ataatcgaattattttaccaataaaaaaccaaattcgaattttagttacagc Fabiola shaleriella ataatcgaattattttacctataaaaaatcaaattcgtattatagttactgc Ethmia monticola ataatcgaattattttaccaataaataatcaaattcgaattttagtcactgc Ethmia trifurcella ataatcgtattattctacctataaataatcaaattagaattttagttacagc Nites sp 1 ataatcgaattattttaccaataaataaccaaattcgaattatagttacggc Depressaria pastinacella ataaccgaattgttatacctataaataaccaaattcgaattatagttacagc Agonpterix robinella ataatcgaattattttaccaataaataatcaaattcgaattttagttacagc Pigritia sp 1 ataaccgtattattcttcctataaataaccaaattcgaattatagttacagc Hypatopa sp 3 ------Blastobasis sp 3 ataatcgtattattttacccataaataatcaaattcgaattatagttacagc Calosima sp 1 ataatcgaattattttaccaataaataaccaaattcgaattttagtaacagc

372

832 837 842 847 852 857 862 867 872 877 882 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis tactgatgtaattcattcatgaactattccttctttaggagtaaaagttgat Batrachedra praeangusta aacagatgtaattcattcttgaactattccttctttaggagttaaagttgat Batrachedra calator aactgatgtaattcattcatgaactattccttctttaggagtaaaagttgat Coleophora ericoides ------Coleophora new species tacagatgttattcattcatgaactattccttctttaggggttaaagtagat Mompha circumscriptella ttcagatgtaattcactcatgaactatccctcattaggagt-aaaagtagac Mompha eloisella tacagatgtgattcactcttgaactattcatctttaggggtaaaaa-ttgat Mompha new species ------Pterolonche sp 1 tacagatgtcattcactcatgaactattccttcattaggagtaaaagttgat Triclonella pergandeella tacagatgttattcattcatgaaccgttccctctctaggggtaaaagtagat Eteobalea serratella aacagatgttattcactcttgaactattccatcattaggtgtaaaaacagat Cosmopterix sp 1 aacagatgtaattcattcatgaacagtaccttgtttaggtgtaaaagtagat Periploca laetella tactgatgttattcattcatgaactattccttcattaggagtaaaagttgat Walshia miscecolorella ------Sitotroga cerealella tacagatgtaattcattcttgaactattccttctttaggaattaaagttgat Gelechiinae 2 tactgatgttattcactcatgaactattccctcattaggagtgaaagtagat Gelechia sp 1 aactgatgttattcattcttgaactattccatctttaggagtaaaagtagat Gelechia sp 4 tactgatgttattcattcatgaacaattccatcattaggagtcaaagttgat Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 aactgatgtaattcattcatgaactgtaccttctttaggtgtaaaagtagat Asymmetura sp 1 cactgatgttattcattcttgaactattccatcattaggtattaaaattgat Scythris limbella ------Antaeotricha schlaegeri aactgatgtaattcattcatgaactattccatccttaggaattaaagtagat Oegoconia quadripuncta tacagatgtaatccattcttgaactattccatccctaggaattaaagttgat Decantha boraesella tacagatgtaatccattcttgaactattccatccctaggaattaaagttgat Psilocorsis reflexella aactgatgttattcactcatgaacaattcctgcattaggagttaaaattgat Mathildana newmanella ------Epicallima argenticinctella aacagatgttattcattcatgaacaattccagccctgggggtaaaagtagat Fabiola shaleriella tacagatgtaattcactcttgaactatcccatccttagggattaaagtagac Ethmia monticola tacagatgtaatccactcatgaacaatcccctcattaggggtaaaagttgat Ethmia trifurcella tacagatgttattcattcctgaacagtaccatctttaggaattaaagtagat Nites sp 1 tacagatgtaattcattcatgaactatcccatctttaggggttaaagtagat Depressaria pastinacella aactgatgtaattcattcatgaactgtaccatctttaggggtaaaagtagat Agonpterix robinella aacagatgtaattcattcttgaactatcccttctttaggagtaaaagtagac Pigritia sp 1 cacagatgttattcactcttgaactattccttctctaggtgtaaaaattgac Hypatopa sp 3 ------Blastobasis sp 3 cacagatgtaattcattcctgaactattccttccttaggggtaaaaattgat Calosima sp 1 tacagatgttattcattcatgaactattccttctttaggggttaaagtagat

