Advances in and Systematics of ()

Dissertation

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

By

Charuwat Taekul, M.S.

Graduate Program in Evolution, Ecology, and Organismal Biology

*****

The Ohio State University

2012

Dissertation Committee:

Dr. Norman F. Johnson, Advisor

Dr. Johannes S. H. Klompen

Dr. John V. Freudenstein

Dr. Marymegan Daly

Copyright by

Charuwat Taekul

2012

ABSTRACT

Wasps, , , and one of the most familiar and important , are scientifically categorized in the order Hymenoptera. Hymenoptera display some of the most advanced biology of the order. Platygastroidea, one of the significant groups of parasitoid , attacks host eggs more than 7 orders. Despite its success and importance, an understanding of this group is still unclear. I present here the world systematic revisions of two genera in Platygastroidea: Platyscelio Kieffer and

Oxyteleia Kieffer, as well as introduce the first comprehensive molecular study of the most important subfamily in platygastroids as biological control benefit, .

For the systematic study of two Old World genera, I address the taxonomic history of the genus, identification key to species, as well as review the existing concepts and propose descriptive new species. Four new species of Platyscelio are discovered from South

Africa, Western Australia, Botswana and Zimbabwe. Four species are considered to be junior synonyms of P. pulchricornis. Fron nine valid species of Oxyteleia, the new species are discovered throughout Indo-Malayan and Australasian regions in total of twenty-seven species. The genus Merriwa Dodd, 1920 is considered to be a new synonym. To better understanding of the relationships of subfamily Telenominae, I reconstruct the phylogeny based on approximately 3.7 kb of DNA sequence from 4

ii molecular markers (18S, 28S, COI and EF1-α) expanding 80 terminals: 6 genera (11 species groups), 5 broad-ranged out group genera. Results are drawn from both parsimony and statistical analyses (Bayesian and Maximum likelihood), and from 6 character coding and partitioning schemes. The molecular evidence showed that the subfamily is not monophyletic: one clade, the Psix group of genera, forms a monophyletic group with species of the tribe Gryonini, subfamily .

Monophyletic clades were recovered with strong support including

(Psix+Paratelenomus) and Gryon; Telenominae, s.str. (without Gryon and Psix group of genera); Phanuromyia and crassiclava species group; Telenomus laricis species group; and Telenomus longicornis species group. The genera Eumicrosoma and

Platytelenomus are nested within Telenomus californicus species complex.

Pentatomoidea () is the plesiomorphic host. The species in the clade

Phanuromyia + Telenomus crassiclava species group all shared the same host group,

Auchenorrhyncha (Hemiptera: and ). The monophyly of the major genera Telenomus and is not supported; the interrelationships of their component species are largely unresolved.

iii

Dedication

This work is dedicated to all of those in the United States and Thailand who gave me the opportunity and inspiration to work hard, overcome every kind of scientific and cultural

obstacle, and see this project through to a successful conclusion.

iv

ACKNOWLEDGMENTS

I have been passionate for detail, organizing, illustration, and nature since I was young. I am very lucky that my characters are related to my professional career. The dream of becoming systematist, however, would not have come true without the involvement of several people and I am most deeply indebted to my mentors, colleagues, friends, and family.

First and foremost, I would like to express my sincere gratitude to my advisor, Dr.

Norman F. Johnson, for the intellectual support, encouragement, and the hours of his dedicated works making me a better scientist. I am thankful for his patience with my insanity, as well as for the logic and intellect that he impressed upon me. He truly is the role model.

I would also like to express my gratitude to the members of my committee, Dr.

Johannes S. H. Klompen, Dr. John V. Freudenstein, and Dr. Marymegan Daly, for providing guidance and support throughout my graduate education. I am grateful to

Dr.Hans to put up with a whining student in the molecular lab. Thanks to John V.F. giving me many unforgettable memories, the discussion at MBD lunch table in particular.

I thanks to Dr. Meg’s enthusiasm, giving me more energy. I also thank to Dr. John

Wenzel, my former committee member, for advice and encouragement.

v One chapter of this research, the molecular phylogeny of Telenominae, could not have succeeded with the primary help from two people: Dr. Alejandro A. Valerio

(molecular technique and discussion) and Joseph R. Cora (statistical analyses). I am indebted to their support.

This research would not be complete without the help of other taxonomists, from whom the specimens of my study have been borrowed, allowing me to examine approximately more than 1500 loaned specimens. Thanks also must be given to the curators of the museums indicated in the publications.

I am grateful to the officemate I have made during my stay, Elijah J. Talamas, for giving me stimulating discussions, moral support and encouragement in both life and study. I would finally like to thank the following members of the Ohio State University

Insect Collection, who made my life here more enjoyable: Dr. Luciana Musetti, Dr.

Roger A. Burks, Joseph R. Cora, and Sara E. Hemly.

My intellectual and philosophical knowledge could not be improved without surrounding by several scholars at MBD. I thanks to these people: Mike Broe, Ryan Folk,

Ryan Caesar, Glene Mynhardt, Jeff Rose, Brandon Sinn, Mesfin Tadesse, Ryan Kitko,

Paul Larson, Erin Morris, Elijah Talamas, Alejandro Valerio, and Matt Yoder.

My graduate study was supported by the Agricultural Research Development

Agency, Ministry of Agriculture and Cooperatives, Thailand, and by the National Science

Foundation, USA.

vi On a personal note, I thank to Carey K. S., Havely M., Khunying Prapaisri

Pitakprivan, and Dr. Manita Kongchuensin for moral support and always believing in me.

I am pleased to thank my parents, Dumrong and Jarunee Taekul, and both my elder brother and sister, Natthakarn and Tanate Taekul, for moral support.

The last six years in my graduate program have meant more to me than the study of systematics, describing species, molecular systematics, and mastering the English language. I now have a better understanding of the importance of passions, sciences, nature, and thus have grown to love the flora and fauna of the world more than ever before.

vii

VITA

December 18, 1975 ...... Born – Krabi, Thailand March 1998 ...... B.S. Entomology (Honors) Kasetsart University, Thailand 1999 – Present ...... Official Government, Department of Agriculture, Ministry of Agriculture and Cooperatives, Thailand. 2008...... M.S. Entomology, The Ohio State University, USA 2010 – 2011...... Research Aide, Department of Evolution, Ecology and Organismal Biology, The Ohio State University

PUBLICATIONS

1. Taekul, C., Johnson, N.F., Masner, L., Polaszek, A. & Rajmohana, K. (2010) World species of the genus Platyscelio Kieffer (Hymenoptera: ). Zookeys, 50:97 – 126.

2. Taekul, C., Johnson, N.F., Masner,L., Rajmohana K., & Shu–Pei, C. (2008) Revision of the world species of the genus Fusicornia Risbec (Hymenoptera: Platygastridae, Scelioninae). Zootaxa, 1966:1–52

viii

FIELD OF STUDY

Major Field: Evolution, Ecology, and Organismal Biology

Areas of Emphasis: Entomology, Taxonomy, Systematics, and Molecular Phylogeny

ix

TABLE OF CONTENTS

Page

Abstract ...... ii Dedication ...... iv Acknowledgments...... v Vita ...... viii List of Tables ...... xv List of Figures ...... xvii

Chapters:

1. General Introduction ...... 1 Hymenoptera ...... 1 Parasitoid Hymenoptera ...... 2 Superfamily Platygastroidea ...... 3 Taxonoic history ...... 3 Diversity and geographic distribution ...... 4 Fossil taxa ...... 5 Monophyly of the superfamily...... 5 Biology ...... 6 Impediments and project description ...... 8

2. World species or the genus Platyscelio Kieffer (Hymenoptera: Platygastroidea) ...... 13 Introduction ...... 13

x Materials and methods ...... 16 Materials ...... 16 Terminology and data handling ...... 17 Genus Platyscelio Kieffer Taxonomic reviews ...... 19 Diagnosis...... 20 Description of the genus ...... 22 Distribution ...... 22 Key to species of Platyscelio ...... 23 Species description...... 25 Platyscelio africanus Risbec ...... 25 Platyscelio arcuatus Taekul & Johnson, new species ...... 28 Platyscelio mysterium Taekul & Johnson, new species ...... 30 Platyscelio pulchricornis Kieffer ...... 32 Platyscelio striga Taekul & Johnson, new species ...... 36 Platyscelio umzantsi Taekul & Johnson, new species...... 38 Conclusion ...... 41

3. World species exploration of the genus Oxyteleia Kieffer (Hymenoptera: Platygastroidea) ...... 55 Introduction ...... 55 Materials and methods ...... 58 Materials ...... 58 Terminology and data handling ...... 59 Genus Oxyteleia Kieffer ...... 61 Taxonomic reviews ...... 62 Description of the genus ...... 62 Diagnosis...... 65 Distribution ...... 65

xi Key to species of Oxyteleia...... 66 Species description ...... 66 Oxyteleia inthanona Taekul, new species ...... 75 Oxyteleia aenigma Taekul, new species ...... 77 Oxyteleia oecus Taekul, new species ...... 81 Oxyteleia pangasugana Taekul, new species ...... 84 Oxyteleia angularis Taekul, new species ...... 86 Oxyteleia rugosa Taekul, new species...... 89 Oxyteleia bimucronata Taekul, new species ...... 92 Oxyteleia epidermosa Taekul, new species ...... 94 Oxyteleia quadridentata (Dodd) ...... 99 Oxyteleia alba Taekul, new species ...... 102 Oxyteleia arcuata Taekul, new species ...... 104 Oxyteleia talamasi Taekul, new species ...... 107 Oxyteleia lux Taekul, new species...... 110 Oxyteleia scopulus Taekul, new species ...... 113 Oxyteleia solomona Taekul, new species ...... 115 Oxyteleia insula Taekul, new species ...... 118 Oxyteleia lenisa Taekul, new species ...... 120 Oxyteleia striga Taekul, new species ...... 123 Oxyteleia ponticulus Taekul, new species ...... 126 Oxyteleia arcuza Taekul, new species...... 129 Oxyteleia apektos Taekul, new species ...... 132 Oxyteleia johnsoni Taekul, new species ...... 134 Oxyteleia lagunasis Taekul, new species ...... 137 Oxyteleia primata Taekul, new species ...... 139 Oxyteleia astricta Taekul, new species ...... 142 Oxyteleia catheta Taekul, new species ...... 144 Oxyteleia bidentata (Dodd) ...... 147

xii Oxyteleia obscura Taekul, new species ...... 150 Oxyteleia fasciata Taekul, new species ...... 152 Conclusion ...... 156

4. Molecular phylogeny of the subfamily Telenominae (Platygastridae, Platygastroidea, Hymenoptera) ...... 195 Introduction ...... 195 Materials and methods ...... 202 Taxonomic sampling and specimen vouchering ...... 202 DNA extraction, amplification, and sequencing ...... 203 Sequence alignment ...... 204 Data coding and partitioning ...... 205 Phylogenetic analyses ...... 206 Maximum parsimony ...... 207 Maximum likelihood ...... 207 Bayesian analysis ...... 208 Results ...... 209 Data properties ...... 209 Phylogenetic relationships of Telenominae ...... 210 The monophyly of subfamily ...... 211 Psix group of genera ...... 211 Telenomus and Trissolcus ...... 212 Telenomus californicus species complex ...... 212 The placement of Eumicrosoma ...... 212 Problematic taxa...... 212 Discussion ...... 213 Telenominae is not monophyletic...... 214 Genus Gryon ...... 214 The monophyly of genus Psix and Paratelenomus...... 216

xiii Telenomus crassiclava species group and Phanuromyia...... 217 The genera Trissolcus and Telenomus ...... 217 species group...... 219 Conclusion ...... 221

Bibliography ...... 237

xiv

LIST OF TABLES

Table Page

1.1 The valid synonyms of the family Platygastridae and its inner taxa, Sceliotrachelinae, , Scelioninae, , and Telenominae (the bold text indicates an original description) ...... 9

1.2 The relationship between subfamily Scelioninae, Teleasinae, and Telenominae, and associated host groups (Austin et al. 2005) ...... 10

1.3 The relationship between subfamilies Sceliotrachelinae and Platygastrinae and associated host groups (Austin et al. 2005) ...... 11

1.4 The application of platygastroid wasps in biological control approach (Orr 1988) ...... 12

4.1 List of taxa and gene-amplified regions for each species included in the telenomine phylogenetic analysis ...... 223

4.2 Primer descriptions ...... 226

4.3 Gene partitions and alignment summary for subfamily Telenominae and outgroups. Hypervariable regions and introns were removed manually. Percent parsimony informative sites were calculated using WinClada ver. 1.00.08 (Nixon 2002). Models chosen for data partitions were selected using the Akaike Information Criterion (AICc) as calculated in jModelTest 0.1.1 (Posada 2008)...... 227

4.4 Definitions for coding strategies and partitioning schemes ...... 228

xv 4.5 Support values for selected clade across the analyses. Dataset are based on nucleotide character coding and partitioning schemas shown on Table 4. Support values are reported as posterior probability (PP) for Bayesian inference and bootstrap support (BS) for maximum likelihood (RAxML) and jackknife support (JK) for parsimony TNT. The support values for all analyses are reported in percent above 50. Abbreviations used in text: Tr., Trissolcus; Te., Telenomus; sp., species; gr., group taxa; com., complex; y=yes group recovered but support less than 50 percent; n=no, not monophyletic; - , taxa excluded; +, taxa included...... 229

4.6 Host records of Gryon and Telenominae ...... 231

xvi

LIST OF FIGURES

Figure Page

2.1 Platyscelio africanus Risbec, female (OSUC 250659). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Fore wing marginal vein, dorsal view; E, Head, dorsal view; F, Mesoscutellum, Propodeum and T1, dorsal view. Scale bars in millimeters...... 42

2.2 Platyscelio africanus Risbec, female (OSUC 207985). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Fore wing marginal vein, dorsal view; E, Head, dorsal view; F, Mesoscutellum, Propodeum and T1, dorsal view. Scale bars in millimeters...... 43

2.3 Platyscelio arcuatus, n.sp., holotype female (OSUC 250635). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, dorsal view; F, Propodeum and T1, dorsal view. Scale bars in millimeters...... 44

2.4 Platyscelio mysterium, n.sp., holotype female (OSUC 171372). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, dorsal view; F, Cublateral carina on T2–T4, dorsal view. Scale bars in millimeters...... 45

2.5 Platyscelio pulchricornis Kieffer, holotype female (MCSN 0004). A, Dorsal habitus; B, Lateral habitus; C, Head and mesosoma, dorsal view; D, Head and mesosoma, lateral view; E, Head, dorsal view; F, Metasoma, dorsal view. Scale bars in millimeters...... 46

2.6 Platyscelio pulchricornis Kieffer, female (OSUC 207837). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Head and mesosoma, lateral view; E, Head, dorsal view; F, Fore and hind wing, lateral view. Scale bars in millimeters...... 47

xvii 2.7 Platyscelio pulchricornis Kieffer, female (OSUC 250628). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Propodeum and T1, dorsal view; E, Head, dorsal view; F, Fore wing, dorsal view. Scale bars in millimeters...... 48

2.8 Platyscelio abnormis Crawford, holotype male (USNM Type No. 12895). A, Dorsal habitus; B, Lateral habitus; C, Head and mesosoma, dorsal view; D, Head and mesosoma, lateral view; E, Head, dorsal view; F, Mesoscutellum and propodeum, dorsal view. Scale bars in millimeters...... 49

2.9 Platyscelio dunensis Mukerjee, holotype male (NZSI 0001). A, Mesosoma and Propodeum, dorsal view; B, Metasoma, dorsal view; C, Head, dorsal view; D, T1 and T2, dorsal view. Platyscelio punctatus Kieffer, syntype male (OSUC 207855). E, Mesosoma, dorsal view; F, T1 and T2, dorsal view. Scale bars in millimeters...... 50

2.10 Platyscelio wilcoxi Fullaway, holotype female (USNM Type No. 26186). A, Dorsal habitus; B, Lateral habitus; C Propodeum and T1, dorsal view; D, Head and mesosoma, lateral view; E, Head, dorsal view; F, Metasoma, dorsal view. Scale bars in millimeters...... 51

2.11 Platyscelio striga, n.sp., holotype female (OSUC 250630). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Propodeum and T1, dorsal view; E, Head, dorsal view; F, Antennae, dorsal view. Scale bars in millimeters...... 52

2.12 Platyscelio umzantsi, n.sp., holotype female (OSUC 243790). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Fore wing marginal vein, dorsal view; E, Head, dorsal view; F, Propodeum and T1, dorsal view. Scale bars in millimeters...... 53

2.13 Platyscelio host eggs. A, Platyscelio pulchricornis and host egg collected from sugarcane, dorsal view (OSUC 207839); B, Host egg and specimen label (OSUC 207839); C, Platyscelio pulchricornis host eggs collected from rice (BMNH(E)#790194); 76, Host eggs collected from sugarcane (BMNH(E)#790205). Scale bars in millimeters...... 54

xviii 3.1 Oxyteleia inthanona, n.sp., holotype female (OSUC 266812). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Antennae, lateral view. Scale bars in millimeters...... 158

3.2 Oxyteleia aenigma, n.sp., holotype female (OSUC 173884). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Antennae, lateral view. Scale bars in millimeters...... 159

3.3 Oxyteleia oecus, n.sp., holotype female (OSUC 266674). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 160

3.4 Oxyteleia oecus, n.sp., female (OSUC 253800). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters. ... 161

3.5 Oxyteleia pangasugana, n.sp., holotype female (OSUC 276527). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Mesonotum, dorsal view. Scale bars in millimeters...... 162

3.6 Oxyteleia angularis, n.sp., holotype female (OSUC 234583). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, antennae. Scale bars in millimeters...... 163

3.7 Oxyteleia rugosa, n.sp., holotype female (OSUC 234104). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, antennae. Scale bars in millimeters...... 164

3.8 Oxyteleia bimucronata, n.sp., holotype female (OSUC 253618). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Mesoscutum, dorsal view. Scale bars in millimeters...... 165

xix 3.9 Oxyteleia epidemosa, n.sp., holotype female (OSUC 266804). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Ocellar triangle, dorsal view. Scale bars in millimeters...... 166

3.10 Oxyteleia epidemosa, n.sp., (OSUC 276496). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 167

3.11 Oxyteleia epidemosa, n.sp., (OSUC 266601). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 168

3.12 Merriwa quadridentata Dodd, Holotype (B.M. TYPE HYM. 9.465). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Type label. Scale bars in millimeters...... 169

3.13 Oxyteleia quadridentata, (OSUC 253786). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Antennae. Scale bars in millimeters...... 170

3.14 Oxyteleia quadridentata, (OSUC 276580). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Antennae. Scale bars in millimeters...... 171

3.15 Oxyteleia alba, n.sp. holotype female (OSUC 276610). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Wings. Scale bars in millimeters...... 172

3.16 Oxyteleia acuata, n.sp. holotype female (OSUC 256842). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Ocellar triangle. Scale bars in millimeters...... 173

3.17 Oxyteleia talamasi, n.sp. holotype female (OSUC 276557). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Gena. Scale bars in millimeters...... 174

3.18 Oxyteleia lux, n.sp. holotype female (OSUC 276526). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Antennae. Scale bars in millimeters...... 175

xx 3.19 Oxyteleia scopulus, n.sp. holotype female (FBA 143133). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 176

3.20 Oxyteleia solomona, n.sp. holotype female (OSUC 234112). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 177

3.21 Oxyteleia insula, n.sp. holotype female (OSUC 266722). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 178

3.22 Oxyteleia lenisa, n.sp. holotype female (OSUC 276562). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Antennae. Scale bars in millimeters...... 179

3.23 Oxyteleia striga, n.sp. holotype female (OSUC 266746). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Mesoscutellum & horn on T1, dorsal view. Scale bars in millimeters...... 180

3.24 Oxyteleia ponticulus, n.sp. holotype female (OSUC 234040). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Horn. Scale bars in millimeters...... 181

3.25 Oxyteleia arcuza, n.sp. holotype female (OSUC 266740). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Gena. Scale bars in millimeters...... 182

3.26 Oxyteleia arcuza, n.sp. (OSUC 266710). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Gena. Scale bars in millimeters...... 183

3.27 Oxyteleia apektos, n.sp. holotype female (OSUC 266743). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma,

xxi lateral view; E, Head, anterior view; F, Mesoscutellum. Scale bars in millimeters...... 184

3.28 Oxyteleia johnsoni, n.sp. holotype female (FBA014399). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 185

3.29 Oxyteleia lagunasis, n.sp. holotype female (OSUC 276576). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Gena. Scale bars in millimeters...... 186

3.30 Oxyteleia primata, n.sp. holotype female (OSUC 246586). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Mesoscutellum, dorsal view. Scale bars in millimeters...... 187

3.31 Oxyteleia astricta, n.sp. holotype female (OSUC 234135). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Mesoscutellum & horn on T1, dorsal view; F, Metasoma, dorsal view. Scale bars in millimeters...... 188

3.32 Oxyteleia catheta, n.sp. holotype female (OSUC 234033). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Mesoscutellum & T1, dorsal view. Scale bars in millimeters...... 189

3.33 Oxyteleia bidentata Kieffer, Holotype (MCSN 0003). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view. Scale bars in millimeters...... 190

3.34 Oxyteleia bidentata (OSUC 266733). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Mesoscutellum & T1, dorsal view. Scale bars in millimeters...... 191

3.35 Oxyteleia bidentata (OSUC 234125). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Horn on T1. Scale bars in millimeters...... 192

3.36 Oxyteleia obscura, n.sp. holotype female (OSUC 266742). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma,

xxii lateral view; E, Head, anterior view; F, Metasoma, dorsal view. Scale bars in millimeters...... 193

3.37 Oxyteleia fasciata, n.sp. holotype female (OSUC 234050). A, Dorsal habitus; B, Lateral habitus; C, Mesosoma, dorsal view; D, Mesosoma, lateral view; E, Head, anterior view; F, Mesoscutellum, dorsal view. Scale bars in millimeters...... 194

4.1 Telenominae relationships based on Bayesian analysis of aligned, concatenated all nucleotides, degenerated protein coding genes. Ribosomal sequences aligned using secondary structure with hypervariable regions excluded. Analysis executed on MrBayes 3.2.1 with 1 partition parameter set (Degen1) (two runs of four chains each,14.13 M gen, convergence diagnostic hit stop value, 0.0099) of combined COI, 18S, 28S and EF-1a (3767 bp). Posterior probability (PP) indicated on branches (in percent); branches with PP below 50% collapsed...... 232

4.2 Relationships of the subfamily Telenominae derived from Maximum likelihood analysis of combined COI, 18S, 28S and EF-1a (3767 bp), executed on RAxML (1000 standard bootstrap replicates) with a single partition (Degen1). Data matrix from degenerated protein coding genes and manually aligned ribosomal sequences based on secondary structure with the exclusion of ambiguious regions. Bootstrap values above 50% indicated on branches...... 233

4.3 Phylogeny of subfamily Telenominae based on Bayesian analysis (two runs of four chains each, 6.57 M gen, convergence diagnostic hit stop value, 0.009) with 4 partitions. Character set from combined COI, 18S, 28S and EF-1a (3767 bp) with degenerated protein coding genes and aligned ribosomal sequences using secondary structure with hypervariable regions excluded. Posterior probability (PP) indicated on branches (in percent); branches with PP below 50% collapsed...... 234

4.4 Telenominae relationships derived from Parsimony analysis (TNT, New Technology Search) of combined COI, 18S, 28S and EF-1a (3767 bp). Strict consensus based on 4 trees of length 7076 steps, CI 0.34, RI 0.53. Numbers under branches indicate jackknife support values (JK) based on 1,000 pseudoreplicates (values below 50% not shown)...... 235

xxiii 4.5 Telenominae relationships derived from Parsimony analysis (TNT, New Technology Search) of combined COI, 18S, 28S and EF-1a: A, nt12 (3106 bp) and B, Degen (3765 bp). A(nt12): strict consensus based on 16 trees of length 2370 steps, CI 0.41, RI 0.65. B(degen): strict consensus based on 174 trees of length 2265 steps, CI 0.47, RI 0.70. Numbers under branches indicate jackknife support values (JK) based on 1,000 pseudoreplicates (values below 50% not shown)...... 236

xxiv

CHAPTER 1

GENERAL INTRODUCTION

The order Hymenoptera is thought to be one of the most important groups among insects. This large, megadiverse group contains four groups familiar to most people: the wasps, bees, ants, and sawflies. The number of species is estimated at over 115,000

(LaSalle and Gauld 1993). In addition, Sharkey (2007) claimed that the species richness of the order may constitute 10 percent of the species of life and estimated that the number of species may reach 1,000,000. The best available hypothesis of the phylogenetic position of this order is that it is sister to all other Holometabola (Sharkey 2007, Savard et al. 2006). Hymenoptera are traditionally divided into two suborders, the paraphyletic

Symphyta (sawflies, woodwasps) and the monophyletic , the latter containing the monophyletic and polyphyletic (Heraty et al. 2011; Sharkey et al.

2012). The biology of Hymenoptera is diverse and includes both phytophagous and entomophagous groups. Entomophagous groups may be either or predators.

Eusociality is also common in this group. In comparison with other insect orders,

Hymenoptera is considered to have the greatest number of beneficial species. Bees are

1 important in pollination of flowering plants, as well as producers of commercial goods: honey and beeswax. Aculeata and Parasitica are economically important as biological control agents, especially parasitoid Hymenoptera. Out of 393 species of parasitoids of all groups introduced in classical biological control programs, 87 percent of the parasitoid wasps were successfully established and achieved pest control (Greathead 1986, Lasalle and Gauld 1993).

Parasitoid Hymenoptera— Parasitoid wasps display some of the most advanced biology of the order. They feed on a single host, either externally (ectoparasitoids) or internally (endoparasitoids), and ultimately kill the host as a result. This biology is different from predators and parasites; a predator directly kills and eats numerous prey, a parasite obtains nourishment from its host, inflicting minimal to severe ill effects, but normally does not kill the host. The number of immature parasitoids within a host may differ, these being referred to as solitary and gregarious parasitoids. Some parasitoid wasps are hyperparasitoids, in which one parasitoid attacks another that is on or within a primary host. Parasitoid wasps are important elements in the ecosystem. They sustain the diversity of nature. Parasitoids attack and kill their hosts, thereby affecting their density.

Parasitoid Hymenoptera may act as biological indicators to reveal the diversity of , since they are representatives of the hosts that they attack. The use of parasitoids as biological control agents has been proven to be an effective approach to control agricultural and forestry pests as well as vectors of human and disease.

The level of parasitoid populations indicates that of the pests and whether the pests are

2 approaching the economic threshold. This may help determine whether to apply pesticides or alternative approaches in an integrated pest management (IPM). With effective conservation biological control, the parasitoids are maintained in the agricultural crops over years without application of chemical insecticides. Parasitoids are very sensitive to ecological perturbation such as pesticides, and thus they are indicators for conservation. These beneficial wasps can have a significant impact in promoting environmental quality. Seven families of egg parasitoids have been used in classical biological control, and one genus – Trichogramma – has been commercially mass-reared and effectively used in augmentative biological control (Mills 2010).

THE SUPERFAMILY PLATYGASTROIDEA

Taxonomic History—The first species now classified in superfamily

Platygastroidea was published in the tenth edition of Linnaeus’s Systema Naturae in

1758. In the book he proposed the name, Ichneumon ovulorum, which is actually a species of Telenomus reared from lepidopteran eggs (Linnaeus 1758). Genera currently recognized as platygastroids were typically placed within the superfamily

Proctotrupoidea, also sometimes referred to as Serphoidea or Oxyura. The family-group name Proctotrupii was originally proposed by Latreille (1802), and the first platygastroid genera to be placed within it were Scelio (Latreille 1805), Sparasion (Latreille 1805), and

Teleas (Latreille 1809). The family group name Platygastridae (as Platygastres) was originally proposed by Haliday (1833), and Scelionidae also by Haliday (1839). These two concepts, sometimes treated separately, sometimes with platygastrids within the

3 Scelionidae, remained grouped within the heterogeneous until the second half of the 20th century. Masner (1956) suggested that the two families should be treated as a separate superfamily, the Scelionoidea, a name later changed to Platygastroidea in recognition of the latter’s priority. A brief summary of family-group names used for subfamilies and families is presented in Table 1.1 Recently, Sharkey (2007) has combined the families Scelionidae and Platygastridae under the single family

Platygastridae. The reasons for the synonymy are (1) the absence of conspicuous synapomorphies defining the scelionids as a monophyletic group, a conclusion based on the phylogenetic analysis of Murphy et al. (2007); and (2) Sharkey’s perception that the superfamily is morphologically homogeneous. In this research, the rank and monophyly of the previously recognized subfamilies were not discussed. Thus, the review of literature here will discuss Platygastrinae, Sceliotrachelinae, Scelioninae, Teleasinae, and

Telenominae, all as subfamilies of Platygastridae.

Diversity and Geographic Distribution—Platygastroidea is the third largest parasitoid superfamily after and Chacidoidea (Austin et al., 2005). The superfamily is placed as a monophyletic group with Prototrupoidea and

(Sharkey 2007). The former two families, Platygastridae and Scelionidae were catalogued

(Johnson 1992, Vlug 1995) and the taxonomic information is kept updated in Johnson

(2012). Currently, 264 genera and 5,514 species of Platygastroidea are considered valid: i.e., Platygastrinae (44 genera, 1640 species), Sceliotrachelinae (28 genera, 132 species),

Scelioninae (155 genera, 2,331 species), Teleasinae (13 genera, 506 species), and

Telenominae (20 genera, 895 species).

4 The geographic coverage of the superfamily is widespread in both Old and New

World. With several recent studies of the relative scale of platygastroid diversity, the areas that remain largely unstudied are tropical and southern continents (Austin et al.

2005). There appears to be little overlap in species composition, indicating a high level of regional endemism (Iqbal and Austin 1997).

Fossil Taxa—Most described fossils of the superfamily Platygastroidea are from

Baltic amber inclusions. Grimaldi and Engel (2005) claimed it to be the dominant group of Hymenoptera in the , and that the superfamily probably had its greatest diversity during the Cretaceous, dwindling during the Cenozoic. In fact, merely 5 species have been so far described from the Cretaceous period, and the number of platygastroid fossil species dramatically increased after the Paleocene epoch, based on the list of fossil and subfossil species of Platygastroidea by Johnson et al. (2012). The platygastroid fossil and subfossils known include 5 genera in the Cretaceous, 1 genus in the Paleocene, 23 genera in the Eocene, 4 genera in the Oligocene, and 4 genera in the Miocene (Johnson et al. 2012).

Monophyly of the Superfamily—The monophyly of Platygastroidea is supported by the modified structure and function of the abdomen and ovipositor, as well as the paired basiconic sensilla on the apical segments of the female antenna (Masner 1976;

Austin et al. 2005),. The function of the ovipositor in plastygastroids is complex. Unlike most other parasitic Hymenoptera, the proximal elements of the ovipositor, i. e., the gonocoxae and gonapophyses, are disassociated from the posterior segment of the

5 abdomen. (Quicke et al. 1999; Vilhelmsen 2003; Austin et al. 2005). Platygastroid wasps are generally 0.5 to 12 mm in body length, but most are less than 2.5 mm long. In comparison with other wasps, most have greatly reduced wing venation, the antennae are inserted close together just above the mouth, they lack a prepectus behind the lateral pronotum, and are well-sclerotized and often intricately sculptured. Platygastrinae and

Sceliotrachelinae (the Platygastridae in the traditional sense) are distinguished from other subfamilies by reduced wing venation, the forewings with at most a single tubular vein; and few antennomeres (usually 7-10). In Scelioninae, Teleasinae, and Telenominae, the forewing venation is somewhat more elaborate, consisting of the submarginal, marginal, stigmal and postmarginal veins and the antenna usually has 11-12 antennomeres (Masner, in Goulet and Huber, 1993).

The Biology of Platygastroidea—Predation and parasitism are two antagonistic ecological interactions in which one species takes advantage of another species. A predator kills and eats another, its prey. Parasites obtain nourishment via sub-lethal consumtion of another organism; this typically results in some harm to the host, but not death. Parasitoids, generally in their immature stages, feed on only a single “host”, but in the process kill it.

A number of families within parasitic Hymenoptera are egg parasitoids of pests of agriculture and forestry as well as of vectors of human and animal disease. Numerous groups within Chalcidoidea are egg parasitoids, e.g., the and

Mymaridae, and are known for wide host ranges (Huber 1986; Nagarkatti 1977). These

6 species are keystone species in their role as biological control agents (Lasalle 1993).

Austin et al. (2005) stated that all members of subfamilies Scelioninae, Teleasinae, and

Telenominae are endoparasitoids of the eggs of insects or spiders (Table 1.2). They interpreted this to be the ground plan biology for the superfamily. In addition to parasitizing host eggs, species of Platygastrinae and Sceliotrachelinae also attack immature stages of sessile hosts such as , , and . Most species of these two subfamilies, however, parasitize gall flies (Diptera: Cecidomyiidae), either in the egg or in an early larval stage (Table 1.3).

With their hypodermic-like ovipositor, female platygastroids pierce the chorion of a host egg and lay their own single egg, or sometimes several eggs, within. The larva that hatches derives its nourishment from the host egg and pupates within it. The hosts of Platygastridae comprise species of Odonata, Orthoptera, Mantodea, Embiidina,

Hemiptera, Neuroptera, Coleoptera, Diptera, , and spiders (Austin & Field.

1997, Masner 1976; Austin et al. 2005).

Despite the lack of complete knowledge of the diversity of taxa within this group, platygastroid wasps are considered to be valuable potential biological control agents (Orr

1988). This is attributed to several characteristics, including high searching abilities and reproductive rates, lack of hyperparasitoids, synchrony with host populations, positive host-density responsiveness, simple adult diets, and the ease with which they can be reared in the laboratory. Although only 30 species have been used in classical biological control attempts, several of these have produced excellent results (Orr 1988).

7 Platygastroid wasps can be used in all biological control practices, including classical control, conservation, and augmentation (Table 1.4). Because scelionids parasitize the eggs of insect pests, they are very useful in concert with other management tactics such as integrated pest management and conservation practice.

Despite the fact that Platygastroidea is one of the most important parasitoid wasps, a better understanding for this group is still needed. The taxonomy and relationships of this group are still unclear. Species richness is the major impediment to investigate the taxonomy and systematics of Platygastroidea. My contributions are in two areas;

1) I present world species revisions of two Old World tropical genera of

Platygastridae, Platyscelio Keiffer and Oxyteleia Kieffer (Chapter 2 and 3). I document taxonomic history of the genus, develop identification key to the species, revise existing taxonomic concepts, and propose new species concepts.

2) I investigate the phylogeny of subfamily Telenominae (Chapter 4) to better understand the relationships among its genera and species groups, address the position of problematic taxa, and provide a framework for interpreting the relationships between parasitoids and their host.

8

Table 1.1 The synonyms of the family Platygastridae and its child taxa, Sceliotrachelinae, Platygastrinae, Scelioninae, Teleasinae, and Telenominae. (Bold text indicates an original description).

Platygastridae Platygastres Haliday, 1833 Teleadidae Walker, 1836 Platygastrides Westwood, 1840 Scelionidae Haliday, 1839 Platygasteriens Brullé, 1846 Scelionoidae Förster, 1856 Platygastroidae Förster, 1856 Sparasionidae Dahlbom, 1858 Platygastrini Thomson, 1859 Scelionini Thomson, 1859 Platygasterides Desmarest, 1860 Muscidides Motschoulsky, 1863 Platygasteridae Walker 1873 Scelioninae Howard, 1886 Platygasterinae Howard, 1886 Platygastrinae Morley, 1923 Platygasterini Jansson, 1939

Sceliotrachelinae Scelioninae Sceliotrachelinae, Brues, 1908 Baeini Ashmead 1893 Amitini Szabó, 1959 Baeinae Ashmead 1903 Scelionini Ashmead, 1893 Platygastrinae Scelionides Morley, 1929 Inostemmini, Ashmead, 1893 Baeides Morley, 1929 Platygasterini Ashmead, 1893 Scelioniniens Risbec, 1950 Platygasterinae Ashmead, 1903 Inostemminae Ashmead, 1903 Teleasinae Inostemmatinae Kozlov, 1970 Teleasini Ashmead, 1893 Teleasinae Ashmead, 1903 Teleasides Morley, 1929

Telenominae Telenomini Thomson, 1860 Telenomides Morley, 1929 Telenominiens Risbec, 1950

9

Table 1.2 The relationship between subfamily Scelioninae, Teleasinae, and Telenominae, and associated host groups (Austin et al. 2005).

Kozlov 1970 Masner 1976 Austin & Field 1997 Host groups

Scelioninae Scelioninae Scelioninae — Sparasionini Sparasionini Tettigoniidae Caloteleini Nixonini Nixonini Tettigoniidae Baryconini Baryconini Baryconini Tettigoniidae Caloteleini Calliscelionini Calliscelionini Gryllidae Psilanteridini Psilanteridini Psilanteridini Gryllidae Aradophagini Aradophagini Aradophagini Unknown — — Neoscelionini Unknown Platyscelionini Platyscelionini Platyscelionini Tettigoniidae — Doddiellini Doddiellini Unknown — Cremastobaeini Cremastobaeini Gryllidae Mantibarini Mantibarini Mantibarini Mantodea Gryonini Gryonini Gryonini , Lepidoptera Embidobiini Embidobiini Embidobiini Embiidina Pseudanteridini — — Unknown Thoronini Thoronini Thoronini Baeini Baeini Baeini Araneae Idrini Idrini — Teleasinae Teleasinae Teleasinae Teleasini Teleasini Teleasini Coleoptera Xenomerini Xenomerini Xenomerini Coleoptera Telenominae Telenominae Telenominae Telenomini Telenomini Telenomini Heteroptera, , Lepidoptera, Neuroptera, Diptera Tiphodytini — —

10

Table 1.3 The relationship between subfamilies Sceliotrachelinae and Platygastrinae and associated host groups (Austin et al. 2005).

Kozlov 1970 Masner & Huggert Austin & Field 1997 Host groups 1989 (based on type genera) Sceliotrachelinae Sceliotrachelinae Sceliotrachelinae Sceliotrachelini — — Fidobiini Fidobia-cluster Fidiobia-cluster Coleoptera Aphanomerini Aphanomerus- Aphanomerus-cluster Fulgoroidea cluster — Isolia-cluster Isolia-cluster Unknown — Amitus-cluster Amitus-cluster Aleyrodidae Allotropini -cluster Allotropa-cluster Pseudococcidae Platygastrinae Platygastrinae Platygastrinae Metaclisiini — — — Proplatygaster- Proplatygaster-cluster Cecidomyiidae cluster Inostemmatini Isostasius-cluster Isostasius-cluster Cecidomyiidae Inostemmatini -cluster Inostemma-cluster Cecidomyiidae — Allostemma-cluster Allostemma-cluster Unknown Platygastrini — -cluster Cecidomyiidae Synopeadini — Synopeus-cluster Cecidomyiidae Coelopeltini — —

11

Table 1.4 Platygastroid wasps used successfully in biological control (Orr 1988).

Successful Classical Augmentation Conservation biological control biological control biological control Gryon muscaeforme Telenomus proditor Trissolcus flavipes Telenomus alsophilae Trissolcus semistriatus Telenomus gifuensis Telenomus monilicornis Trissolcus nigribasalis Trissolcus basalis Telenomus chloropus Trissolcus mitsukurii Trissolcus grandis Telenomus podisi Telenomus chloropus Trissolcus semistriatus Telenomus californicus Telenomus cyrus Trissolcus remus Telenomus remus Psix lacunatus Telenomus olsenni Telenomus sechellensis Telenomus proditor Telenomus applanatus Telenomus alsophilae

12

CHAPTER 2

WORLD SPECIES OF THE GENUS PLATYSCELIO KIEFFER

(HYMENOPTERA: PLATYGASTRIDAE)1

INTRODUCTION

Species of Platyscelio (Hymenoptera: Platygastroidea, Platygastridae) are morphologically unique among the known Scelioninae by a number of characters, most distinctively the extremely flat body, the broad hypostomal bridge, and the absence of a netrion. The genus was originally described by Kieffer (1905) with a single species,

Platyscelio pulchricornis from Dilo in British New Guinea. To date, six species-group taxa have been described; five species are recorded in Asia, Australia and Oceania: P. pulchricornis Kieffer, P. abnormis Crawford, P. dunensis Mukerjee, P. mirabilis Dodd, and P. punctatus Kieffer. Only one species is known from Africa, P. africanus Risbec, described from Cameroon.

1 This chapter was originally published in Taekul, C., Johnson, N. F., Masner, L., Polaszek, A. & Rajmohana, K. (2010) World species of the genus Platyscelio Kieffer (Hymenoptera: Platygastridae). Zookeys, 50:97–126.

13 Platyscelio was classified within the subfamily Scelioninae of the family

Scelionidae by Kieffer (1905), but was not placed in any tribe until Kozlov (1970) erected the monobasic tribe Platyscelionini. Sharkey (2007) has subsequently combined the families Scelionidae and Platygastridae under the single family Platygastridae, but he did not address the status of their respective subfamilies or tribes. Kozlov (1970) asserted that Platyscelionini is close to the tribe Scelionini, but did not explain this hypothesis.

Masner (1976) noted that the genus is distant from other Scelioninae due to its lack of a netrion, the greatly reduced palpi, and the expansion of the female antennal scape into a flat, almost triangular piece armed laterally with a sharp spine. Austin & Field (1997) examined the ovipositor structure and concluded that Platyscelio possesses a Scelio-type system. They also commented on two unusual features of the genus: the second gonocoxae are developed as broad membranous plates, and the lateral apodemes of the sixth metasomal sternum protrude proximally past the telescopic tube at rest. Platyscelio was not included as a taxon in the most comprehensive attempt to infer relationships within the Platygastroidea published to date (Murphy et al. 2007).

The known hosts of species in the subfamilies Scelioninae, Teleasinae, and

Telenominae are the eggs of insects and spiders (Austin et al. 2005). In the original description of Platyscelio africanus, Risbec (1956) mentioned that the 15 females and 3 males were obtained from eggs of “Locustidae.” Specimens of Platyscelio pulchricornis from the National Museum of Natural History (Washington, DC; OSUC 207839) include unidentified host eggs (Figs. 2.13 A, and B). Two reared series of Platyscelio pulchricornis in the Natural History Museum, London are from eggs identified as

14 “?Conocephalus sp” collected from sugarcane (BMNH(E)#790205) and rice in Papua

New Guinea (BMNH(E)#790194) (Figs, 13 C and D). Agyen-Sampong (1980, cited in

Heinrichs and Barrion 2004) reported Platyscelio sp. as a parasitoid of Conocephalus conocephalus (Linnaeus) (Orthoptera: Tettigoniidae). Agyen-Sampong conducted all his observations in West Africa, mostly Sierra Leone. These data, albeit fragmentary, suggest that Conocephalus eggs are at least among the hosts of Platyscelio spp, across the range of the genus from West Africa to Australasia. Kozlov (1970) asserted that the host of the genus must be the flattened eggs of Phaneropterinae (Orthoptera: Tettigoniidae), and also suggested that the strongly flattened body of Platyscelio may indicate that the species are phoretic in habit.

In more than a century since its original description, Platyscelio has never been comprehensively reviewed or revised. Our goal of this paper is to present a taxonomic revision of the world species of genus Platyscelio, as well as to expand the biogeographic data associated with these species. The taxonomic history of the genus is summarized and existing concepts are reviewed. Four new species are proposed, two from Western

Australia, two from southern Africa.

15

MATERIALS AND METHODS

Materials —This work is based upon specimens in the following collections, with abbreviations used in the text;

AEIC American Entomological Institute, Gainesville, FL

ANIC Australian National Insect Collection, Canberra, Australia

BMNH The Natural History Museum, London, UK

CNCI Canadian National Collection of Insects, Ottawa, Canada

EMEC Essig Museum of Entomology, Berkeley, CA

ISNB Institut Royal des Sciences Naturelles de Belgique, Belgium

MCSN Museo Civico di Storia Naturale "Giacomo Doria," Genova, Italy

MZLU Lund University, Lund, Sweden

NZSI Zoological Survey of India, North Regional Station,

Uttaranchal, India

OSUC C.A. Triplehorn Insect Collection, Columbus, OH

RMNH Nationaal Natuurhistorisch Museum, Leiden, Netherlands

ROME Royal Ontario Museum, Toronto, Canada

SAMA South Australian Museum, Adelaide, Australia

SAMC Iziko Museums of South Africa, Cape Town, South Africa

SANC South African National Collection of Insects,

Pretoria, South Africa

SCAU South China Agricultural University, Guangzhou, China

16 TARI Taiwan Agricultural Research Institute - Entomology,

Taichung, Taiwan

UASK Ukrainian Academy of Sciences, Kiev, Ukraine

UCDC University of California, Davis, CA

USNM National Museum of Natural History, Washington, DC

WINC Waite Insect and Nematode Collection, Adelaide, Australia

ZMAS Zoological Museum, Academy of Sciences, St. Petersburg, Russia

Terminology and Data Handling — Abbreviations and morphological terms used in text: A1, A2, ... A12: antennomere 1, 2, … 12; claval formula: distribution of the large, multiporous basiconic sensilla on the underside of apical antennomeres of the female, with the segment interval specified followed by the number of sensilla per segment (Bin 1981); POL: posterior ocellar line, the shortest distance between inner margins of posterior ocelli; OOL: ocular ocellar line, the shortest distance from inner orbit and outer margin of lateral ocellus (Masner 1980); T1, T2, ... T7: metasomal tergite

1, 2, ... 7. Morphological terminology follows Masner (1980) and Mikó et al. (2007).

In the Material Examined the numbers prefixed with “OSUC” are unique identifiers for the individual specimens. Since the label data for all specimens have been georeferenced and recorded in the Hymenoptera On-Line database, details on the data associated with these specimens can be accessed at the following link, purl.oclc.org/NET/hymenoptera/hol, and entering the identifier in the form. Note the space between the acronym and the number.

17 Data associated with specimens can be accessed at http://purl.oclc.org/NET/hymenoptera/hol?id=485. Species descriptions were generated using a database application, vSysLab (http://vsyslab.osu.edu), designed to facilitate the production of a taxon by character data matrix, and to integrate this with the existing taxonomic and specimen-level database. Data are exported in both text format and as input files for other applications. The text output for descriptions is in the format of

“Character: Character state (s). Images and measurements were made using

AutoMontage and Cartograph extended-focus software, using JVC KY-F75U digital camera, Leica Z16 APOA microscope, and 1X objectve lens. Images are archived at

Specimage (http://specimage.osu.edu), the image database at The Ohio State University.

In this article we have followed the precedent of Pyle et al. (2008) and Johnson et al. (2008) in the implementation of biodiversity informatics standards within a taxonomic publication. The electronic version of the paper contains hyperlinks to external resources.

Insofar as possible the external information conforms to standards developed and maintained through the organization Biodiversity Information Standards (Taxonomic

Database Working Group). All new species have been prospectively registered with

Zoobank (Polaszek et al. 2005), and other taxonomic names, where appropriate, have been retrospectively registered. The external hyperlinks are explicitly cited in the footnotes so that users of the printed version of this article have access to the same resources. Life sciences identifiers, LSIDs, may be resolved at the specified URLs or at lsid.tdwg.org. This work is conducted as part of the Platygastroidea Planetary

Biodiversity Inventory.

18

RESULTS

GENUS PLATYSCELIO KIEFFER

(HYMENOPTERA: PLATYGASTRIDAE: SCELIONINAE)

Taxonomic reviews — Platyscelio Kieffer, 1905:11 Original description. Type:

Platyscelio pulchricornis Kieffer, by monotypy. Brues, 1908: 27, 40 (diagnosis, list of species, keyed); Kieffer, 1908: 113 (keyed); Kieffer, 1910: 62, 66 (description, list of species, keyed); Dodd, 1913: 130 (keyed); Kieffer, 1913: 222 (description); Brues, 1922:

21 (diagnosis, figure); Kieffer, 1926: 265, 553 (description, keyed, key to species); Mani,

1941: 29 (catalog of species of India); Muesebeck & Walkley, 1956: 386 (citation of type species); Baltazar, 1966: 186 (catalog of species of the Philippines); Masner, 1976: 55

(description); Mani & Sharma, 1982: 190 (description); Galloway & Austin, 1984: 9, 75

(diagnosis, keyed); Johnson, 1992: 461 (catalog of world species); Austin & Field, 1997:

34, 68 (structure of ovipositor system, discussion of phylogenetic relationships); Lê,

2000: 31 (keyed); Rajmohana K., 2006: 117, 128 (description, keyed); Kononova &

Kozlov, 2008: 25, 319 (description, keyed).

Diagnosis —Platyscelio is distinguished from most other genera of Scelioninae sensu Masner (1976) by the strongly dorsoventrally depressed, flat, and foliaceous body.

A small number of other groups, such as Aradophagus Ashmead and Synoditella

19 Muesebeck, are also more or less strongly depressed. Platyscelio may be distinguished from them by the absence of a netrion; the mandibles are tridentate, with the middle tooth small; the postmarginal vein is very short or absent, clearly shorter than the stigmal vein; and the R in the hind wing is complete, extending to the hamuli on the costal margin of the wing.

Description— Moderate-sized, length 3.0–5.6 mm; head prognathous, flattened anteroposteriorly, mesosoma and metasoma strongly dorsoventrally depressed; body black; macropterous.

Head in dorsal view strongly transverse; vertex laterad of posterior ocellus smooth or with few faint striae, between posteriorly ocelli finely longitudinally striate; hyperoccipital carina present as fine ridge on vertex between compound eyes; occipital carina absent; posterior ocellus distinctly separated from inner orbit of compound eyes,

OOL > diameter of ocellus; compound eye large, appearing glabrous; frons without depression, shallowly convex, with median longitudinal sulcus bifurcating dorsally near median ocellus and ventrally near toruli; interantennal process well-developed, narrow; torular triangle present; submedian carina sometimes present; orbital carina sometimes present; lower frons, including cheek, with weak fanlike striae arising from mandibular condyle; shortest distance on frons between eyes less than eye height; inner orbits weakly diverging ventrally; postclypeus strongly projecting above anteclypeus, subtriangular, anteclypeus short, longest medially, lateral corners not produced; malar sulcus present; gena variably expanded, smooth to longitudinally striate or with few faint striae; labrum

20 hidden by clypeus; mandible short, apex with tridentate, middle tooth distinctly shortest, teeth arranged transversely; maxillary palpus 2-segmented, all segments cylindrical; labial palpus 1-segmented, very short; antenna 12-merous in both sexes; radicle very broad, inserted apically into A1, nearly parallel to longitudinal axis of A1, with small lateral spine; A1 almost triangular and expanded outwardly into spine, particularly in female; A2 distinctly shorter than A3; gustatory sensilla on female antenna arranged in longitudinal pairs on apical antennomeres; clava laterally compressed, claval formula

A12–A8: 1-2-2-2-1; male antenna with tyloid on A5.

Mesosoma in dorsal view longer than wide, in lateral view extremely longer than high; pronotum in dorsal view strongly narrowed laterally, anterolateral corner weakly angulate; transverse pronotal carina absent; vertical epomial carina absent; dorsal epomial carina absent; lateral face of pronotum concave; netrion absent; anterior margin of mesoscutum strongly flexed ventrally to meet pronotum; mesoscutum semicircular in outline, posterolateral corner rounded; parapsidal lines visible, some species absent or faint (P. pulchricornis, P. striga, P. umzantsi); notauli variable, if absent create strong carina (P.mysterium); skaphion absent; prespecular sulcus and posterior mesepimeral sulcus present; speculum smooth, rarely longitudinally striate (P. arcuatus) transscutal articulation well-developed; mesoscutellum rectangular in outline, truncate, sculpture smooth to longitudinally striate; axilla small; posterior scutellar sulcus interrupted medially, complete; metanotum narrow, metascutellum clearly differentiated, size of metascutellum variable; dorsal surface of propodeum variable, faintly setation posteriorly; median propodeal sulcus present; plicae well-developed; propodeal

21 projections reduced; sternaulus absent; metapleural pit reduced; metapleural sulcus present separate dorsal and ventral metaplural area; setation of posterior half of ventral metapleural area variable; legs elongate; posterior surface of hind coxa smooth, glabrous to dense setose; trochantellus present; tibial spur formula 1-1-1; tarsal formula 5-5-5; tarsomeres cylindrical, broadening apically; pretarsal claw simple; apex of fore wing extending slightly beyond apex of S5 in female and S6 in male rarely extending S5

(P.striga), hyaline or infuscate; R straight, extending slightly beyond basal half of length of wing, without strong, elongate bristles, gradually approaching costal margin apically, contiguous with costal margin for distance clearly exceeding length of r-rs (i.e., marginal vein elongate); r-rs straight; R1 usually absent, reduced, stumplike in some species (P. arcuatus and P. umzantsi) (i.e., postmarginal vein); bulla absent; Rs+M (basal vein) weakly to clearly indicated, nebulous; hind wing with R extending from base of wing to hamuli; three hamuli present.

Metasoma elongate, parallel-sided, strongly flattened dorsoventrally; female with six visible terga and sterna, male with seven visible terga and sterna; second to fifth segments equal in length, third and fourth widest, subequal to each other in size; submarginal ridge developed, defined by narrow laterotergites to form deep submarginal rim; sublateral carina well-developed on tergites, rarely absent (P. striga); no spiracles visible; sculpture on T1–T4 variable; S1 not laterally compressed; felt field absent.

Distribution— The genus Platyscelio is a widespread group throughout the Old

World tropics, extending from West Africa to Queensland. Four species have relatively

22 restricted distribution: P. arcuatus and P. striga (Western Australia), P. mysterium

(Zimbabwe, Botswana, South Africa), and P. umzantsi (South Africa). The known distribution also suggests two species are widespread; P. africanus in the Afrotropical realm from Senegal east to Yemen and south to South Africa; and P. pulchricornis is found from India, Southeast Asia to Papua New Guinea and eastern Australia.

Key to species of Platyscelio

1. Striae within ocellar triangle sparse (fewer than 20); distance between anterior

ocellus and posterior ocellar line in frontal view greater than or equal to 0.5× POL

(Figs. 2.3E, 2.12E)……………………………………………………………….. 2

– Striae within ocellar triangle dense (more than 20); distance between anterior

ocellus and posterior ocellar line in frontal view less than 0.5× POL (Figs. 1.1E,

2.2E, 2.6E, 2.7E) …………………………………………………………………. 3

2. Sculpture on T1 longitudinally striate, interstices with coriacious microsculpture

(Fig. 2.3F); postmarginal vein absent (Fig. 2.3D); notaulus with mesal margin

arched, lateral margin straight (Fig. 2.3C); legs beyond coxae yellow (Figs. 2.3A,

2.3B); female outer lateral apex of scape sharply pointed (Fig. 2.3E) (Western

Australia)……………………..…… Platyscelio arcuatus Taekul & Johnson, n.sp.

– Sculpture on T1 longitudinally striate throughout (Fig. 2.12F); postmarginal vein

present, stumplike (Fig. 2.12D); notaulus with both mesal and lateral margins

arched (Fig 2.12C); legs beyond coxae brown (Figs. 2.12A); female outer lateral

23 apex of scape bluntly rounded (Fig. 2.12E) (South Africa)…………………………

…………………………………….Platyscelio umzantsi Taekul & Johnson, n.sp.

3. Frontal sculpture between inner orbit and central keel longitudinally striate, striae

either extending through most of length of frons, or with few (4–5) striae restricted

to upper half of frons (Figs. 2.4E, 2.11E)………………………………………….. 4

– Frons between inner orbit and central keel smooth (Figs. 2.1E, 2.2E, 2.5E, 2.6E,

2.7E) ………………………………………………………………………………...5

4. Mesoscutum with one lateral carina; notaulus absent (Fig 2.4C); sculpture on T1

longitudinally striate laterally, uniformly setigerous punctate medially (Fig. 2.2D)

(Zimbabwe, Botswana, South Africa) ……………………………………………….

…………………………………..Platyscelio mysterium Taekul & Johnson, n.sp.

– Mesoscutum with two lateral carinae; notaulus present (Fig. 2.11C); sculpture on

T1 longitudinally striate throughout (Fig 2.11D) (Western Australia)…….…………

………………………………..………. Platyscelio striga Taekul & Johnson, n.sp.

5. Posterior scutellar sulcus complete (Figs 2.1C, 2.1F, 2.2C, 2.2F); margin of

propodeum longitudinally striate laterally, rugulose posteriorly (Figs 2.1F, 2.2F);

vertex between inner orbit and posterior ocellus densely striate (Figs 2.1E, 2.2E)

(Benin, Central African Republic, Ivory Coast, Nigeria, Cameroon, Yemen,

Tanzania, Guinea, Sierra Leone, Guinea-Bissau, Kenya, Mozambique,

Uganda)………………………………………………. Platyscelio africanus Risbec

24 – Posterior scutellar sulcus interrupted medially (Figs. 2.5C, 2.6C, 2.7C, 2.8C,

2.10C); margin of propodeum smooth laterally, longitudinally striate to rugulose

posteriorly vertex between inner orbit and posterior ocellus smooth (Figs. 2.5E,

2.6E, 2.7E) (India, Indonesia, Guam, Malaysia, Papua New Guinea, Philippines,

Thailand) ...... Platyscelio pulchricornis Kieffer

Species Descriptions

Platyscelio africanus Risbec

(Figures 2.1A–F, 2.2A–F)

Platyscelio africanus Risbec, 1956: 105 (original description); Masner, 1976: 55

(description).

Description. Female body length: 3.46–4.28 mm (n=20). Male body length: 2.95–4.35 mm (n=20).

Head: Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: densely striate. Frontal sculpture between inner orbit and central keel: smooth. Submedial ventral area of head anterior to fossa: smooth, finely longitudinally striate posteriorly. Malar sulcus: absent. Orbital carina: absent. Sculpture of malar region: smooth, faintly longitudinal striae with limited microsculpture near eye.

25 Color of female antenna: A1–A7 yellow to light brown, A8–A12 dark brown to black.

Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–

A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: yellow to light brown throughout.

Mesosoma: Sculpture on medial lobe of mesoscutum: longitudinally striate with elongate punctures. Setation of medial lobe of mesoscutum: moderately dense, even.

Notaulus form: mesal and lateral margin arched. Length of notaulus: abbreviated, clearly not reaching anterior margin of mesoscutum. Width of notaulus anteriorly: parallel-sided.

Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: absent. Medial carina of mesoscutum: absent. Notaulus: present. Parapsidial line: present. Posterior scutellar sulcus: complete. Setation of posterior half of ventral metapleural area: sparsely

(less than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: longitudinally striate or with few reticulations. Median propodeal sulcus: widened posteriorly. Sculpture of submedian propodeal field: longitudinally striate. Posterolateral margin of propodeum: longitudinally striate laterally, rugulose posteriorly. Color of legs: coxae dark brown to black, otherwise yellow throughout, rarely hind femur dark brown.

Wings: Female Postmarginal vein: absent. Forewing: hyaline.

Metasoma: Sculpture on T1: longitudinally striate with setigerous punctures medially. Sublateral carina on T2–T4: present anteriorly, absent posteriorly. Sculpture on

T2–T4: setigerous punctures throughout, longitudinally striate anteriorly.

26 Diagnosis. Platyscelio africanus is similar to P. pulchricornis in the lack of sculpture on the frons between the inner orbit and the central keel. It may be distinguished by the complete posterior scutellar sulcus, and the sculpture on the margin of the propodeum is longitudinally striate laterally and rugulose posteriorly.