373

884 889 894 899 904 909 914 919 924 929 934 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis gcaaatccaggtcgtttaaatcaaactaatttttttattaatcgacctggaa Batrachedra praeangusta gctaatcctggtcgtttaaatcaaactaatttttttataaatcgcccaggaa Batrachedra calator gctaacccaggtcgtttaaatcaaactaatttttttataaatcgaccaggaa Coleophora ericoides ------Coleophora new species gctaatcctggtcgtcttaatcaaataagattttttattaatcgtcctggta Mompha circumscriptella gctaatcctggacgactaaaccaaactaacttctttattaaccgtccaggta Mompha eloisella gctaatcctggccgattaaatcaaactaacttttttatcaatcgacctggga Mompha new species ------Pterolonche sp 1 gctaatcctggacgattaaatcaaactaatttctttttaaatactccaggaa Triclonella pergandeella gctaatccaggacgattaaatcaatcaagtttttttattaaccgaccaggaa Eteobalea serratella gctaatcctggtcgtcttaatcaatctagatttttcattaatcgtcctggaa Cosmopterix sp 1 gctaatccaggacgattaaatcaaaccaattttttcttaaatcgacctggaa Periploca laetella gctaatcctggccgacttaatcaaactagctttcttattaatcgaccaggaa Walshia miscecolorella ------Sitotroga cerealella gctaatcctggtcgt-taaatcaaac------Gelechiinae 2 gcaaatccaggacgtctcaatcaaactaatttttttattaatcgaccaggaa Gelechia sp 1 gctaatcctggccgactaaatcaaacaaacttttttattaatcgacccggaa Gelechia sp 4 gctaatcctggacgtttaaatcaaactaatttttttattaatcgtccaggta Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 gctaatccaggacgtcttaatcaaactaattttttcattaatcgcccaggaa Asymmetura sp 1 gctaatccaggtcgattaaatcaaacaaatttttttattaatcgacctggaa Scythris limbella ------Antaeotricha schlaegeri gctaatccaggtcgattaaatcaaacttccttttttattaatcgaccaggta Oegoconia quadripuncta gctaatcctggtcgtcttaatcaaactagcttttttattaatcgaccaggaa Decantha boraesella gctaatcctggtcgtcttaatcaaactagcttttttattaatcgaccaggaa Psilocorsis reflexella gctaatccaggtcgactaaatcaaactaatttttttattaaccgacctggaa Mathildana newmanella ------Epicallima argenticinctella gctaatccaggacgtttaaatcaaacaaattttttcattaatcgaccaggaa Fabiola shaleriella gctaatcctggtcgattaaatcaaactaatttttttattaatcgacctggta Ethmia monticola gccaatcctggtcgactaaatcaaactaatttttttattaaccgaccaggaa Ethmia trifurcella gctaatccaggacgtcttaatcaaactagattttttattaatcgaccaggaa Nites sp 1 gctaatccaggtcgattaaatcaaacaaatttttttattaatcgacctggaa Depressaria pastinacella gctaatccaggtcgactcaatcaaacaaatttttttattaaccgacctggtg Agonpterix robinella gctaatcctggtcgtttaaatcaaacaaatttttttattaatcgaccaggaa Pigritia sp 1 gctaatccaggacgtctcaatcaaacaagattttttattaatcgacctggaa Hypatopa sp 3 ------Blastobasis sp 3 gctaatcctggccgattaaatcaaactagattttttattaatcggccaggaa Calosima sp 1 gctaatcctggtcgtcttaatcaaataagattttttattaatcgtcctggta

374

936 941 946 951 956 961 966 971 976 981 986 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis tttttt-atggtcaatgttcagaaatctgcggagcaaa----tcatagtttt Batrachedra praeangusta ttttct-atggtcaatgctcagaaatttgtggagctaa----tcatagtttt Batrachedra calator ttttttt-tggtcaatgctctgaaatttgtggagcaaa----tcatagattt Coleophora ericoides ------Coleophora new species tttttta-tgggcaatgttctgaaatttgtggagcaaa----tcatagtttt Mompha circumscriptella ttttttt-tggacaatgttcagaaatttgtggagcaaa----tcatagattt Mompha eloisella tttttta-tggtcaatgctcagaaatttgtggggctaa----tcatagattt Mompha new species ------Pterolonche sp 1 tttttta-tggacaatgttcagaaatttgtggtgcaaa----tcatagattt Triclonella pergandeella tttttta-tggtcaatgttcagaaatttgtggagctaa----tcatagattt Eteobalea serratella tttatt-ttggacaatgttcagaaatttgtggagctaa----tcatagtttt Cosmopterix sp 1 tttttta-tggacaatgttcagaaatttgtggagcaaa----tcacagattt Periploca laetella tttttgtatggacaatgttcagcacaattgtgtggggcaagatcatagtttt Walshia miscecolorella ------Sitotroga cerealella ------Gelechiinae 2 tttttta-tggccaatgttctgaaatttgtggggcaaa----tcatagattt Gelechia sp 1 ttttttt-tgggcaatgttctgaaatttgtggagctaa----ccatagtttt Gelechia sp 4 ttttctt-tggtcaatgttctgaaatttgtggtgctaa----tcatagtttt Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 tttttta-tggtcaatgttcagaaatttgtggagctaa----tcatagattt Asymmetura sp 1 tttattt-tggacaatgttctgaaatttgtggagcaaa----tcatagattt Scythris limbella ------Antaeotricha schlaegeri tttttt-atgggcaatgttcagaaatttgtggagcaaa----tcatagattt Oegoconia quadripuncta ttttttt-tggtcaatgttctgaaatttgcggagctaa----tcatagtttt Decantha boraesella ttttttt-tggtcaatgttctgaaatttgcggagctaa----tcatagtttt Psilocorsis reflexella ttttttt-cggacaatgttcagaaatttgtggagcaaa----tcatagtttt Mathildana newmanella ------Epicallima argenticinctella tttatta-tggacaatgctcagaaatttgtggagctaa----tcatagtttt Fabiola shaleriella ttttttt-tgggcaatgttcagaaatttgtggggccaa----tcatagtttt Ethmia monticola tttttta-cggacaatgttctgaaatttgtggagctaa----tcatagattt Ethmia trifurcella tttttta-tggacaatgttcagaaatttgtggagctaa----tcatagtttt Nites sp 1 ttttcta-tggtcaatgttcagaaatttgtggagcaaa----tcatagattt Depressaria pastinacella tatttta-tggtcaatgttcagagatttgtggagcaaa----tcatagattt Agonpterix robinella tttttta-tggtcaatgttcagaaatttgtggagcaaa----tcatagtttt Pigritia sp 1 tttttt-atggtcaatgttcagaaatttgtggtgcaaa----tcatagtttt Hypatopa sp 3 ------Blastobasis sp 3 tttttta-tggccaatgttcagaaatttgtggggcaaa----tcatagattc Calosima sp 1 tttttta-tgggcaatgttctgaaatttgtggagcaaa----tcatagtttt