Material Examined. Holotype female: CAMEROON: Garoua (deposited in

MNHN). Other material: (76 females, 55 males) BENIN: 13 females, 5 males, OSUC

207951–207968 (CNCI). CAMEROON: 17 females, 6 males, BMNH(E)#790211,

848520–848536 (BMNH); OSUC 250657–250661 (CNCI). CENTRAL AFRICAN

REPUBLIC: 1 female, 1 male, OSUC 176086, 247778 (SAMC). GHANA: 2 males,

BMNH(E)#790201, 848510 (BMNH). GUINEA: 1 female, 1 male, OSUC 207895,

250625 (CNCI). GUINEA-BISSAU: 1 female, OSUC 253728 (MZLU). IVORY

COAST: 22 females, 13 males, OSUC 207977–207983, 207986–208007 (CNCI); OSUC

58731–58736 (OSUC). KENYA: 1 female, 7 males, OSUC 207969–207976

(CNCI). MOZAMBIQUE: 3 males, OSUC 207948–207950 (CNCI). NIGERIA: 9 females, 4 males, BMNH(E)#790200 (BMNH); OSUC 250639–250650

(CNCI). SIERRA LEONE: 2 females, 5 males, BMNH(E)#790195, 848506 (BMNH);

OSUC 253722–253726 (MZLU). SOUTH AFRICA: 2 females, 5 males, OSUC 207938,

207943–207946, 250663 (CNCI); OSUC 253727 (MZLU). TANZANIA: 3 females, 1 male, OSUC 253741–253744 (SAMC). TOGO: 1 female, OSUC 253754

(CNCI). UGANDA: 1 female, 1 male, OSUC 207984–207985 (CNCI). YEMEN: 1 female, 1 male, OSUC 250651–250652 (CNCI).ZIMBABWE: 1 female,

BMNH(E)#790209 (BMNH).

27 Comments. This species is widespread in the Afrotropical realm, extending from east Africa to Yemen and south to the north of South Africa. The color of the female antenna is variable: the scape is yellow to light brown, but in some specimens is dark brown to black (OSUC 207985, 207972; Figs 2.2A, 2.2E). The color variability is also seen on the legs: coxae are dark brown to black, otherwise the legs are yellow throughout in most specimens, but in some the hind femur is dark brown (OSUC 207985, 207954,

207955).

Platyscelio arcuatus Taekul & Johnson, new species

(Figures 2.3A–F)

Description. Female body length: 3.48–3.97 mm (n=3). Male body length: 3.87 mm (n=1).

Head: Length between anterior ocellus and posterior ocellar line in frontal view: greater than or equal to 0.5 times POL. Striae within ocellar triangle: sparse (equal to or less than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae. Frontal sculpture between inner orbit and central keel: smooth.

Submedial ventral area of head anterior to fossa: longitudinally striate throughout. Malar sulcus: absent. Orbital carina: absent. Sculpture of malar region: longitudinally striate or with few faint striae.

Antennae: Color of female antenna: dark brown to black throughout. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 concave,

28 closely fitting basal margin of following antennomere. Color of male antenna: dark brown to black throughout.

Mesosoma: Sculpture on medial lobe of mesoscutum: longitudinally striate.

Setation of medial lobe of mesoscutum: sparse to glabrous. Notaulus form: mesal margin arched, lateral margin straight. Length of notaulus: percurrent or nearly so. Width of notaulus anteriorly: narrowed anteriorly. Pilosity of notaulus: present. Number of lateral carinae on mesoscutum: absent. Medial carina of mesoscutum: present. Notaulus: present. Parapsidial line: present. Posterior scutellar sulcus: interrupted medially, complete. Setation of posterior half of ventral metapleural area: sparsely (less than 25 setae). Metascutellum size: narrow, metanotum lateral to metascutellum with 4–6 foveae.

Sculpture on ventral metapleural area: longitudinally striate, coriaceous microsculpture within interstices, strongly reticulate rugose with foveolae. Median propodeal sulcus: narrow throughout length. Sculpture of submedian propodeal field: longitudinally striate, in male interstices with coriacious microsculpture. Posterolateral margin of propodeum: reticulate rugose with foveolae throughout. Color of legs: coxae dark brown to black, otherwise yellow throughout.

Wings: Postmarginal vein: reduced, stumplike. Forewing: hyaline.

Metasoma: Sculpture on T1: longitudinally striate, coriaceous microsculpture within interstices. Sublateral carina on T2–T4: present, percurrent. Sculpture on T2–T4: longitudinally striate with coriaceous microsculpture within interstices.

29 Diagnosis. Platyscelio arcuatus may be separated from Platyscelio striga

(Western Australia) by the less densely striate sculpture within the ocellar triangle (20 striae or fewer).

Etymology. The epithet arcuatus, Latin for bent like a bow, refers to the shaep of the notaulus.

Material Examined. Holotype female: AUSTRALIA: Western Australia,

Keystone Rd., 3 km W Walpole, 34°59.01'S 116°40.76'E, George, Hawks, Munro, YPT,

OSUC 250635 (deposited in CNCI). Paratypes: AUSTRALIA: 2 females, 1 male,

OSUC 250633, 250634, 250636 (CNCI).

Platyscelio mysterium Taekul & Johnson, new species

(Figures 2.4A–F)

Description. Female body length: 3.14–4.23 mm (n=8). Male body length: 3.46–4.2 mm

(n=7).

Head: Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: densely striate. Frontal sculpture between inner orbit and central keel: longitudinally striate, striae extending through most of length of frons. Submedial ventral area of head anterior to fossa: smooth, finely longitudinally striate posteriorly. Malar sulcus: present. Orbital carina: present. Sculpture of malar region: longitudinally striate or with few faint striae.

30 Antennae: Color of female antenna: A1–A7 yellow to light brown, A8–A12 dark brown to black. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: brown , dark brown to black throughout.

Mesosoma: Sculpture on medial lobe of mesoscutum: longitudinally striate with elongate punctures. Setation of medial lobe of mesoscutum: moderately dense, even.

Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: one. Medial carina of mesoscutum: absent. Notaulus: absent. Parapsidial line: present. Posterior scutellar sulcus: complete. Setation of posterior half of ventral metapleural area: sparsely

(less than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: smooth anteriorly, coarsely foveolate punctate posteriorly. Median propodeal sulcus: narrow throughout length. Sculpture of submedian propodeal field: smooth throughout or with few faint striae. Posterolateral margin of propodeum: smooth laterally, longitudinally striate to rugulose posteriorly. Color of legs: coxae dark brown to black, otherwise variable.

Wings: Postmarginal vein: absent. Forewing: hyaline.

Metasoma: Sculpture on T1: longitudinally striate laterally, uniformly setigerous punctate medially. Sublateral carina on T2–T4: present anteriorly, absent posteriorly.

Sculpture on T2–T4: setigerous punctures throughout, longitudinally striate anteriorly.

Diagnosis. Platyscelio mysterium is distinguished from other species by the presence of only a single lateral carina on the mesoscutum, the lack of a notaulus, and the presence of orbital carinae on the frons (Figs 2.4C, 2.4E).

31 Etymology. The epithet mysterium, Latin for mystery, refers to the interpretation of the mesoscutal carinae.

Material Examined. Holotype female: BOTSWANA: Serowe, Farmer's

Brigade, 22.3833°S 26.7167°E, May 1989, Malaise trap, P. Forchhammer, OSUC

171372 (deposited in USNM). Paratypes: (7 females, 14 males) BOTSWANA: 1 female,

OSUC 250665 (CNCI). SOUTH AFRICA: 3 females, 13 males, 1 unknown,

BMNH(E)#790187–790189, 790196–790197, 790199, 848507–848509 (BMNH); OSUC

207935–207937, 207939, 207942, 207947 (CNCI); OSUC 230254

(OSUC). ZIMBABWE: 3 females, 1 male, OSUC 250653–250656 (CNCI).

Comments. Some specimens show variability in the prominence of the sculpture between inner orbit and central keel on the frons.

Platyscelio pulchricornis Kieffer

Figures (2.5A–2.10F)

Platyscelio pulchricornis Kieffer, 1905: 13 (original description); Kieffer, 1926: 553

(description, keyed); Bin, 1974: 458 (type information)

Platyscelio abnormis Crawford, 1910: 126 (original description), new synonymy.

Platyscelio dunensis Mukerjee, 1993: 78 (original description), new synonymy.

Platyscelio mirabilis Dodd, 1913: 132 (original description), new synonymy.

Platyscelio punctatus Kieffer, 1913: 321 (original description), new synonymy.

32 Platyscelio wilcoxi Fullaway, 1913: 283 (original description); Kieffer, 1926: 553, 555

(description, keyed); Yasumatsu, 1941: 76 (junior synonym of Platyscelio abnormis

Crawford); Masner & Muesebeck, 1968: 42 (type information).

Description. Female body length: 3.31–5.59 mm (n=20). Male body length: 3.24–4.71 mm (n=20).

Head: Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae.

Frontal sculpture between inner orbit and central keel: smooth. Submedial ventral area of head anterior to fossa: smooth, finely longitudinally striate posteriorly. Malar sulcus: absent. Orbital carina: absent. Sculpture of malar region: smooth, longitudinally striate or with few faint striae.

Antennae: Color of female antenna: dark brown to black throughout, A1–A7 yellow to light brown, A8–A12 dark brown to black, or A1–A7 brown, A8–A12 dark brown to black, in some antennae sequencially darker from scape to apex. Female outer lateral apex of scape: sharply pointed. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: brown.

Mesosoma: Sculpture on medial lobe of mesoscutum: longitudinally striate with elongate punctures. Setation of medial lobe of mesoscutum: moderately dense, even.

Notaulus form: mesal and lateral margin arched. Length of notaulus: percurrent or nearly so, abbreviated, clearly not reaching anterior margin of mesoscutum. Width of notaulus

33 anteriorly: parallel-sided. Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: absent. Medial carina of mesoscutum: absent. Notaulus: present. Parapsidial line: absent or faint. Posterior scutellar sulcus: interrupted medially. Setation of posterior half of ventral metapleural area: dense (more than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: smooth to faintly logitudinally striate, longitudinally striate or with few reticulations. Median propodeal sulcus: narrow throughout length, widened posteriorly.

Sculpture of submedian propodeal field: smooth throughout or with few faint striae, longitudinally striate. Posterolateral margin of propodeum: smooth laterally, longitudinally striate to rugulose posteriorly. Color of legs: coxae dark brown to black, otherwise brown, or coxae dark brown to black, otherwise yellow throughout.

Wings: Postmarginal vein: absent. Forewing: hyaline, infuscate.

Metasoma: Sculpture on T1: longitudinally striate laterally, uniformly setigerous punctate medially. Sublateral carina on T2–T4: present, percurrent. Sculpture on T2–T4: uniformly setigerous punctate.

Diagnosis. Platyscelio pulchricornis can be separated from Platyscelio africanus by the medial absence of the posterior scutellar sulcus and the smoothness of submarginal propodeal field (Fig 2.6C, 2.7C, 2.8C, 2.8F).

Material Examined. Holotype female, Platyscelio pulchricornis: PAPUA NEW

GUINEA: Madang Prov., Dilo, VI.1890–VII.1890, none specified, Loria, MCSN 0004

(deposited in MCSN). Holotype male, Platyscelio abnormis: PHILIPPINES: R.E.

Brown, (deposited in USNM Cat. No. 12895). Holotype male, Platyscelio dunensis:

34 INDIA: Uttarakhand, Rishiskesh, 24.VI.1991, Mukerjee & party (deposited in USNM).

Holotype female, Platyscelio mirabilis: AUSTRALIA: Queensland, Nelson, 16.II.1912, sweeping open forest (deposited in SAMA). Syntype male, Platyscelio punctatus:

PHILIPPINES: Los Baños (deposited in USNM). Holotype male, Platyscelio wilcoxi:

GUAM: USNM No. 26186 (deposited in USNM).

Other material: (105 females, 51 males, 3 unknowns) AUSTRALIA: 1 female, 4 males, BMNH(E)#790202-790203, 790208 (BMNH); OSUC 250637 (CNCI); OSUC

141949 (OSUC). BANGLADESH: 1 female, OSUC 173855 (CNCI). CHINA: 2 females, 4 males, BMNH(E)#848537-848540 (BMNH); OSUC 321841-321842 (CNCI).

GUAM: 3 females, 1 male, OSUC 207850-207851, 207856, 207858 (USNM). INDIA: 7 females, 2 males, BMNH(E)#790185, 790206-790207, 848518-848519 (BMNH); OSUC

230648 (OSUC); OSUC 207838-207840 (USNM). INDONESIA: 13 females, 9 males,

BMNH(E)#790210 (BMNH); OSUC 207859-207874, 207876, 207893-207894, 207896

(CNCI); OSUC 204851 (UCDC). JAPAN: 1 female, OSUC 173082 (UASK).

MALAYSIA: 9 females, 5 males, BMNH(E)#790193, 790204 (BMNH); OSUC 207877-

207883, 207885-207887, 207892 (CNCI); OSUC 207884 (SCAU). PAPUA NEW

GUINEA: 9 females, 5 males, BMNH(E)#790205, 848511-848517 (BMNH); OSUC

207897, 250626-250628 (CNCI); OSUC 160037-160038 (EMEC). PHILIPPINES: 2 females, 6 males, BMNH(E)#790184 (BMNH); OSUC 207875 (CNCI); OSUC 207842-

207847 (USNM). SOLOMON ISLANDS: 1 female, BMNH(E)#848543 (BMNH).

TAIWAN: 18 females, 11 males, OSUC 173803-173831 (TARI). THAILAND: 36 females, 2 males, 2 unknowns, BMNH(E)#790190, 790192, 790194, 848502-848505,

35 848541-848542 (BMNH); OSUC 207888-207890 (CNCI); OSUC 207837, 215797,

253701-253721, 253760-253762, 253764 (OSUC); OSUC 207891 (SCAU).

VANUATU: 2 males, 1 unknown, BMNH(E)#790183, 790186, 790198 (BMNH).

VIETNAM: 2 females, OSUC 277708-277709 (RMNH).

Comments. Some specimens show variability in antenna and leg color. Because of the uniformity of the principal characters – notaulus form and length, interrupted posterior scutellar sulcus, and the propodeal sculpture – we consider these specimens to be conspecific with Platyscelio pulchricornis.

Platyscelio striga Taekul & Johnson, new species

(Figures 2.11 A–F)

Description. Female body length: 2.89–23.4 mm (n=2). Male body length: 3.05–

3.22 mm (n=2).

Head: Length between anterior ocellus and posterior ocellar line in frontal view: less than 0.5 times POL. Striae within ocellar triangle: dense (greater than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae.

Frontal sculpture between inner orbit and central keel: with few (4–5) striae, striae limited to upper half of frons. Submedial ventral area of head anterior to fossa: longitudinally striate throughout. Malar sulcus: absent. Orbital carina: absent. Sculpture of malar region: longitudinally striate or with few faint striae.

36 Antennae: Color of female antenna: A1–A7 brown, A8–A12 dark brown to black, in some antennae sequencially darker from scape to apex. Female outer lateral apex of scape: sharply pointed. Claval shape: apicali margin of A9–A11 straight, following antennomere distinctly separated. Color of male antenna: dark brown to black throughout.

Mesosoma: Sculpture on medial lobe of mesoscutum: longitudinally striate.

Setation of medial lobe of mesoscutum: sparse to glabrous. Notaulus form: mesal margin arched, lateral margin straight. Length of notaulus: abbreviated, clearly not reaching anterior margin of mesoscutum. Width of notaulus anteriorly: narrowed anteriorly.

Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: two. Medial carina of mesoscutum: absent. Notaulus: present. Parapsidial line: absent or faint.

Posterior scutellar sulcus: complete. Setation of posterior half of ventral metapleural area: sparsely (less than 25 setae). Metascutellum size: wide, metanotum lateral to metascutellum reduced, with 0–3 foveae. Sculpture on ventral metapleural area: strongly reticulate rugose with foveolae . Median propodeal sulcus: narrow throughout length.

Sculpture of submedian propodeal field: smooth throughout or with few faint striae.

Posterolateral margin of propodeum: longitudinally striate laterally, rugulose posteriorly.

Color of legs: coxae dark brown to black, otherwise yellow in female and brown in male.

Wings: Postmarginal vein: absent. Forewing: hyaline.

Metasoma: Sculpture on T1: longitudinally striate throughout. Sublateral carina on T2–T4: absent. Sculpture on T2–T4: longitudinally striate throughout, uniformly setigerous punctate within interstices on T3–T4.

37 Diagnosis. Platyscelio striga is unique in the genus with the possession of two lateral carina on mesoscutum (Fig 2.11C), and the absence of sublateral carina on T2–T4

(Figs. 2.11A, 2.11D).

Etymology. The epithet striga, Latin for swath, refers to the distinct striae of the frons.

Material Examined. Holotype female: AUSTRALIA: Western Australia, Cape

Arid N.P., 33.5189°S 123.4342°E (WDPA-UN), 30.XII–3.I.1987, Malaise trap, J.S.

Noyes, OSUC 250630 (deposited in CNCI). Paratypes: AUSTRALIA: 1 female, 2 males,

OSUC 250629, 250631, 250632 (CNCI).

Platyscelio umzantsi Taekul & Johnson, new species

(Figures 2.12 A–F)

Description. Female body length: 3.14–4.23 mm (n=20). Male body length:

3.21–4.53 mm (n=20).

Head: Length between anterior ocellus and posterior ocellar line in frontal view: greater than or equal to 0.5 times POL. Striae within ocellar triangle: sparse (equal to or less than 20). Vertex sculpture between inner orbit and posterior ocellus: smooth or with few faint striae. Frontal sculpture between inner orbit and central keel: smooth.

Submedial ventral area of head anterior to fossa: longitudinally striate throughout. Malar sulcus: absent. Orbital carina: absent. Sculpture of malar region: longitudinally striate or with few faint striae.

38 Antennae: Color of female antenna: dark brown to black throughout. Female outer lateral apex of scape: bluntly rounded. Claval shape: apical margin of A9–A11 concave, closely fitting basal margin of following antennomere. Color of male antenna: dark brown to black throughout.

Mesosoma: Sculpture on medial lobe of mesoscutum: longitudinally striate.

Setation of medial lobe of mesoscutum: moderately dense, even. Notaulus form: mesal and lateral margin arched. Length of notaulus: percurrent or nearly so. Width of notaulus anteriorly: parallel-sided. Pilosity of notaulus: absent. Number of lateral carinae on mesoscutum: absent. Medial carina of mesoscutum: absent. Notaulus: present. Parapsidial line: present, absent or faint. Posterior scutellar sulcus: interrupted medially. Setation of posterior half of ventral metapleural area: sparsely (less than 25 setae). Metascutellum size: narrow, metanotum lateral to metascutellum with 4–6 foveae. Sculpture on ventral metapleural area: longitudinally striate or with few reticulations. Median propodeal sulcus: narrow throughout length. Sculpture of submedian propodeal field: longitudinally striate. Posterolateral margin of propodeum: margined by coarsely punctate furrow. Color of legs: coxae dark brown to black, otherwise brown.

Wings: Postmarginal vein: reduced, stumplike. Forewing: hyaline, infuscate.

Metasoma: Sculpture on T1: longitudinally striate throughout. Sublateral carina on T2–T4: present, percurrent. Sculpture on T2–T4: T2 longitudinally striate throughout,

T3–T4 longitudinally reticulate laterally, smooth medially.

Diagnosis. Platyscelio umzansi may be separated from other African species by the sparse striae within ocellar triangle (20 or fewer), the narrow metascutellum, and the

39 presence of 4–6 foveae on the metanotum laterad of the metascutellum (Figs 2.12E,

2.12F). Some specimens have the sculpture on the submedian propodeal field strongly effaced.

Etymology. The epithet umzantsi, Xhosa for south, is a reference to the the collecting locality.

Material Examined. Holotype female: SOUTH AFRICA:

34°27.414'S19°21.393'E Western Cape, Walker Bay Nat. Res., 57 m, Site1, Malaise trap,

S. Coast Strandveld, 4.X–1.XI.1997, S.van Noort, WB97–M11, OSUC 243790

(deposited in SAMC). Paratypes: SOUTH AFRICA: 63 females, 31 males, OSUC

202440–202441 (AEIC); OSUC 207908–207934, 207941, 250662, 250664 (CNCI);

OSUC 266101–266102 (MZLU); OSUC 188488–188489, 207830–207836, 207898–

207907, 207940, 213995, 226020–226023, 237213–237217, 243454–243458, 243506–

243507, 243790–243791, 253729–253740, 253745–253753 (SAMC).

40

CONCLUSION

The genus Platyscelio Kieffer (Hymenoptera: Platygastridae, Scelioninae) is a widespread group in the Old World, found from West Africa to Queensland, Australia.

The species concepts are revised and a key to world species is presented. The genus is comprised of 6 species, including 2 known species which are redescribed: P. africanus

(Benin, Central African Republic, Ivory Coast, Nigeria, Cameroon, Yemen, Tanzania,

Guinea, Sierra Leone, Guinea-Bissau, Kenya, Mozambique, Uganda); and P. pulchricornis (India, Indonesia, Guam, Malaysia, Papua New Guinea, Philippines,

Thailand). Four species-group names are considered to be junior synonyms of P. pulchricornis: P. abnormis Crawford, P. dunensis Mukerjee, P. mirabilis Dodd, and P. punctatus Kieffer. The following species are hypothesized and described as new taxa: P. arcuatus Taekul & Johnson, n.sp. (Western Australia); P. mysterium Taekul & Johnson, n.sp. (Zimbabwe, Botswana, South Africa); P. striga Taekul & Johnson, n.sp. (Western

Australia); and P. umzantsi Taekul & Johnson, n.sp. (South Africa).

41 42 43 44 45 46 47 48 49 50 51 52 53 54

CHAPTER 3

WORLD SPECIES EXPLORATION OF THE GENUS OXYTELEIA KIEFFER

(HYMENOPTERA: PLATYGASTRIDAE)

INTRODUCTION

The genus Oxyteleia was originally described by Kieffer (1908) on the basis of the single species Caloteleia bidentata Kieffer 1905, from Papua New Guinea. In 1908 he also transferred three additional Caloteleia species, so that the genus originally contained four species, O.bidentata and O. subdentata (Kieffer) from Papua New Guinea,

O. punctata (Ashmead) and O.nigriceps (Ashmead) collected from Saint Vincent and

Grenada respectively (Kieffer 1908). Masner (1976) later transferred O. nigriceps to the genus Probaryconus. Kieffer (1914) also described the genus Dilapitha for two species from the Philippines and later in 1926 he presented identification keys for both genera, at that time placing six species in Oxyteleia and three species in Dilapitha (Kieffer 1926).

Masner (1976) synonymized Dilapitha with Oxyteleia. Six species presently are recorded

55 in Asia, Australia and Oceania: O. albipes (Kieffer), O. nitida (Kieffer), O.variipennis

(Kieffer), O. panchganii (Mani), O.bidentata (Kieffer) and O. subdentata (Kieffer). Only two species are recorded from Neotropical realm, O. aenea (Ashmead), and O. punctata

(Ashmead), from Saint Vincent (Johnson 1992).

Masner (1976) classified Oxyteleia (Kieffer), Merriwa (Dodd), and Nyleta (Dodd) in the tribe Psilanteridini. The genus Oxyeteleia was distinguished from other two by the absence of the skaphion, complete notauli, and the presence of two lateral spines on mesoscutellum in some species. The genus Nyleta differs by the absence of both notauli and lateral mesoscutellar spines. Merriwa possesses complete notauli and mesoscutellar spines. Masner’s commented that these genera are different only in few characters and may result in misidentification. He also suggested that based on morphology, Oxyteleia is surrounded by several closely related genera of Scelioninae in Indo-Australia. Dr. L.

Masner (personal communication) has suggested that Merriwa should be considered to be a junior synonym of Oxyteleia.

Two species, O. aenea and O. punctata were originally described by Ashmead

(1894) from the New World. This generic assignment of the former dates to Kieffer

(1926), and the distribution of these species strongly contrasts that of all other known species of the genus. Dr. L. Masner (Canadian National Collection of Insects, Ottawa,

Canada) and Dr. A. Polaszek (The Natural History Museum, London, UK) will revise

Ashmead’s type material, and thus the scope of this research will not cover the examination of the type specimens of these species.

56 In the 107 years since the original description, Oxyteleia has never been comprehensively reviewed or revised. The known hosts of species in the subfamilies

Scelioninae, Teleasinae, and Telenominae are the eggs of insects and spiders. The host of

Oxyteleia, however, is still unclear. The goal of this research is to produce a systematic revision of the world species of genus Oxyteleia. This will include a description of the genus, an identification key to species, descriptions or redescriptions of all species, examination of all primary types, a literature review, databasing of all available specimen records.

57

MATERIALS AND METHODS

Materials —This work is based upon specimens in the following collections, with abbreviations used in the text;

AEIC American Entomological Institute, Gainesville, FL

AMNH American Museum of Natural History, New York, NY

ANIC Australian National Insect Collection, Canberra, Australia

BMNH The Natural History Museum, London, UK

BPBM Bernice P. Bishop Museum, Honolulu, HI

CASC California Academy of Sciences, San Francisco, CA

CNCI Canadian National Collection of Insects, Ottawa, Canada

MCSN Museo Civico de Storia Naturale, Genoa, Italy

MCZC Museum of Comparative Zoology, Harvard University

Cambridge, MA

MZLU Lund Museum of Zoology, Lund University, Lund, Sweden

OSUC C.A. Triplehorn Insect Collection, The Ohio State University,

Columbus, OH

QMBA Queensland Museum, Brisbane, Australia

RMNH Nationaal Natuurhistorisch Museum, Leiden, Netherlands

ROME Royal Ontario Museum, Toronto, Canada

SCAU South China Agricultural University, Guangzhou, China

58 TARI Taiwan Agricultural Research Institute - Entomology,

Taichung, Taiwan

USNM National Museum of Natural History, Washington, DC

WINC Waite Insect and Nematode Collection, Adelaide, Australia

Terminology and Data Handling — Abbreviations and morphological terms used in text: A1, A2,... A12: antennomere 1, 2, … 12; claval formula: distribution of the large, multiporous basiconic sensilla on the underside of apical antennomeres of the female, with the segment interval specified followed by the number of sensilla per segment (Bin

1981); POL: posterior ocellar line, the shortest distance between inner margins of posterior ocelli; OOL: ocular ocellar line, the shortest distance from inner orbit and outer margin of lateral ocellus (Masner 1980); T1, T2,... T7: metasomal tergite 1, 2,... 7.

Morphological terminology follows Masner (1980) and Mikó et al. (2007).

In the Material Examined the numbers prefixed with “OSUC” are unique identifiers for the individual specimens. Since the label data for all specimens have been georeferenced and recorded in the Hymenoptera On-Line database, details on the data associated with these specimens can be accessed at the following link, purl.oclc.org/NET/hymenoptera/hol, and entering the identifier in the form. Note the space between the acronym and the number.

Data associated with specimens of Oxyteleia can be accessed at http://purl.oclc.org/NET/hymenoptera/hol?id=485. Species descriptions were generated using a database application, vSysLab (http://vsyslab.osu.edu), designed to facilitate the

59 production of a taxon by character data matrix, and to integrate this with the existing taxonomic and specimen-level database. Data may be exported in both text format and as input files for other applications. The text output for descriptions is in the format of

“Character: Character state (s). Polymorphic characters have their multiple character states separated by semicolons. Images and measurements were made using

AutoMontage and Cartograph extended-focus software, using JVC KY-F75U digital camera, Leica Z16 APOA microscope, and 1X objectve lens. Images are archived at

Specimage (http://specimage.osu.edu), the image database at The Ohio State University.

In this article we have followed the precedent of Pyle et al. (2008) and Johnson et al. (2008) in the implementation of biodiversity informatics standards within a taxonomic publication. Insofar as possible the external information conforms to standards developed and maintained through the organization Biodiversity Information Standards (Taxonomic

Database Working Group). The external hyperlinks are explicitly cited in the footnotes so that users of the printed version of this article have access to the same resources. Life sciences identifiers, LSIDs, may be resolved at the specified URLs or at lsid.tdwg.org.

This work is conducted as part of the Platygastroidea Planetary Biodiversity Inventory.

60

RESULTS

GENUS OXYTELEIA KIEFFER

(HYMENOPTERA: PLATYGASTRIDAE: SCELIONINAE)

Oxyteleia Kieffer urn:lsid:biosci.ohio-state.edu:osuc_concepts:530

(Figures 3.1 A–F –3.37 A–F)

Oxyteleia Kieffer, 1908: 118 (original description. Type: Caloteleia bidentata

Kieffer, designated by Kieffer (1926). Keyed); Kieffer, 1910: 64, 78 (description, list of species, keyed); Kieffer, 1913: 221, 228 (description); Kieffer, 1926: 269, 516

(description, keyed, key to species); Dodd, 1927: 72 (diagnosis); Muesebeck & Walkley,

1956: 378 (citation of type species); Masner, 1976: 48 (description, key to Merriwa

Dodd, Oxyteleia Kieffer, and Nyleta Dodd); Masner, 1976: 9, 48 (keyed, synonymy); De

Santis, 1980: 312 (catalog of species of Brazil); Mani & Sharma, 1982: 189 (description);

Galloway & Austin, 1984: 49 (the only species from Australia transferred to Neoparidris

Galloway); Johnson, 1992: 455 (catalog of world species).

61 Dilapitha Kieffer, 1914: 293 (original description. Type: Dilapitha albipes Kieffer, by

original designation. Synonymized by Masner (1976); Kieffer, 1926: 269, 404

(description, keyed, key to species); Muesebeck & Walkley, 1956: 346 (citation

of type species); Baltazar, 1966: 179 (catalog of species of the Philippines);

Masner, 1976: 48 (junior synonym of Oxyteleia Kieffer).

Merriwa Dodd, 1920: 332 (Original description, Type: Merriwa quadridentata Dodd, by

monotypy and original designation; Muesebeck & Walkley, 1956: 368 (Citation

of type species); Masner, 1976: 48 (Key to Merriwa Dodd, Oxyteleia Kieffer, and

Nyleta Dodd); Johnson, 1992: 437 (Catalog of world species); Austin & Field,

1997: 26, 68 (Structure of ovipositor system, discussion of phylogenetic

relationships); Lê, 2000: 32. (Keyed). New synonymy.