375

988 993 998 1003 1008 1013 1018 1023 1028 1033 1038 | | | | | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis atacct-attgtaatcgaaagaatttc-aattaaaaattttattaat-tgaa Batrachedra praeangusta atacct-attgtaatcgaaagaatttcagattaaaagtttcattaat-tgaa Batrachedra calator atacct-attgtaattgaaagaatttc-aattaaaaaattcattaat-tgaa Coleophora ericoides ------Coleophora new species atacct-attgtaattgaaagaatttc-aattaaaaattttattaat-tgaa Mompha circumscriptella atacct-attttaattgaaagaatttc-aattaaaaattttattaat-tgaa Mompha eloisella atacca-attatagtagaaagaattcc-aattaaaaattttattaat-tgaa Mompha new species ------Pterolonche sp 1 atacca-atcgtaattgaaagtattca-tattaaaaattttattaat-tgaa Triclonella pergandeella atacct-attgttattgaaagtattcc-tataataaattttattaat-tgaa Eteobalea serratella atacct-attgtaattgaaagaatttc-aattaaaaactttattaaa-tgaa Cosmopterix sp 1 atacct-attgtaattgagagagtatc-aataagaaatttcattaaa-tgaa Periploca laetella atacctcattgtaattgaaagaatttc-aattatt------Walshia miscecolorella ------Sitotroga cerealella ------Gelechiinae 2 atacca-attgtaattgaaagtatttc-aattaaaaattttattaat-tgaa Gelechia sp 1 atacca-attgtaattgaaagaatttc-aattaataactttattaat-tgag Gelechia sp 4 atacct-attgtaattgaaagtatttc-aattaaaaattttattaat-tgaa Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 atacct-attgtaattgaaagaatttc-aattaaaaattttattaat-tgaa Asymmetura sp 1 atacct-attgtaattgaaagaattcc-tattaaaaattttattaat-tgaa Scythris limbella ------Antaeotricha schlaegeri atacct-attataattgaaagaatttc-aattaaaaactttattaaa-tgaa Oegoconia quadripuncta ataccc-attataattgaaagtatctc-aattaaaaattttattaaa-tgaa Decantha boraesella ataccc-attataattgaaagtatctc-aattaaaaattttattaaa-tgaa Psilocorsis reflexella atacct-attgtaattgaaagaatttc-aattaaaaattttattaat-tgaa Mathildana newmanella ------Epicallima argenticinctella atgcct-atagtagtagaaagaattca-tattaaaaattttattaat-tgaa Fabiola shaleriella atgcct-attgtaattgaaagaattta-tattaaaaattttattaac-tgaa Ethmia monticola atacct-attgtaattgaaagaatttc-tattaaaaatttcattaat-tgaa Ethmia trifurcella atacct-attgtaattgaaagaatctc-aatcaaaaactttattaatatgaa Nites sp 1 atacca-attgtaattgaaagaatctc-aattaaaaattttattaat-tgaa Depressaria pastinacella atacct-attgtaattgaaagaatctc-aattaaaaattttattaat-tgaa Agonpterix robinella atacct-attgtaattgaaagaatttc-aattaaaaattttattaat-tgaa Pigritia sp 1 atacct-attgt------Hypatopa sp 3 ------Blastobasis sp 3 atacct-atcgtaattgaaagtatctc-aattaaaaatttcattaat-tgaa Calosima sp 1 atacct-attgtaattgaaagaatttc-aattaaaaattttattaat-tgaa

376

1040 1045 1050 1055 1060 1065 1070 | | | | | | | Yponomeuta multipunctella ------Batrachedra enormis ttaataatttttcttcat------Batrachedra praeangusta ttaataatt------Batrachedra calator ttaataattattct-tcattagatgactgaaa Coleophora ericoides ------Coleophora new species ttaataactattct-tcattagatgactgaaa Mompha circumscriptella ttaataa------Mompha eloisella ttaataattattca-tcatta------Mompha new species ------Pterolonche sp 1 ttaaaaattactct-tcattagatgactgaaa Triclonella pergandeella ttaaaaactattca-tcattagatgactgaaa Eteobalea serratella ttaataattattcttcatt------Cosmopterix sp 1 ttaacaattattct-tcattagatgactgaaa Periploca laetella ------cttcattagatgactgaaag Walshia miscecolorella ------Sitotroga cerealella ------Gelechiinae 2 ttaataattattct-tcattagatgactgaaa Gelechia sp 1 ttaataattattct-tcattagatgactgaaa Gelechia sp 4 ttaataattactcc-tcattagatgactgaaa Teleiodini sp 1 ------Filatima sp 1 ------Paralita group sp 1 ------Dichomeris ligulella ------Gelechia sp 3 ttaataattattct-tcattagatgactgaaa Asymmetura sp 1 ttaataattattct-tcattagatgactgaaa Scythris limbella ------Antaeotricha schlaegeri ttaataattattaaatgaattaataatta--- Oegoconia quadripuncta ttaataatta------Decantha boraesella ttaataattattca-tcattagatgactgaaa Psilocorsis reflexella ttaataattattca-tcattagatgactgaaa Mathildana newmanella ------Epicallima argenticinctella ttaataattattca-tcattagatgactgaaa Fabiola shaleriella ttaacaattactca-tcattagatgactgaaa Ethmia monticola ttaataattattca-tcattagatgactgaaa Ethmia trifurcella ttaataattattct-tcattagatgactgaaa Nites sp 1 ttaataactattcc-tcattagatgactgaaa Depressaria pastinacella ttaataattattct-tcattagatgactgaaa Agonpterix robinella ttaataattattca-tcattagatgactg--- Pigritia sp 1 ------Hypatopa sp 3 ------Blastobasis sp 3 ttaataattattcc-tcattagatgactgaaa Calosima sp 1 ttaataactattct-tcattagatgactgaaa

377

SUMMARY PERCENTAGES

MISSING DATA (?): 0 cells, <1 percent of matrix.

DASHES (-): 12655 cells, 28 percent of matrix.

TOTAL POLYMORPHISM ($, *): 4 cells, <1 percent of matrix.

TOTAL FULL AMBIGUITY (?, -): 12655 cells, 28 percent of matrix.

TOTAL FULL + PARTIAL AMBIGUITY (?, -, *, $): 12659 cells, 28 percent of matrix.

STATE (A): 10634 cells, 23 percent of matrix.

STATE (A) EMBEDDED IN POLYMORPHISM: 4 cells, 0 percent of matrix.

STATE (C): 4481 cells, 9 percent of matrix.

STATE (C) EMBEDDED IN POLYMORPHISM: 4 cells, 0 percent of matrix.

STATE (G): 4268 cells, 9 percent of matrix.

STATE (G) EMBEDDED IN POLYMORPHISM: 4 cells, 0 percent of matrix.

STATE (T): 12982 cells, 28 percent of matrix.

STATE (T) EMBEDDED IN POLYMORPHISM: 4 cells, 0 percent of matrix.

378

APPENDIX D.

LEPIDOPTERA MOLECULAR PHYLOGENETICS

Gene Taxa Age Author Results 16s rDNA Spodoptera; Early tertiary; 1 tRNAVal and ND1most tRNALeu,Val ~65 mya – 100 conserved ND1 mt DNA mya ND 1 useful below the superfamily level 18s rDNA Ditrysia Early 2 28B (4048 – 4348 of 28s 28s rDNA Mesozoic; rDNA) variation differs ~250 mya among comparable age superfamilies, recovers test clades 18s rDNA Holometabolous 3 18s supports the monophyly Insects (362-289 mya) of Amphiesmenoptera + Diptera sister relationships ND1 mtDNA Noctuoidea Early tertiary; 4 ND1 did not recover deep 28B of 28s rDNA ~65 mya – 100 splits mya 28B did not seem to have strong signal combined gene trees show some signal CO I mtDNA Heliconius; ~65 mya – 100 5 CO I and II informative at CO II mtDNA Nymphalidae mya generic level ND 1 mtDNA Macrolepidoptera Early tertiary; 6 ND 1 useful in resolving 16s rDNA ~65 mya – 100 family level relationships 28s rDNA mya EF-1α nDNA Heliothinae; ~65 mya – 100 7 slowly evolving gene Noctuidae mya recovered recent divergences ND 1 mtDNA Papilionini; ~65 mya – 100 8 Informative 16s rDNA Papilionidae mya PEPCK Lepidoptera (moths) Mesozoic 9 Useful for Mesozoic-aged lineages