Description. Female body length: 2.30-5.11 mm (n=173). Male body length:

3.15-4.53 mm (n=12).

Head shape in dorsal view: transverse. Hyperoccipital carina: absent; present.

Occipital carina: present, complete medially; present laterally, broadly interrupted medially. Occipital carina sculpture: crenulate; punctate; unsculptured. OOL: lateral ocellus nearly contiguous with inner orbits, OOL < 0.5 OD. Upper frons: convex, without frontal shelf. Scrobe shape: frons with shallow unmargined depression above toruli.

Submedian carina: absent. Orbital carina: present. Inner orbits: diverging ventrally.

IOS/EH: IOS distinctly less than EH. Interantennal process: rounded, strongly developed.

62 Central keel: present, extending through entire frontal depression. Torulus opening: laterally on interantennal process. Lower frons striae: present. Malar sulcus: present.

Compound eye size: very large, dominating lateral portion of head. Compound eye setation: glabrous; sparsely setose. Gena: narrow, weakly convex, receding behind posterior orbit. Apical margin of clypeus: straight. Labrum: not visible. Mandible shape: moderate. Mandibular teeth: apex with 3, acute, subequal teeth. Arrangement of mandibular teeth: transverse. Shape of maxillary palpomeres: cylindrical.

Number of antennomeres in female: 12. Number of antennomeres in male: 12.

Insertion of radicle into A1: parallel to longitudinal axis of A1. Shape of A1: more or less cylindrical, not flattened. Length of A3 of female: distinctly longer than A2. Number of clavomeres in female antenna: 6. Claval formula of female antenna: A12-A7/1-2-2-2-2-1.

Arrangement of doubled multiporous plate sensilla on female clava: in longitudinal pairs.

Tyloid distribution on male antenna: none visible. Shape of male flagellum: filiform.

Mesosoma shape in dorsal view: longer than wide. Mesosoma shape in lateral view: longer than high; quadrate. Medial portion of transverse pronotal carina: absent.

Posterior apex of pronotum in dorsal view: straight, bifid apically to articulate with tegula. Vertical epomial carina: present. Dorsal epomial carina (corresponding to lateral portion of transverse pronotal carina of Vilhelmsen et al. 2010) : present; absent. Anterior face of pronotum: vertical, not visible in dorsal view. Lateral face of pronotum: weakly concave below position of dorsal epomial carina. Netrion: present. Netrion shape: moderately wide, open ventrally. Anterior margin of mesoscutum: vertical, flexed ventrally to meet pronotum. Mesoscutum shape: pentagonal in outline, posterolateral

63 corner rounded. Skaphion: absent; present. Notauli: present, percurrent; absent; present, abbreviated. Parapsidal lines: present; absent. Admedial lines: present; absent.

Transscutal articulation: well-developed, narrow. Shape of mesoscutellum: quadrate to trapezoidal; semielliptical; pentagonal, pointed posteriorly. Armature of mesoscutellum: absent; axillula produced posteriorly into short, broad spines. Surface of mesoscutellum: convex throughout. Median longitudinal furrow on mesoscutellum: absent. Shape of axillula: small, dorsal margin sinuate. Metascutellum: clearly differentiated.

Metascutellar armature: bidentate, teeth widely separated. Metascutellar setation: glabrous. Metapostnotum: not defined externally. Extent of metasomal depression of propodeum: percurrent, extending anteriorly to anterior margin of propodeum. Lateral propodeal projection: absent. Mesopleural carina: present, extending at least to sternaulus. Mesal course of acetabular carina: straight. Mesopleural pit: present.

Sternaulus: absent. Posterodorsal corner of mesopleuron: rounded anteriorly.

Number of mid tibial spurs: 1. Number of hind tibial spurs: 1. Dorsal surface of hind coxa: smooth. Hind tibia shape: cylindrical, ecarinate.

Wing development of female: macropterous. Wing development of male: macropterous. Tubular veins in fore wing: present. Bulla of fore wing R: absent. Extent of marginal venation of fore wing: distinct marginal or postmarginal veins developed.

Origin of r-rs in fore wing: arising from marginal vein along costal margin. Development of basal vein (Rs+M) in fore wing: nebulous, weakly pigmented; nebulous, strongly pigmented. Development of R in hind wing: elongate, extending to costal margin.

64 Number of external terga in female: 6. Number of external sterna in female: 6.

Number of external terga in male: 7. Number of external sterna in male: 7. Shape of metasoma: acuminate, widest submedially. Laterotergites: present, narrow.

Laterosternites: present. T1 of female: produced medially into cylindrical or elliptical horn housing ovipositor. Relative size of metasomal segments: T3 distinctly largest.

Terga with basal crenulae: T1-T2. Sublateral carinae on tergites: present on T1-T4; present on T1-T3. Median longitudinal carina on metasomal terga: absent. Shape of female T6: flattened. Shape of posterior margin of male T7: straight. Distribution of felt fields: absent. Ovipositor type: Scelio-type (Austin & Field 1997).

Diagnosis. Oxyteleia is distinguished from most other genera of Scelioninae by the glabrous and extremely large eyes, narrow temples and cheeks, lateral ocelli nearly contiguous with the inner orbits, and the long postmarginalis (much longer than stigmalis) (Masner 1976).

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=530]

65 Key to species of Oxyteleia

1. Skaphion present (Figs. 3.1–3.15C); gena reticulate punctate and with fanlike

striae ventrally ...... 2

– Skaphion absent (Figs. 3.16–3.37C); sculpture of gena variable ...... 11

2. Notaulus present; parapsidal line present (Figs. 3.3C, 3.4C); occipital carina

usually complete medially ...... 4

– Notaulus absent; parapsidal line absent; occipital carina absent dorsally (Figs.

3.1C, 3.2C) ...... 3

3. Mesoscutellum semicircular, posterior margin rounded (Fig. 3.1C); subdivided

cells within metapleural triangle, poorly defined; sculpture of dorsal half of frons

in female reticulate rugose with sparse setigerous punctures; color of female A2-

A5 sequentially lighter from dark brown to light yellow, A6 dark brown (3.1F)

(Thailand) ...... O. inthanona Taekul n.sp.

– Mesoscutellum shape pentagonal, posterior margin converging to slight point

medially (Fig. 3.2C); subdivided cells within metapleural triangle, well defined;

sculpture of dorsal half of frons in female predominantly transversely strigose,

with granular microsculpture within interstices, sculpture attenuating ventrally;

color of female funicle A2-A6 antennae sequentially lighter from dark brown to

light yellow (Fig. 3.2F); (Thailand, Laos, Viet Nam) ..... O. aenigma Taekul n.sp.

4. Mesoscutellum pentagonal, posterior margin converging to slight point medially

(Figs. 3.8C 3.9C 3.10C, 3.11C) ...... 6

66 – Mesoscutellum strongly transverse, posterior margin deeply concave usually

housing horn of T1 (Figs. 3.3C, 3.4C), or transverse with posterior margin straight

(rectangular shape) (Figs. 3.5) ...... 5

5. Sculpture of posterior vertex and occiput reticulate rugose; sculpture of medial

lobe of mesoscutum completely covered with fine dense rugulae; mesoscutellum

shape strongly transverse, posterior margin deeply concave housing horn of T1;

medial carina of mesoscutum present percurrent or present posteriorly (Figs.

3.3C, 3.4C) (Thailand, Laos, Malaysia, Indonesia, Malaysia) O. oecus Taekul n.sp.

– Sculpture of posterior vertex and occiput transversely strigose, setigerous

punctures ventrally; sculpture of medial lobe of mesoscutum smooth throughout,

or at most with sparse puncture; mesoscutellum shape transverse with posterior

margin straight, rectangular; medial carina of mesoscutum absent (Fig. 3.5C)

(Philippines) ...... O. pangasugana Taekul n.sp.

6. Occipital carina complete; medial lobe of mesoscutum completely covered with

fine dense rugulae or dense setigerous punctures; sculpture of mesoscutellum

almost entirely covered by microsculpture (Figs 3.8C,F, 3.9C, 3.10C, 3.11C,

3.13C) ...... 8

– Occipital carina absent dorsally; medial lobe of mesoscutum smooth throughout,

or at most with sparse puncture; mesoscutellum predominantly smooth, or at most

with sparse puncture (Figs 3.6C, 3.7C) ...... 7

67 7. Ocellar triangle reticulate rugose with granular microsculpture within interstices

(Fig 3.6C) (Malaysia) ...... O. angularis Taekul n.sp.

– Ocellar triangle reticulate rugose, microsculpture absent (Fig 3.7C) (Philippines)

...... O. rugosa Taekul n.sp.

8. Posterior vertex and occiput transversely strigose with microsculpture within

interstices, densely punctate ventrally (Figs. 3.8C,F, 3.9C,F);sculpture of dorsal

half of frons in female predominantly transversely strigose, with microsculpture

within interstices, sculpture attenuating ventrally (Figs. 3.8E, 3.9E, 3.10E, 3.11E);

sculpture of ocellar triangle reticulate rugose with microsculpture within

interstices; axillular spine elongate, length distinctly greater than width (Figs.

3.8F 3.9C 3.11C) ...... 9

– Posterior vertex and occiput transversely strigose, setigerous punctures ventrally

(Figs. 3.13C 3.14C); sculpture of dorsal half of frons in female transversely

carinate ventrally, rugulose dorsally or transverse carinate throughout (Figs. 3.14E

3.15E); sculpture of ocellar triangle reticulate rugose; axillular spine short to

almost absent ...... 10

9. Posterior margin of skaphion convex, with two submedial points (Figs. 3.8C,

3.8F); parapsidal line present as broad groove (Fig. 3.8F); female A2 – A4 dark

brown, A5, A6 white (Fig. 3.8B) (Malaysia, Indonesia) ......

...... O. bimucronata Taekul n.sp.

68 – Posterior margin of skaphion concave (Figs. 3.9CF, 3.10C, 3.11C); parapsidal line

present, indicated as carina attenuating dorsally(Figs. 3.9C, 3.10C, 3.11C); female

A2-A6 yellow to dark brown throughout or sometimes A2-A6 sequentially lighter

from dark brown to light yellow (Fig. 3.9B 3.10B) (Thailand, India, Indonesia,

Laos, Malaysia, Taiwan, Vietnam) ......

...... O. epidemosa Taekul n.sp.

10. Forewing membrane generally hyaline throughout or moderately infuscate

throughout (Figs. 3.13A, 3.14A); female A1 white to light yellow (Figs. 3.13F

3.14F), A2-A6 sequentially lighter from dark brown to light brown; medial carina

of mesoscutum present percurrent or sometimes present posteriorly (Figs. 3.12–

3.14C) (Indonesia, Malaysia) ...... O. quadridentata (Dodd)

– Forewing membrane hyaline basally, brown infuscate apically, with strong white

streak below marginal vein and at distal apex (Figs. 3.15A, 3.15F); female A1

dark brown (Fig. 3.15B), A2-A6 antennae sequentially lighter from dark brown to

light yellow; medial carina of mesoscutum absent (Fig. 3.15C); (Malaysia)

...... O. alba Taekul n.sp.

11. Posterior vertex and occiput entirely smooth, or predominantly smooth with

sparse punctures, in some with weak strigae medially (Figs. 3.16C,F, 3.17C,

3.18C) ...... 12

– Sculpture of posterior vertex and occiput variable ...... 20

69 12. Sculpture of mesoscutellum predominantly smooth, or at most with sparse

punctures (Fig. 3.16C); gena predominantly smooth, with weak crenulae adjacent

to eye and ocipital carina, dorsally with fine microsculpture (Fig. 3.16D); notauli

percurrent, indicated as weak narrow grooves, converging posteriorly (Figs. 3.16–

3.18C); clava (A7–A12) bicolored, dark brown to light yellow (Figs 3.16B,

3.17A,B, 3.18B,F) ...... 13

– Mesoscutellum setigerously punctate throughout (Figs. 3.19C); gena smooth

throughout; notaulus variable; clava (A7–A12) dark brown to black throughout

(Figs. 3.19–3.21B) ...... 15

13. Ocellar triangle transversely concentrically strigose (Fig. 3.16F); axillular spine

elongate, length distinctly greater than width (Fig. 3.16C); netrion glabrous (Fig.

3.16D) (Philippines) ...... O. arcuata Taekul n.sp.

– Ocellar triangle predominantly smooth, or rugulose forming small puncture;

length of axillular spine short to almost absent;netrion setose ...... 14

14. Sculpture of dorsal half of frons in female smooth with densely white setation on

antennal scrobe (Fig. 3.17E); mesoscutellum more or less semicircular, posterior

margin rounded (Fig. 3.17C) (Indonesia) ...... O. talamasi Taekul n.sp.

– Sculpture of dorsal half of frons in female with transverse carinae throughout (Fig

3.18E), without dense white setae on scrobe; mesoscutellum pentagonal, posterior

margin converging to slight point medially (Fig. 3.18C) (Philippines) O. lux Taekul n.sp.

70 15. Coxae yellow (Figs 3.19–3.21B); metasoma in female bicolored, dark brown to

black and yellow to amber-orange with variable patterns (Figs. 3.19–3.21F);

sublateral carina on T3 absent ...... 16

– Coxae brown to black (Figs 3.22–3.23B); metasoma in female dark brown to

black (Figs 3.22–3.24A); sublateral carina on T3 present ...... 18

16. Posterior vertex sharply truncate, anterior divided from posterior by sharp

transverse carina (Fig. 3.19C) (Fiji) ...... O. scopulus Taekul n.sp.

– Posterior vertex more or less evenly rounded from anterior to posterior (Figs.

3.20C, 3.21C) ...... 17

17. Medial lobe of mesoscutum setigerously punctate throughout (Fig 3.20C);

sublateral carina on T2 absent (Fig. 3.20F); medial area of T2-T3 defined by

carina or striga absent (Fig. 3.20F) (Solomon Islands) ......

...... O. solomona Taekul n.sp.

– Medial lobe of mesoscutum smooth throughout, or at most with sparse punctures

(Fig 3.21C); sublateral carina on T2 present (Fig 3.21F); medial area of T2-T3

defined by carina or striga present (Fig. 3.21F); Papua New Guinea

...... O. insula Taekul n.sp.

18. Sculpture of dorsal half of frons in female entirely smooth with sparse punctures,

in some ventral half transversely carinate (Fig. 3.22E); female A2-A6 sequentially

lighter from dark brown to light yellow (Fig. 3.22F) (Indonesia) ......

...... O. lenisa Taekul n.sp.

71 – Sculpture of dorsal half of frons in female transversely carinate ventrally,

rugulose dorsally (Figs 3.23E, 3.24E); female funicle (A2-A6) dark brown to

black throughout (Fig. 3.23B) ...... 19

19. T1 horn in female longitudinally striate, striae present at least in basal half, in

some striae reaching apex of horn (Fig. 3.23F); sculpture of T2 longitudinally

rugose (Papua New Guinea, Indonesia) ...... O. striga Taekul n.sp.

– Basal half of T1 horn in female transversely carinate, apical half smooth (Fig.

3.24F); T2 predominantly rugose with punctures within interstices, longitudinally

striate (Indonesia) ...... O. ponticulus Taekul n.sp.

20. Genal carina behind compound eyes present (Figs. 3.25F, 3.26F, 3.27D) ...... 21

– Genal carina behind compound eyes absent (Figs. 3.29D,F, 3.30D) ...... 22

21. Posterior vertex and occiput concentrically rugose, in some rugae reduced with

surface somewhat smoother (Figs. 3.25C, 3.26C); medial lobe of mesoscutum

smooth throughout, at most with sparse punctures (Figs. 3.25C), or with faint

rugulae emerging from medial carina, sculpture attenuating apically, sparsely

punctate anteriorly (Fig. 3.26C) (Papua New Guinea, Indonesia, Malaysia,

Philippines) ...... O. arcuza Taekul n.sp.

– Posterior vertex and occiput transversely strigose, with setigerous punctures

ventrally (Fig. 3.27C); medial lobe of mesoscutum longitudinally strigose,

reticulate medially, with setigerous punctures within interstices (Fig. 3.27C);

Papua New Guinea ...... O. apektos Taekul n.sp.

72 22. Gena behind compound eyes smooth throughout, or predominantly smooth, with

weak crenulae adjacent eye and occipital carina, dorsally with fine microsculpture

(Figs. 3.30D, 3.31D) ...... 24

– Gena behind compound eyes densely punctate with interstices setose (Figs.

3.28D, 3.29D, F) ...... 23

23. Speculum canaliculate throughout (Fig 3.28D); (Fiji) .... O. johnsoni Taekul n.sp.

– Speculum crenulate along posterior margin (Fig. 3.29) (Philippines) ......

...... O. lagunasis Taekul n.sp.

24. Notaulus absent; lateral pronotal area adjacent to netrion punctate with crenulae

ventrally (Figs. 3.30–32C) ...... 25

– Notaulus present; lateral pronotal area adjacent to netrion smooth with sparse

setation (Figs. 3.33–37C) ...... 27

25. Malar region with fanlike striae (Fig. 3.31D); female A1 white to light yellow

sometimes bicolored white and dark brown; fore coxa yellow (Figs. 3.31B,

3.32B) ...... 26

– Malar region smooth (Fig 3.30D); female A1 dark brown to black; fore coxa

brown (Fig. 3.30B, D) (Papua New Guinea, Philippines) ......

...... O. primata Taekul n.sp.

26. Posterior margin of mesoscutellum deeply concave housing horn of T1 (Fig.

3.31C); transverse carina present on T1 horn (Fig 3.31E) (Solomon Islands) ......

73 ...... O. astricta Taekul n.sp.

– Posterior margin straight, rectangular (Fig. 3.32C); transverse carina of horn on

T1 absent (Fig. 3.32F) (Solomon Islands) ...... O. catheta Taekul n.sp.

27. Occipital carina absent dorsally (Figs. 3.36C, 3.37C); sculpture of posterior vertex

and occiput variable; female A1 dark brown; medial area of T2-T3 not

demarcated by raised carina (Figs. 3.36F) ...... 28

– Occipital carina complete (Figs. 3.33–35C); posterior vertex and occiput

concentrically rugose, in some rugae reduced with surface somewhat smoother

(Figs. 3.33–35C); female A1 bicolored, white and dark brown apically; medial

area of T2-T3 defined by carina or striga (Papua New Guinea) ......

...... O. bidentata (Dodd)

28. Medial lobe of mesoscutum weakly transversely rugose with punctures within

interstices (Fig. 3.36C); malar region with fanlike striae (Fig. 3.36D); posterior

vertex and occiput transversely strigose, with setigerous punctures ventrally (Fig.

3.36C) (Papua New Guinea) ...... O. obscura Taekul n.sp.

– Medial lobe of mesoscutum longitudinally striate, striae attenuating apically (Fig.

3.37C, 3.37F); malar region smooth (Fig. 3.37D); posterior vertex and occiput

predominantly smooth with sparse punctures, transversely strigose dorsally (Fig.

3.37C); (Indonesia, Papua New Guinea) ...... O. fasciata Taekul n.sp

74 Species Descriptions

Oxyteleia inthanona Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307026

(Figures 3.1 A–F)

Description. Female body length: 3.61-3.84 mm (n=3). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: transversely strigose with granulose microsculpture within interstices, densely punctate ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: reticulate rugose with granular microsculpture within interstices. Sculpture of dorsal half of frons in female: reticulate rugose with sparse setigerous punctures. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: reticulate punctate with fanlike striae ventrally. Setation of gena: densely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A5 sequentially lighter from dark brown to light yellow, A6 dark brown. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: rugulose.

Skaphion: present. Shape of posterior margin of skaphion: evenly arched. Notaulus:

75 absent. Parapsidal line: absent. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: completely covered with fine dense rugulae. Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae. Sculpture of mesoscutellum: almost entirely covered by reticulate microsculpture. Medial carina of mesoscutum: absent.

Mesoscutellum shape: semicircular, posterior margin rounded. Length of axillular spine: elongate, length distinctly greater than width. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent.

Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose.

Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: punctate. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent.

Sculpture of T3: with longitudinal interconnected rugae. Sublateral carina on T3: absent.

76 Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: longitudinally striate, coriaceous microsculpture within interstices. Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: absent.

Diagnosis. Oxyteleia inthanona is distinguishable from Oxyteleia species with a skaphion by the color of the female antenna in which A2–A5 are sequentially lighter from dark brown to light yellow, and A6 is dark brown to black

Etymology. The epithet inthanona, refers to the collecting locality, Doi Inthanon

National Park, Chang Mai, Thailand. Doi Inthanon is the highest mountain in Thailand; the name given in honor of the King Inthawichayanon, one of the last kings of Chiang

Mai before merging with Siam or Thailand. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307026]

Material Examined. Holotype, female: THAILAND: Chiang Mai Prov., checkpoint 2, T1909, Doi Inthanon National Park, 18°31.554'N 98°29.940'E, 1700m,

14.XI-15.XI.2006, pan trap, Y. Areeluck, OSUC 266812 (deposited in OSUC).

Paratypes: THAILAND: 2 females, OSUC 266799-266800 (CNCI).

Oxyteleia aenigma Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307025

Figures (Figures 3.2 A–F)

77 Description. Female body length: 3.10-4.08 mm (n=20). Male body length: 3.25 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: transversely strigose with granulose microsculpture within interstices, densely punctate ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: reticulate rugose with granular microsculpture within interstices. Sculpture of dorsal half of frons in female: predominantly transversely strigose, with granular microsculpture within interstices, sculpture attenuating ventrally.

Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate.

Sculpture of gena: reticulate punctate with fanlike striae ventrally. Setation of gena: densely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A6 antennae sequentially lighter from dark brown to light yellow. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: present. Shape of posterior margin of skaphion: evenly arched. Notaulus: absent.

Parapsidal line: absent; present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: completely covered with fine dense rugulae. Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae. Sculpture of

78 mesoscutellum: almost entirely covered by reticulate microsculpture. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: elongate, length distinctly greater than width. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined.

Dorsal metapleural sulcus: absent.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent.

Sculpture of T3: with longitudinal interconnected rugae. Sublateral carina on T3: absent.

Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: longitudinally striate. Sublateral carina on T2: absent.

79 Diagnosis. Two species of Oxyteleia possess the skaphion, but lack notauli: O. inthanona and O. aenigma. Oxyteleia aenigma is distinguishable from O. inthanona by the color of the female antenna: A2-A6 is sequentially lighter from dark brown to light yellow (with A6 light yellow).

Etymology. The epithet aenigma, Latin for obscure, riddle, mystery, is a reference to the complex characters of antennae funicle and mesoscutellum shape which may mislead the diagnosis to O. inthanona. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307025]

Material Examined. Holotype, female: THAILAND: Chiang Mai Prov., nature trail, T2829, Doi Chiang Dao Wildlife Sanctuary, 19°24.187'N 98°55.312'E, 491m,

31.VII-7.VIII.2007, malaise trap, S. Jagsu & A. Watwanich, OSUC 173884 (deposited in

OSUC). Paratypes: (26 females, 2 males) LAOS: 9 females, OSUC 234022, 234461,

234532, 234545, 234554, 234557 (BPBM); OSUC 266675, 266699, 276630 (CNCI).

THAILAND: 9 females, 1 male, OSUC 173885, 173888, 173892, 173898, 256812,

266810, 280644, 359301, 359377, 359380 (OSUC). VIETNAM: 8 females, 1 male,

OSUC 234015-234017 (BPBM); OSUC 266852, 277536, 277539, 277661, 277683-

277684 (RMNH).

Comment. The mesoscutellum in O. aenigma normally is pentagonal with the posterior margin converging to a slight point medially. There are two female specimens

(OSUC 359301, OSUC 173898), however, in which the shape of the mesoscutellum is more semicircular.

80 Oxyteleia oecus Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307027

(Figures 3.3A – F, 3.4A – F)

Description. Female body length: 3.10-3.50 mm (n=21). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: reticulate rugose. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose, interstices smooth.

Sculpture of dorsal half of frons in female: with oblique strigae near anterior ocellus, smooth ventrally. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: reticulate punctate with fanlike striae ventrally. Setation of gena: densely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A6 antennae sequentially lighter from dark brown to light yellow. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: rugulose.

Skaphion: present. Shape of posterior margin of skaphion: evenly arched. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal

81 humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: deeply foveolate. Sculpture of medial lobe of mesoscutum: completely covered with fine dense rugulae. Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae. Sculpture of mesoscutellum: almost entirely covered by reticulate microsculpture. Medial carina of mesoscutum: present posteriorly; present, percurrent. Mesoscutellum shape: strongly transverse, posterior margin deeply concave housing horn of T1. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: glabrous. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous anteriorly, with sparse setation posteriorly. Setation of metapleural triangle: sparsely to densely setose.

Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: deeply sulcate.

Coxae color: all coxae dark brown basally, light brown to yellow apically.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: longitudinally strigose with granular microsculpture within interstices. Sculpture of T3:

82 with longitudinal interconnected rugae. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: longitudinally striate. Sublateral carina on T2: absent.

Diagnosis. Oxyteleia oecus is unique among the species with a skaphion in the possession of a medial carina on the mesoscutum and the strongly transverse shape of the mesoscutellum, sometimes with the posterior margin deeply concave to house the horn of

T1(Fig 3.3C).

Etymology. The epithet oecus, Latin for room or house, refers to the strongly transverse mesoscutellum. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307027]

Material Examined. Holotype, female: THAILAND: Trang Prov., Na Yong

Dist., Khao Chong Mountain, 07°33'18"N 99°47'22"E, 75m, 15.I-20.I.2005, malaise trap,

M. Sharkey, OSUC 266674 (deposited in CNCI). Paratypes: (44 females) INDONESIA:

37 females, OSUC 253602-253603, 253605-253606, 253608, 253772-253775, 253780,

253782-253783, 253785, 253789-253800, 266683, 266685, 276564, 276567, 276577-

276579, 276582-276584, 276587, 276634 (CNCI). LAOS: 1 female, OSUC 234549

(BPBM). MALAYSIA: 2 females, OSUC 234560 (BPBM); OSUC 276540 (CNCI).

THAILAND: 4 females, OSUC 266665-266666 (CNCI); OSUC 359375, 361724

(OSUC).

Comments. There are two groups sharing the distinctive characters in this species: 1) specimens from Thailand with the mesoscutellum strongly transverse, its posterior margin deeply concave and having the medial carina on the mesoscutum (Fig

83 3.3C); and 2) specimens collected from Indonesia with the rectangular mesoscutellum and the medial carina on the medial lobe of mesoscutum present posteriorly (Fig 3.4C).

Oxyteleia pangasugana Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307031

(Figures 3.5 A – F)

Description. Female body length: 3.15 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: entirely transversely strigose, lacking microsculpture between striae, setigerous punctures ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose, interstices smooth. Sculpture of dorsal half of frons in female: smooth with sparse setigerous punctures dorsally. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: reticulate punctate with fanlike striae ventrally. Setation of gena: densely setose.

Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A6 antennae sequentially lighter from dark brown to light yellow.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area:

84 narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: present. Shape of posterior margin of skaphion: evenly arched. Notaulus: present. Shape of notaulus: indicated as weak narrow groove. Course of notauli: converging posteriorly.

Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: deeply foveolate. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: transverse, posterior margin straight. Length of axillular spine: short to almost absent.

Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae dorsally. Netrion: setose. Mesopleural carina: present.

Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: deeply sulcate.

Coxae color: all coxae dark brown basally, light brown to yellow apically.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae

85 present only at base. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent. Sculpture of T3: punctate, punctures attenuating medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially.

Sublateral carina on T2: absent.

Diagnosis. Oxyteleia pangasugana is most similar to O. oecus and may be distinguished from it by the absence of a medial carina on the mesoscutum and the rectangular mesoscutellum.

Etymology. The epithet pangasugana, refers to the collecting locality Mount

Pangasugan. Mount Pangasugan is located to the north of the town of Baybbay. The mountain is renowned in the number of plant and animal species. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307031]

Material Examined. Holotype, female: PHILIPPINES: Leyte Prov., nr. Baybay,

Mount Pangasugan, 10°45'N 124°50'E, 250m, 28.V-30.V.1987, D. C. Darling, OSUC

276527 (deposited in CNCI).

Oxyteleia angularis Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307030

(Figures 3.6 A – F)

86 Description.Female body length: 2.97 mm (n=1). Body color: bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: entirely transversely strigose, lacking microsculpture between striae, setigerous punctures ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: reticulate rugose with granular microsculpture within interstices. Sculpture of dorsal half of frons in female: predominantly transversely strigose, with granular microsculpture within interstices, sculpture attenuating ventrally.

Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate.

Sculpture of gena: smooth throughout. Setation of gena: glabrous to sparsely setose.

Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A6 antennae sequentially lighter from dark brown to light yellow. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: amber-orange throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: smooth throughout. Skaphion: present.

Shape of posterior margin of skaphion: evenly arched. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel. Parapsidal line: absent. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to

87 sparsely punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: glabrous. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: smooth throughout. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: glabrous. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: fore coxa brown, mid and hind coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: finely longitudinally striate with faint area medially, microsculpture absent. Sculpture of T3: longitudinally striate laterally, smooth medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: smooth with only sparse punctures. Sublateral carina on T2: absent.

88 Diagnosis. Oxyteleia angularis is distinguishable from species with a skaphion by the smooth sculpture of the medial lobe of mesoscutum, and it may be separated from O. sepia by the reticulate rugose sculpture and the granular microsculpture of the ocellar triangle.

Etymology. The epithet angularis, Latin for angle or corner, refers to the sculpture on ocellar triangle. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307030]

Material Examined. Holotype, female: MALAYSIA: Sabah St., Borneo Isl.,

Sandakan, 20.XI.1958, T. C. Maa, OSUC 234583 (deposited in BPBM).