379

Gene Taxa Age Author Results EF-1α nDNA Noctuoidea Early tertiary; 10 Signal strength for basal ~65 mya – 100 divergences is weak mya CO I mtDNA Heliconius ~65 mya – 100 11 Supports traditional views of CO II mtDNA mya phylogeny Wingless nDNA

DDC nDNA Noctuoidea Early tertiary; 12 Parsimony, distance, and ~65 mya – 100 maximum-likelihood mya analyses recover nearly all “test clades” supported by morphological synapomorphies DDC nDNA Heliothinae ~65 mya – 100 12 Resolves relationships that mya are largely concordant with prior evidence from EF-1 alpha, morphology, and allozymes

COII mtDNA Josiini; Early tertiary; 13 best supported hypothesis is 18s rDNA ; ~65 mya – 100 derived from combined 28s rDNA Dioptinae mya analysis Period nDNA Lepidoptera (mostly Late K/T 14 Supported groupings at the macros and some family level and below outgroups) Wingless nDNA Nymphalidae ~65 mya – 100 15 Wg becomes saturated more mya slowly than mtDNA CO 1 and wg appear to be evolving at similar rates Informative for more recent divergences CO I Ostrinia () Early tertiary; 16 some species relationships ~65 mya – 100 remain uncertain mya CO I mtDNA Argyrotaenia; Early tertiary; 17 some species relationships CO II mtDNA Tortricidae ~65 mya – 100 remain uncertain tRNALUE mya Wingless nDNA , ~65 mya – 100 18 robust support for and mya Riodinidae and Lycaenidae, Nymphalidae not for Nymphalidae noisy 3rd position? Wingless nDNA Nymphalidae ~65 mya – 100 19 useful source of mya phylogenetic data provided there is good taxon sampling 18s rDNA Main basal lineages Mesozoic (250 20 18s highly informative about PEPCK nDNA of Lepidoptera –65 mya) basal lineages PEPCK added resolution

380

Gene Taxa Age Author Results 16s rDNA tribe Melitaeini; ~65 mya 21 CO I informative at species CO I mtDNA Nymphalinae level 16s alignment sensitive and has small regions of variation COI mtDNA (Euphydryas); ~65 mya 22 all three genes useful for ND1 mtDNA Nymphalidae recovering recent 16s rDNA divergences DDC (Nt3) nDNA Noctuoidea Early tertiary; 23 Compared to utility of EF- ~65 mya – 100 1α = faster evolving, fewer mya homoplasious sites DDC provides phylogenetic signal at all levels in Noctuoidea (family and higher)

DDC nDNA Heteroneuran Middle 24 DDC carries signal at family Lepidoptera Mesozoic; level and higher ~200 mya Failed to sequence DDC for primitive taxa/did not resolve basal divergences DDC maybe informative within DDC nDNA Noctuoidea Early tertiary; 25 Addition of taxa may not EF-1α nDNA ~65 mya – 100 resolve problems mya Support for EF-1α lower than Support for EF-1α + DDC Resolved family/subfamily level relationships

ITS2 species in the genus Early 26 although rapidly evolving, Wiseana; Mesozoic; ITS2 was not useful in ~250 mya determining species level phylogeny alone useful when combined with morphology and CO I & CO II 16s rDNA Satyridae ~65 mya – 100 27 incongruence found between ND 1 mtDNA mya morphological and molecular analyses CO II mtDNA Chrysoritis; ~65 mya 28 supports traditional Aphnaeini; morphology Lycaenidae EF-1α nDNA Early tertiary; 29 phylogenetic signal was ~65 mya – 100 strong mya

381

Gene Taxa Age Author Results CO I mtDNA Nymphalidae ~65 mya – 100 30 EF-1α is the slowest CO II mtDNA Bicyclus genus mya evolving EF-1α nDNA group CO I is the fastest evolving EF-1α is as useful as CO I and II for resolving relationships at the tips (species?) OPS1 nDNA (Heliconius); ~65 mya 31 OPS1 phylogeny consistent (eyeball gene) Nymphalidae with Brower et al 1997 COI, COII, and wingless tree CO II mt DNA Subtribe Oligocene 32 Informative at the generic Cyt b mtDNA Mycalesina; level Satyrinae ND 1 mtDNA Subfamilies of Early tertiary; 33 Used because informative at 28s rDNA (D1, D2) Geometridae ~65 mya – 100 subfamily and below mya Useful in resolving family/subfamily level relationships DDC nDNA ~65 mya – 100 34 Useful in resolving EF-1α nDNA mya family/subfamily level relationships Cyt-b mtDNA Genera of Arctiidae Early tertiary; 35 Cyt-b gives signal across the CO 1 mtDNA Insecta ~65 mya – 100 tree but has most 28s rDNA (D1) mya substitutions at the terminals CO I gives resolution at both the inter and intra specific levels D1 not very phylogenetically informative (only 10 informative characters) Much variation in Cyt-b in Insecta 18s rRNA Arthropoda 36 Reveal Class and Ordinal 28s rRNA explosion (570 level relationships H3 n – 509 mya) U2 nrRNA EF-1α POL CO I mtDNA 16s rRNA

CO I mtDNA Papilionidae tribes ~65 mya – 100 37 data partitions disagree? CO II mtDNA mya EF1α nDNA CO I mtDNA Archips; Tortricidae 38 useful in determining species group and species phylogeny allozymes Oeneis chryxus ~65 mya 39 results ambiguous CO I mtDNA complex; Satyridae

382

Gene Taxa Age Author Results ndl mtDNA Polygonia, Early tertiary; 40 useful at the generic level Wingless nDNA Nymphalis, other ~65 mya – 100 Nymphalidae mya CO I mtDNA Xylostella species 41 useful in resurrecting group of Archips; Archippus Tortricidae intron 3 Heliothine Early tertiary; 42 intron sequences evolved at triosephosphate Butterflies ~65 mya – 100 rates similar to those of isomerase (Tpi) mya mitochondrial coding intron 3 mannose- sequences phosphate isomerase (Mpi) CO I mtDNA CO II mtDNA Allozymes Lycaeninae ~65 mya 43 useful EF1α nDNA Saturniinae ~65 mya 44 weak resolution of deeper DDC nDNA divergences in Saturnini CO I mtDNA Hemiluca ~65 mya 45 two-gene phylogeny EF1α nDNA suggests that wing morphology is homoplastic CO I mtDNA Anartia ~65 mya 46 CO I and II useful in CO II mtDNA (Nymphalidae) determining interspecific tRNALUE relationships surprising results!! CO I mtDNA Acrodipsas ~65 mya 47 Morphology has evolved CO II mtDNA (Lycaenidae) faster than molecular data CO I mtDNA Lycaenidae ~65 mya 48 mDNA reveals cryptic CO II mtDNA species CO I mtDNA Phycodes ~65 mya 49 CO I useful in species (Nymphalidae) delineations of haplotypes CO I mtDNA Agrodiaetus ~65 mya 50 Agrodiaetus is monophyletic CO II mtDNA (Lycaenidae) EF1α nDNA CO I mtDNA Arhopala; ~65 mya 51 confirms monophyly and CO II mtDNA Lycaenidae; relationships; incongruence tRNALUE Theclinae with previous taxonomic wingless nDNA studies

Nuclear genes – 18s rRNA (SSU) and 28s rRNA (LSU), EF-1 alpha (protein coding), wingless, DDC, H3 (protein coding), LSU RNA polymerase II, ITS2 rRNA, PEPCK

Mitochondrial genes – 12s rRNA and 16s rRNA, ND1, ND2, CO1, CO2, Cyt-b,

383

Citations

1 Pashley, D.P. and Ke, L.D. (1992) Molecular Biology and Evolution 9, 1061-1075 2 Weller, S.J., Friedlander, T.P., Martin, J.A. and Pashley, D.P. (1992) Molecular Phylogenetics and Evolution 4, 312-337 3 Pashley, D.P. and Ke, L.D. (1993) Molecular Biology and Evolution 10, 268-268 4 Weller, S.J., Pashley, D.P., Martin, J.A. and Constable, J.L. (1994) Systematic Biology 43, 194-211 5 Brower, A.V.Z. (1994) Molecular Phylogenetics and Evolution 3, 159-174 6 Weller, S.J. and Pashley, D.P. (1995) Molecular Phylogenetics and Evolution 4, 235-246 7 Cho, S.W. et al. (1995) Molecular Biology and Evolution 12, 650-656 8 Aubert, J., Barascud, B., Descimon, H. and Michel, F. (1996) Comptes Rendus De L Academie Des Sciences Serie Iii-Sciences De La Vie-Life Sciences 319, 647-651 9 Friedlander, T.P., Regier, J.C., Mitter, C. and Wagner, D.L. (1996) Molecular Biology and Evolution 13, 594-604 10 Mitchell, A. et al. (1997) Molecular Biology and Evolution 14, 381-390 11 Brower, A.V.Z. and Egan, M.G. (1997) Proceedings of the Royal Society of London Series B-Biological Sciences 264, 969-977 12 Fang, Q.Q. et al. (1997) Systematic Biology 46, 269-283 13 Miller, J.S., Brower, A.V.Z. and DeSalle, R. (1997) Biological Journal of the Linnean Society 60, 297-316 14 Regier, J.C. et al. (1998) Molecular Biology and Evolution 15, 1172-1182 15 Brower, A.V.Z. and DeSalle, R. (1998) Insect Molecular Biology 7, 73-82 16 Kim, C.G., Hoshizaki, S., Huang, Y.P., Tatsuki, S. and Ishikawa, Y. (1999) Applied Entomology and Zoology 34, 405-412 17 Landry, B., Powell, J.A. and Sperling, F.A.H. (1999) Annals of the Entomological Society of America 92, 40-46 18 Campbell, D.L., Brower, A.V.Z. and Pierce, N.E. (2000) Molecular Biology and Evolution 17, 684-696 19 Brower, A.V.Z. (2000) Proceedings of the Royal Society of London Series B- Biological Sciences 267, 1201-1211 20 Wiegmann, B.M. et al. (2000) Molecular Phylogenetics and Evolution 15 21 Wahlberg, N. and Zimmermann, M. (2000) Cladistics-the International Journal of the Willi Hennig Society 16, 347-363 22 Zimmermann, M., Wahlberg, N. and Descimon, H. (2000) Annals of the Entomological Society of America 93, 347-355 23 Fang, Q.Q. et al. (2000) Molecular Phylogenetics and Evolution 15, 473-486 24 Friedlander, T.P., Regier, J.C., Mitter, C., Wagner, D.L. and Fang, Q.Q. (2000) Zoological Journal of the Linnean Society 130, 213-234 25 Mitchell, A., Mitter, C. and Regier, J.C. (2000) Systematic Biology 49, 202-224 26 Brown, B., Emberson, R. and Paterson, A. (2000) New Zealand Journal of Zoology 27, 1-14