Oxyteleia rugosa Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307029

(Figures 3.7 A – F)

Description. Female body length: 3.28-3.32 mm (n=2). Body color: bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: concentrically rugose, rugae sometimes reduced with surface somewhat smoother. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: reticulate rugose, interstices smooth. Sculpture of dorsal half of frons in female: transversely carinate ventrally, rugulose dorsally. Dense setae within antennal

89 scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A4 dark brown, A5-A6 white. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: smooth throughout.

Skaphion: present. Shape of posterior margin of skaphion: evenly arched. Notaulus: present. Shape of notaulus: indicated as weak narrow groove. Course of notauli: converging posteriorly. Parapsidal line: absent. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent.

Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially.

Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, attenuated dorsally. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: glabrous. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: smooth throughout. Sculpture of ventral

90 mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: glabrous. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: fore coxa brown, mid and hind coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: amber-orange to light brown. Transverse carina of horn on T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent. Sculpture of T3: punctate, punctures attenuating medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially.

Sublateral carina on T2: absent.

Diagnosis. Oxyteleia rugosa is one of the Oxyteleia species with smooth sculpture on medial lobe of mesoscutum. It may be distinguished from O. angularis by the reticulate rugose sculpture of the ocellar triangle.

Etymology. The epithet rugosa, Latin for wrinkled or rough, refers to the reticulate rugae of ocellar triangle. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307029]

91 Material Examined. Holotype, female: PHILIPPINES: Ifugao Prov., Mayoyao,

1200-1500m, 3.IX.1966, H. M. Torrevillas, OSUC 234104 (deposited in BPBM).

Paratype: PHILIPPINES: 1 female, OSUC 234106 (BPBM).

Oxyteleia bimucronata Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307052

(Figures 3.8 A – F)

Description. Female body length: 4.60-5.11 mm (n=3). Male body length: 4.00-

4.53 mm (n=4). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: transversely strigose with granulose microsculpture within interstices, densely punctate ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose with granular microsculpture within interstices. Sculpture of dorsal half of frons in female: predominantly transversely strigose, with granular microsculpture within interstices, sculpture attenuating ventrally.

Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate.

Sculpture of gena: reticulate punctate with fanlike striae ventrally. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A4 dark brown, A5-A6 white. Claval color (A7-A12) : dark brown to black throughout.

92 Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: present. Shape of posterior margin of skaphion: sinuous, with two submedial points. Notaulus: present.

Shape of notaulus: indicated as broad groove. Course of notauli: parallel. Parapsidal line: present, indicated as broad groove. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: completely covered with fine dense rugulae. Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae. Sculpture of mesoscutellum: almost entirely covered by reticulate microsculpture. Medial carina of mesoscutum: absent.

Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially.

Length of axillular spine: elongate, length distinctly greater than width. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae brown to black; all coxae yellow.

Color of fore wing membrane: infuscate throughout.

93 Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent.

Sculpture of T3: finely punctate reticulate throughout. Sublateral carina on T3: absent.

Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: present.

Diagnosis. Oxyteleia bimucronata is unique in among species with a skaphion and can be distinguished in both sexes by the two submedial points on the posterior margin of the skaphion.

Etymology. The epithet bimucronata, mucronata Latin for pointed, refers to the submedial points on the posterior margin of the skaphion. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307052]

Material Examined. Holotype, female: MALAYSIA: Sabah St., Borneo Isl.,

Liwagu River, Kinabalu National Park, 1500m, 23.V.1987, screen sweeping, A. Smetana,

OSUC 253618 (deposited in CNCI). Paratypes: (2 females, 6 males) INDONESIA: 1 male, OSUC 266689 (CNCI). MALAYSIA: 2 females, 5 males, OSUC 234537, 234584,

234587 (BPBM); OSUC 253621, 276561 (CNCI); OSUC 266764-266765 (USNM).

Oxyteleia epidemosa Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307053

94 (Figures 3.9 A–F, 3.10 A–F, 3.11 A–F)

Description. Female body length: 3.31-4.47 mm (n=20). Body color: bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: transversely strigose with granulose microsculpture within interstices, densely punctate ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose with granular microsculpture within interstices. Sculpture of dorsal half of frons in female: predominantly transversely strigose, with granular microsculpture within interstices, sculpture attenuating ventrally.

Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate.

Sculpture of gena: reticulate punctate with fanlike striae ventrally. Setation of gena: densely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: unicolorous, yellow to dark brown throughout; A2-A6 antennae sequentially lighter from dark brown to light yellow. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: orange with dark brown patch on posteromedial mesoscutum, dark brown color sometimes extending to mesoscutellum. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: present. Shape of posterior margin of skaphion: evenly arched. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli:

95 parallel. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: completely covered with fine dense rugulae. Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae. Sculpture of mesoscutellum: almost entirely covered by reticulate microsculpture. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: elongate, length distinctly greater than width. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally. Netrion: setose.

Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent.

96 Sculpture of T3: finely punctate reticulate throughout. Sublateral carina on T3: absent.

Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: present.

Diagnosis. Oxyteleia epidemosa and O. bimucronata are separated from other skaphion species by the sculpture of posterior vertex and occiput which are transversely strigose with granular microsculpture within the interstices. Oxyteleia epidemosa is distinguished from O.bimucronata by the yellow coxa and the concave posterior margin of the skaphion.

Etymology. The epithet epidemosa, Greek for prevalent, spreading through a large number, refers to the widespread distribution and high variability of this species.

The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307053]

Material Examined. Holotype, female: THAILAND: Loei Prov., Hua Dong

Tham Sun, T1124, Phu Rua National Park, 17°29.540'N 101°20.995'E, 1130m, 26.XI-

2.XII.2006, malaise trap, P. Tumtip, OSUC 266804 (deposited in OSUC). Paratypes:

(187 females, 1 unknown) INDIA: 1 female, OSUC 266680 (CNCI). INDONESIA: 35 females, OSUC 234078 (BPBM); OSUC 266681-266682, 266684, 266686-266687,

276551, 276589-276595, 276597-276604, 276611, 276619-276621, 276626 (CNCI);

OSUC 173893, 173895-173897, 256843, 283061-283062, 361779 (ROME). LAOS: 9 females, OSUC 233002, 234407, 234548, 234550-234551, 234553, 234559 (BPBM);

OSUC 266700, 266704 (CNCI). MALAYSIA: 22 females, OSUC 234006, 234009,

234544, 234569-234570, 234579, 234603 (BPBM); OSUC 253620, 276541, 276569-

97 276571, 276607-276608, 276627-276628, 276633 (CNCI); OSUC 359373-359374

(MZLU); OSUC 381326-381327, 414887 (OSUC). TAIWAN: 56 females, OSUC

234028 (BPBM); CASENT 2137985 (CASC); OSUC 253622, 253625-253640, 253667,

266690-266692, 266695, 276487-276505, 276507-276512, 276514-276516, 276520-

276521, 276536, 276539 (CNCI). THAILAND: 44 females, OSUC 266664, 266667-

266673, 266801, 276573 (CNCI); OSUC 173875-173877, 173880-173881, 173883,

173886, 173889-173891, 173894, 237598-237599, 237601, 237659, 254598-254599,

266802-266803, 266805-266809, 266811, 266813-266814, 359298-359300, 359302,

359376, 359378-359379 (OSUC). VIETNAM: 20 females, 1 unknown, OSUC 234011,

234014, 234020 (BPBM); OSUC 276606, 276617-276618, 276623, 276635 (CNCI);

OSUC 266851, 266860-266862, 277364, 277520, 277543, 277553, 277697 (RMNH);

OSUC 232270, 240071, 240073, 240075 (ROME). Other material: TAIWAN: 1 female,

OSUC 276506 (CNCI).

Comments. Oxyteleia epidemosa is the most widespread species in the genus distributed from Taiwan to India, Vietnam, Laos, Thailand, Malaysia and Indonesia.The general characters of the specimens vary geographically: specimens from Taiwan (3.3–

3.5 mm body length, mesosoma and metasoma orange amber), Thailand (3.6–3.9 mm. body length, mesosoma bicolored with yellow with dark brown pattern dorsally, metasoma some specimens entirely black, in some bicolored: black and yellow),

Malaysia and Indonesia (4.2-4.5 mm body length, meso and metasoma entirely black).

Despite these differences in the general characters, they are otherwise identical. I consider these specimens all to be conspecific.

98 Oxyteleia quadridentata Taekul, new combination urn:lsid:biosci.ohio-state.edu:osuc_concepts:4911

(Figures 3.12 A–F, 3.13 A–F, 3.14 A–F)

Merriwa quadridentata Dodd, 1920: 333 (original description); Masner, 1965: 85 (type information).

Description. Female body length: 2.61-3.72 mm (n=19). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: entirely transversely strigose, lacking microsculpture between striae, setigerous punctures ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose, interstices smooth. Sculpture of dorsal half of frons in female: transversely carinate throughout; transversely carinate ventrally, rugulose dorsally. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: A2-A6 antennae sequentially lighter from dark brown to light brown. Claval color (A7-A12) : dark brown to black throughout; sequentially lighter from dark brown to light yellow.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area:

99 narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: present. Shape of posterior margin of skaphion: evenly arched; sinuous, with two submedial points. Notaulus: present. Shape of notaulus: indicated as weak narrow groove.

Course of notauli: converging posteriorly. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: densely setigerously punctate with reticulate rugae posteriorly. Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: present posteriorly; present, percurrent.

Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially.

Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth.

Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: sparsely to densely setose.

Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: all coxae brown to black.

100 Color of fore wing membrane: generally hyaline throughout; infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex; with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent. Sculpture of T3: longitudinally striate laterally, smooth medially. Sublateral carina on T3: absent.

Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: present.

Diagnosis. Oxyteleia quadridentata is unique among species with a skaphion in having medial carina on mesoscutum present at least posteriorly.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=4911].

Material Examined. Holotype, male: INDONESIA: Jawa Barat Prov., Java Isl.,

Cibodas (Tjibodas), 5000-7000ft, VIII-1913, Koningsberger, B.M. TYPE HYM. 9.465

(deposited in BMNH). Other material: (27 females, 1 unknown) INDONESIA: 25 females, 1 unknown, BMNH(E) #790167 (BMNH); OSUC 253601, 253604, 253611,

253765-253766, 253768-253770, 253776-253779, 253781, 253784, 253786, 276548-

276550, 276552-276553, 276565-276566, 276580, 276586, 276622 (CNCI).

MALAYSIA: 2 females, OSUC 276560 (CNCI); OSUC 376764 (MCZC)

Comments. Some specimens show variability in the development of the medial carina on the mesoscutum: it may be percurrent, present posteriorly, or developed as fine

101 longitudinal striae attenuating anteriorly. The clava may be sequentially dark brown to light yellow, as is seen in three specimens: OSUC 276580, OSUC 276622, and OSUC

253784.

Oxyteleia alba Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307024

(Figures 3.15 A – F) Morphbank

Description. Female body length: 3.72 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: entirely transversely strigose, lacking microsculpture between striae, setigerous punctures ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose, interstices smooth. Sculpture of dorsal half of frons in female: transversely carinate ventrally, rugulose dorsally. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: reticulate punctate with fanlike striae ventrally. Setation of gena: densely setose.

Genal carina behind compound eye: absent.

Color of female A1: dark brown. Color of female A2-A6: A2-A6 antennae sequentially lighter from dark brown to light yellow. Claval color (A7-A12) : dark brown to black throughout.

102 Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: strongly punctate. Skaphion: present. Shape of posterior margin of skaphion: evenly arched.

Notaulus: present. Shape of notaulus: indicated as weak narrow groove. Course of notauli: converging posteriorly. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: densely setigerously punctate with faint punctures posteriorly, sometimes with weak longitudinally striae posteriorly.

Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae.

Sculpture of mesoscutellum: almost entirely covered by reticulate microsculpture. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: short to almost absent.

Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin.

Sculpture of ventral mesepisternum below mesopleural carina: punctate. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose.

Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: deeply sulcate.

Coxae color: all coxae brown to black.

103 Color of fore wing membrane: hyaline basally, brown infuscate half, with white band below marginal vein and white at distal apex.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent.

Sculpture of T3: finely punctate reticulate throughout. Sublateral carina on T3: absent.

Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: absent.

Diagnosis. Oxyteleia alba is unique in the genus in the possession of white streak below marginal vein and distal apex of forewing.

Etymology. The epithet alba, Latin for white, refers to the white streak on forewing. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307024]

Material Examined. Holotype, female: MALAYSIA: Sabah St., Borneo Isl.,

Kinabalu National Park, VIII-1999, malaise trap, D. Quicke, OSUC 276610 (deposited in

CNCI).

Oxyteleia arcuata Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307033

(Figures 3.16 A – F)

104 Description. Female body length: 2.88 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: transversely concentrically strigose, without microsculpture. Sculpture of dorsal half of frons in female: transversely carinate throughout. Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: dark brown. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : sequentially lighter from dark brown to light yellow.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as weak narrow groove. Course of notauli: converging posteriorly. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel.

Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to sparsely

105 punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: elongate, length distinctly greater than width. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete.

Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally.

Netrion: glabrous. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: sparsely to densely setose.

Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae brown to black.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae throughout, microsculpture absent. Sculpture of T3: punctate, punctures attenuating medially.

Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: absent.

106 Diagnosis. Oxyteleia arcuata is in the group without a skaphion and having a smooth mesoscutellum. It can be distinguished from O. talamasi and O. lux by the transversely concentric strigose sculpture on the ocellar triangle.

Etymology. The epithet arcuata, Latin for arch, refers to the sculpture on ocellar triangle. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307033]

Material Examined. Holotype, female: PHILIPPINES: Negros Oriental Prov.,

7km W Valencia, 1° forest edge, ROM 873065, Cuernos de Negros Mountain, 09°17'N

123°15'E, 700m, 31.VII-7.VIII.1987, malaise trap/pan trap, D. C. Darling & E. Mayordo,

OSUC 256842 (deposited in ROME).

Oxyteleia talamasi Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307034

Figures 3.17 (A – F)

Description. Female body length: 2.65-2.84 mm (n=7). Body color: bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose.

Sculpture of dorsal half of frons in female: entirely smooth with sparse punctures, in

107 some ventral half with faint transverse carinae. Dense setae within antennal scrobe: present. Sculpture of malar region: smooth. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: densely setose.

Genal carina behind compound eye: absent.

Color of female A1: dark brown. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : A7 dark brown to black, A8-A12 white to light yellow.

Color of mesosoma in female: amber-orange throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: smooth throughout. Skaphion: absent.

Notaulus: present. Shape of notaulus: indicated as weak narrow groove. Course of notauli: converging posteriorly. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel.

Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent.

Mesoscutellum shape: semicircular, posterior margin rounded. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: short, only slightly longer than wide. Vertical epomial carina: present, attenuated dorsally. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: smooth

108 throughout. Sculpture of ventral mesepisternum below mesopleural carina: smooth throughout. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: sparsely to densely setose.

Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae yellow.

Color of fore wing membrane: generally hyaline throughout.

Color of metasoma in female: amber-orange to light brown. Transverse carina of horn on T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae throughout, microsculpture absent. Sculpture of T3: finely longitudinally striate laterally, reticulate punctate medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially.

Sublateral carina on T2: absent.

Diagnosis. The smooth sculpture of dorsal half of the frons and the dense white setation on the antennal scrobe is unique to Oxyteleia talamasi and will immediately distinguish this species.

Etymology. The epithet talamasi, refers to Elijah Talamas, an officemate who has shared the good and bad times as well as providing moral support during the graduate career with CT. The epithet is used as a noun in the genitive case.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307034]

109 Material Examined. Holotype, female: INDONESIA: Sulawesi Utara Prov.,

Lake Danau, Kotamobagu, 1300m, V-1985, malaise trap, J. Noyes, OSUC 276557

(deposited in CNCI). Paratypes: INDONESIA: 8 females, OSUC 253787-253788,

276558, 276568, 276581, 276585, 276588 (CNCI); OSUC 58674 (OSUC).

Oxyteleia lux Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307035

(Figures 3.18 A – F)

Description. Female body length: 2.50-3.14 mm (n=7). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: rugulose, otherwise with sparse setigerous punctures on smooth background. Sculpture of dorsal half of frons in female: transversely carinate throughout. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

110 Color of female A1: dark brown; bicolored, white with dark brown apex. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : sequentially lighter from dark brown to light yellow.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as weak narrow groove. Course of notauli: converging posteriorly. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to sparsely punctate.

Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture.

Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, attenuated dorsally. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous anteriorly, with sparse setation

111 posteriorly. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: fore coxa brown, mid and hind coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae anteriorly, reticulate posteriorly, microsculpture absent. Sculpture of T3: punctate, punctures attenuating medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially.

Sublateral carina on T2: absent.

Diagnosis. Oxyteleia lux is separated from O. arcuata and O. talamasi by the transverse carinae on the frons and the clava which is sequentially lighter in color from dark brown to light yellow.

Etymology. The epithet lux, Latin for light, refers to the bright color of the clava.

The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307035]

Material Examined. Holotype, female: PHILIPPINES: Leyte Prov., nr. Baybay,

Mount Pangasugan, 10°45'N 124°50'E, 250m, 28.V-30.V.1987, D. C. Darling, OSUC

276526 (deposited in CNCI). Paratypes: PHILIPPINES: 6 females, OSUC 276524-

276525, 276535, 276575, 276605, 276612 (CNCI).

112 Oxyteleia scopulus Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307032

(Figures 3.19 A – F)

Description. Female body length: 3.23-3.88 mm (n=5). Male body length: 3.33 mm (n=1). Body color: bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: entirely smooth. Transition from vertex to occiput: sharply angled, anterior divided from posterior by sharp transverse carina.

Occipital carina: absent dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose. Sculpture of dorsal half of frons in female: entirely smooth with sparse punctures, in some ventral half with faint transverse carinae. Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth. Sculpture of gena: smooth throughout.

Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: bicolored, white with dark brown apex. Color of female A2-

A6: A2-A4 dark brown, A5-A6 white. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: orange with dark brown patch on posteromedial mesoscutum, dark brown color sometimes extending to mesoscutellum; amber-orange throughout. Dorsal pronotal area: absent. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: strongly punctate. Skaphion: absent.

Notaulus: absent. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal

113 humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: setigerously punctate throughout. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: transverse, posterior margin straight; pentagonal, posterior margin converging to slight point medially. Length of axillular spine: elongate, length distinctly greater than width.

Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: glabrous. Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: deeply sulcate.

Coxae color: all coxae yellow.

Color of fore wing membrane: generally hyaline throughout.

Color of metasoma in female: bicolored, dark brown to black and patterned with yellow to amber-orange. Transverse carina of horn on T1: absent. Sculpture of horn on

T1 in female: longitudinally striate, striae converging posteriorly to form medial carina.

Sculpture of T2: with straight longitudinal striae throughout, microsculpture absent.

Sculpture of T3: longitudinally striate laterally, smooth medially. Sublateral carina on T3:

114 absent. Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: finely punctate throughout. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: absent.

Diagnosis. The sharply truncate posterior vertex is unique to Oxyteleia scopulus.

Etymology. The epithet scopulus, Latin for cliff, refers to the truncate vertex which is separated from the occiput by a sharp transverse carina. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307032]

Material Examined. Holotype, female: FIJI: Central Div., Rewa Prov., Viti

Levu Isl., 4km WSW Suva Hill (Colo-i-Suva) Village, MT3, Mount Nakobalevu,

18.055°S 178.424°E, 372m, 17.III-9.IV.2003, malaise trap, Schlinger & Tokota'a,

FBA143133 (deposited in CNCI). Paratypes: FIJI: 4 females, 1 male, OSUC 234109,

234120 (BPBM); FBA055087, FBA101573, OSUC 266698 (CNCI).

Oxyteleia solomona Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307049

(Figures 3.20 A – F)

Description. Female body length: 2.81-3.82 mm (n=4). Body color: bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent

115 dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose.

Sculpture of dorsal half of frons in female: dorsally smooth with sparsely setigerous punctures, with transverse or arcuate carinate ventrally. Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth. Sculpture of gena: smooth throughout.

Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: bicolored, white with dark brown apex. Color of female A2-

A6: A2-A4 dark brown, A5-A6 white to light yellow, A6 usually dark brown to black apically. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: smooth throughout. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: setigerously punctate throughout. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: transverse, posterior margin straight. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: short, only slightly longer than wide. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: absent medially.

116 Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined.

Dorsal metapleural sulcus: absent.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: bicolored, dark brown to black and patterned with yellow to amber-orange. Transverse carina of horn on T1: absent. Sculpture of horn on

T1 in female: longitudinally striate, striae converging posteriorly to form medial carina.

Sculpture of T2: with longitudinal intersecting rugae. Sculpture of T3: with longitudinal interconnected rugae. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially.

Sublateral carina on T2: absent.

Diagnosis. Oxyteleia solomona can be separated from O. insula by the absence of carinae or strigae separating off the medial area of T2-T3.

Etymology. The epithet solomona, refers to the Solomon Islands, to which this species is endemic. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307049]

Material Examined. Holotype, female: SOLOMON ISLANDS: Western Prov.,

New Georgia Isl., Munda, 0-100m, XI-1980, N. L. H. Krauss, OSUC 234112 (deposited

117 in BPBM). Paratypes: SOLOMON ISLANDS: 4 females, OSUC 232999, 234035,

234038, 234133 (BPBM).

Oxyteleia insula Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307047

(Figures 3.21 A –F)

Description. Female body length: 3.09 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose.

Sculpture of dorsal half of frons in female: dorsally smooth with sparsely setigerous punctures, with transverse or arcuate carinate ventrally. Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth. Sculpture of gena: smooth throughout.

Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: bicolored, white with dark brown apex. Color of female A2-

A6: A2-A4 dark brown, A5-A6 white to light yellow, A6 usually dark brown to black apically. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area:

118 enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: smooth throughout. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: transverse, posterior margin straight. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width.

Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: glabrous. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: bicolored, dark brown to black and patterned with yellow to amber-orange. Transverse carina of horn on T1: absent. Sculpture of horn on

119 T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of

T2: finely longitudinally striate with faint area medially, microsculpture absent. Sculpture of T3: with longitudinal interconnected rugae. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: finely punctate throughout. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: present.

Diagnosis. Oxyteleia insula can be separated from O. solomona by the presence of carinae or strigae defining the medial area of T2-T3.

Etymology. The epithet insula, Latin for island, refers to the medial area of T2-

T3. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307047]

Material Examined. Holotype, female: PAPUA NEW GUINEA: East New

Britain Prov., DPI base camp, Baining Mountains, 04°26.36'S 151°49.02'E, 30.VI-

15.VII.1999, flight intercept trap, A. Mararuai & M. Kalamen, OSUC 266722 (deposited in CNCI).

Oxyteleia lenisa Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307050

(Figures 3.22 A – F)

Description. Female body length: 3.10-3.50 mm (n=16). Body color: entirely dark, dark brown to black throughout.

120 Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose.

Sculpture of dorsal half of frons in female: entirely smooth with sparse punctures, in some ventral half with faint transverse carinae. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: smooth throughout.

Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: dark brown. Color of female A2-A6: A2-A6 antennae sequentially lighter from dark brown to light yellow. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel.

Mesoscutal suprahumeral sulcus: deeply foveolate. Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular

121 spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: rugose throughout. Netrion: setose.

Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: smooth throughout. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae brown to black.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae anteriorly, becoming irregulary longitudinally rugose posteriorly, microsculpture absent. Sculpture of T3: reticulate rugose medially with fine reticulations laterally. Sublateral carina on T3: present. Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: reticulate punctate. Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: present.

Diagnosis. Oxyteleia lenisa can be separated from O. striga and O.poniculus by the fanlike striae on the malar region and the smooth sculpture of the dorsal half of the frons.

122 Etymology. The epithet lenisa, Latin for soft or smooth, refers to the smooth frons. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307050]

Material Examined. Holotype, female: INDONESIA: Maluku Prov., Ceram

(Seram) Isl., Solea, VIII-1987, malaise trap, M. C. Day, OSUC 276562 (deposited in

CNCI). Paratypes: INDONESIA: 15 females, OSUC 253613-253615, 266688, 276542-

276547, 276563, 276613-276616 (CNCI).

Oxyteleia striga Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307036

(Figures 3.23 A – F)

Description. Female body length: 4.20-4.72 mm (n=19). Male body length: 3.86-

4.46 mm (n=4). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose.

Sculpture of dorsal half of frons in female: transversely carinate ventrally, rugulose dorsally. Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth.

Sculpture of gena: smooth throughout. Setation of gena: glabrous to sparsely setose.

Genal carina behind compound eye: absent.

123 Color of female A1: dark brown. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as broad groove. Course of notauli: parallel. Parapsidal line: present, indicated as broad groove. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: densely setigerously punctate with faint punctures posteriorly, sometimes with weak longitudinally striae posteriorly. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: elongate, length distinctly greater than width. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, attenuated dorsally. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: sparsely to densely setose.

124 Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: all coxae brown to black.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: irregularly longitudinally rugose throughout. Sculpture of T3: reticulate rugose medially with fine reticulations laterally. Sublateral carina on T3: present. Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: finely punctate throughout. Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: present.

Diagnosis. Oxyteleia striga shares the character of the grooved notaulus with

O.ponticulus, but can be separated by the longitudinally striate sculpture of the horn on

T1.

Etymology. The epithet striga, Latin for channel or groove, refers to the character of notaulus. The epithet is used as a noun in apposition.

Associations. collected on Castanopsis (Fagales: Fagaceae)

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307036]

Material Examined. Holotype, female, O. strigis (MS) : PAPUA NEW

GUINEA: East New Britain Prov., DPI base camp station, Baining Mountains, 28.IV-

13.V.1999, yellow pan trap/flight intercept trap, L. Leblanc, OSUC 266746 (deposited in

125 CNCI). Paratypes: (32 females, 4 males) INDONESIA: 7 females, OSUC 234096,

234404, 234409, 234463, 234465-234466, 234469 (BPBM). PAPUA NEW GUINEA:

25 females, 4 males, OSUC 367545 (ANIC); OSUC 234048, 234051, 234065, 234069,

234071, 234079, 234082-234084, 234092, 234113, 234115, 234129 (BPBM); OSUC

266706-266707, 266709, 266711-266715, 266717, 266721, 266723-266724, 266731,

266737, 266747 (CNCI).

Oxyteleia ponticulus Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307039

(Figures 3.24 A – F)

Description. Female body length: 4.76 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose.

Sculpture of dorsal half of frons in female: transversely carinate ventrally, rugulose dorsally. Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth.

Sculpture of gena: smooth throughout. Setation of gena: glabrous to sparsely setose.

Genal carina behind compound eye: absent.

126 Color of female A1: dark brown. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as broad groove. Course of notauli: parallel. Parapsidal line: present, indicated as broad groove. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: deeply foveolate. Sculpture of medial lobe of mesoscutum: densely setigerously punctate with faint punctures posteriorly, sometimes with weak longitudinally striae posteriorly.

Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent.

Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially.

Length of axillular spine: elongate, length distinctly greater than width. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: smooth.

Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth.

Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: smooth throughout. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural

127 triangle: well defined. Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: all coxae brown to black.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: present. Sculpture of horn on T1 in female: posterior half transversely carinate throughout, anterior half smooth. Sculpture of T2: with straight longitudinal striae anteriorly, becoming irregulary longitudinally rugose posteriorly, microsculpture absent.

Sculpture of T3: reticulate rugose medially with fine reticulations laterally. Sublateral carina on T3: present. Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: reticulate punctate. Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: present.

Diagnosis. Oxyteleia ponticulus is separated from O. striga by the transverse carinae present on the basal half of the horn on T1.

Etymology. The epithet ponticulus, Latin for bridge, refers to the transverse carinae on the horn on T1. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307039]

Material Examined. Holotype, female: INDONESIA: Papua Barat Prov.,

Vogelkop Peninsula, W of Manokwari, under dead bark, Kebar Valley, 550m, 4.I-

31.I.1962, L. W. Quate, OSUC 234040 (deposited in BPBM).

128 Oxyteleia arcuza Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:305782

(Figures 3.25 A – F, 3.26 A – F)

Description. Female body length: 2.36-3.49 mm (n=20). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: concentrically rugose, rugae sometimes reduced with surface somewhat smoother. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose, interstices smooth. Sculpture of dorsal half of frons in female: transversely carinate throughout. Dense setae within antennal scrobe: absent.

Sculpture of malar region: fanlike striate. Sculpture of gena: strongly crenulate along genal carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: present.

Color of female A1: dark brown. Color of female A2-A6: A2-A4 dark brown, A5-

A6 white; A2-A3, base of A4, apex of A6 dark brown, apex of A4, A5, base ofA6 white to light yellow. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel; converging posteriorly. Parapsidal line: absent;

129 present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: deeply foveolate.

Sculpture of medial lobe of mesoscutum: smooth throughout, or at most with sparse punctures; with weak rugulae emerging from medial carina, becoming sparsely punctate anteriorly. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent; present, percurrent. Mesoscutellum shape: semicircular, posterior margin rounded. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: short, only slightly longer than wide. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: smooth throughout. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose; glabrous anteriorly, with sparse setation posteriorly. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined.

Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: all coxae brown to black.

Color of fore wing membrane: generally hyaline throughout; infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae

130 present only at base. Sculpture of T2: with straight longitudinal striae throughout, microsculpture absent. Sculpture of T3: longitudinally striate laterally, smooth medially; with longitudinal interconnected rugae. Sublateral carina on T3: present; absent.

Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: present; absent.

Diagnosis. The concentrically rugose sculpture of posterior vertex and occiput are unique to Oxyteleia arcuza and immidieately distinguish it.