384

27 Martin, J.F., Gilles, A. and Descimon, H. (2000) Molecular Phylogenetics and Evolution 15, 70-82 28 Rand, D.B., Heath, A., Suderman, T. and Pierce, N.E. (2000) Molecular Phylogenetics and Evolution 17, 85-96 29 Regier, J.C., Mitter, C., Peigler, R.S. and Friedlander, T.P. (2000) Insect Systematics & Evolution 31, 179-186 30 Monteiro, A. and Pierce, N.E. (2001) Molecular Phylogenetics and Evolution 18, 264-281 31 Hsu, R., Briscoe, A.D., Chang, B.S.W. and Pierce, N.E. (2001) Biological Journal of the Linnean Society 72, 435-449 32 Torres, E. et al. (2001) Molecular Phylogenetics and Evolution 20, 460-473 33 Abraham, D. et al. (2001) Molecular Phylogenetics and Evolution 20, 65-77 34 Regier, J.C., Mitter, C., Friedlander, T.P. and Peigler, R.S. (2001) Molecular Phylogenetics and Evolution 20, 311-316 35 Simmons, R.B. and Weller, S.J. (2001) Molecular Phylogenetics and Evolution 20, 196-210 36 Giribet, G., Edgecombe, G.D. and Wheeler, W.C. (2001) Nature 413, 157-161 37 Caterino, M.S., Reed, R.D., Kuo, M.M. and Sperling, F.A.H. (2001) Systematic Biology 50, 106-127 38 Kruse, J.J. and Sperling, F.A.H. (2001) Annals of the Entomological Society of America 94, 166-173 39 Nice, C.C. and Shapiro, A.M. (2001) Molecular Phylogenetics and Evolution 20, 111-123 40 Nylin, S. et al. (2001) Zoological Journal of the Linnean Society 132, 441-468 41 Kruse, J.J. and Sperling, F.A.H. (2002) Annals of the Entomological Society of America 95, 288-301 42 Beltran, M. et al. (2002) Molecular Biology and Evolution 19, 2176-2190 43 Pratt, G.F. and Wright, D.M. (2002) Pan-Pacific Entomologist 78, 219-229 44 Regier, J.C., Mitter, C., Peigler, R.S. and Friedlander, T.P. (2002) Insect Systematics & Evolution 33, 9-21 45 Rubinoff, D. and Sperling, F.A.H. (2002) Molecular Phylogenetics and Evolution 25, 70-86 46 Blum, M.J., Bermingham, E. and Dasmahapatra, K. (2003) Molecular Phylogenetics and Evolution 26, 46-55 47 Eastwood, R. and Hughes, J.M. (2003) Molecular Phylogenetics and Evolution 27, 93-102 48 Eastwood, R. and Hughes, J.M. (2003) Australian Journal of Zoology 51, 331- 340 49 Wahlberg, N., Oliveira, R. and Scott, J.A. (2003) Systematic Entomology 28, 257- 274 50 Kandul, N.P. et al. (2004) Systematic Biology 53, 278-298 51 Megens, H.J., Van Nes, W.J., Van Moorsel, C.H.M., Pierce, N.E. and De Jong, R. (2004) Systematic Entomology 29, 115-131

385

APPENDIX E

FEATURES THAT DEFINE HODGES SPECIES GROUPS AS CHARACTERIZED

BY BUCHELI AND PASSOA (UNPUBLISHED) IN CHAPTER 5:

NORTH AMERICAN FLAT-BODY MOTHS (ELACHISTIDAE: DEPRESSARIINAE:

DEPRESSARIA HAWORTH): MORPHOLOGICAL EVOLUTION, HOST-PLANT

SELECTION, AND GEOGRAPHIC DISTRIBUTION

0 5 | | Agonopterix 0000000 Exaeretia 0000000 Artemisiae group 1010101 Pastinacella group 1010111 Thomaniella group 1100020 Betina group 1111020 Douglasella group 1111030

0. Pupal coxae: exposed = 0; hidden = 1.

1. Aedeagus: with flange = 0; lacking flange = 1.

2. Ductus bursa meets corpus bursa: at right angle = 0; straight or curved = 1.

3. Cornuti: present = 0; absent = 1.

4. If cornuti present: fine = 0; stout = 1.

386

5. Saccal process: without basal or distal process = 0; with basal process = 1; with distal

process = 2; with both basal and distal process = 3. [nonadditive].

6. Distal process: present = 0; absent = 1.

387

APPENDIX F

COMPLETE MORPHOLOGICAL MATRIX FOR SPECIES OF DEPRESSARIA AND

OUTGROUPS USED IN THIS ANALYSIS USED TO GENERATE PHYLOGENIES

IN CHAPTER6: NORTH AMERICAN FLAT-BODY MOTHS (ELACHISTIDAE:

DEPRESSARIINAE: DEPRESSARIA HAWORTH): MORPHOLOGICAL

EVOLUTION, HOST-PLANT SELECTION, AND GEOGRAPHIC DISTRIBUTION

.

0 5 10 15 20 25 30 35 40 45 | | | | | | | | | | Psilocorsis reflexella ?????????021011?11011-1------0-0----1000-00-1101 0????1?0000-200?11010-1------0-100-00200-00-0101 Himmacea huachucella 0????0000021001?02011-1------0-100-00100-00-0010 Bibarrambla allenella 11000011101-110104010-1------0-100-00000-00-0010 Apachea barberella 11000101011-000113012-1------0-100-00100-00-1?10 Nites maculatella 010001010120010113010-1------0-11--00100-0101100 Agonopterix flavicomella 10010111110-200110010-1------0-100-00100-1101000 Agonopterix gelidella 10010111110-200110010-1------0-100-00100-0021000 Exaeretia fulvus 100110111010200100010-1------0-11---0100-0101000 Exaeretia lutosella 100110111010200112010-1------0-11---0100-1111000 atrostrigella 11001111011020011201101------110----010100101001 artemisiae 1100111101102001120110000----110----010101101001 absynthiella 1100111101102001120111000----100----010100101001 haydenii 11001111011020011201101------100----01?100101000 chaerophylli 11001111011020011101111------110----01?100111001 depressana 11001111011020011201111------110----010100101010 pastinacella 11101111010-200112011001101000-0----0200-1101000 cinereocostella 111011110110200101011001101110-0----0100-0110010 juliella 111011110110200101011001101110-0----0100-00-0010 daucella 111011110110200101011001101110-0----0100-00-0010 eleanorae 111011110110200101011001101110-0----0100-00-0010 ultimella 111011110110200101011102101100-0----0100-0100010 rubricella 111011110110200101011001101110-0----0100-0110010 bupleurella 111011110110200101011002101000-0----010100111100 pimpinellae 111011110110200112011002101000-0----010100111100 libanotidella 111011110110200112011002101000-0----010100111100 388

0 5 10 15 20 25 30 35 40 45 | | | | | | | | | | silesiaca 11101111011020011201100010101110----011110111000 badiella 111011110110200111011001101010-0----01?110110110 velox 111011110110200112011001101010-0----010110110110 alienella 11001111011020000211011------0-100000100-0111010 artimisiella 11001111011020000211011------0-100000100-0111010 betina 11001111011020010010--1------0-100001000-0100010 constancei 11001111011020010010--1------0-100001000-0100010 whitmani 11001111011020010020--02110100-101000100-0110001 schellbachi 11001111011020010020--02110100-101000100-0110001 angelicivora 11001111011020010020--02110100-101000100-0110100 leptotaeniae 11001111011020011220--02110100-101000100-0111000 yakimae 11001111011020011220--02110100-101000100-0111100 multifidae 11001111011020010220--02110100-101000100-0111001 moya 11001111011020011220--02110100-100-00110-0101000 pteryxiphaga 11001111011020010220--02110000-100-00110-0111000 besma 11001111011020011220--02110100-100-00110-0111000 angustati 11001111011020010020--02110100-101000100-0110000 togata 11001111011020010020--02110100-101000100-0110000 armata 11001111011020010020--02110000-101000100-0110000 douglasella 11001111011020010020--01110000-100-10010-0121010 weirella 11001111011020010020--01110000-100-10010-0121010 nemolella 11001111011020010220--02110000-100-10010-0120010 beckmanni 11001111011020010020--01110000-100-10010-0120010 pulcherrimella 11001111011020010220--02110000-100-10010-0120010 emeritella 11001111011020011120--01110000-100-00110-0110010 hofmanni 11001111011020010120--01110000-101000110-2110000 albipunta 11001111011020010020--01110100-101100110-2110010 olerella 11001111011020010020--01110100-101100110-2110010 ululana 11001111011020010020--01110000-102000110-2110000 leucocephala 11001111011020010101201------0-10--00110-0110001 cervicella 11001111011020010101201------0-10--00110-0110011 gallicella 11001111011020010101201------0-10--00010-0110011 discipunctella 11001111011020010101201------0-10--00110-0110010

SUMMARY PERCENTAGES

MISSING (?): 25 cells, <1 percent of matrix.

DASHES (-): 422 cells, 14 percent of matrix.

389