Etymology. The epithet arcuza, based on the Latin word for arch, refers to the concentrically rugose sculpture of posterior vertex and occiput. The epithet is used as an noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=305782]

Material Examined. Holotype, female: PAPUA NEW GUINEA: Chimbu Prov., nr. Goroka, Tangeu, 05°54'S 144°29'E, no date, malaise trap, T. Anderson, OSUC

266740 (deposited in CNCI). Paratypes: (52 females) INDONESIA: 20 females, OSUC

234041, 234045, 234047, 234053, 234064, 234074, 234077, 234101, 234111, 234128,

234405, 234457-234458, 234462, 234471, 234474, 234529-234530, 234561, 234567

(BPBM). MALAYSIA: 1 female, OSUC 234580 (BPBM). PAPUA NEW GUINEA: 30 females, OSUC 367544, 367546, 367548-367549 (ANIC); OSUC 232998, 234052,

234062, 234070, 234073, 234086, 234093-234094, 234117, 234119, 234131, 234576

(BPBM); CASENT 2137984 (CASC); OSUC 266708, 266710, 266718-266719, 266725-

266726, 266730, 266739, 266741, 266744-266745, 266748-266749 (CNCI).

PHILIPPINES: 1 female, OSUC 376763 (MCZC).

131 Oxyteleia apektos Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307043

(Figures 3.27 A – F)

Description. Female body length: 2.92-2.96 mm (n=2). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: entirely transversely strigose, lacking microsculpture between striae, setigerous punctures ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose, interstices smooth. Sculpture of dorsal half of frons in female: transversely carinate throughout. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: strongly crenulate along genal carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: present.

Color of female A1: dark brown. Color of female A2-A6: A2-A4 dark brown, A5-

A6 white. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel. Parapsidal line: present, indicated as broad groove. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel.

132 Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: longitudinally strigose, reticulate medially, with setigerous puncture within interstices. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent. Mesoscutellum shape: semicircular, posterior margin rounded. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: short, only slightly longer than wide. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: smooth, reticulate rugose anterioly, transversely striate ventrally. Setation of dorsal metapleural area: setose dorsally. Setation of ventral metapleural area: densely setose. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae brown to black.

Color of fore wing membrane: generally hyaline throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae throughout, microsculpture absent. Sculpture of T3: finely punctate reticulate throughout. Sublateral carina on T3: present. Sculpture of medial area of T2-T3: merging with sculpture

133 laterally. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on

T2: present.

Diagnosis. Oxyteleia apektos is very similar to O.arcuza; both having a genal carina behind the compound eye. Oxyteleia apektos can be distinguished from O. aruza by the longitudinal reticulation on the medial lobe of mesoscutum.

Etymology. The epithet apektos, Greek for confusion, refers to the longitudinally reticulate striae on the medial lobe of mesoscutum. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307043]

Material Examined. Holotype, female: PAPUA NEW GUINEA: Northern

Prov., Mount Suckling, 350m, VII-1972, T. L. Fenner, OSUC 266734 (deposited in

CNCI). Paratype: PAPUA NEW GUINEA: 1 female, OSUC 266735 (CNCI).

Oxyteleia johnsoni Taekul, new species

urn:lsid:biosci.ohio-state.edu:osuc_concepts:307023

(Figures 3.28 A – F)

Description. Female body length: 2.30-3.39 mm (n=5). Male body length: 3.15-

3.42 mm (n=2). Body color: entirely dark, dark brown to black throughout; bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: with dense setigerous punctures throughout. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle:

134 predominantly smooth to faintly rugulose. Sculpture of dorsal half of frons in female: entirely smooth with sparse punctures, in some ventral half with faint transverse carinae.

Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth. Sculpture of gena: strongly punctate. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: bicolored, white with dark brown apex. Color of female A2-

A6: A2-A4 dark brown, A5-A6 white. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: strongly punctate.

Skaphion: absent. Notaulus: absent. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: setigerously punctate throughout.

Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent. Mesoscutellum shape: semicircular, posterior margin rounded; transverse, posterior margin straight. Length of axillular spine: short to almost absent.

Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: rugose throughout. Netrion: setose. Mesopleural carina: absent medially.

Sculpture of femoral depression: punctate. Sculpture of speculum: canaliculate

135 throughout. Sculpture of ventral mesepisternum below mesopleural carina: punctate.

Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: deeply sulcate.

Coxae color: all coxae yellow.

Color of fore wing membrane: generally hyaline throughout.

Color of metasoma in female: dark brown to black; bicolored, dark brown to black and patterned with yellow to amber-orange. Transverse carina of horn on T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae anteriorly, otherwise predominantly smooth with sparse punctures posteriorly, microsculpture absent.

Sculpture of T3: smooth. Sublateral carina on T3: absent. Sculpture of medial area of T2-

T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: smooth and shiny. Sculpture of T4: smooth with only sparse punctures. Sublateral carina on T2: absent.

Diagnosis. The canaliculate sculpture of the speculum is unique for Oxyteleia johnsoni and will immediately distinguish this species.

Etymology. The epithet johnsoni, refers to Norman F. Johnson, CT’s Professor, named in the memory of CT’s graduate career. The epithet is used as a noun in the genitive case.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307023]

136 Material Examined. Holotype, female: FIJI: Western Div., Ba Prov., Viti Levu

Isl., Eteni, FJ_11B, Navai, 17°37'S 177°59'E, 700m, 6.VI-15.VII.2003, malaise trap, E.

Schlinger, M. Irwin & Tokota'a, FBA014399 (deposited in CNCI). Paratypes: FIJI: 4 females, 2 males, FBA014386, FBA042191, FBA058997, FBA089253, FBA095107,

OSUC 266751 (CNCI).

Oxyteleia lagunasis Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307048

(Figures 3.29 A – F); Morphbank

Description. Female body length: 3.85 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: with dense setigerous punctures throughout. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: reticulate rugose, interstices smooth. Sculpture of dorsal half of frons in female: transversely carinate throughout. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: strongly punctate. Setation of gena: densely setose. Genal carina behind compound eye: absent.

Color of female A1: dark brown. Color of female A2-A6: A2-A4 dark brown, A5-

A6 white to light yellow, A6 usually dark brown to black apically. Claval color (A7-A12)

: dark brown to black throughout.

137 Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: absent. Parapsidal line: absent. Mesoscutal humeral sulcus: crenulate.

Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: setigerously punctate throughout. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout.

Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: short, only slightly longer than wide.

Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae dorsally. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin.

Sculpture of ventral mesepisternum below mesopleural carina: punctate. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: densely setose.

Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae brown to black.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with straight longitudinal striae anteriorly,

138 reticulate posteriorly, microsculpture absent. Sculpture of T3: finely punctate reticulate throughout. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on T2 and T3: reticulate punctate.

Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: present.

Diagnosis. Oxyteleia lagunasis shares the character of the densely punctuate genus with setose interstices with Oxyteleia johnsoni, but can be distinguished by the absence of canaliculate sculpture on the speculum.

Etymology. The epithet lagunasis, refers to the collecting locality, Laguna

Province in the Philippines. The epithet is used as a noun in apposition.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307048]

Material Examined. Holotype, female: PHILIPPINES: Laguna Prov., Mount

Makiling (Maquiling), 8.XII-19.XII.1990, pan trap, I. Nabi, OSUC 276576 (deposited in

CNCI).

Oxyteleia primata Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307046

(Figures 3.30 A – F)

Description. Female body length: 2.79-2.87 mm (n=2). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: entirely transversely strigose, lacking microsculpture between striae, setigerous punctures ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent

139 dorsally. Sculpture of ocellar triangle: predominantly smooth to faintly rugulose.

Sculpture of dorsal half of frons in female: dorsally smooth with sparsely setigerous punctures, with transverse or arcuate carinate ventrally. Dense setae within antennal scrobe: absent. Sculpture of malar region: smooth. Sculpture of gena: smooth throughout.

Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: dark brown. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: rugulose.

Skaphion: absent. Notaulus: absent. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: setigerously punctate throughout.

Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent.

Mesoscutellum shape: transverse, posterior margin straight. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural

140 area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: poorly defined. Dorsal metapleural sulcus: deeply sulcate.

Coxae color: all coxae brown to black; fore coxa brown, mid and hind coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present at least on basal half of horn, sometimes striae reaching apex. Sculpture of T2: finely longitudinally striate with faint area medially, microsculpture absent. Sculpture of

T3: finely longitudinally striate laterally, reticulate punctate medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally.

Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: absent.

Diagnosis. Oxyteleia primata can be separated from other species lacking notauli by the dark brown to black color of A1.

Etymology. The epithet primata, Latin for first or chief, refers to A1 character.

The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307046]

Material Examined. Holotype, female: PAPUA NEW GUINEA: East New

Britian Prov., New Britain Isl., 10km E Keravat, Vunabakan, 180m, 16.XI-20.XI.1959,

T. C. Maa, OSUC 246586 (deposited in BPBM). Paratypes: (2 females) PAPUA NEW

141 GUINEA: 1 female, OSUC 234098 (BPBM). PHILIPPINES: 1 female, OSUC 376768

(MCZC).

Oxyteleia astricta Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307044

(Figures 3.31 A – F)

Description. Female body length: 3.39-3.82 mm (n=3). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: with dense setigerous punctures throughout. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: rugulose, otherwise with sparse setigerous punctures on smooth background. Sculpture of dorsal half of frons in female: transversely carinate throughout. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: white to light yellow. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: enlarged, triangular, and visible dorsally. Sculpture of dorsal pronotal area: rugulose.

142 Skaphion: absent. Notaulus: absent. Parapsidal line: absent. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: densely setigerously punctate with faint punctures posteriorly, sometimes with weak longitudinally striae posteriorly. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: strongly transverse, posterior margin deeply concave housing horn of T1. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to netrion: punctate with rugulae ventrally. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: present. Sculpture of horn on T1 in female: longitudinally striate, striae converging posteriorly to form medial carina. Sculpture of T2: finely longitudinally striate with faint area medially, microsculpture absent. Sculpture of T3: finely longitudinally striate

143 laterally, reticulate punctate medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: absent.

Diagnosis. Oxyteleia astricta is one of the species with neither skaphion and notaulus, and can be easily distinguished by the strongly transverse mesoscutellum and excavate posterior margin housing the horn of T1.

Etymology. The epithet astricta, Latin for narrow or tight, refers to the strongly transverse mesoscutellum shape. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307044]

Material Examined. Holotype, female: SOLOMON ISLANDS: Western Prov.,

Ghizo (Gizo) Isl., Gizo, 0-200m, XII-1975, N. L. H. Krauss, OSUC 234135 (deposited in

BPBM). Paratypes: SOLOMON ISLANDS: 2 females, OSUC 234121, 234136 (BPBM).

Oxyteleia cathetus Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307045

(Figures 3.32 A – F)

Description. Female body length: 3.18-3.81 mm (n=2). Body color: bicolored, head dark brown to black, mesosoma or metasoma with at least some areas yellow.

Sculpture of posterior vertex and occiput: with dense setigerous punctures throughout. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: rugulose,

144 otherwise with sparse setigerous punctures on smooth background. Sculpture of dorsal half of frons in female: transversely carinate throughout. Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: bicolored, white with dark brown apex. Color of female A2-

A6: dark brown to black throughout. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: orange with dark brown patch on posteromedial mesoscutum, dark brown color sometimes extending to mesoscutellum. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: absent. Parapsidal line: absent. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: densely setigerously punctate with faint punctures posteriorly, sometimes with weak longitudinally striae posteriorly. Sculpture of lateral mesoscutum: covered with dense punctures to fine dense rugulae. Sculpture of mesoscutellum: setigerously punctate throughout. Medial carina of mesoscutum: absent. Mesoscutellum shape: transverse, posterior margin straight. Length of axillular spine: short to almost absent.

Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent. Sculpture of lateral pronotal area adjacent to

145 netrion: punctate with rugulae ventrally. Netrion: setose. Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined.

Dorsal metapleural sulcus: deeply sulcate.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: bicolored, dark brown to black and patterned with yellow to amber-orange. Transverse carina of horn on T1: absent. Sculpture of horn on

T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of

T2: with longitudinal intersecting rugae. Sculpture of T3: finely longitudinally striate laterally, reticulate punctate medially. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: delimited by longitudinal carinae. Sculpture of sablateral area on

T2 and T3: reticulate punctate. Sculpture of T4: punctate with faint punctures medially.

Sublateral carina on T2: present.

Diagnosis. Oxyteleia catheta can be separated from O. astricta by the straight posterior margin of the mesoscutellum.

Etymology. The epithet catheta, derived from the Latin word for a straight line, refers to the rectangular shape of mesoscutellum. The epithet is used as a noun in apposition.

146 Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307045]

Material Examined. Holotype, female: SOLOMON ISLANDS: Guadalcanal

Prov., Guadalcanal Isl., Gold Ridge - Suta, Mount Chaunapaho (Jonapau), 1100m,

26.VII.1956, J. L. Gressitt, OSUC 234033 (deposited in BPBM). Paratype: SOLOMON

ISLANDS: 1 female, OSUC 234037 (BPBM).

Oxyteleia bidentata Kieffer urn:lsid:biosci.ohio-state.edu:osuc_concepts:5039

(Figures 3.33 A – F, 3.34 A – F, 3.35 A – F)

Caloteleia bidentata Kieffer, 1905: 16 (original description). Oxyteleia bidentata

(Kieffer) : Kieffer, 1908: 118 (generic transfer); Kieffer, 1926: 516, 518 (description, keyed); Bin, 1974: 457 (type information); Masner, 1976: 49 (description).

Description. Female body length: 3.60-4.82 mm (n=2). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: concentrically rugose, rugae sometimes reduced with surface somewhat smoother. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: complete. Sculpture of ocellar triangle: reticulate rugose, interstices smooth; predominantly smooth to faintly rugulose. Sculpture of dorsal half of frons in female: transversely carinate ventrally, rugulose dorsally; ventrally with arcuate carinae parallel to dorsal margin of antennal scrobe, with oblique strigae dorsally. Dense setae within antennal scrobe: absent.

147 Sculpture of malar region: fanlike striate. Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: bicolored, white with dark brown apex. Color of female A2-

A6: dark brown to black throughout; A2-A5 sequentially lighter from dark brown to light yellow, A6 dark brown. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as broad groove. Course of notauli: parallel. Parapsidal line: present, indicated as broad groove. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: deeply foveolate. Sculpture of medial lobe of mesoscutum: densely setigerously punctate with faint punctures posteriorly, sometimes with weak longitudinally striae posteriorly; densely setigerously punctate with reticulate rugae posteriorly. Sculpture of lateral mesoscutum: smooth to sparsely punctate; covered with dense punctures to fine dense rugulae. Sculpture of mesoscutellum: setigerously punctate throughout; rugose with punctures within interstices. Medial carina of mesoscutum: absent. Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina:

148 absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: all coxae yellow.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: present; absent. Sculpture of horn on T1 in female: longitudinally striate laterally, otherwise smooth apically, base with flanking transverse rugulae. Sculpture of T2: with straight longitudinal striae anteriorly, becoming irregulary longitudinally rugose posteriorly, microsculpture absent; irregularly longitudinally rugose throughout.

Sculpture of T3: longitudinally rugose with crenulae within interstices. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: delimited by longitudinal carinae.

Sculpture of sablateral area on T2 and T3: finely punctate throughout; reticulate punctate.

Sculpture of T4: punctate with faint punctures medially; reticulate punctate throughout.

Sublateral carina on T2: present.

Diagnosis. Oxyteleia bidentata is similar to O. fasciata and O. obscura, and can be easily distinguished by the complete occipital carina and the presence of carinae or strigae delimiting the medial area of T2 and T3.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=5039]

149 Material Examined. Holotype, female, C. bidentata: PAPUA NEW GUINEA:

Central Prov., Moroka, 1300m, VII-1893 - XI-1893, Loria, MCSN 0003 (deposited in

MCSN). Other material: PAPUA NEW GUINEA: 3 females, OSUC 367543 (ANIC);

OSUC 234044 (BPBM); OSUC 266733 (CNCI).

Oxyteleia obscura Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307040

(Figures 3.36 A – F)

Description. Female body length: 2.56 mm (n=1). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: entirely transversely strigose, lacking microsculpture between striae, setigerous punctures ventrally. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: rugulose, otherwise with sparse setigerous punctures on smooth background. Sculpture of dorsal half of frons in female: dorsally smooth with sparsely setigerous punctures, with transverse or arcuate carinate ventrally.

Dense setae within antennal scrobe: absent. Sculpture of malar region: fanlike striate.

Sculpture of gena: predominantly smooth, with weak crenulae adjacent to eye and occipital carina. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

150 Color of female A1: dark brown. Color of female A2-A6: A2-A5 sequentially lighter from dark brown to light yellow, A6 dark brown. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as sulcus margined medially by weak carina. Course of notauli: parallel. Parapsidal line: present, indicated as weak, narrow sulcus. Mesoscutal humeral sulcus: crenulate. Mesoscutal suprahumeral sulcus: finely crenulate. Sculpture of medial lobe of mesoscutum: weakly transversely rugose with puncture within interstices. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum: predominantly smooth, or at most with sparse puncture. Medial carina of mesoscutum: absent. Mesoscutellum shape: semicircular, posterior margin rounded. Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: short, only slightly longer than wide. Vertical epomial carina: present, complete. Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose. Mesopleural carina: present. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: setose dorsally. Setation of ventral metapleural area: glabrous anteriorly, with sparse setation posteriorly. Setation of metapleural triangle: sparsely to

151 densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: absent.

Coxae color: all coxae brown to black.

Color of fore wing membrane: generally hyaline throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: with longitudinal intersecting rugae. Sculpture of

T3: finely punctate reticulate throughout. Sublateral carina on T3: absent. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: punctate with faint punctures medially. Sublateral carina on T2: present.

Diagnosis. Oxyteleia obscura can be separated from O. fasciata by the weak transversely rugose sculpture on the mesoscutum.

Etymology. The epithet obscura, Latin for faint or dim, refers to the faint sculpture of mesoscutum. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307040]

Material Examined. Holotype, female: PAPUA NEW GUINEA: Morobe Prov., primary forest, Mount Missim, 27.V.1982, P. Grootaert, OSUC 266742 (deposited in

CNCI).

Oxyteleia fasciata Taekul, new species urn:lsid:biosci.ohio-state.edu:osuc_concepts:307041

(Figures 3.37 A – F)

152

Description. Female body length: 2.45-3.91 mm (n=2). Body color: entirely dark, dark brown to black throughout.

Sculpture of posterior vertex and occiput: predominantly smooth with sparse punctures, sometimes with weak transverse strigae medially. Transition from vertex to occiput: more or less evenly rounded from anterior to posterior. Occipital carina: absent dorsally. Sculpture of ocellar triangle: rugulose, otherwise with sparse setigerous punctures on smooth background. Sculpture of dorsal half of frons in female: transversely carinate ventrally, rugulose dorsally. Dense setae within antennal scrobe: absent.

Sculpture of malar region: smooth. Sculpture of gena: smooth throughout. Setation of gena: glabrous to sparsely setose. Genal carina behind compound eye: absent.

Color of female A1: dark brown. Color of female A2-A6: dark brown to black throughout. Claval color (A7-A12) : dark brown to black throughout.

Color of mesosoma in female: dark brown to black throughout. Dorsal pronotal area: present, defined by transverse pronotal carina. Anterior dorsal pronotal area: narrow, not visible dorsally. Sculpture of dorsal pronotal area: rugulose. Skaphion: absent. Notaulus: present. Shape of notaulus: indicated as broad groove. Course of notauli: parallel. Parapsidal line: present, indicated as broad groove. Mesoscutal humeral sulcus: indicated as uninterrupted smooth channel. Mesoscutal suprahumeral sulcus: indicated as uninterrupted channel connected to mesoscutal humeral sulcus. Sculpture of medial lobe of mesoscutum: longitudinally striate, striae attenuating anteriorly. Sculpture of lateral mesoscutum: smooth to sparsely punctate. Sculpture of mesoscutellum:

153 setigerously punctate throughout. Medial carina of mesoscutum: present posteriorly.

Mesoscutellum shape: pentagonal, posterior margin converging to slight point medially.

Length of axillular spine: short to almost absent. Length of lateral metascutellar spine: moderately elongate, length distinctly greater than width. Vertical epomial carina: absent.

Sculpture of lateral pronotal area adjacent to netrion: smooth. Netrion: setose.

Mesopleural carina: absent medially. Sculpture of femoral depression: smooth. Sculpture of speculum: crenulate along posterior margin. Sculpture of ventral mesepisternum below mesopleural carina: reticulate punctate anteriorly, otherwise smooth. Setation of dorsal metapleural area: glabrous. Setation of ventral metapleural area: glabrous. Setation of metapleural triangle: sparsely to densely setose. Subdivided cells within metapleural triangle: well defined. Dorsal metapleural sulcus: indicated as uninterrupted smooth channel.

Coxae color: all coxae brown to black.

Color of fore wing membrane: infuscate throughout.

Color of metasoma in female: dark brown to black. Transverse carina of horn on

T1: absent. Sculpture of horn on T1 in female: with parallel longitudinal striae, striae present only at base. Sculpture of T2: irregularly longitudinally rugose throughout.

Sculpture of T3: reticulate rugose medially with fine reticulations laterally. Sublateral carina on T3: present. Sculpture of medial area of T2-T3: merging with sculpture laterally. Sculpture of T4: reticulate punctate throughout. Sublateral carina on T2: present.

154 Diagnosis. Oxyteleia fasciata is similar to O. obscura in the absence of occipital carina and the color of A1. It may be distinguished by the longitudinal striae on the posterior area of the mesoscutum.

Etymology. The epithet fasciata, Latin for stripe or band, refers to the longitudinal sculpture on mesoscutum. The epithet is used as an adjective.

Link to Distribution Map. [http://hol.osu.edu/map-full.html?id=307041]

Material Examined. Holotype, female: INDONESIA: Papua Prov., W New

Guinea Isl., Central Mts., Archbold Lake, 760m, 26.XI-3.XII.1961, L. W. Quate, OSUC

234050 (deposited in BPBM). Paratype: PAPUA NEW GUINEA: 1 female, OSUC

367547 (ANIC).

155

CONCLUSION

The genus Oxyteleia Kieffer (Hymenoptera: Platygastridae, Scelioninae) is a widespread group in the Indo-Malayan and Australasian regions, distributed from north

India to Fiji. Despite the fact that all scelionines are egg parasitoids of arthropods, the host of Oxyteleia is not yet known. The species concepts are revised and a key to world species is presented. The genus comprises 36 species of which 27 are new to science. The genus Merriwa Dodd, 1920 is considered to be a new synonym and its type species, O. quadridentata (Dodd) n.comb. (Indonesia, Malaysia), is redescribed.. Two species, O. aenea (Ashmead), and O, punctata (Ashmead) are transferred to genus Paridris. The following species are hypothesized and described as new taxa: O. inthanona Taekul, n.sp. (Thailand); O. aenigma Taekul, n.sp. (Thailand, Laos, Viet Nam); O. oecus Taekul, n.sp. (Thailand, Laos, Malaysia, Indonesia, Malaysia); O. pangasugana Taekul, n.sp.

(Philippines); O. angularis Taekul, n.sp. (Malaysia); O. rugosa Taekul, n.sp.

(Philippines); O. bimucronata Taekul, n.sp. (Malaysia, Indonesia); O. epidemosa Taekul, n.sp. (Thailand, India, Indonesia, Laos, Malaysia, Taiwan, Vietnam); O. alba Taekul, n.sp. (Malaysia); O. arcuata Taekul, n.sp. (Philippines); O. talamasi Taekul, n.sp.

(Indonesia); O. lux Taekul, n.sp. (Philippines); O. scopulus Taekul n.sp. (Fiji); O. solomona Taekul, n.sp. (Solomon Islands); O. insula Taekul, n.sp. (Papua New Guinea)

O. lenisa Taekul, n.sp. (Indonesia); O. striga Taekul, n.sp. (Papua New Guinea,

156 Indonesia); O. ponticulus Taekul, n.sp. (Indonesia); O. arcuza Taekul, n.sp. (Papua New

Guinea, Indonesia, Malaysia, Philippines); O. apektos Taekul, n.sp. (Papua New Guinea)

O. johnsoni Taekul, n.sp. (Fiji); O. lagunasis Taekul, n.sp. (Phhilippines); O. primata

Taekul, n.sp. (Papua New Guinea, Philippinev); O. astricta Taekul, n.sp. (Solomon

Islands); O. catheta Taekul, n.sp. (Solomon Islands); O. obscura Taekul n.sp. (Papua

New Guinea); O. fasciata Taekul n.sp. (Indonesia, Papua New Guinea).

157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194

CHAPTER 4

MOLECULAR PHYLOGENY OF THE SUBFAMILY TELENOMINAE

(PLATYGASTRIDAE, PLATYGASTROIDEA, HYMENOPTERA)

INTRODUCTION

Parasitoid Hymenoptera has been proved as one the most beneficial insects for society. To minimize the use of chemical pesticides, parasitoids have been used to control economic insect pests of agriculture and forestry as well as human and animal disease vectors. Of 393 species of parasitoid Hymenoptera introduced in classical biological control programs, 87 % successfully established and achieved pest control (Greathead

1986, LaSalle & Gauld 1993). The subfamily Telenominae, the most economically important group of the superfamily Platygastroidea, falls in this category.

The monophyletic superfamily Platygastroidea is sister to Cynipoidea (gall wasps) nested together and basal to the remaining Proctotrupomorpha (Proctotrupoidea,

Diaprioidea, , and Chalcidoidea) (Sharkey 2007, Heraty et al. 2011,

195 Sharkey et al. 2012). Two families, Platygastridae and Scelionidae, were recognized

(Johnson 1992, Vlug 1995) until Sharkey (2007) combined them under the single family

Platygastridae sensu lato. The reasons for this synonymy were (1) the absence of synapomorphies defining the scelionids as a monophyletic group, a conclusion based on the phylogenetic analysis of Murphy et al. (2007) and Austin et al. (2005); and (2)

Sharkey’s perception that the superfamily is morphologically homogeneous.

Platygastridae in this sense contains five subfamilies: Platygastrinae, Sceliotrachelinae,

Scelioninae, Teleasinae, and Telenominae with the latter two being demonstratably monophyletic (Murphy et al. 2007). Telenomines are distinguished from other platygastroid subfamilies based on the following combination of characters: 1) metasoma with wide laterotergites loosely attached to the corresponding sternites, no laterosternites, and therefore no impressed submarginal ridge, 2) T2 by far the largest of all metasomatic tergites, and 3) the female antennae are 10–11 segmented, while those of the male – with one known exception – have 12 segments (Masner 1976).

The subfamily Telenominae constitutes 20 genera and 894 species currently treated as valid, distributed all around the world (Johnson 2012). However, Bin (1982) suggested that the number of described species in this subfamily represents only 10–25 % of the number actually in existence, and the group may therefore contain as many as 6000 species. Most described fossils of the superfamily Platygastroidea are from Baltic amber inclusions, and are clearly assignable to modern genera. Five species of platygastroids have so far been described from the Cretaceous period, and the number of fossil species dramatically increased after the Paleocene, based on the list of fossil and subfossils

196 species of Platygastroidea by Johnson et al. (2012). Very few telenomine fossils have been described. Fossils of the Telenominae only appear from the late Oligocene-

Miocene, collected from Dominican Republic (Johnson 1984). One genus with a single species, Sinoprotelenomus, was described from Shanwan, Shandong, China (Zhang 1989;

Johnson 2012).

The ground plan biology of the superfamily Platygastroidea is endoparasitism of the eggs of insects and spiders (Austin et al. 2005). Most species in the subfamilies

Platygastrinae and Sceliotrachelinae, however, either attack later stages of sessile hosts such as planthoppers, whiteflies, and aphids, or they attack gall flies (Diptera:

Cecidomyiidae), either in the egg or early larva of the host. With their hypodermic needle-like ovipositor, female platygastroids pierce the chorion of a host egg and lay their own single egg, or sometimes several, within. The wasp larva that hatches derives its nourishment from the host egg and pupates within it. The host eggs of platygastroid wasps are in the orders Odonata, Orthoptera, Mantodea, Embiidina, Hemiptera,

Neuroptera, Coleoptera, Diptera, and Lepidoptera as well as spiders (Austin & Field

1997; Masner 1976; Austin et al. 2005). Telenomine species attack half of these insect orders, primarily the Lepidoptera and Hemiptera, but also some species of Diptera and

Neuroptera (Johnson 2012).

Despite the inadequacy of the complete understanding of the diversity and relationships, telenomine wasps have been proven to be potential biological control agents (Orr 1988). This is attributed to several characteristics, including high searching

197 abilities and reproductive rates, lack of hyperparasitoids, synchrony with host populations, positive host-density responsiveness, simple adult diets, and they can be reared easily. These parasitoids can be applied to all biological control practices: classical, conservation, and augmentative control. Orr (1988) reported that 30 species were used in classical biological control attempts, and several of these have produced successful results.

Telenominae is traditionally divided into two genera: Telenomus and Trissolcus.

Together these currently encompass 820 species (Trissolcus: 176, Telenomus: 644), which comprise approximately 92% of all valid telenomine species (Johnson 2012). The rest is made up of (1) fossils representing only a single genus, Sinoprotelenomus (2)

“satellite” genera containing a very small number of morphologically aberrant species or sometimes based on vague concepts, e.g., Aradoctonus Masner, Latonius Kononova,

Protelenomus Kieffer, Phlebiaporus Kozlov, Paratrissolcus Mineo, O’Connor & Ashe,

Televiggianus Mineo, Rachelia Mineo, Kozlotelenomus Mineo, O’Connor & Ashe,

Ioseppinella Mineo, O’Connor & Ashe and (3) the “orphan” genus, Bruchiola Kieffer.

There are some intramural disputes about what should be recognized, and the validation of constituent genera are not presented in this paper. Kozlov (1970) and Kozlov &

Kononova (1983) recognized three tribes in the subfamily: Tiphodytini, Aradophagini, and Telenomini. Masner (1976), however, placed the former two tribes in Scelioninae, arguing that they were assigned to Telenominae on the basis of convergent characters, results corroborated by Murphy et al. (2007). Here we focus on Telenominae sensu

Masner, i.e., including only the tribe Telenomini.

198 Johnson (1984) divided the genus Telenomus into informal species groups because it possesses a huge number of species and revisions at the species level are impractical. These putatively monophyletic taxa are informally named by using the name of an included species, for example, the T. podisi species group.

The original description of genus Psix was published by Kozlov & Lê (1976) from a short series of specimens from Afghanistan. The genus was later expanded by

Johnson & Masner (1985) into 21 species. Psix is primarily restricted to Africa, Asia, and

Australia. However one species of the genus, P. tunetanus (Mineo & Szabó), was introduced to the southwestern US and northern South America. Paratelenomus was originally described from an Australian species (Dodd 1914) and later described again as

Archiphanurus by Szabó (1975). The latter name was synonymized by Johnson (1988).

The genus now includes 17 species restricted to southern Europe, Africa, Asia, Australia,

Vanuatu and Hawaii (Johnson 2012). Nirupama, a closely related genus to

Paratelenomus and Psix (Johnson 1985), was originally described by Nixon (1935). The distribution of the genus is restricted to the Afrotropical realm. Mudigere was originally described by Johnson (1988). The distribution of the genus is known thus far from southern India and Indochina. Together these four genera are morphologically distinct from the rest of Telenominae. Despite Masner’s (1976) suggestion that Psix is closely related to Trissolcus, this group is distinguished by numerous characters, e.g. the fusion of clypeus and labrum, the bifurcation of the central keel on the frons to pass outside of the antennal insertions, location of the lateral ocelli, fusion of the metapostnotum with the propodeum, the pattern of lines of foveae on the meso- and metapleuron, and the setation

199 pattern on the first metasomal tergite. These genera, with the exception of the one introduced species, are restricted to the Old World (Johnson 1988). Johnson (1985, 1988) discussed the relationship and the position of this Psix-group of genera within

Telenominae. He suggested that these genera form a monophyletic group – Nirupama obviously seemed to be the most closely related to Paratelenomus and Psix. Mudigere was hypothesized to be the sister group of this clade. Johnson (1985, 1988) suggested that this group of genera was possibly related to the genus Phanuromyia, as well as suspected that the characters shared with Telenomus and Trissolcus are parallel or convergent.

Despite two and a half centuries of work on the generic classification of the subfamily, the interrelationships within Telenominae still are poorly understood. The difficulty of reconstructing the phylogeny using morphological data arises from the preponderance of character loss and reduction which can lead to extensive homoplasies.

Within the classification of Platygastroidea, the recognition of monophyletic subfamilies and tribes has been problematic (Austin et al. 2005). Murphy et al. (2007) recorded that many of these reductional characters appear to have evolved convergently. This may result from the issues of character conceptualization, coding criteria, and homology assessments (Scotland et al. 2003). The minuteness and ambiguity of telenomine morphological characters has greatly hindered phylogeny estimation. Therefore, the addition of molecular characters for the reconstruction of the phylogeny of Telenominae seems to hold promise. Our attempt here is to use a multi-gene dataset to infer the phylogenetic relationships across the subfamily Telenominae. We examine the monophyly of Telenominae and its primary genera, examine the phylogeny within

200 Trissocus and Telenomus, and evaluate the placement of the Psix-group of genera within the subfamily. The patterns of evolution of host use are discussed. This work is part of the ongoing Platygastroidea Planetary Biodiversity Inventory.

201

MATERIALS AND METHODS

Taxonomic sampling and specimen vouchering

A total of 80 specimens comprising 62 species were examined: 69 specimens representing 51 species for the ingroup, and 11 specimens representing 11 species in the outgroup. The ingroup, Telenominae, contains 6 genera: Telenomus (8 species groups),

Trissolcus (3 species groups), Psix, Paratelenomus, Phanuromyia, and Eumicrosoma.

The generic names Platytelenomus Dodd and Aholcus Kieffer are still recognized as valid by some authors (e.g., Kononova 2008, Mineo 2011), and representatives of each were also included in the analyses. Eleven genera were not sampled because they are fossils

(e.g. Sinoprotelenomus), are extremely rare, or are doubtfully valid. The taxonomic samples were selected mainly to infer interrelationships within Telenominae and so to better understand the evolution of associations between the parasitoids and their host eggs. Outgroup taxa were chosen from a broad range of Platygastridae, made up of 5 genera of Scelioninae and Teleasinae. In the results of Murphy et al. (2007) the genus

Gryon emerged as the sister group to the Telenominae. Therefore, particular emphasis was placed on representatives of this genus. Material studied was derived from fresh collections from around the world: Australia, Bolivia, Canada, Colombia, Ghana,

Hungary, Italy, Kenya, Mexico, Panama, Thailand, United States (CA, OH, AZ), and

Venezuela.

202 The voucher specimens after non-destructive DNA extraction are deposited in the

C. A. Triplehorn Insect Collection (OSUC). The numbers prefixed with “OSUC” are unique identifiers for the individual specimens. The label data for all specimens have been georeferenced and recorded in the Hymenoptera online database, and details on the data associated with these specimens is publicly accessible at the following link, hol.osu.edu, and entering the identifier in the form (Johnson 2012) (Note the space between the acronym and the number). Table 4.1 lists the specimens, voucher information, and gene markers included in the analyses.

DNA Extraction, Amplification, and Sequencing

Fresh specimens were sorted from bulk material and preserved in 95% ethanol, and then maintained in a freezer at -15 °C. Non-destructive DNA extraction was performed using the DNEasy extraction protocol (Qiagen Inc.) as modified for

Hymenoptera by C.D. Zhu & J.S. Noyes at the British Museum of Natural History.

Sequences in total of approximately 3760 base pairs were obtained from 4 gene regions:

COI, 18S, 28S, and EF-1α. Primers for PCR amplification were drawn from previously published papers listed in Table 4.2. Amplification generally was carried out via polymerase chain reaction following the protocols of Murphy et al. (2007) and Klompen

(2000).

Nested PCRs were employed to amplify the F2 copy of EF-1α. This approach is relatively successful in reducing or eliminating unwanted products, although it dramatically increases sensitivity to contamination. The initial PCR primers were F2F and Cho10 (Danforth et al. 1999) following by a combination of other primers from

203 Heraty et al. (2011): F2F8 [5'–CAA RTA TGC NTG GGY ATT GGY AAG–3'], F2R6

[5'–TTG WGC RGT GAA GTC AGC NGC–3'], and Simon et al. (2010): EF-7 [5'–AAC

AAR ATG GAY TCN ACN GAR CCN CC–3'] and EF-9 [5'–CCN ACN GGB ACH

GTT CCR ATA CC–3']. The choice of second round primers depended on the taxa and product results. Thermocycler conditions for F2F-Cho10, F2F8-F2R6, and For3-Cho10 followed Klompen (2000) except annealing temperatures of 54°C, 54°C, and 58°C were used respectively. The amplification profile for EF-7 and EF-9 followed Simon et al.

(2010). We excluded the F1 copy from the analysis because of the difficulty of amplification and the often incomplete products (Heraty et at. 2011). The homology of positions of F2 amplicons was evaluated based on two methods: 1) the F2 amplicons possessed highly conserved intron patterns located at positions 753/754 and 1029/1030

(Danforth & Ji 1998), 2) comparing against the sequence of Archaeoteleia mellea

(GenBank:GQ410731.1). Contaminants were detected using negative controls in all rounds of PCR, as well as comparisons of the product sequences with each other and with a range of published sequences. PCR products were purified either using the

QIAquick PCR purification kit protocol or done prior to sequencing by Beckman Coulter

– Agencourt. Products were sequenced in both directions and assembled using

Sequencher 4.0.

Sequence Alignment

Sequences of the mitochondrial protein-coding gene cytochrome oxidase I (COI, approximately 857 bp) were aligned initially with MUSCLE (Edgar 2004) and then adjusted manually when necessary. The total of 1124 bp of the F2 copy of the nuclear

204 protein-coding gene elongation factor 1-alpha (EF-1α) were manually aligned against reference taxa and reported intron positions (Heraty et al. 2011; Danforth & Ji 1998).

Alignments of both protein coding genes were translated to amino acids to verify the homology positions and termination codons using Mesquite (Maddison & Maddison

2011). Secondary structural alignments were implemented for ribosomal RNA sequences of 18S and 28S for a total of 1783 bp. The manual alignment conventions followed Kjer

(1995) with slight modifications (Gillespie 2004). The core alignments initially followed published secondary structure models of Hymenoptera (as in Murphy et al. 2007; Heraty et al. 2011; Gillespie et al. 2005; Dean et al. 2006). Ambiguous homology positions were defined across all taxa based on structural criteria of Kjer (1995). These regions were characterized as (1) RAAs for the regions of alignment ambiguity, (2) RSCs for regions of slipped-strand compensation, and (3) RECs for regions of expansion and contraction

(Gillespie 2004; Gillespie et al. 2005).

A total of 42 ambiguous regions (RAAs, RSCs, RECs) (567 bp) encompassing approximately 37.7% of rRNA characters were excluded from the final analyses. The final matrix alignment for analyses contains 3767 bp (COI: 857, 18S: 1003, 28S: 782,

EF-1α: 1125). The alignment summary, gene partitions, nucleotide composition, and percentage of parsimony informative sites are presented in Table 4.3.

Data Coding and Partitioning

Regier et al. (2011) discussed the role of synonymous nucleotide change in protein-coding genes in reducing the performance of phylogenetic analyses and proposed

205 mechanisms to attempt to compensate for this effect. Here we employ three strategies to reduce the effect of synonymous changes. The third codon position (nt3) is the source of most synonymous change and, therefore, one option is to eliminate these data from the analysis. Additionally, synonymous changes in the first codon position are possible for sequences coding for arginine and leucine. The DEGEN approach (Regier et al. 2011) degenerates synonymous nucleotide changes in both the first and third codon positions.

For example the five codons for leucine are represented as YTN (Y: pyrimidines C or T,

N: any nucleotide). The third strategy is to segregate data from nt3 in a separate partition.

We employed three partitioning schemes: 1) use of a single partition for data from all four markers; 2) four partitions, separating data from 18S, 28S, COI and EF-1α; and

3) six partitions for data from 18S, 28S, COI nt1+2, EF-1α nt1+2, COI nt3, and EF-1α nt3. Combining the coding and partitioning strategies resulted in seven data sets for analysis: 1–3) no coding, 1, 4, and 6 partitions; 4–5) nt3 excluded, 1 and 4 partitions; 6–

7) DEGEN coding, 1 and 4 partitions. The models chosen for data partitions and definitions for coding strategies as well as partitioning schemes are shown in Table 4.3 and 4 respectively.

Phylogenetic analyses

Three phylogenetic estimation strategies were implemented to infer relationships: maximum parsimony, maximum likelihood, and Bayesian approaches. Parsimony estimates of phylogeny are based on requiring observed character state changes to explain the data. The statistical approaches employ rates of evolution. Each criterion has different

206 concepts as well as distinctive strengths and weaknesses (Holder & Lewis 2003). Here we explored these three dimensions with data partitions discussed in the previous section.

Maximum Parsimony—The analyses were conducted in TNT version 1.1

(Goloboff et al. 2008). Heuristic tree search algorithms were implemented using the New

Technology Search by default setting with some modifications followed by tree bisection-reconnection (TBR) branch swapping. Some parameters were adjusted: using a sectorial search, treating gaps as missing data, tree drifting of 4 cycles, tree fusing of 5 rounds and best score hit of 20–25 times depending on the matrix size. Branch support was estimated using the Parsimony Jackknife (JK; Farris et al. 1996) executed with the setting 36% probability removal, “emulate JAC” resampling, 1000 replications, “random addition sequences”=1, and “hold trees”=2 (Klompen et al. 2007; Freudenstein et al.

2004). The parsimony jackknife was selected over the alternative, bootstrapping, because it is less sensitive to missing and invariant characters (Farris et al. 1996, Freudenstein et al. 2004).

Maximum Likelihood—Analyses were executed using RAxML v.7.2.6

(Stamatakis 2006; Stamatakis et al. 2008). By using GTRGAMMA for the model of nucleotide substitution (-m), individual α-shape parameters, GTR-rates, and empirical base frequencies were estimated and optimized for each partition. RAxML analyses were implemented using a rapid bootstrap and search for the best-scoring ML tree in one single program run. Heuristic hill-climbing tree searches were performed generated from 1000

207 distinct randomized maximum parsimony starting trees and computed for 1000 repetitions.

Bayesian Analysis—The analyses were performed using the parallel (MPI) version of MrBayes 3.2.1 (Ronquist et al. 2012). The evolutionary models chosen for each data partition were estimated by the Akaike Information Criterion (AICc) as calculated in jModelTest 0.1.1 (Posada 2008), shown in Table 4.2. The Markov Chain

Monte Carlo (MCMC) parameters for combined data were as follows: ngen= 40 000 000 printfreq=1000 samplefreq=1000 nchains=4 (5 if could not meet the convergence value) savebrlens=yes stoprule=yes stopval=0.01. For each gene run, the MCMC was designed as: ngen= 20 000 000 printfreq=1000 samplefreq=1000 nchains=4 savebrlens=yes stoprule=yes stopval=0.01.

208

RESULTS

Data properties

Total DNA sequences of over 3.7 kb from 4 molecular markers were successfully obtained for all 80 specimens: 69 ingroups and 11 outgroups (Table 4.1). Amplified PCR products varied in length: COI: 857 bp, 18S: 1003 bp, 28S: 782 bp, EF-1α F2 copy: 1125 bp (the numbers shown excluding hypervariable regions from ribosomal genes).

Sequence characteristics of the four gene partitions are summarized in Table 4.3. The proportion of parsimony informative sites differs among data sets, with 18S showing the least (4.76%) and protein coding genes nucleotide position 3 representing the highest values (COI nt3: 82.87%, EF-1α nt3: 86.4%). The combined 4 genes, 18S, 28S, EF-1α, and COI markers showed roughly the same level of parsimony informative sites. Of the total characters for the combined 4 genes with different coding schemes (allnt: 3767, nt12: 750, Degen: 3765), the percent parsimony informative sites are 28.67, 15.33, and

11.82 respectively. No differences in nucleotide frequencies were detected in any gene partitions.

Phylogenetic relationships of Telenominae

Summaries of the support for relationships within the subfamily Telenominae derived from 3 analytical criteria, 3 coding schemes, and different partitions (3 partitions for allnt coding and 2 partitions for nt12 and Degen coding) for the combination of the 4

209 markers are presented in Table 4.5. Five phylogenetic results are presented here: Figs.

4.1–4.5: The results of all genes analyzed in a single partition using Bayesian (Fig. 4.1) and maximum likelihood (Fig. 4.2) inference; Fig 4.3 Bayesian result of combined genes for 4 partitions, the phylogeny presenting the difference in placement of the genus

Eumicrosoma which is concordant with the other 5 coding and partitioning schemes,

Figs. 4.4–4.5 parsimony analysis of 3 different nucleotide character coding: allnt, nt12, and Degen. The results of analyses of individual gene partitions are discussed but not figured.

The assessment of homoplasy is made through maximum parsimony. Individual gene partitions vary in the amount of homoplasy: 18S (150 trees, 192 steps, CI=0.21,

RI=0.15), 28S (16 trees, 1025 steps, CI=0.22, RI=0.14), COI (80 trees, 3746 steps,

CI=0.16, RI=0.18), and EF-1α (17 trees, 1873 steps, CI=0.29, RI=0.27). The phylogenetic signal, however, is relatively strong in the combined gene analyses as suggested by the CI, RI and the support values. The phylogenies generated from the combined 4 genes with different coding schemes are 1) all nucleotides: a strict consensus from 4 trees of length 7076 steps, CI=0.34, RI=0.53 (Fig. 4); 2) nt12: a strict consensus from 16 trees of length 2370 steps, CI=0.41, RI=0.65, and 3); Degen: a strict consensus from 174 trees of length 2265 steps, CI=0.47, RI=0.7 (Fig. 4.5).

The statistical model-based topologies in general have greater support than parsimony trees. For all but one of the Bayesian analyses, the average standard deviation of split frequencies (ASDSF) fell below 0.01 in 40 M or fewer generations. For one

210 coding scheme (all nucleotides in a single partition), the ASDSF failed to reach the set stop value, and the analysis was terminated at 40 M generations. At that point the graph of log likelihood and number of generations showed no obvious trend, suggesting that the analysis had reached stationarity.

Telenominae — All analytical techniques, coding and partitioning schemes support the conclusion that the subfamily Telenominae is not monophyletic: the genera

Psix and Paratelenomus are strongly supported as the sister to the six species of the outgroup Gryon (Figs. 4.1–4.5, Table 4.5). The most restricted monophyletic group that includes all telenomines also includes Gryon (PP 98–100 percent, BS 76–91, JK 56–84).

Psix group of genera — Only two genera of the Psix group are represented in the taxon sample. Both Psix (3 specimens of 3 species) and Paratelenomus (6 specimens of one species) are individually monophyletic (Psix: PP 100, BS 100, JK 97–99;

Paratelenomus: PP 100, BS 99–100, JK 100). The two genera are also consistently grouped together in all analyses, coding and partition schemes (PP 100, BS 100, JK 99–

100). Together, Psix, Paratelenomus and Gryon are strongly supported as a monophyletic group (PP 97–99, BS 68–75, JK 54–80 except Degen JK less than 50%).

Telenominae exclusive of Psix group of genera —All of the species in the genera Telenomus, Trissolcus, Eumicrosoma and Phanuromyia form a monophyletic group (PP 98–100, BS 85–98, JK 81–98). The two species of Phanuromyia are monophyletic (PP 89–99, BS 59–89, JK 50–53). The Telenomus crassiclava species group is monophyletic in most analyses (PP 96–100, BS 86–90, JK 80–82); one species is

211 excluded in Maximum Likelihood analyses including all nucleotide positions.

Phanuromyia is consistently the sister to the cluster of Telenomus crassiclava group species (89–99, BS 88–91, JK 61–65). The remaining Telenominae, that is, the species of

Trissolcus, Telenomus (not including the crassiclava group) and Eumicrosoma formed a monophyletic group with relatively strong support in the Bayesian and Maximum

Likelihood results (PP 84–98, BS 54–94). The species of the Telenomus californicus complex, the large group within which are found the parasitoids of the eggs of

Lepidoptera and Diptera, all group together in all analyses. This group, represented by 8 species, is not supported as a distinct clade because of the inclusion of Te. sechellensis

(of the podisi group) and one species of the floridanus group (PP 85–100, BS 71–93, JK

70–80). The Telenomus longicornis species group (PP 99–100, BS 59–96, JK < 50) and the Te. laricis sp. group (PP 100, BS 95–100, JK 61–98) were recovered as monophyletic groups all character coding and partitions schemes.

The representatives of Telenomus that are sometimes placed in the genera Aholcus

(Te. dalmanni) and Platytelenomus always group with the species of the Te. californicus complex. The position of the genus Eumicrosoma is unstable: it may emerge with the californicus complex or as sister to Trissolcus ogyges (Table 4.5).

Problematic taxa. Neither the genera Telenomus nor Trissolcus are resolved as monophyletic groups, even with the exclusion of the crassiclava species group from the former. None of the three species groups of Trissolcus are shown to be monophyletic, nor are the podisi and floridanus species groups of Telenomus.

212

DISCUSSION

The focus of this research is the inference of phylogeny of the Telenominae based on molecular data. Although numerous attempts have been made to characterize groups above the species using morphological data, these have been less than satisfactory or comprehensive. These problems arise from the preponderance of reductional characters, high levels of apparent convergence, and overall relatively small numbers of characters for the task at hand (Austin et al. 2005). Molecular data, while not immune from such issues, do serve as rich sources of largely independent data. As such, they may provide valuable context for interpreting both characters derived from morphology as well as those from biological features, especially host relationships.

The latest phylogenetic study of Platygastroidea inferred from molecular data was carried out by Murphy et al., (2007). This work was based on the analysis of three molecular markers, 18S rDNA, 28S rDNA, and COI. Their results support the hypotheses that Platygastridae s.str., Teleasinae, and Telenominae are monophyletic. The tribe

Gryonini, on the basis of 3 species, is also monophyletic with strong support and is the sister-group to Telenominae. Within telenomines, Phanuromyia (represented by one species) and the Telenomus crassiclava species group form a monophyletic group, and the remaining telenomines – two Trissolcus species and three Telenomus – are also a monophyletic clade. Preliminary results based on the same set of markers, but using an expanded representation of telenomines showed that Gryon fell in the ingroup

213 (Telenominae), and the relationship within the subfamily was mostly unresolved. We expanded the study to include more out groups to test monophyly of subfamily and also added the more slowly evolving nuclear protein-coding gene EF-1α.

Telenominae is not monophyletic — Parsimony, Bayesian and maximum likelihood analyses of the combined data from CO1, 18S and 28S and EF-1α fail to support the hypothesis that the subfamily is monophyletic. This strongly contrasts with all previous work, whether based on morphological or molecular characters. Masner’s

(1976) concept of the subfamily Telenominae, in its restricted sense, i.e., excluding the tribes Aradophagini and Tiphodytini, is recognized by the following combination of characters: metasoma with wide laterotergites loosely attached to the respective sternites of each segment; laterosternites absent, and thus the typical impressed submarginal ridge of Scelioninae and Teleasinae is lacking; the second segment is by far the largest metasomal segment; and the number of female antennomeres is either 10 or 11, while those of the male have 12 segments (with only one known exception). Our analysis, in contrast, separates from other telenomines the monophyletic clade of Psix and

Paratelenomus, these together arising as the sister-group of the scelionine genus Gryon.

Gryon is one of the largest genera in subfamily Scelioninae (Masner 1976) possessing many characters in common with Telenominae: frons usually without margined depression; skaphion absent; metasoma typically very broad and short; and the second segment is usually the largest of the metasomal segments. The hosts attacked by these two groups are also very similar and distinctive within Platygastroidea as a whole.

214 Gryon primarily attacks the eggs of Heteroptera, mainly , ,

Scutelleridae, , and , but also with at least one species confidently reared from the eggs of Lepidoptera. Psix and Paratelenomus attack the eggs of

Pentatomidae, , and Plastaspidae (Table 4.6); Trissolcus is known only to attack eggs of ; and numerous species of Telenomus of the podisi group are also parasitoids of stink bugs and their allies (Johnson 1984).

The results of our analysis may be interpreted in two ways: 1) Gryon, and presumably the genera closely related to it, should be classified within the Telenominae; or 2) Telenominae sensu Masner (1976) is polyphyletic. In either case, our current concept of Telenominae cannot be maintained as a monophyletic group. Our initial expectation was that Gryon would indeed be a close relative of Telenominae, and a range of species of the former were included to better understand with which group of Gryon the Telenominae might be most closely related. Therefore, the range of outgroups beyond

Gryonini that were included in the analysis was fairly restricted. While we are reasonably confident that Psix and Paratelenomus should be excluded from Telenominae, we are far less certain that the sister group relation of these genera with Gryon would be supported with a more extensive range of other genera. As a result, we restrict our interpretation of these results to concluding that telenomines are not monophyletic. Our use of the name

Telenominae s.str. is meant to encompass the subfamily in the sense of Masner (1976), but excluding the Psix group of genera: Psix, Paratelenomus, Nirupama and Mudigere

(see Johnson 1985, 1988). Telenominae s.str., therefore, includes the genera Telenomus,

Trissolcus, Eumicrosoma, Aradoctonus, Ioseppinella, Kozlotelenomus, Latonius,

215 Paratrissolcus, Phanuromyia, Phanuropsis, Phlebiaporus, Protelenomus, Rachelia,

Televiggianus, and (presumably) the fossil genus Sinoprotelenomus. The identity of

Bruchiola is unknown

Monophyly of Psix and Paratelenomus — Johnson (1988) proposed that the Psix group of genera, composed of Psix, Paratelenomus, Nirupama and Mudigere, forms a monophyletic group. Only two of these genera are included in this study, but the results strongly support the hypothesis. Morphologically, at least the first three of these genera are quite distinctive when compared with other telenomines: the clypeus is large and obscures the labrum (alternatively, the labrum is immovably and indistinguishably fused to the clypeus); a pair of orbital carinae extend from the anterior mandibular articulation dorsally along the inner orbits of the eyes to the lateral ocelli; the lateral ocelli are separated from the inner orbits (contiguous with or connected to the inner orbit by a narrow sulcus in other telenomines); the central keel on the frons bifurcates ventrally, with the arms surrounding the antennal insertions; the mesopleuron and metapleuron are coarsely foveate, and the patterns of the foveae strongly differ from telenomines; and the first metasomal tergite lacks differentiated sublateral setae. One possible interpretation of these distinctive characters was that the Psix group of genera arises basally from the

Telenominae. An alternative interpretation arising from these results is that since the two groups are not part of a monophyletic Telenominae, these differences are not exceptional.

The question shifts to interpreting the supposed synapomorphies of the broader concept of Telenominae as having arisen independently. Broad laterotergites are known outside of telenomines in the tribes Aradophagini, Tiphodytini, and Baeini (particularly male

216 Baeus), as well as the subfamilies Platygastrinae and Sceliotrachelinae. Reduction in the number of antennomeres occurs commonly throughout the superfamily. Even within the

Gryonini, the genus Maruzza is characterized by having 11-segmented female antennae.

The relative dominance in size of the second metasomal segment may be a convergent development or a synapomorphy at a higher level within the family.

Telenomus crassiclava species group and Phanuromyia — All phylogenetic results strongly support the hypothesis that the genus Phanuromyia and the Telenomus crassiclava species group together are monophyletic. These two groups are unique in their host relations, attacking the eggs of Auchenorrhyncha (Fulgoroidea) (Table 4.6).

The Telenomus crassiclava species group may usually be separated from other telenomines by the presence of an oblique furrow or line of foveae arising from the dorsal end of the acetabular carina and extending toward the mesopleural pit. This is shared with

Phanuromyia. Additional features characteristic of the two groups (but not universally found among species) are a convex frons, with the antennal scrobe strongly reduced to two small depressions above antennal insertions; the female antennal clava is often enlarged; the ovipositor is commonly exserted from the metasoma (usually retracted in telenomines), and the second metasomal tergite often has a characteristic reticulate sculpture.

The genera Trissolcus and Telenomus — Our results do not support monophyly of either of the genera Telenomus or Trissolcus. The Telenomus crassiclava group forms a monophyletic group with Phanuromyia with strong support, thus rendering the

217 remainder of the genus paraphyletic at best. Further, one species of Telenomus,

Te_FLO1, emerges as the sister of all other Telenomus (excluding the crassiclava group) and Trissolcus. Monophyly of none of the three species groups of Trissolcus is supported.

The Trissolcus basalis group emerge as three separate clades, but those three clades do not group together. The three species of Trissolcus flavipes species group do not group together nor with any other Trissolcus or Telenomus. Interestingly, the species assigned to the Trissolcus thyantae specis group emerge at the base of the main clade of

Telenomus + Trissolcus (excluding the crassiclava species group and Te_FLO1), but do not group together.

Within Telenomus, first, the floridanus species group appears to be massively polyphyletic. Species emerge throughout the Telenomus+Trissolcus clade and none of them appear to be closely related to Eumicrosoma (as suggested by Johnson 1984). Good support is found that the laricis, longicornis and phymatae groups together are monophyletic (Table 4.5) and at least the first two are monophyletic groups themselves.

Johnson (1984) noted that the longicornis group is superficially similar to the scelionine genus Tiphodytes, the species of which attack the eggs of Gerridae (Heteroptera). The species of the longicornis group are also found near water and collected in aquatic emergence traps. He concluded that the hosts of this group are aquatic heteropterans. The laricis group attacks the eggs of . The phymatae group attack the eggs of

Reduviidae and include some species that are important as natural enemies of vectors of

Chagas disease in Latin America.

218 The Telenomus podisi species group parasitize the eggs of stink bugs and their allies (Pentatomoidea). The results do not support the hypothesis that this is a monophyletic group. One species, Telenomus sechellensis, falls within the Telenomus californicus species complex.

The greatest diversity of Telenomus species parasitize the eggs of and butterflies. Many of these are important biological control agents. According to Johnson

(1984), all these lepidopteran parasitoids fall into the californicus species complex which includes four species groups (californicus, arzamae, dalmanni, and tabanivorus), as well as a number of unplaced species. All members of the californicus complex do fall within and make up the majority of members of a single clade, but are joined there by T. sechellensis and an undescribed species of the floridanus group from Kenya.

In the taxon sampling, special effort was made to include representatives of three small taxa. Eumicrosoma attacks the eggs of chinch bugs and has been hypothesized to be closely related to the floridanus group (Johnson 1984). Aholcus was described by J.-J.

Kieffer for species of Telenomus in which the female antennae are 10-segmented (all other Telenomus have 11-segmented antennae in the female). Nixon synonymized

Aholcus with Telenomus, reasoning that the only distinguishing feature was the number of antennal segments and that, since this was only found in one sex, this was a poor basis for formal generic status. Nevertheless, the name is still used by some taxonomists as a subgenus of Telenomus. We included it here, represented by T. dalmanni, to find evidence for its relations with other species. Similarly, Platytelenomus was described for

219 a highly flattened species from Australia. Johnson (1984) synonymized this name with

Telenomus, but its use lingers on.

As noted above, Eumicrosoma does not group with any representatives of the floridanus species, although strong support for its position is still lacking. Aholcus and

Platytelenomus cluster together with the lepidopteran parasitoids of the californicus complex. If either were to be formally recognized as a valid taxon (assuming that with further taxon sampling they proved to be monophyletic themselves), this would mean that the remainder of species of Telenomus would be placed in a paraphyletic group. I do not propose formal nomenclature changes here because of the lack of sufficient support for the resolution of relationships. Progress toward this goal appears to require an increase in the extent of taxon sampling (additional species of Eumicrosoma, Aholcus, and

Platytelenomus) as well as additional markers

220

CONCLUSION

Telenominae is the most important parasitoid subfamily of Platygastridae, the single family of the Platygastroidea. Despite being efficient biological control agents of agricultural pests, the relationships within the subfamily as well as the evolution of host associations are still unclear. We present the first phylogenetic analysis focused on

Telenominae, exploring 80 terminals representing 6 genera and 11 species group taxa from the specimens around the world. The observed data were explored in both parsimony and statistical frameworks (Maximum Likelihood & Bayesian inference), based on approximately 3.7 kb of DNA sequences from 4 molecular markers: 18S, 28S,

COI and EF-1α. The molecular evidence showed that the subfamily is not monophyletic: one clade, the Psix group of genera, forms a monophyletic group with species of the tribe

Gryonini, subfamily Scelioninae. Monophyletic clades were recovered with strong support including (Psix+Paratelenomus) and Gryon; Telenominae, s.str. (without Gryon and Psix group of genera); Phanuromyia and Telenomus crassiclava species group;

Telenomus laricis species group; and Telenomus longicornis species group. The genera

Eumicrosoma and Platytelenomus are nested within the Telenomus californicus species complex. Pentatomoidea (Hemiptera) is the plesiomorphic host. The species in the clade

Phanuromyia + Telenomus crassiclava all shared the same host group, Auchenorrhyncha

(Hemiptera: Fulgoridae and Flatidae). The monophyly of the major genera Telenomus

221 and Trissolcus is not supported; the interrelationships of their component species are largely unresolved.

222 Table 4.1 List of taxa and gene-amplified regions for each species included in the telenomine phylogenetic analysis

Taxa Ref. Voucher no. 18S 28S CO1 EF- code "OSUC" 1α Ingroup Telenominae: Trissolcus Basalis group Trissolcus basalis Tr_basalis 173847 X X X Trissolcus basalis Tr_basalis 173848 X X X X Trissolcus basalis Tr_basalis 173849 X X X Trissolcus latisulcus Tr_latisulcus 248218 X X X X Trissolcus hullensis Tr_hullensis 266782 X X X X Trissolcus sp. Tr_BAS1 266785 X X X Trissolcus sp. Tr_BAS2 181674 X X Trissolcus sp. Tr_BAS3 173865 X X X X Trissolcus sp. Tr_BAS3 173866 X X X X Trissolcus sp. Tr_BAS3 248205 X X X X Flavipes group Trissolcus urichi Tr_urichi 248199 X X X X Trissolcus strabus Tr_strabus 248187 X X X X Trissolcus euschisti Tr_euschisti 181546 X X Thyantae group Trissolcus thyantae Tr_thyantae 266773 X X X X Trissolcus sp. Tr_THY1 266787 X X X Trissolcus sp. Tr_THY2 173870 X X X Not assigned to species group Trissolcus ogyges Tr_ogyges 270812 X X X X Trissolcus ogyges Tr_ogyges 270813 X X X Telenominae: Telenomus Subgenus Aholcus Telenomus dalmanni Te_dalmanni 173871 X Telenomus dalmanni Te_dalmanni 173872 X Telenomus dalmanni Te_dalmanni 173873 X X Californicus complex Telenomus sp. Te_CAC1 248204 X X X Telenomus sp. Te_CAC2 266784 X X X X Telenomus sp. Te_CAC3 173867 X X X X Telenomus sp. Te_CAC4 248191 X X X Telenomus sp. Te_CAC5 173868 X X X Continued

223

Table 4.1 Continued

Crassiclava group Telenomus sp. Te_CRA1 248221 X X X Telenomus sp. Te_CRA2 266780 X X X X Telenomus sp. Te_CRA3 173851 X X X Telenomus sp. Te_CRA4 266769 X X X X Telenomus sp. Te_CRA5 176057 X X X Telenomus sp. Te_CRA6 188473 X X X Floridanus group Telenomus consimilis Te_consimilis 248195 X X X X Telenomus nysivorus Te_nysivorus 266771 X X X X Telenomus sp. Te_FLO1 266768 X X X X Telenomus sp. Te_FLO2 173852 X X X Telenomus sp. Te_FLO3 266774 X X X X Laricis group Telenomus sp. Te_LAR1 265220 X X X X Telenomus sp. Te_LAR2 266781 X X X X Telenomus sp. Te_LAR3 248197 X X X X Longicornis group Telenomus sp. Te_LON1 176014 X X X X Telenomus sp. Te_LON2 248193 X X X X Telenomus sp. Te_LON3 173832 X X X X Telenomus sp. Te_LON3 173833 X X X X Telenomus sp. Te_LON4 173839 X X X X Telenomus sp. Te_LON4 173840 X X X X Phymatae group Telenomus dolichocerus Te_dolichocerus 176036 X X X Telenomus dolichocerus Te_dolichocerus 248192 X X X Podisi group Telenomus podisi Te_podisi 173850 X X X X Telenomus podisi Te_podisi 181680 X X X Telenomus podisi Te_podisi 176037 X X X Telenomus sechellensis Te_sechellensis 266783 X X X Telenomus grenadensis Te_grenadensis 248194 X X X X Tabanivorus group Telenomus sp. Te_TAB1 248198 X X X X Telenomus goniopis Te_goniopidis 248188 X X X X Telenomus goniopis Te_goniopidis 248189 X X X Continued

224

Table 4.1 Continued

Telenominae: other genera Paratelenomus saccharalis Pa_saccharalis 181673 X X X Paratelenomus saccharalis Pa_saccharalis 248183 X X X Paratelenomus saccharalis Pa_saccharalis 248206 X X X X Paratelenomus saccharalis Pa_saccharalis 248207 X X X Paratelenomus saccharalis Pa_saccharalis 270803 X X X X Paratelenomus saccharalis Pa_saccharalis 270804 X X X X Phanuromyia bidentata Ph_bidentata 271001 X X X Phanuromyia sp. Ph_SP1 270805 X X X X Psix watshami Ps_watshami 248092 X X X Psix striaticeps Ps_striaticeps 266777 X X X X Psix tunetanus Ps_tunetanus 266779 X X X X Eumicrosoma sp. Eu_SP1 173874 X X X X Platytelenomus sp. Pl_SP1 270802 X X X outgroup Gryon cultratus Gryon_SP2 176020 X X X Gryon largi Gryon_SP4 176056 X X X Gryon sp. Gryon_SP1 266789 X X X X Gryon sp. Gryon_SP3 266775 X X X X Gryon sp. Gryon_SP5 266778 X X X X Gryon sp. Gryon_SP6 248210 X X X X Oxyteleia sp. Oxyteleia_SP1 266063 X X X Probaryconus Probaryconus_SP1 181541 X X X X Scelio sp. Scelio_SP1 206847 X X X Scelio sp. Scelio_SP2 176075 X X X Trimorus caraborum Trimorus_SP1 173838 X X X

225

Table 4.2 Primer descriptions

Gene Primer Primer Sequences (5'-3') Reference Name 18S ai CCT GAG AAA CGG CTA CCA CAT C Whiting et al. 1997 18S 18S-bi GAG TCT CGT TCG TTA TCG GA Whiting et al. 1997 18S 18S-5R CTT GGC AAA TGC TTT CGC Giribet et al. 1996 28S D2-3 D23F GAG AGT TCA AGA GTA CGT G Whiting et al. 1997 28S D2-3 28Sb TCG GAA GGA ACC AGC TAC TA Whiting et al. 1997 CO1 1F GGA GGA TTT GGA AAT TGR YTW RTT Simon et al. CC 1994 CO1 1R ACT GTA AAT ATR TGA TGW GCT CA Simon et al. 1994 CO1 HCO2198 TAA ACT TCA GGG TGA CCA AAA AAT Folmer et al. CA 1994 CO1 LCO1490 GGT CAA CAA ATC ATA AAG ATA TTG G Folmer et al. 1994 EF-1α F2F GGG YAA AGG WTC CTT CAA RTA TGC Danforth et al.1999 EF-1α F2F8 CAA RTA TGC NTG GGY ATT GGY AAG Heraty et al. 2011 EF-1α For3 GGN GAC AAY GTT GGY TTC AAC Heraty et al. 2011 EF-1α Cho10 ACR GCV ACK GTY TGH CKC ATG TC Danforth et al.1999 EF-1α F2R6 TTG WGC RGT GAA GTC AGC NGC Heraty et al. 2011 EF-1α F7 AAC AAR ATG GAY TCN ACN GAR CCN Simon et al. CC 2010 EF-1α F9 CCN ACN GGB ACH GTT CCR ATA CC Simon et al. 2010

226

Table 4.3 Gene partitions and alignment summary for subfamily Telenominae and outgroups. Hypervariable regions and introns were removed manually. Percent parsimony informative sites were calculated using WinClada ver. 1.00.08 (Nixon 2002). Models chosen for data partitions were selected using the Akaike Information Criterion (AICc) as calculated in jModelTest 0.1.1 (Posada 2008).

Gene partitions Aligned A (%) T(%) C (%) G(%) % AICc base pairs parsimony model informative 18S 1003 24.7 22.3 29.1 23.9 4.76 TIM3+I+Γ 28S 782 19.4 23.1 29.5 27.9 26.34 TVM+Γ COI 857 47.9 10.4 4.1 37.5 43.76 TIM3+I+Γ EF-1α 1125 23.4 29 25.8 21.8 36.26 TIM1+I+Γ 4 Genes 3767 25.9 22.3 21.74 29.9 28.67 GTR + Γ COI nt1+2 571 31.6 15.5 13.7 39 24.52 TIM1+Γ COI nt3 286 48.2 10.3 1.6 40 82.87 HKY+Γ EF-1α nt1+2 750 31.6 22.3 26.8 19.4 11.2 TrN+Γ EF-1α nt3 375 14.7 34.3 28.5 22.6 86.4 TIM2+Γ 4 Genes nt1+2 3106 26.6 21.9 25.7 25.8 15.33 GTR + Γ Degen COI 855 14.62 TIM1+Γ Degen EF-1α 1125 5.87 GTR + Γ Degen 4 Genes 3765 11.82 GTR + Γ

227

Table 4.4 Definitions for coding strategies and partitioning schemes

Coding Definitions allnt-1 A dataset consisting of all nucleotide characters subjected to a single partition allnt-4 All nucleotide characters subjected to 4 partitions, separating data from 18S, 28S, COI, and EF-1α allnt-6 All nucleotide characters subjected to 6 partitions, for data from 18S, 28S, COI nt1+2, EF-1α nt1+2, COI nt3, and EF-1α nt3 nt12-1 A dataset consisting of nucleotide characters position 1 and 2, subjected to a single partition nt12-4 Nucleotide characters position 1 and 2 subjected to 4 partitions, separating data from 18S, 28S, COI, and EF-1α Degen1 A dataset consisting of all nucleotide characters in which codon in protein coding genes (COI and EF-1α) were completely degenerated and subjected to single partition Degen4 Degenerated codons for protein coding genes subjected to 4 partitions, separating data from 18S, 28S, COI, and EF-1α

228 Table 4.5 Support values for selected clade across the analyses. Dataset are based on nucleotide character coding and partitioning schemas shown on Table 4. Support values are reported as posterior probability (PP) for Bayesian inference and bootstrap support (BS) for maximum likelihood (RAxML) and jackknife support (JK) for parsimony TNT. The support values for all analyses are reported in percent above 50. Abbreviations used in text: Tr., Trissolcus; Te., Telenomus; sp., species; gr., group taxa; com., complex; y=yes group recovered but support less than 50 percent; n=no, not monophyletic; - , taxa excluded; +, taxa included.

Clade Bayesian analysis (MrBayes) allnt- allnt- allnt- nt12- nt12- Degen Dege 1 4 6 1 4 -1 n-4 Telenominae n n n n n n n (Gryon + Psix gr.) + Telenominae 98 98 98 100 100 100 100 Psix + Paratelenomus + Gryon 99 99 99 99 99 97 99 Gryon 100 100 100 100 100 100 100 Psix + Paratelenomus 100 100 100 100 100 100 100 Psix 100 100 100 100 100 100 100 Paratelenomus 100 100 100 100 100 100 100 Telenominae s.str. 98 98 98 100 100 100 100 Telenominae - (Psix +Paratelenomus) Phanuromyia + Te. crassiclava sp. 96 100 95 99 100 100 100 gr. Phanuromyia 95 94 95 95 89 99 93 Te. FLO1 + Telenomines s.str. 98 98 98 94 84 87 84 Telenomines - Te. FLO1 - 96 98 96 94 96 99 99 (Phanuromyia + Te. crassiclava sp. gr. Te. crassiclava sp. gr. 96 100 96 99 100 100 100 Te. tabanivorus sp. gr. 95 70 95 99 99 99 92 Te. laricis sp. gr. 100 100 100 100 100 100 100 Te. longicornis sp. gr. 100 100 100 99 100 100 99 Te. phymatae sp. gr. 97 97 97 91 97 95 95 Aholcus 100 100 100 100 100 100 100 Te. laricis sp. gr. + phymatae sp. gr. 90 85 90 n n 91 62 + longicornis sp. gr. Te. californicus sp. com. + 100 100 100 85 98 97 99 Platytelenomus, Te. sechellensis, Te_FLO3 Te. californicus sp. com. + n n n 90 n 93 n Platytelenomus, Te. sechellensis, Te_FLO3 + Eumicrosoma Eumicrosoma + Tr. Ogyges 94 98 94 n 90 n 96 Continued

229 Table 4.5 Continued Clade Maximum Likelihood (RAxML) Parsimony (TNT) all all all nt nt De- De- allnt nt Deg nt-1 nt-4 nt-6 12- 12- gen gen 12 en 1 4 -1 -4 Telenominae n n n n n n n n n n (Gryon + Psix gr.) + 84 76 83 87 87 91 89 84 63 56 Telenominae Psix + Paratelenomus + 75 73 72 76 77 69 68 80 54 y Gryon Gryon 100 100 100 100 100 100 100 100 100 100 Psix + Paratelenomus 100 100 100 100 100 100 100 99 100 100 Psix 100 100 100 100 100 100 100 99 99 97 Paratelenomus 100 100 100 100 100 100 99 100 100 100 Telenominae s.str. i.e. 93 85 94 94 98 97 98 81 98 94 Telenominae - (Psix + Paratelenomus) Phanuromyia + n n n 88 91 88 79 y 65 61 Te. crassiclava sp. gr. Phanuromyia 89 59 76 81 69 83 80 50 y 53 Te. FLO1 + Telenomines 93 85 94 61 55 54 59 y n n s.str. Telenomines - Te. FLO1 y y y y y 60 64 y y y - (Phanuromyia + Te. crassiclava sp. gr.) Te. crassiclava sp. gr. n n n 90 89 89 86 y 80 82 Te. tabanivorus sp. gr. 80 84 74 79 75 87 76 63 72 85 Te. laricis sp. gr. 95 98 96 98 100 99 99 61 62 98 Te. longicornis sp. gr. 96 64 59 91 95 63 85 y y y Te. phymatae sp. gr. 59 96 97 64 65 85 65 87 86 74 Aholcus 100 100 100 100 100 100 100 y 99 98 Te. laricis sp. gr. + y y y n y y y n n y phymatae sp. gr. + longicornis sp. gr. Te. californicus sp. com. 71 87 93 80 80 80 79 y 80 70 + Platytelenomus, Te. sechellensis, Te_FLO3 Te. californicus sp. com. n n n y y y n n n n + Platytelenomus, Te. sechellensis, Te_FLO3 + Eumicrosoma Eumicrosoma + Tr. n n n n n n y y 51 y ogyges

230 Table 4.6 Host records of Gryon and Telenominae (Johnson 2012)

Telenomine genera or species group Host records

Phanuropsis Pentatomidae (Hemiptera) Trissolcus basalis group Pentatomidae (Hemiptera) Trissolcus thyantae group Pentatomidae (Hemiptera) Protelenomus Coreidae (Hemiptera) Telenomus podisi group Pentatomidae, Scutelleridae (Hemiptera)

Psix Pentatomidae, Scutelleridae (Hemiptera) Nirupama Unknown (Hemiptera) Paratelenomus , Pentatomidae (Hemiptera) Mudigere Unknown

Phanuromyia Fulgoridae (Hemiptera) Telenomus crassiclava group Fulgoridae, Flatidae (Hemiptera)

Eumicrosoma (Heteroptera) Telenomus floridanus group Blissidae (Heteroptera)

Aradoctonus Telenomus longicornis group Unknown, presumably aquatic Telenomus laricis group Miridae (Hemiptera) Telenomus nigrocoxalis group (Hemiptera) Telenomus phymatae group Reduviidae (Hemiptera) Telenomus tabanivorus group Tabanidae, Asilidae, Stratiomyidae (Diptera) Telenomus dalmanni group Telenomus californicus group Telenomus arzamae group

Gryon Coreidae (Hemiptera) Pentatomidae (Hemiptera) Reduviidae Lymantriidae (Lepidoptera)

231 Probaryconus_SP1 Trimorus_SP1

52 Oxyteleia_SP1 93 Scelio_SP1 100 Scelio_SP2 Gryon_SP1 100 Gryon_SP2 Gryon_SP3 A 100 100 Gryon_SP4 Gryon 100 Gryon_SP5 100 97 Gryon_SP6 Ps_watshami 100 Ps_striaticeps Psix 100 Ps_tunetanus B 100 Pa_saccharalis 99 Pa_saccharalis 100 Pa_saccharalis 94 Pa_saccharalis. Paratelenomus

100 100 Pa_saccharalis 96 Pa_saccharalis 99 Ph_bidentata Ph_SP1 Phanuromyia C 100 Te_CRA1

100 Te_CRA2 Te_CRA3 Telenomus 62 Te_CRA4 Crassiclava 100 94 Te_CRA5 99 Te_CRA6 sp. group Te_FLO1 Tr_thyantae 87 Tr_THY1 Tr_THY2 99 Tr_strabus Tr_urichi 68 Tr_euschisti Tr_BAS1 100 Tr_latisulcus Trissolcus Tr_hullensis D Tr_BAS2 Basalis 100 Tr_basalis 61 sp. group 100 Tr_basalis 99 Tr_basalis Tr_ogyges A Parasitize Hemiptera: 100 Tr_ogyges 64 Tr_BAS3 Coreidae, Pentatomidae 100 Tr_BAS3 Trissolcus basalis Reduviidae 99 Tr_BAS3 & Lepidoptera Eu_SP1 Eumicrosoma Te_sechellensis E B Parasitize Hemiptera: 93 Te_CAC1 59 Pentatomidae, Scutelleridae, Te_dalmanni 97 Te_dalmanni Plataspidae 100 Aholcus Te_dalmanni Telenomus Te_CAC2 C 75 Parasitize Hemiptera: Te_CAC3 Californicus Fulgoridae, Flatidae Te_FLO3 sp. complex 61 Te_CAC4 D Parasitize Hemiptera: 75 Platytelenomus 79 Pl_SP1 Pentatomidae 69 Te_CAC5 E Eumicrosoma: Hemiptera Te_TAB1 Telenomus Tabanivorus 99 Te_goniopidis (Lygaeidae) 100 Te_goniopidis sp. group Telenomus Californicus complex: Te_nysivorus Lepidoptera Te_grenadensis Platytelenomus: Lepidoptera F Te_consimilis Te_FLO2 Telenomus Tabanivorus group: 93 Te_podisi Diptera (Tabanidae, Asilidae, 60 Te_podisi Telenomus Podisi sp. group Stratiomyidae) 67 Te_podisi Te_LAR1 F Telenomus species group 100 Te_LAR2 Telenomus Laricis sp. group * Podisi: Hemiptera (Pentatomidae, 76 Te_LAR3 Te_dolichocerus Scutelleridae) 91 96 Telenomus Phymatae sp. group * Laricis group: Hemiptera (Miridae) Te_dolichocerus 95 Te_LON1 * Phymatae: Hemiptera (Reduviidae) Te_LON2 100 * Longicornis: Hemiptera Te_LON3 Telenomus Longicornis (aquatic insects) 100 99 Te_LON4 Te_LON5 sp. group 100 Te_LON6

0.02

232 Probaryconus_SP1 Trimorus_SP1 58 Oxyteleia_SP1 76 Scelio_SP1 100 Scelio_SP2 Gryon_SP1 100 Gryon_SP2 Gryon_SP4 100 97 Gryon_SP3 Gryon 95 Gryon_SP5 93 Gryon_SP6 69 Psix_watshami 100 Psix_striaticeps Psix 93 Psix_tunetanus 100 Pa_saccharalis 100 Pa_saccharalis 100 Pa_saccharalis 67 Pa_saccharalis Paratelenomus 100 Pa_saccharalis 91 91 Pa_saccharalis Ph_bidentata 83 Ph_SP1 Phanuromyia 88 Te_CRA1 Te_CRA3 89 Te_CRA2 Telenomus 50 Te_CRA4 Crassiclava 57 Te_CRA5 A 77 Te_CRA6 sp. group 97 Te_FLO1 Tr_thyantae B Tr_hullensis Tr_BAS2 75 100 Tr_basalis 54 Tr_basalis 100 Tr_basalis C Tr_THY2 Tr_THY1 Tr_euschisti 60 87 Tr_BAS1 Tr_latisulcus Trissolcus Basalis Tr_strabus sp. group Tr_urichi Tr_ogyges 100 Tr_ogyges Tr_BAS3 100 Tr_BAS3 Trissolcus basalis 99 Tr_BAS3 Eu_SP1 Te_sechellensis Te_CAC1 Te_dalmanni 80 100 Te_dalmanni Telenomus 97 Te_dalmanni Te_FLO3 Californicus Te_CAC4 Te_CAC2 sp.complex Pl_SP1 D A Telenomus sp. Te_CAC5 Te_CAC3 Floridanus group Te_TAB1 Telenomus Tabanivorus 87 Te_goniopidis B Trissolcus thyantae 99 Te_goniopidis sp. group Thyantae group Te_nysivorus Te_grenadensis Te_consimilis C Trissolcus Te_FLO2 64 Te_podisi D Telenomus + Eumicrosoma 40Te_podisi Telenomus Podisi sp. group + Platytelenomus) - Crassiclava group 69 Te_podisi Te_LAR1 Te_LAR3 99 Telenomus Laricis sp. group 91 Te_LAR2 63 Te_dolichocerus Te_dolichocerus Telenomus Phymatae sp. group Te_LON1 91 Te_LON3 85 Te_LON3 Te_LON2 Telenomus Longicornis 90 Te_LON4 sp. group 97 Te_LON4

0.03

233 Probaryconus_SP1 Trimorus_SP1 Oxyteleia_SP1

96 Scelio_SP1 100 100 Scelio_SP2 Gryon_SP1 100 Gryon_SP2 Gryon_SP3 100 100 Gryon_SP4 Gryon 100 Gryon_SP5 100 Gryon_SP6 55 99 Ps_watshami 100 Ps_striaticeps Psix 100 Ps_tunetanus 100 Pa_saccharalis 100 Pa_saccharalis 99 Pa_saccharalis 94 Pa_saccharalis Paratelenomus

100 Pa_saccharalis 100 93 Pa_saccharalis 93 Ph_bidentata Ph_SP1 Phanuromyia Te_CRA2 100 Te_CRA1 100 56 Te_CRA3 Telenomus Te_CRA4 Crassiclava 90 Te_CRA5 100 94 Te_CRA6 sp. group Te_FLO1 Tr_thyantae Tr_THY1 84 56 Tr_THY2 Tr_strabus 99 Tr_urichi Tr_euschisti Tr_BAS1 84 100 Tr_latisulcus Trissolcus Tr_hullensis Basalis Tr_BAS2 100 Tr_basalis sp. group 68 100 Tr_basalis 99 Tr_basalis 100 Tr_BAS3 Tr_BAS3 Trissolcus basalis 69 99 Tr_BAS3 Eu_SP1 Eumicrosoma 96 Tr_ogyges 100 Tr_ogyges Trissolcus ogyges Te_CAC1 Te_dalmanni Te_dalmanni 100 99 Te_dalmanni Telenomus Te_CAC5 Te_CAC2 Californicus Pl_SP1 90 sp. complex Te_FLO3 64 Te_CAC4 69 Te_sechellensis 99 Te_CAC3 Te_TAB1 96 Telenomus Tabanivorus 92 Te_goniopidis 99 Te_goniopidis sp. group Te_nysivorus Te_grenadensis 55 Te_consimilis Te_FLO2 Te_podisi Telenomus Podisi Te_podisi 91 Te_podisi sp.group 99 100 Te_LAR1 Te_LAR2 Telenomus Laricis sp.group 66 Te_LAR3 62 95 Te_dolichocerus Te_dolichocerus Telenomus Phymatae sp. group

95 Te_LON1 99 Te_LON3 0.2 99 Te_LON3 Telenomus 99 Te_LON2 55 Te_LON4 Longicornis sp. group 100 Te_LON4

234 Probaryconus_SP1 Trimorus_SP1 Oxyteleia_SP1 Scelio_SP1 100 Scelio_SP2 Gryon_SP1 100 100 Gryon_SP2 Gryon_SP3 100 100 Gryon_SP4 Gryon 100 Gryon_SP5 56 Gryon_SP6 80 Ps_watshami 99 Ps_striaticeps Psix 100 Psix tunetanus 99 Pa_saccharalis Pa_saccharalis 100 100 Pa_saccharalis 96 Pa_saccharalis. Paratelenomus 84 100 Pa_saccharalis 62 Pa_saccharalis Te_FLO1 Tr_thyantae Ph_bidentata Phanuromyia 50 Ph_SP1 Te_CRA1 81 Te_CRA3 Telenomus Te_CRA2 Te_CRA4 Crassiclava Te_CRA5 sp. group Te_CRA6 Te_FLO3 Pl_SP1 Tel_CAC5 Te_sechellensis Te_CAC3 Te_CAC1 Te_dalmanni 98 Te_dalmanni Te_dalmanni Te_CAC2 71 Te_CAC4 Te_TAB1 63 Te_goniopidis 100 Te_goniopidis Te_dolichocerus Telenomus sp. Te_dolichocerus Te_LON1 Te_LON4 87 99 Te_LON4 Te_LON2 Te_LON3 99 Te_LON3 Tel_consimilis Te_grenadensis 68 Te_podisi Te_FLO2 Te_LAR3 61 Te_LAR2 72 Te_LAR1 Te_podisi 99 Te_podisi Te_nysivorus Tr_hullensis Tr_BAS2 Tr_basalis 100 Tr_basalis 99 Tr_basalis Trissolcus sp. Tr_THY1 Tr_strabus Tr_THY2 Tr_BAS1 87 Tr_latisulcus Eu_SP1 Eumicrosoma Tr_ogyges 100 Tr_ogyges Tr_urichi Tr_euschisti Trissolcus sp. Tr_BAS3 2.0 100 Tr_BAS3 60 Tr_BAS3

235 Probaryconus_SP1 Probaryconus_SP1 Trimorus_SP1 Trimorus_SP1 Oxyteleia_SP1 Oxyteleia_SP1 Scelio_SP1 58 Scelio_SP1 100 Scelio_SP2 100 Scelio_SP2 Gryon_SP1 Gryon_SP1 100 Gryon_SP2 100 Gryon_SP2 100 Gryon_SP3 100 Gryon_SP3 100 100 96 Gryon_SP4 92 Gryon_SP4 75 Gryon_SP5 79 Gryon_SP5 58 Gryon_SP6 71 Gryon_SP6 54 Ps_watshami Ps_watshami 63 99 Ps_striaticeps 97 Ps_striaticeps 100 Psix tunetanus 97 Psix tunetanus 100 Pa_saccharalis 100 Pa_saccharalis Pa_saccharalis Pa_saccharalis 100 97 Pa_saccharalis 100 92 Pa_saccharalis 93 Pa_saccharalis. 71 Pa_saccharalis. 100 Pa_saccharalis 100 Pa_saccharalis Pa_saccharalis 63 Pa_saccharalis 63 Ph_bidentata 56 Ph_bidentata Ph_SP1 53 Ph_SP1 Te_CRA1 65 61 Te_CRA1 Te_CRA3 80 82 Te_CRA3 Te_CRA2 Te_CRA2 Te_CRA4 Te_CRA4 Te_CRA5 55 Te_CRA5 67 Te_CRA6 67 Te_CRA6 Te_FLO1 Te_FLO1 Tr_thyantae 98 94 Tr_thyantae Tr_strabus Tr_THY1 Tr_THY2 Tr_strabus Tr_euschisti Tr_urichi Tr_BAS3 Tr_THY2 100 Tr_BAS3 Tr_euschisti 96 Tr_BAS3 Tr_BAS1 Eu_SP1 90 Tr_latisulcus 63 57 51 Tr_ogyges Tr_BAS3 100 Tr_ogyges 100 Tr_BAS3 Tr_urichi 64 Tr_BAS3 Tr_THY1 Eu_SP1 Tr_BAS1 Tr_ogyges 69 Tr_latisulcus 100 Tr_ogyges Tr_hullensis Tr_hullensis Tr_BAS2 Tr_BAS2 Tr_basalis Tr_basalis 100 Tr_basalis 100 Tr_basalis 64 Tr_basalis 65 Tr_basalis Te_CAC1 Te_CAC1 Te_CAC5 Te_dalmanni Te_dalmanni Te_dalmanni 80 98 99 Te_dalmanni Te_dalmanni Te_dalmanni Te_FLO3 Te_FLO3 Te_CAC2 Pl_SP1 Te_CAC4 Te_CAC2 Pl_SP1 66 Te_CAC4 Te_CAC5 Te_sechellensis Te_sechellensis 60 Te_CAC3 70 Te_CAC3 Te_TAB1 Te_TAB1 72 Te_goniopidis 85 Te_goniopidis 99 Te_goniopidis 99 Te_goniopidis Te_nysivorus Te_nysivorus Te_podisi Te_dolichocerus 56 89 Te_podisi Te_grenadensis Te_grenadensis Te_consimilis Te_dolichocerus Te_dolichocerus Te_consimilis Te_podisi Te_dolichocerus Te_FLO2 Te_podisi Te_podisi Te_FLO2 55 Te_podisi A Te_LAR1 B Te_LAR1 Te_LAR2 97 98 Te_LAR2 62 Te_LAR3 60 Te_LAR3 Te_LON1 Te_LON1 Te_LON3 Te_LON2 86 91 Te_LON3 74 Te_LON3 Te_LON2 73 Te_LON3 2.0 81 Te_LON4 Te_LON4 2.0 97 Te_LON4 92 Te_LON4

236

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